ANATOMY MIDTERM 2

CHAPTER 7

The skeletal system includes all of the bones, cartilages, and ligaments of the body that support and give shape to the body and body structures.
The skeleton consists of the bones of the body. For adults, there are 206 bones in the skeleton. Younger individuals have higher numbers of bones
because some bones fuse together during childhood and adolescence to form an adult bone. The primary functions of the skeleton are to provide a rigid,
internal structure that can support the weight of the body against the force of gravity, and to provide a structure upon which muscles can act to produce
movements of the body. The lower portion of the skeleton is specialized for stability during walking or running. In contrast, the upper skeleton has
greater mobility and ranges of motion, features that allow you to lift and carry objects or turn your head and trunk.

In addition to providing for support and movements of the body, the skeleton has protective and storage functions. It protects the internal organs,
including the brain, spinal cord, heart, lungs, and pelvic organs. The bones of the skeleton serve as the primary storage site for important minerals such
as calcium and phosphate. The bone marrow found within bones stores fat and houses the blood-cell producing tissue of the body.

The skeleton is subdivided into two major divisions—the axial and appendicular. The axial skeleton forms the vertical, central axis of the body and
includes all bones of the head, neck, chest, and back (Figure 7.2). It serves to protect the brain, spinal cord, heart, and lungs. It also serves as the
attachment site for muscles that move the head, neck, and back, and for muscles that act across the shoulder and hip joints to move their corresponding
limbs.

The axial skeleton of the adult consists of 80 bones, including the skull, the vertebral column, and the thoracic cage. The skull is formed by 22 bones.
Also associated with the head are an additional seven bones, including the hyoid bone and the ear ossicles (three small bones found in each middle
ear). The vertebral column consists of 24 bones, each called a vertebra, plus the sacrum and coccyx. The thoracic cage includes the 12 pairs of ribs,
and the sternum, the flattened bone of the anterior chest.

Figure 7.2 Axial and Appendicular Skeleton The axial

skeleton supports the head, neck, back, and chest and thus
forms the vertical axis of the body. It consists of the skull,
vertebral column (including the sacrum and coccyx), and the
thoracic cage, formed by the ribs and sternum. The
appendicular skeleton is made up of all bones of the upper and
lower limbs.

The Appendicular Skeleton

The appendicular skeleton includes all bones of the upper

and lower limbs, plus the bones that attach each limb to the
axial skeleton. There are 126 bones in the appendicular
skeleton of an adult. The bones of the appendicular skeleton
are covered in a separate chapter.

The cranium (skull) is the skeletal structure of

the head that supports the face and protects the
brain. It is subdivided into the facial bones and
the brain case, or cranial vault (Figure 7.3). The
facial bones underlie the facial structures, form
the nasal cavity, enclose the eyeballs, and
support the teeth of the upper and lower jaws.
The rounded brain case surrounds and protects
the brain and houses the middle and inner ear
structures. In the adult, the skull consists of 22
individual bones, 21 of which are immobile and
united into a single unit. The 22nd bone is
the mandible (lower jaw), which is the only
moveable bone of the skull.

Figure 7.3 Parts of the Skull The skull consists

of the rounded brain case that houses the brain
and the facial bones that form the upper and
lower jaws, nose, orbits, and other facial
structures.

The orbit is the bony socket that houses the

eyeball and muscles that move the eyeball or
open the upper eyelid. The upper margin of the
anterior orbit is the supraorbital margin.
Located near the midpoint of the supraorbital margin is a small opening called the supraorbital foramen. This provides for passage of a sensory nerve
to the skin of the forehead. Below the orbit is the infraorbital foramen, which is the point of emergence for a sensory nerve that supplies the anterior
face below the orbit.

Figure 7.4 Anterior View of Skull An anterior view of the skull shows the bones that form the forehead, orbits (eye sockets), nasal cavity, nasal
septum, and upper and lower jaws.
 Inside the nasal area of the skull, the nasal cavity is divided into halves by the nasal septum. The
upper portion of the nasal septum is formed by the perpendicular plate of the ethmoid bone and
the lower portion is the vomer bone. Each side of the nasal cavity is triangular in shape, with a broad
inferior space that narrows superiorly. When looking into the nasal cavity from the front of the skull,
two bony plates are seen projecting from each lateral wall. The larger of these is the inferior nasal
concha, an independent bone of the skull. Located just above the inferior concha is the middle nasal
concha, which is part of the ethmoid bone. A third bony plate, also part of the ethmoid bone, is
the superior nasal concha. It is much smaller and out of sight, above the middle concha. The
superior nasal concha is located just lateral to the perpendicular plate, in the upper nasal cavity.

Lateral View of Skull

A view of the lateral skull is dominated by the large, rounded brain case above and the upper and
lower jaws with their teeth below (Figure 7.5). Separating these areas is the bridge of bone called the
zygomatic arch. The zygomatic arch is the bony arch on the side of skull that spans from the area of
the cheek to just above the ear canal. It is formed by the junction of two bony processes: a short
anterior component, the temporal process of the zygomatic bone (the cheekbone) and a longer
posterior portion, the zygomatic process of the temporal bone, extending forward from the
temporal bone. Thus the temporal process (anteriorly) and the zygomatic process (posteriorly) join
together, like the two ends of a drawbridge, to form the zygomatic arch. One of the major muscles
that pulls the mandible upward during biting and chewing arises from the zygomatic arch.

On the lateral side of the brain case, above the level of the zygomatic arch, is a shallow space
called the temporal fossa. Below the level of the zygomatic arch and deep to the vertical
portion of the mandible is another space called the infratemporal fossa. Both the temporal
fossa and infratemporal fossa contain muscles that act on the mandible during chewing.
Figure 7.5 Lateral View of Skull The lateral skull shows the large rounded brain case,
zygomatic arch, and the upper and lower jaws. The zygomatic arch is formed jointly by the
zygomatic process of the temporal bone and the temporal process of the zygomatic bone. The
shallow space above the zygomatic arch is the temporal fossa. The space inferior to the
zygomatic arch and deep to the posterior mandible is the infratemporal fossa.

Bones of the Brain Case

The brain case contains and protects the brain. The interior space that is almost completely
occupied by the brain is called the cranial cavity. This cavity is bounded superiorly by the
rounded top of the skull, which is called the calvaria (skullcap), and the lateral and posterior
sides of the skull. The bones that form the top and sides of the brain case are usually referred to as the “flat” bones of the skull.

The floor of the brain case is referred to as the base of the skull. This is a complex area that varies in depth and has numerous openings for the passage
of cranial nerves, blood vessels, and the spinal cord. Inside the skull, the base is subdivided into three large spaces, called the anterior cranial
fossa, middle cranial fossa, and posterior cranial fossa (fossa = “trench or ditch”) (Figure 7.6). From anterior to posterior, the fossae increase in
depth. The shape and depth of each fossa corresponds to the shape and size of the brain region that each houses. The boundaries and openings of the
cranial fossae (singular = fossa) will be described in a later section. Figure 7.6 Cranial Fossae The bones of the brain case surround and protect the
brain, which occupies the cranial cavity. The base of the brain case, which forms the floor of cranial cavity, is subdivided into the shallow anterior cranial
fossa, the middle cranial fossa, and the deep posterior cranial fossa.

The brain case consists of eight bones. These include the paired parietal and temporal bones, plus the unpaired frontal, occipital, sphenoid, and ethmoid
bones.
Parietal Bone

The parietal bone forms most of the upper lateral side of the skull (see Figure 7.5). These are paired bones, with the right and left parietal bones joining
together at the top of the skull. Each parietal bone is also bounded anteriorly by the frontal bone, inferiorly by the temporal bone, and posteriorly by the
occipital bone.
Temporal Bone

The temporal bone forms the lower lateral side of the skull (see Figure 7.5). Common wisdom has it that the temporal bone (temporal = “time”) is so
named because this area of the head (the temple) is where hair typically first turns gray, indicating the passage of time.

The temporal bone is subdivided into several regions (Figure 7.7). The flattened, upper portion is the squamous portion of the temporal bone. Below this
area and projecting anteriorly is the zygomatic process of the temporal bone, which forms the posterior portion of the zygomatic arch. Posteriorly is the
mastoid portion of the temporal bone. Projecting inferiorly from this region is a large prominence, the mastoid process, which serves as a muscle
attachment site. The mastoid process can easily be felt on the side of the head just behind your earlobe. On the interior of the skull, the petrous portion
of each temporal bone forms the prominent, diagonally oriented petrous ridge in the floor of the cranial cavity. Located inside each petrous ridge are
small cavities that house the structures of the middle and inner ears.

Figure 7.7 Temporal Bone A lateral view of the isolated temporal bone shows the squamous, mastoid, and zygomatic portions of the temporal bone.
Important landmarks of the temporal bone, as shown in Figure 7.8, include the following:

 External acoustic meatus (ear canal)—This is the large opening on the lateral side of the skull that is associated with the ear.
 Internal acoustic meatus—This opening is located inside the cranial cavity, on the medial side of the petrous ridge. It connects to the middle
and inner ear cavities of the temporal bone.
 Mandibular fossa—This is the deep, oval-shaped depression
located on the external base of the skull, just in front of the
external acoustic meatus. The mandible (lower jaw) joins with the
skull at this site as part of the temporomandibular joint, which
allows for movements of the mandible during opening and closing
of the mouth.
 Articular tubercle—The smooth ridge located immediately
anterior to the mandibular fossa. Both the articular tubercle and
mandibular fossa contribute to the temporomandibular joint, the
joint that provides for movements between the temporal bone of
the skull and the mandible.
 Styloid process—Posterior to the mandibular fossa on the
external base of the skull is an elongated, downward bony
projection called the styloid process, so named because of its
resemblance to a stylus (a pen or writing tool). This structure
serves as an attachment site for several small muscles and for a
ligament that supports the hyoid bone of the neck. (See
also Figure 7.7.)
 Stylomastoid foramen—This small
opening is located between the styloid
process and mastoid process. This is the
point of exit for the cranial nerve that
supplies the facial muscles.
 Carotid canal—The carotid canal is a zig-
zag shaped tunnel that provides passage
through the base of the skull for one of the
major arteries that supplies the brain. Its
entrance is located on the outside base of
the skull, anteromedial to the styloid
process. The canal then runs
anteromedially within the bony base of the
skull, and then turns upward to its exit in
the floor of the middle cranial cavity,
above the foramen lacerum.

Figure 7.8 External and Internal Views of Base of

Skull (a) The hard palate is formed anteriorly by the
palatine processes of the maxilla bones and
posteriorly by the horizontal plate of the palatine
bones. (b) The complex floor of the cranial cavity is
formed by the frontal, ethmoid, sphenoid, temporal,
and occipital bones. The lesser wing of the sphenoid
bone separates the anterior and middle cranial
fossae. The petrous ridge (petrous portion of
temporal bone) separates the middle and posterior
cranial fossae.
Frontal Bone

The frontal bone is the single bone that forms the

forehead. At its anterior midline, between the
eyebrows, there is a slight depression called
the glabella (see Figure 7.5). The frontal bone also
forms the supraorbital margin of the orbit. Near the
middle of this margin, is the supraorbital foramen,
the opening that provides passage for a sensory
nerve to the forehead. The frontal bone is thickened
just above each supraorbital margin, forming
rounded brow ridges. These are located just behind
your eyebrows and vary in size among individuals,
although they are generally larger in males. Inside
the cranial cavity, the frontal bone extends
posteriorly. This flattened region forms both the roof
of the orbit below and the floor of the anterior cranial
cavity above (see Figure 7.8b).
 Occipital Bone

The occipital bone is the single bone that forms the posterior skull
and posterior base of the cranial cavity (Figure 7.9; see also Figure
7.8). On its outside surface, at the posterior midline, is a small
protrusion called the external occipital protuberance, which serves
as an attachment site for a ligament of the posterior neck. Lateral to
either side of this bump is a superior nuchal line (nuchal = “nape” or
“posterior neck”). The nuchal lines represent the most superior point
at which muscles of the neck attach to the skull, with only the
scalp covering the skull above these lines. On the base of
the skull, the occipital bone contains the large opening of
the foramen magnum, which allows for passage of the
spinal cord as it exits the skull. On either side of the foramen
magnum is an oval-shaped occipital condyle. These
condyles form joints with the first cervical vertebra and thus
support the skull on top of the vertebral column.

Figure 7.9 Posterior View of Skull This view of the

posterior skull shows attachment sites for muscles and joints
that support the skull.
Sphenoid Bone

The sphenoid bone is a single, complex bone of the central

skull (Figure 7.10). It serves as a “keystone” bone, because it
joins with almost every other bone of the skull. The sphenoid
forms much of the base of the central skull (see Figure 7.8)
and also extends laterally to contribute to the sides of the
skull (see Figure 7.5). Inside the cranial cavity, the right and
left lesser wings of the sphenoid bone, which resemble the
wings of a flying bird, form the lip of a prominent ridge that
marks the boundary between the anterior and middle cranial
fossae. The sella turcica (“Turkish saddle”) is located at the
midline of the middle cranial fossa. This bony region of the
sphenoid bone is named for its resemblance to the horse saddles
used by the Ottoman Turks, with a high back and a tall front. The
rounded depression in the floor of the sella turcica is the hypophyseal (pituitary) fossa, which houses the pea-sized pituitary (hypophyseal) gland.
The greater wings of the sphenoid bone extend laterally to either side away from the sella turcica, where they form the anterior floor of the middle
cranial fossa. The greater wing is best seen on the outside of the lateral skull, where it forms a rectangular area immediately anterior to the squamous
portion of the temporal bone.

On the inferior aspect of the skull, each half of the sphenoid bone forms two thin, vertically oriented bony plates. These are the medial pterygoid
plate and lateral pterygoid plate (pterygoid = “wing-shaped”). The right and left medial pterygoid plates form the posterior, lateral walls of the nasal
cavity. The somewhat larger lateral pterygoid plates serve as attachment sites for chewing muscles that fill the infratemporal space and act on the
mandible.

Figure 7.10 Sphenoid Bone Shown in isolation in (a) superior and (b) posterior views, the sphenoid bone is a single midline bone that forms the
anterior walls and floor of the middle cranial fossa. It has a pair of lesser wings and a pair of greater wings. The sella turcica surrounds the hypophyseal
fossa. Projecting downward are the medial and lateral pterygoid plates. The sphenoid has multiple openings for the passage of nerves and blood
vessels, including the optic canal, superior orbital fissure, foramen rotundum, foramen ovale, and foramen spinosum.
Ethmoid Bone

The ethmoid bone is a single, midline bone that forms the roof and lateral walls of the upper nasal cavity, the upper portion of the nasal septum, and
contributes to the medial wall of the orbit (Figure 7.11 and Figure 7.12). On the interior of the skull, the ethmoid also forms a portion of the floor of the
anterior cranial cavity (see Figure 7.8b).

Within the nasal cavity, the perpendicular plate of the ethmoid bone forms the upper portion of the nasal septum. The ethmoid bone also forms the
lateral walls of the upper nasal cavity. Extending from each lateral wall are the superior nasal concha and middle nasal concha, which are thin, curved
projections that extend into the nasal cavity (Figure 7.13).

In the cranial cavity, the ethmoid bone forms a small area at the midline in the floor of the anterior cranial fossa. This region also forms the narrow roof of
the underlying nasal cavity. This portion of the ethmoid bone consists of two parts, the crista galli and cribriform plates. The crista galli (“rooster’s comb
or crest”) is a small upward bony projection located at the midline. It functions as an anterior attachment point for one of the covering layers of the brain.
To either side of the crista galli is the cribriform plate (cribrum = “sieve”), a small, flattened area with numerous small openings termed olfactory
foramina. Small nerve branches from the olfactory areas of the nasal cavity pass through these openings to enter the brain.

The lateral portions of the ethmoid bone are located between the orbit and upper nasal cavity, and thus form the lateral nasal cavity wall and a portion of
the medial orbit wall. Located inside this portion of the ethmoid bone are several small, air-filled spaces that are part of the paranasal sinus system of the
skull.
 Figure 7.11 Sagittal Section of Skull This midline view of
the sagittally sectioned skull shows the nasal septum.

Figure 7.12 Ethmoid Bone The unpaired ethmoid bone is

located at the midline within the central skull. It has an
upward projection, the crista galli, and a downward
projection, the perpendicular plate, which forms the upper
nasal septum. The cribriform plates form both the roof of
the nasal cavity and a portion of the anterior cranial fossa
floor. The lateral sides of the ethmoid bone form the lateral
walls of the upper nasal cavity, part of the medial orbit wall,
and give rise to the superior and middle nasal conchae.
The ethmoid bone also contains the ethmoid air cells.

Figure 7.13 Lateral Wall of Nasal Cavity The three nasal conchae

are curved bones that project from the lateral walls of the nasal
cavity. The superior nasal concha and middle nasal concha are parts
of the ethmoid bone. The inferior nasal concha is an independent bone of
the skull.

Sutures of the Skull

A suture is an immobile joint between adjacent bones of the skull. The

narrow gap between the bones is filled with dense, fibrous connective
tissue that unites the bones. The long sutures located between the bones
of the brain case are not straight, but instead follow irregular, tightly twisting
paths. These twisting lines serve to tightly interlock the adjacent bones,
thus adding strength to the skull for brain protection.

The two suture lines seen on the top of the skull are the coronal and
sagittal sutures. The coronal suture runs from side to side across the
skull, within the coronal plane of section (see Figure 7.5). It joins the frontal
bone to the right and left parietal bones. The sagittal suture extends
posteriorly from the coronal suture, running along the midline at the top of
the skull in the sagittal plane of section (see Figure 7.9). It unites the right
and left parietal bones. On the posterior skull, the sagittal suture terminates by joining the lambdoid suture. The lambdoid suture extends downward
and laterally to either side away from its junction with the sagittal suture. The lambdoid suture joins the occipital bone to the right and left parietal and
temporal bones. This suture is named for its upside-down "V" shape, which resembles the capital letter version of the Greek letter lambda (Λ).
The squamous suture is located on the lateral skull. It unites the squamous portion of the temporal bone with the parietal bone (see Figure 7.5). At the
intersection of four bones is the pterion, a small, capital-H-shaped suture line region that unites the frontal bone, parietal bone, squamous portion of the
temporal bone, and greater wing of the sphenoid bone. It is the weakest part of the skull. The pterion is located approximately two finger widths above
the zygomatic arch and a thumb’s width posterior to the upward portion of the zygomatic bone.

DISORDERS OF THE...
Skeletal System

Head and traumatic brain injuries are major causes of immediate death and disability, with bleeding and infections as possible additional complications.
According to the Centers for Disease Control and Prevention (2010), approximately 30 percent of all injury-related deaths in the United States are
caused by head injuries. The majority of head injuries involve falls. They are most common among young children (ages 0–4 years), adolescents (15–19
years), and the elderly (over 65 years). Additional causes vary, but prominent among these are automobile and motorcycle accidents.

Strong blows to the brain-case portion of the skull can produce fractures. These may result in bleeding inside the skull with subsequent injury to the
brain. The most common is a linear skull fracture, in which fracture lines radiate from the point of impact. Other fracture types include a comminuted
fracture, in which the bone is broken into several pieces at the point of impact, or a depressed fracture, in which the fractured bone is pushed inward. In
a contrecoup (counterblow) fracture, the bone at the point of impact is not broken, but instead a fracture occurs on the opposite side of the skull.
Fractures of the occipital bone at the base of the skull can occur in this manner, producing a basilar fracture that can damage the artery that passes
through the carotid canal.

A blow to the lateral side of the head may fracture the bones of the pterion. The pterion is an important clinical landmark because located immediately
deep to it on the inside of the skull is a major branch of an artery that supplies the skull and covering layers of the brain. A strong blow to this region can
fracture the bones around the pterion. If the underlying artery is damaged, bleeding can cause the formation of a hematoma (collection of blood)
between the brain and interior of the skull. As blood accumulates, it will put pressure on the brain. Symptoms associated with a hematoma may not be
apparent immediately following the injury, but if untreated, blood accumulation will exert increasing pressure on the brain and can result in death within a
few hours.

INTERACTIVE LINK

View this animation to see how a blow to the head may produce a contrecoup (counterblow) fracture of the basilar portion of the occipital bone on the
base of the skull. Why may a basilar fracture be life threatening?

Facial Bones of the Skull

The facial bones of the skull form the upper and lower jaws, the nose, nasal cavity and nasal septum, and the orbit. The facial bones include 14 bones,
with six paired bones and two unpaired bones. The paired bones are the maxilla, palatine, zygomatic, nasal, lacrimal, and inferior nasal conchae bones.
The unpaired bones are the vomer and mandible bones. Although classified with the brain-case bones, the ethmoid bone also contributes to the nasal
septum and the walls of the nasal cavity and orbit.
Maxillary Bone

The maxillary bone, often referred to simply as the maxilla (plural = maxillae), is one of a pair that together form the upper jaw, much of the hard palate,
the medial floor of the orbit, and the lateral base of the nose (see Figure 7.4). The curved, inferior margin of the maxillary bone that forms the upper jaw
and contains the upper teeth is the alveolar process of the maxilla (Figure 7.14). Each tooth is anchored into a deep socket called an alveolus. On the
anterior maxilla, just below the orbit, is the infraorbital foramen. This is the point of exit for a sensory nerve that supplies the nose, upper lip, and anterior
cheek. On the inferior skull, the palatine process from each maxillary bone can be seen joining together at the midline to form the anterior three-
quarters of the hard palate (see Figure 7.8a). The hard palate is the bony plate that forms the roof of the mouth and floor of the nasal cavity, separating
the oral and nasal cavities.

Figure 7.14 Maxillary Bone The maxillary bone forms the upper jaw and supports the upper teeth. Each maxilla also forms the lateral floor of each orbit
and the majority of the hard palate.
Palatine Bone

The palatine bone is one of a pair of irregularly shaped bones that

contribute small areas to the lateral walls of the nasal cavity and the medial
wall of each orbit. The largest region of each of the palatine bone is
the horizontal plate. The plates from the right and left palatine bones join
together at the midline to form the posterior quarter of the hard palate
(see Figure 7.8a). Thus, the palatine bones are best seen in an inferior view
of the skull and hard palate.

HOMEOSTATIC IMBALANCES
Cleft Lip and Cleft Palate

During embryonic development, the right and left maxilla bones come
together at the midline to form the upper jaw. At the same time, the muscle
and skin overlying these bones join together to form the upper lip. Inside
the mouth, the palatine processes of the maxilla bones, along with the
horizontal plates of the right and left palatine bones, join together to form
the hard palate. If an error occurs in these developmental processes, a birth
defect of cleft lip or cleft palate may result.

Cleft lip is a common development defect that affects approximately 1:1000 births, most of which are male. This defect involves a partial or complete
failure of the right and left portions of the upper lip to fuse together, leaving a cleft (gap).

A more severe developmental defect is cleft palate, which affects the hard palate. The hard palate is the bony structure that separates the nasal cavity
from the oral cavity. It is formed during embryonic development by the midline fusion of the horizontal plates from the right and left palatine bones and
the palatine processes of the maxilla bones. Cleft palate affects approximately 1:2500 births and is more common in females. It results from a failure of
the two halves of the hard palate to completely come together and fuse at the midline, thus leaving a gap between them. This gap allows for
communication between the nasal and oral cavities. In severe cases, the bony gap continues into the anterior upper jaw where the alveolar processes of
the maxilla bones also do not properly join together above the front teeth. If this occurs, a cleft lip will also be seen. Because of the communication
between the oral and nasal cavities, a cleft palate makes it very difficult for an infant to generate the suckling needed for nursing, thus leaving the infant
at risk for malnutrition. Surgical repair is required to correct cleft palate defects.
Zygomatic Bone

The zygomatic bone is also known as the cheekbone. Each of the paired zygomatic bones forms much of the lateral wall of the orbit and the lateral-
inferior margins of the anterior orbital opening (see Figure 7.4). The short temporal process of the zygomatic bone projects posteriorly, where it forms
the anterior portion of the zygomatic arch (see Figure 7.5).
Nasal Bone

The nasal bone is one of two small bones that articulate (join) with each other to form the bony base (bridge) of the nose. They also support the
cartilages that form the lateral walls of the nose (see Figure 7.11). These are the bones that are damaged when the nose is broken.
Lacrimal Bone

Each lacrimal bone is a small, rectangular bone that forms the anterior, medial wall of the orbit (see Figure 7.4 and Figure 7.5). The anterior portion of
the lacrimal bone forms a shallow depression called the lacrimal fossa, and extending inferiorly from this is the nasolacrimal canal. The lacrimal fluid
(tears of the eye), which serves to maintain the moist surface of the eye, drains at the medial corner of the eye into the nasolacrimal canal. This duct
then extends downward to open into the nasal cavity, behind the inferior nasal concha. In the nasal cavity, the lacrimal fluid normally drains posteriorly,
but with an increased flow of tears due to crying or eye irritation, some fluid will also drain anteriorly, thus causing a runny nose.
Inferior Nasal Conchae

The right and left inferior nasal conchae form a curved bony plate that projects into the nasal cavity space from the lower lateral wall (see Figure 7.13).
The inferior concha is the largest of the nasal conchae and can easily be seen when looking into the anterior opening of the nasal cavity.
Vomer Bone

The unpaired vomer bone, often referred to simply as the vomer, is triangular-shaped and forms the posterior-inferior part of the nasal septum
(see Figure 7.11). The vomer is best seen when looking from behind into the posterior openings of the nasal cavity (see Figure 7.8a). In this view, the
vomer is seen to form the entire height of the nasal septum. A much smaller portion of the vomer can also be seen when looking into the anterior
opening of the nasal cavity.
Mandible

The mandible forms the lower jaw and is the only moveable bone of the skull. At the time of birth, the mandible consists of paired right and left bones,
but these fuse together during the first year to form the single U-shaped mandible of the adult skull. Each side of the mandible consists of a horizontal
body and posteriorly, a vertically oriented ramus of the mandible (ramus = “branch”). The outside margin of the mandible, where the body and ramus
come together is called the angle of the mandible (Figure 7.15).

The ramus on each side of the mandible has two upward-going bony projections. The more anterior projection is the flattened coronoid process of the
mandible, which provides attachment for one of the biting muscles. The posterior projection is the condylar process of the mandible, which is topped
by the oval-shaped condyle. The condyle of the mandible articulates (joins) with the mandibular fossa and articular tubercle of the temporal bone.
Together these articulations form the temporomandibular joint, which allows for opening and closing of the mouth (see Figure 7.5). The broad U-shaped
curve located between the coronoid and condylar processes is the mandibular notch.

Important landmarks for the mandible include the following:

 Alveolar process of the mandible—This is the upper border of the mandibular body and serves to anchor the lower teeth.
 Mental protuberance—The forward projection from the inferior margin of the anterior mandible that forms the chin (mental = “chin”).
 Mental foramen—The opening located on each side of the anterior-lateral mandible, which is the exit site for a sensory nerve that supplies
the chin.
 Mylohyoid line—This bony ridge extends along the inner aspect of the mandibular body (see Figure 7.11). The muscle that forms the floor of
the oral cavity attaches to the mylohyoid lines on both sides of the mandible.
 Mandibular foramen—This opening is located on the medial side of the ramus of the mandible. The opening leads into a tunnel that runs
down the length of the mandibular body. The sensory
nerve and blood vessels that supply the lower teeth
enter the mandibular foramen and then follow this tunnel.
 Thus, to numb the lower teeth prior to dental work, the dentist must inject anesthesia into the lateral wall of the oral cavity at a point prior to
where this sensory nerve enters the mandibular foramen.
 Lingula—This small flap of bone is named for its shape (lingula = “little tongue”). It is located immediately next to the mandibular foramen, on
the medial side of the ramus. A ligament that anchors the mandible during opening and closing of the mouth extends down from the base of
the skull and attaches to the lingula. Figure 7.15 Isolated Mandible The mandible is the only moveable bone of the skull.

The Orbit

The orbit is the bony socket that houses the eyeball and contains the muscles that move the eyeball or open the upper eyelid. Each orbit is cone-
shaped, with a narrow posterior region that widens toward the large anterior opening. To help protect the eye, the bony margins of the anterior opening
are thickened and somewhat constricted. The medial walls of the two orbits are parallel to each other but each lateral wall diverges away from the
midline at a 45° angle. This divergence provides greater lateral peripheral vision.

The walls of each orbit include contributions from seven skull bones (Figure 7.16). The frontal bone forms the roof and the zygomatic bone forms the
lateral wall and lateral floor. The medial floor is primarily formed by the maxilla, with a small contribution from the palatine bone. The ethmoid bone and
lacrimal bone make up much of the medial wall and the sphenoid bone forms the posterior orbit.

At the posterior apex of the orbit is the opening of the optic canal, which allows for passage of the optic nerve from the retina to the brain. Lateral to this
is the elongated and irregularly shaped superior orbital fissure, which provides passage for the artery that supplies the eyeball, sensory nerves, and the
nerves that supply the muscles involved in eye movements.

Figure 7.16 Bones of the Orbit Seven skull bones

contribute to the walls of the orbit. Opening into the
posterior orbit from the cranial cavity are the optic canal and
superior orbital fissure.

The Nasal Septum and Nasal Conchae

The nasal septum consists of both bone and cartilage

components (Figure 7.17; see also Figure 7.11). The upper
portion of the septum is formed by the perpendicular plate
of the ethmoid bone. The lower and posterior parts of the
septum are formed by the triangular-shaped vomer bone. In
an anterior view of the skull, the perpendicular plate of the
ethmoid bone is easily seen inside the nasal opening as the
upper nasal septum, but only a small portion of the vomer is seen as the inferior septum. A better view of the vomer bone is seen when looking into the
posterior nasal cavity with an inferior view of the skull, where the vomer forms the full height of the nasal septum. The anterior nasal septum is formed by
the septal cartilage, a flexible plate that fills in the gap between the perpendicular plate of the ethmoid and vomer bones. This cartilage also extends
outward into the nose where it separates the right and left nostrils. The septal cartilage is not found in the dry skull.

Attached to the lateral wall on each side of the nasal cavity are the superior, middle, and inferior nasal conchae (singular = concha), which are named
for their positions (see Figure 7.13). These are bony plates that curve downward as they project into the space of the nasal cavity. They serve to swirl
the incoming air, which helps to warm and moisturize it before the air moves into the delicate air sacs of the lungs. This also allows mucus, secreted by
the tissue lining the nasal cavity, to trap incoming dust, pollen, bacteria, and viruses. The largest of the conchae is the inferior nasal concha, which is an
independent bone of the skull. The middle concha and the superior conchae, which is the smallest, are both formed by the ethmoid bone. When looking
into the anterior nasal opening of the skull, only the inferior and middle conchae can be seen. The small superior nasal concha is well hidden above and
behind the middle concha.

Figure 7.17 Nasal Septum The nasal septum is formed by the perpendicular plate of the ethmoid bone and the vomer bone. The septal cartilage fills
the gap between these bones and extends into the nose.

Cranial Fossae

Inside the skull, the floor of the cranial cavity is subdivided into three cranial fossae (spaces), which increase in depth from anterior to posterior
(see Figure 7.6, Figure 7.8b, and Figure 7.11). Since the brain occupies these areas, the shape of each conforms to the shape of the brain regions that
it contains. Each cranial fossa has anterior and posterior boundaries and is divided at the midline into right and left areas by a significant bony structure
or opening.
Anterior Cranial Fossa

The anterior cranial fossa is the most anterior and the shallowest of the three cranial fossae. It overlies the orbits and contains the frontal lobes of the
brain. Anteriorly, the anterior fossa is bounded by the frontal bone, which also forms the majority of the floor for this space. The lesser wings of the
sphenoid bone form the prominent ledge that marks the boundary between the anterior and middle cranial fossae. Located in the floor of the anterior
cranial fossa at the midline is a portion of the ethmoid bone, consisting of the upward projecting crista galli and to either side of this, the cribriform plates.
Middle Cranial Fossa

The middle cranial fossa is deeper and situated posterior to the anterior fossa. It extends from the lesser wings of the sphenoid bone anteriorly, to the
petrous ridges (petrous portion of the temporal bones) posteriorly. The large, diagonally positioned petrous ridges give the middle cranial fossa a
butterfly shape, making it narrow at the midline and broad laterally. The temporal lobes of the brain occupy this fossa. The middle cranial fossa is divided
at the midline by the upward bony prominence of the sella turcica, a part of the sphenoid bone. The middle cranial fossa has several openings for the
passage of blood vessels and cranial nerves (see Figure 7.8).Openings in the middle cranial fossa are as follows:

 Optic canal—This opening is located at the anterior lateral corner of the sella turcica. It provides for passage of the optic nerve into the orbit.
 Superior orbital fissure—This large, irregular opening into the posterior orbit is located on the anterior wall of the middle cranial fossa, lateral
to the optic canal and under the projecting margin of the lesser wing of the sphenoid bone. Nerves to the eyeball and associated muscles, and
sensory nerves to the forehead pass through this opening.
 Foramen rotundum—This rounded opening (rotundum = “round”) is located in the floor of the middle cranial fossa, just inferior to the superior
orbital fissure. It is the exit point for a major sensory nerve that supplies the cheek, nose, and upper teeth.
 Foramen ovale of the middle cranial fossa—This large, oval-shaped opening in the floor of the middle cranial fossa provides passage for a
major sensory nerve to the lateral head, cheek, chin, and lower teeth.
 Foramen spinosum—This small opening, located posterior-lateral to the foramen ovale, is the entry point for an important artery that supplies
the covering layers surrounding the brain. The branching pattern of this artery forms readily visible grooves on the internal surface of the skull
and these grooves can be traced back to their origin at the foramen spinosum.
 Carotid canal—This is the zig-zag passageway through which a major artery to the brain enters the skull. The entrance to the carotid canal is
located on the inferior aspect of the skull, anteromedial to the styloid process (see Figure 7.8a). From here, the canal runs anteromedially
within the bony base of the skull. Just above the foramen lacerum, the carotid canal opens into the middle cranial cavity, near the posterior-
lateral base of the sella turcica.
 Foramen lacerum—This irregular opening is located in the base of the skull, immediately inferior to the exit of the carotid canal. This opening
is an artifact of the dry skull, because in life it is completely filled with cartilage. All the openings of the skull that provide for passage of nerves
or blood vessels have smooth margins; the word lacerum (“ragged” or “torn”) tells us that this opening has ragged edges and thus nothing
passes through it.

The posterior cranial fossa is the most posterior and deepest portion of the cranial cavity. It contains the cerebellum of the brain. The posterior fossa is
bounded anteriorly by the petrous ridges, while the occipital bone forms the floor and posterior wall. It is divided at the midline by the large foramen
magnum (“great aperture”), the opening that provides for passage of the spinal cord.

Located on the medial wall of the petrous ridge in the posterior cranial fossa is the internal acoustic meatus (see Figure 7.11). This opening provides for
passage of the nerve from the hearing and equilibrium organs of the inner ear, and the nerve that supplies the muscles of the face. Located at the
anterior-lateral margin of the foramen magnum is the hypoglossal canal. These emerge on the inferior aspect of the skull at the base of the occipital
condyle and provide passage for an important nerve to the tongue.
Immediately inferior to the internal acoustic meatus is the large, irregularly shaped jugular foramen (see Figure 7.8a). Several cranial nerves from the
brain exit the skull via this opening. It is also the exit point through the base of the skull for all the venous return blood leaving the brain. The venous
structures that carry blood inside the skull form large, curved grooves on the inner walls of the posterior cranial fossa, which terminate at each jugular
foramen.

The paranasal sinuses are hollow, air-filled spaces located within certain bones of the skull (Figure 7.18). All of the sinuses communicate with the nasal
cavity (paranasal = “next to nasal cavity”) and are lined with nasal mucosa. They serve to reduce bone mass and thus lighten the skull, and they also
add resonance to the voice. This second feature is most obvious when you have a cold or sinus congestion. These produce swelling of the mucosa and
excess mucus production, which can obstruct the narrow passageways between the sinuses and the nasal cavity, causing your voice to sound different
to yourself and others. This blockage can also allow the sinuses to fill with fluid, with the resulting pressure producing pain and discomfort.

The paranasal sinuses are named for the skull bone that each occupies. The frontal sinus is located just above the eyebrows, within the frontal bone
(see Figure 7.17). This irregular space may be divided at the midline into bilateral spaces, or these may be fused into a single sinus space. The frontal
sinus is the most anterior of the paranasal sinuses. The largest sinus is the maxillary sinus. These are paired and located within the right and left
maxillary bones, where they occupy the area just below the orbits. The maxillary sinuses are most commonly involved during sinus infections. Because
their connection to the nasal cavity is located high on their medial wall, they are difficult to drain. The sphenoid sinus is a single, midline sinus. It is
located within the body of the sphenoid bone, just anterior and inferior to the sella turcica, thus making it the most posterior of the paranasal sinuses.
The lateral aspects of the ethmoid bone contain multiple small spaces separated by very thin bony walls. Each of these spaces is called an ethmoid air
cell. These are located on both sides of the ethmoid bone, between the upper nasal cavity and medial orbit, just behind the superior nasal conchae.

Figure 7.18 Paranasal Sinuses The paranasal sinuses are hollow, air-filled spaces named for the skull bone that each occupies. The most anterior is
the frontal sinus, located in the frontal bone above the eyebrows. The largest are the
maxillary sinuses, located in the right and left maxillary bones below the orbits. The
most posterior is the sphenoid sinus, located in the body of the sphenoid bone, under
the sella turcica. The ethmoid air cells are multiple small spaces located in the right
and left sides of the ethmoid bone, between the medial wall of the orbit and lateral wall
of the upper nasal cavity.

Hyoid Bone

The hyoid bone is an independent bone that does not contact any other bone and thus
is not part of the skull (Figure 7.19). It is a small U-shaped bone located in the upper
neck near the level of the inferior mandible, with the tips of the “U” pointing posteriorly.
The hyoid serves as the base for the tongue above, and is attached to the larynx
below and the pharynx posteriorly. The hyoid is held in position by a series of small
muscles that attach to it either from above or below. These muscles act to move the
hyoid up/down or forward/back. Movements of the hyoid are coordinated with
movements of the tongue, larynx, and pharynx during swallowing and speaking.
Figure 7.19 Hyoid Bone The hyoid bone is located in the upper neck and does not join with any other bone. It provides attachments for muscles that
act on the tongue, larynx, and pharynx.

The vertebral column is also known as the spinal column or spine (Figure 7.20). It consists of a sequence of
vertebrae (singular = vertebra), each of which is separated and united by an intervertebral disc. Together,
the vertebrae and intervertebral discs form the vertebral column. It is a flexible column that supports the
head, neck, and body and allows for their movements. It also protects the spinal cord, which passes down
the back through openings in the vertebrae.

Figure 7.20 Vertebral Column The adult vertebral column consists of 24 vertebrae, plus the sacrum and
coccyx. The vertebrae are divided into three regions: cervical C1–C7 vertebrae, thoracic T1–T12 vertebrae,
and lumbar L1–L5 vertebrae. The vertebral column is curved, with two primary curvatures (thoracic and
sacrococcygeal curves) and two secondary curvatures (cervical and lumbar curves).

Regions of the Vertebral Column

The vertebral column originally develops as a series of 33 vertebrae, but this number is eventually reduced
to 24 vertebrae, plus the sacrum and coccyx. The vertebral column is subdivided into five regions, with the
vertebrae in each area named for that region and numbered in descending order. In the neck, there are
seven cervical vertebrae, each designated with the letter “C” followed by its number. Superiorly, the C1
vertebra articulates (forms a joint) with the occipital condyles of the skull. Inferiorly, C1 articulates with the
C2 vertebra, and so on. Below these are the 12 thoracic vertebrae, designated T1–T12. The lower back
contains the L1–L5 lumbar vertebrae. The single sacrum, which is also part of the pelvis, is formed by the
fusion of five sacral vertebrae. Similarly, the coccyx, or tailbone, results from the fusion of four small
coccygeal vertebrae. However, the sacral and coccygeal fusions do not start until age 20 and are not
completed until middle age.

An interesting anatomical fact is that almost all mammals have seven cervical vertebrae, regardless of body size. This means that there are large
variations in the size of cervical vertebrae, ranging from the very small cervical vertebrae of a shrew to the greatly elongated vertebrae in the neck of a
giraffe. In a full-grown giraffe, each cervical vertebra is 11 inches tall.

Curvatures of the Vertebral Column

The adult vertebral column does not form a straight line, but instead has four curvatures along its length (see Figure 7.20). These curves increase the
vertebral column’s strength, flexibility, and ability to absorb shock. When the load on the spine is increased, by carrying a heavy backpack for example,
the curvatures increase in depth (become more curved) to accommodate the extra weight. They then spring back when the weight is removed. The four
adult curvatures are classified as either primary or secondary curvatures. Primary curves are retained from the original fetal curvature, while secondary
curvatures develop after birth.

During fetal development, the body is flexed anteriorly into the fetal position, giving the entire vertebral column a single curvature that is concave
anteriorly. In the adult, this fetal curvature is retained in two regions of the vertebral column as the thoracic curve, which involves the thoracic
vertebrae, and the sacrococcygeal curve, formed by the sacrum and coccyx. Each of these is thus called a primary curve because they are retained
from the original fetal curvature of the vertebral column.

A secondary curve develops gradually after birth as the child learns to sit upright, stand, and walk. Secondary curves are concave posteriorly, opposite
in direction to the original fetal curvature. The cervical curve of the neck region develops as the infant begins to hold their head upright when sitting.
Later, as the child begins to stand and then to walk, the lumbar curve of the lower back develops. In adults, the lumbar curve is generally deeper in
females.

Disorders associated with the curvature of the spine include kyphosis (an excessive posterior curvature of the thoracic region), lordosis (an excessive
anterior curvature of the lumbar region), and scoliosis (an abnormal, lateral curvature, accompanied by twisting of the vertebral column).

DISORDERS OF THE...
Vertebral Column

Developmental anomalies, pathological changes, or obesity can enhance the normal vertebral column curves, resulting in the development of abnormal
or excessive curvatures (Figure 7.21). Kyphosis, also referred to as humpback or hunchback, is an excessive posterior curvature of the thoracic region.
This can develop when osteoporosis causes weakening and erosion of the anterior portions of the upper thoracic vertebrae, resulting in their gradual
collapse (Figure 7.22). Lordosis, or swayback, is an excessive anterior curvature of the lumbar region and is most commonly associated with obesity or
late pregnancy. The accumulation of body weight in the abdominal region results an anterior shift in the line of gravity that carries the weight of the body.
This causes in an anterior tilt of the pelvis and a pronounced enhancement of the lumbar curve.

Scoliosis is an abnormal, lateral curvature, accompanied by twisting of the vertebral column. Compensatory curves may also develop in other areas of
the vertebral column to help maintain the head positioned over the feet. Scoliosis is the most common vertebral abnormality among girls. The cause is
usually unknown, but it may result from weakness of the back muscles, defects such as differential growth rates in the right and left sides of the vertebral
column, or differences in the length of the lower limbs. When present, scoliosis tends to get worse during adolescent growth spurts. Although most
individuals do not require treatment, a back brace may be recommended for growing children. In extreme cases, surgery may be required.

Excessive vertebral curves can be identified while an individual stands in the anatomical position. Observe the vertebral profile from the side and then
from behind to check for kyphosis or lordosis. Then have the person bend forward. If scoliosis is present, an individual will have difficulty in bending
directly forward, and the right and left sides of the back will not be level with each other in the bent position.

Figure 7.21 Abnormal Curvatures of the Vertebral Column (a) Scoliosis is an abnormal lateral bending of the vertebral column. (b) An excessive
curvature of the upper thoracic vertebral column is called kyphosis. (c) Lordosis is an excessive curvature in the lumbar region of the vertebral column.

Figure 7.22 Osteoporosis Osteoporosis is an age-related disorder that causes the gradual

loss of bone density and strength. When the thoracic vertebrae are affected, there can be a
gradual collapse of the vertebrae. This results in kyphosis, an excessive curvature of the
thoracic region.

INTERACTIVE LINK

Osteoporosis is a common age-related bone disease in which bone density and strength is
decreased. Watch this video to get a better understanding of how thoracic vertebrae may
become weakened and may fracture due to this disease. How may vertebral osteoporosis
contribute to kyphosis?

General Structure of a Vertebra

Within the different regions of the vertebral column, vertebrae vary in size and shape, but
they all follow a similar structural pattern. A typical vertebra will consist of a body, a vertebral arch, and seven processes (Figure 7.23).

The body is the anterior portion of each vertebra and is the part that supports the body weight. Because of this, the vertebral bodies progressively
increase in size and thickness going down the vertebral column. The bodies of adjacent vertebrae are separated and strongly united by an intervertebral
disc.

The vertebral arch forms the posterior portion of each vertebra. It consists of four parts, the right and left pedicles and the right and left laminae.
Each pedicle forms one of the lateral sides of the vertebral arch. The pedicles are anchored to the posterior side of the vertebral body.
Each lamina forms part of the posterior roof of the vertebral arch. The large opening between the vertebral arch and body is the vertebral foramen,
which contains the spinal cord. In the intact vertebral column, the vertebral foramina of all of the vertebrae align to form the vertebral (spinal) canal,
which serves as the bony protection and passageway for the spinal cord down the back. When the vertebrae are aligned together in the vertebral
column, notches in the margins of the pedicles of adjacent vertebrae together form an intervertebral foramen, the opening through which a spinal
nerve exits from the vertebral column (Figure 7.24).

Seven processes arise from the vertebral arch. Each paired transverse process projects laterally and arises from the junction point between the
pedicle and lamina. The single spinous process (vertebral spine) projects posteriorly at the midline of the back. The vertebral spines can easily be felt
as a series of bumps just under the skin down the middle of the back. The transverse and spinous processes serve as important muscle attachment
sites. A superior articular process extends or faces upward, and an inferior articular process faces or projects downward on each side of a
vertebrae. The paired superior articular processes of one vertebra join with the corresponding paired inferior articular processes from the next higher
vertebra. These junctions form slightly moveable joints between the adjacent vertebrae. The shape and orientation of the articular processes vary in
different regions of the vertebral column and play a major role in determining the type and range of motion available in each region.
Figure 7.23 Parts of a Typical Vertebra A typical vertebra consists of a body and a vertebral arch. The arch is formed by the paired pedicles and
paired laminae. Arising from the vertebral arch are the transverse, spinous, superior articular, and inferior articular processes. The vertebral foramen
provides for passage of the spinal cord. Each spinal nerve exits through an intervertebral foramen, located between adjacent vertebrae. Intervertebral
discs unite the bodies of adjacent vertebrae.
Figure 7.24 Intervertebral Disc The bodies of adjacent vertebrae are separated and united by an intervertebral disc, which provides padding and
allows for movements between adjacent vertebrae. The disc consists of a fibrous outer layer called the anulus fibrosus and a gel-like center called the
nucleus pulposus. The intervertebral foramen is the opening formed between adjacent vertebrae for the exit of a spinal nerve.

Regional Modifications of Vertebrae

In addition to the general characteristics of a typical vertebra described above, vertebrae also display characteristic size and structural features that vary
between the different vertebral column regions. Thus, cervical vertebrae are smaller than lumbar vertebrae due to differences in the proportion of body
weight that each supports. Thoracic vertebrae have sites for rib attachment, and the vertebrae that give rise to the sacrum and coccyx have fused
together into single bones.
Cervical Vertebrae

Typical cervical vertebrae, such as C4 or C5, have several characteristic features that differentiate them from thoracic or lumbar vertebrae (Figure
7.25). Cervical vertebrae have a small body, reflecting the fact that they carry the least amount of body weight. Cervical vertebrae usually have a bifid
(Y-shaped) spinous process. The spinous processes of the C3–C6 vertebrae are short, but the spine of C7 is much longer. You can find these vertebrae
by running your finger down the midline of the posterior neck until you encounter the prominent C7 spine located at the base of the neck. The transverse
processes of the cervical vertebrae are sharply curved (U-shaped) to allow for passage of the cervical spinal nerves. Each transverse process also has
an opening called the transverse foramen. An important artery that supplies the brain ascends up the neck by passing through these openings. The
superior and inferior articular processes of the cervical vertebrae are flattened and largely face upward or downward, respectively.

The first and second cervical vertebrae are further modified, giving each a distinctive appearance. The first cervical (C1) vertebra is also called the atlas,
because this is the vertebra that supports the skull on top of the vertebral column (in Greek mythology, Atlas was the god who supported the heavens on
his shoulders). The C1 vertebra does not have a body or spinous process. Instead, it is ring-shaped, consisting of an anterior arch and a posterior
arch. The transverse processes of the atlas are longer and extend more laterally than do the transverse processes of any other cervical vertebrae. The
superior articular processes face upward and are deeply curved for articulation with the occipital condyles on the base of the skull. The inferior articular
processes are flat and face downward to join with the superior articular processes of the C2 vertebra.

The second cervical (C2) vertebra is called the axis, because it serves as the axis for rotation when turning the head toward the right or left. The axis
resembles typical cervical vertebrae in most respects, but is easily distinguished by the dens (odontoid process), a bony projection that extends upward
from the vertebral body. The dens joins with the inner aspect of the anterior arch of the atlas, where it is held in place by transverse ligament.
 Figure 7.25 Cervical Vertebrae A typical cervical vertebra has a small
body, a bifid spinous process, transverse processes that have a transverse
foramen and are curved for spinal nerve passage. The atlas (C1 vertebra)
does not have a body or spinous process. It consists of an anterior and a
posterior arch and elongated transverse processes. The axis (C2 vertebra)
has the upward projecting dens, which
articulates with the anterior arch of the atlas.
Thoracic Vertebrae

The bodies of the thoracic vertebrae are larger

than those of cervical vertebrae (Figure 7.26).
The characteristic feature for a typical
midthoracic vertebra is the spinous process,
which is long and has a pronounced downward
angle that causes it to overlap the next inferior
vertebra. The superior articular processes of
thoracic vertebrae face anteriorly and the inferior
processes face posteriorly. These orientations
are important determinants for the type and
range of movements available to the thoracic
region of the vertebral column.

Thoracic vertebrae have several additional

articulation sites, each of which is called a facet,
where a rib is attached. Most thoracic vertebrae
have two facets located on the lateral sides of
the body, each of which is called a costal
facet (costal = “rib”). These are for articulation
with the head (end) of a rib. An additional facet
is located on the transverse process for
articulation with the tubercle of a rib.

Figure 7.26 Thoracic Vertebrae A typical

thoracic vertebra is distinguished by the spinous
process, which is long and projects downward to
overlap the next inferior vertebra. It also has
articulation sites (facets) on the vertebral body
and a transverse process for rib attachment.

Figure 7.27 Rib Articulation in Thoracic

Vertebrae Thoracic vertebrae have superior and
inferior articular facets on the vertebral body for
articulation with the head of a rib, and a
transverse process facet for articulation with the
rib tubercle.
Lumbar Vertebrae

Lumbar vertebrae carry the greatest amount of body weight and are thus characterized by the large size and thickness of the vertebral body (Figure
7.28). They have short transverse processes and a short, blunt spinous process that projects posteriorly. The articular processes are large, with the
superior process facing backward and the inferior facing forward.

Figure 7.28 Lumbar Vertebrae Lumbar vertebrae are characterized by having a large, thick body and a short, rounded spinous process.
Sacrum and Coccyx

The sacrum is a triangular-shaped bone that is thick and wide across its superior base where it is weight bearing and then tapers down to an inferior,
non-weight bearing apex (Figure 7.29). It is formed by the fusion of five sacral vertebrae, a process that does not begin until after the age of 20. On the
anterior surface of the older adult sacrum, the lines of vertebral fusion can be seen as four transverse ridges. On the posterior surface, running down the
midline, is the median sacral crest, a bumpy ridge that is the remnant of the fused spinous processes (median = “midline”; while medial = “toward, but
not necessarily at, the midline”). Similarly, the fused transverse processes of the sacral vertebrae form the lateral sacral crest.

The sacral promontory is the anterior lip of the superior base of the sacrum. Lateral to this is the roughened auricular surface, which joins with the ilium
portion of the hipbone to form the immobile sacroiliac joints of the pelvis. Passing inferiorly through the sacrum is a bony tunnel called the sacral canal,
which terminates at the sacral hiatus near the inferior tip of the sacrum. The anterior and posterior surfaces of the sacrum have a series of paired
openings called sacral foramina (singular = foramen) that connect to the sacral canal. Each of these openings is called a posterior (dorsal) sacral
foramen or anterior (ventral) sacral foramen. These openings allow for the anterior and posterior branches of the sacral spinal nerves to exit the
sacrum. The superior articular process of the sacrum, one of which is found on either side of the superior opening of the sacral canal, articulates with
the inferior articular processes from the L5 vertebra.
The coccyx, or tailbone, is derived from the fusion of four very small coccygeal vertebrae (see Figure 7.29). It articulates with the inferior tip of the
sacrum. It is not weight bearing in the standing position, but may receive some body weight when sitting.

Figure 7.29 Sacrum and Coccyx The sacrum is formed from the fusion of five sacral vertebrae, whose lines of fusion are indicated by the transverse
ridges. The fused spinous processes form the median sacral crest, while the lateral sacral crest arises from the fused transverse processes. The coccyx
is formed by the fusion of four small coccygeal vertebrae.

Intervertebral Discs and Ligaments of the Vertebral Column

The bodies of adjacent vertebrae are strongly anchored to each other by an intervertebral disc. This structure provides padding between the bones
during weight bearing, and because it can change shape, also allows for movement between the vertebrae. Although the total amount of movement
available between any two adjacent vertebrae is small, when these movements are summed together along the entire length of the vertebral column,
large body movements can be produced. Ligaments that extend along the length of the vertebral column also contribute to its overall support and
stability.
Intervertebral Disc

An intervertebral disc is a fibrocartilaginous pad that fills the gap between adjacent vertebral bodies (see Figure 7.24). Each disc is anchored to the
bodies of its adjacent vertebrae, thus strongly uniting these. The discs also provide padding between vertebrae during weight bearing. Because of this,
intervertebral discs are thin in the cervical region and thickest in the lumbar region, which carries the most body weight. In total, the intervertebral discs
account for approximately 25 percent of your body height between the top of the pelvis and the base of the skull. Intervertebral discs are also flexible
and can change shape to allow for movements of the vertebral column.

Each intervertebral disc consists of two parts. The anulus fibrosus is the tough, fibrous outer layer of the disc. It forms a circle (anulus = “ring” or
“circle”) and is firmly anchored to the outer margins of the adjacent vertebral bodies. Inside is the nucleus pulposus, consisting of a softer, more gel-like
material. It has a high water content that serves to resist compression and thus is important for weight bearing. With increasing age, the water content of
the nucleus pulposus gradually declines. This causes the disc to become thinner, decreasing total body height somewhat, and reduces the flexibility and
range of motion of the disc, making bending more difficult.

The gel-like nature of the nucleus pulposus also allows the intervertebral disc to change shape as one vertebra rocks side to side or forward and back in
relation to its neighbors during movements of the vertebral column. Thus, bending forward causes compression of the anterior portion of the disc but
expansion of the posterior disc. If the posterior anulus fibrosus is weakened due to injury or increasing age, the pressure exerted on the disc when
bending forward and lifting a heavy object can cause the nucleus pulposus to protrude posteriorly through the anulus fibrosus, resulting in a herniated
disc (“ruptured” or “slipped” disc) (Figure 7.30). The posterior bulging of the nucleus pulposus can cause compression of a spinal nerve at the point
where it exits through the intervertebral foramen, with resulting pain and/or muscle weakness in those body regions supplied by that nerve. The most
common sites for disc herniation are the L4/L5 or L5/S1 intervertebral discs, which can cause sciatica, a widespread pain that radiates from the lower
back down the thigh and into the leg. Similar injuries of the C5/C6 or C6/C7 intervertebral discs, following forcible hyperflexion of the neck from a
collision accident or football injury, can produce pain in the neck, shoulder, and upper limb.

Figure 7.30 Herniated Intervertebral Disc Weakening of the anulus fibrosus can result in herniation (protrusion) of the nucleus pulposus and
compression of a spinal nerve, resulting in pain and/or muscle weakness in the body regions supplied by that nerve.

Adjacent vertebrae are united by ligaments that run the length of the vertebral column along both its posterior and anterior aspects (Figure 7.31). These
serve to resist excess forward or backward bending movements of the vertebral column, respectively.

The anterior longitudinal ligament runs down the anterior side of the entire vertebral column, uniting the vertebral bodies. It serves to resist excess
backward bending of the vertebral column. Protection against this movement is particularly important in the neck, where extreme posterior bending of
the head and neck can stretch or tear this ligament, resulting in a painful whiplash injury. Prior to the mandatory installation of seat headrests, whiplash
injuries were common for passengers involved in a rear-end automobile collision.

The supraspinous ligament is located on the posterior side of the vertebral column, where it interconnects the spinous processes of the thoracic and
lumbar vertebrae. This strong ligament supports the vertebral column during forward bending motions. In the posterior neck, where the cervical spinous
processes are short, the supraspinous ligament expands to become the nuchal ligament (nuchae = “nape” or “back of the neck”). The nuchal ligament
is attached to the cervical spinous processes and extends upward and posteriorly to attach to the midline base of the skull, out to the external occipital
protuberance. It supports the skull and prevents it from falling forward. This ligament is much larger and stronger in four-legged animals such as cows,
where the large skull hangs off the front end of the vertebral column. You can easily feel this ligament by first extending your head backward and
pressing down on the posterior midline of your neck. Then tilt your head forward and you will fill the nuchal ligament popping out as it tightens to limit
anterior bending of the head and neck.

Additional ligaments are located inside the vertebral canal, next to the spinal cord, along the length of the vertebral column. The posterior longitudinal
ligament is found anterior to the spinal cord, where it is attached to the posterior sides of the vertebral bodies. Posterior to the spinal cord is
the ligamentum flavum (“yellow ligament”). This consists of a series of short, paired ligaments, each of which interconnects the lamina regions of
adjacent vertebrae. The ligamentum flavum has large numbers of elastic fibers, which have a yellowish color, allowing it to stretch and then pull back.
Both of these ligaments provide important support for the vertebral column when bending forward.

Figure 7.31 Ligaments of Vertebral Column The anterior longitudinal ligament runs the length of the vertebral column, uniting the anterior sides of the
vertebral bodies. The supraspinous ligament connects the spinous processes of the thoracic and lumbar vertebrae. In the posterior neck, the
supraspinous ligament enlarges to form the nuchal ligament, which attaches to the cervical spinous processes and to the base of the skull.

CAREER CONNECTION
Chiropractor

Chiropractors are health professionals who use nonsurgical techniques to help patients with musculoskeletal system problems that involve the bones,
muscles, ligaments, tendons, or nervous system. They treat problems such as neck pain, back pain, joint pain, or headaches. Chiropractors focus on the
patient’s overall health and can also provide counseling related to lifestyle issues, such as diet, exercise, or sleep problems. If needed, they will refer the
patient to other medical specialists.

Chiropractors use a drug-free, hands-on approach for patient diagnosis and treatment. They will perform a physical exam, assess the patient’s posture
and spine, and may perform additional diagnostic tests, including taking X-ray images. They primarily use manual techniques, such as spinal
manipulation, to adjust the patient’s spine or other joints. They can recommend therapeutic or rehabilitative exercises, and some also include
acupuncture, massage therapy, or ultrasound as part of the treatment program. In addition to those in general practice, some chiropractors specialize in
sport injuries, neurology, orthopaedics, pediatrics, nutrition, internal disorders, or diagnostic imaging.

To become a chiropractor, students must have 3–4 years of undergraduate education, attend an accredited, four-year Doctor of Chiropractic (D.C.)
degree program, and pass a licensure examination to be licensed for practice in their state. With the aging of the baby-boom generation, employment for
chiropractors is expected to increase.

The thoracic cage (rib cage) forms the thorax (chest) portion of the body. It consists of the 12 pairs of ribs with their costal cartilages and the sternum
(Figure 7.32). The ribs are anchored posteriorly to the 12 thoracic vertebrae (T1–T12). The thoracic cage protects the heart and lungs.

Figure 7.32 Thoracic Cage The thoracic cage

is formed by the (a) sternum and (b) 12 pairs of
ribs with their costal cartilages. The ribs are
anchored posteriorly to the 12 thoracic
vertebrae. The sternum consists of the
manubrium, body, and xiphoid process. The ribs
are classified as true ribs (1–7) and false ribs
(8–12). The last two pairs of false ribs are also
known as floating ribs (11–12).

Sternum

The sternum is the elongated bony structure that

anchors the anterior thoracic cage. It consists of
three parts: the manubrium, body, and xiphoid
process. The manubrium is the wider, superior
portion of the sternum. The top of the
manubrium has a shallow, U-shaped border
called the jugular (suprasternal) notch. This
can be easily felt at the anterior base of the
neck, between the medial ends of the clavicles. The clavicular notch is the shallow depression located on either side at the superior-lateral margins of
the manubrium. This is the site of the sternoclavicular joint, between the sternum and clavicle. The first ribs also attach to the manubrium.

The elongated, central portion of the sternum is the body. The manubrium and body join together at the sternal angle, so called because the junction
between these two components is not flat, but forms a slight bend. The second rib attaches to the sternum at the sternal angle. Since the first rib is
hidden behind the clavicle, the second rib is the highest rib that can be identified by palpation. Thus, the sternal angle and second rib are important
landmarks for the identification and counting of the lower ribs. Ribs 3–7 attach to the sternal body.

The inferior tip of the sternum is the xiphoid process. This small structure is cartilaginous early in life, but gradually becomes ossified starting during
middle age.
Ribs

Each rib is a curved, flattened bone that contributes to the wall of the thorax. The ribs articulate posteriorly with the T1–T12 thoracic vertebrae, and most
attach anteriorly via their costal cartilages to the sternum. There are 12 pairs of ribs. The ribs are numbered 1–12 in accordance with the thoracic
vertebrae.
Parts of a Typical Rib

The posterior end of a typical rib is called the head of the rib (see Figure 7.27). This region articulates primarily with the costal facet located on the body
of the same numbered thoracic vertebra and to a lesser degree, with the costal facet located on the body of the next higher vertebra. Lateral to the head
is the narrowed neck of the rib. A small bump on the posterior rib surface is the tubercle of the rib, which articulates with the facet located on the
transverse process of the same numbered vertebra. The remainder of the rib is the body of the rib (shaft). Just lateral to the tubercle is the angle of the
rib, the point at which the rib has its greatest degree of curvature. The angles of the ribs form the most posterior extent of the thoracic cage. In the
anatomical position, the angles align with the medial border of the scapula. A shallow costal groove for the passage of blood vessels and a nerve is
found along the inferior margin of each rib.
Rib Classifications

The bony ribs do not extend anteriorly completely around to the sternum. Instead, each rib ends in a costal cartilage. These cartilages are made of
hyaline cartilage and can extend for several inches. Most ribs are then attached, either directly or indirectly, to the sternum via their costal cartilage
(see Figure 7.32). The ribs are classified into three groups based on their relationship to the sternum.

Ribs 1–7 are classified as true ribs (vertebrosternal ribs). The costal cartilage from each of these ribs attaches directly to the sternum. Ribs 8–12 are
called false ribs (vertebrochondral ribs). The costal cartilages from these ribs do not attach directly to the sternum. For ribs 8–10, the costal cartilages
are attached to the cartilage of the next higher rib. Thus, the cartilage of rib 10 attaches to the cartilage of rib 9, rib 9 then attaches to rib 8, and rib 8 is
attached to rib 7. The last two false ribs (11–12) are also called floating ribs (vertebral ribs). These are short ribs that do not attach to the sternum at all.
Instead, their small costal cartilages terminate within the musculature of the lateral abdominal wall.

The axial skeleton begins to form during early embryonic development. However, growth, remodeling, and ossification (bone formation) continue for
several decades after birth before the adult skeleton is fully formed. Knowledge of the developmental processes that give rise to the skeleton is
important for understanding the abnormalities that may arise in skeletal structures.

Development of the Skull

During the third week of embryonic development, a rod-like structure called the notochord develops dorsally along the length of the embryo. The tissue
overlying the notochord enlarges and forms the neural tube, which will give rise to the brain and spinal cord. By the fourth week, mesoderm tissue
located on either side of the notochord thickens and separates into a repeating series of block-like tissue structures, each of which is called a somite. As
the somites enlarge, each one will split into several parts. The most medial of these parts is called a sclerotome. The sclerotomes consist of an
embryonic tissue called mesenchyme, which will give rise to the fibrous connective tissues, cartilages, and bones of the body.

The bones of the skull arise from mesenchyme during embryonic development in two different ways. The first mechanism produces the bones that form
the top and sides of the brain case. This involves the local accumulation of mesenchymal cells at the site of the future bone. These cells then
differentiate directly into bone producing cells, which form the skull bones through the process of intramembranous ossification. As the brain case bones
grow in the fetal skull, they remain separated from each other by large areas of dense connective tissue, each of which is called a fontanelle (Figure
7.33). The fontanelles are the soft spots on an infant’s head. They are important during birth because these areas allow the skull to change shape as it
squeezes through the birth canal. After birth, the fontanelles allow for continued growth and expansion of the skull as the brain enlarges. The largest
fontanelle is located on the anterior head, at the junction of the frontal and parietal bones. The fontanelles decrease in size and disappear by age 2.
However, the skull bones remained separated from each other at the sutures, which contain dense fibrous connective tissue that unites the adjacent
bones. The connective tissue of the sutures allows for continued growth of the skull bones as the brain enlarges during childhood growth.

The second mechanism for bone development in the skull produces the facial bones and floor of the brain case. This also begins with the localized
accumulation of mesenchymal cells. However, these cells differentiate into cartilage cells, which produce a hyaline cartilage model of the future bone. As
this cartilage model grows, it is gradually converted into bone through the process of endochondral ossification. This is a slow process and the cartilage
is not completely converted to bone until the skull achieves its full adult size.

At birth, the brain case and orbits of the skull are disproportionally large compared to the bones of the jaws and lower face. This reflects the relative
underdevelopment of the maxilla and mandible, which lack teeth, and the small sizes of the paranasal sinuses and nasal cavity. During early childhood,
the mastoid process enlarges, the two halves of the mandible and frontal bone fuse together to form single bones, and the paranasal sinuses enlarge.
The jaws also expand as the teeth begin to appear. These changes all contribute to the rapid growth and enlargement of the face during childhood.
Figure 7.33 Newborn Skull The bones of the newborn skull are not fully ossified and are separated by large areas called fontanelles, which are filled
with fibrous connective tissue. The fontanelles allow for continued growth of the skull after birth. At the time of birth, the facial bones are small and
underdeveloped, and the mastoid process has not yet formed.

Development of the Vertebral Column and Thoracic cage

Development of the vertebrae begins with the accumulation of mesenchyme cells from each sclerotome around the notochord. These cells differentiate
into a hyaline cartilage model for each vertebra, which then grow and eventually ossify into bone through the process of endochondral ossification. As
the developing vertebrae grow, the notochord largely disappears. However, small areas of notochord tissue persist between the adjacent vertebrae and
this contributes to the formation of each intervertebral disc.

The ribs and sternum also develop from mesenchyme. The ribs initially develop as part of the cartilage model for each vertebra, but in the thorax region,
the rib portion separates from the vertebra by the eighth week. The cartilage model of the rib then ossifies, except for the anterior portion, which remains
as the costal cartilage. The sternum initially forms as paired hyaline cartilage models on either side of the anterior midline, beginning during the fifth
week of development. The cartilage models of the ribs become attached to the lateral sides of the developing sternum. Eventually, the two halves of the
cartilaginous sternum fuse together along the midline and then ossify into bone. The manubrium and body of the sternum are converted into bone first,
with the xiphoid process remaining as cartilage until late in life.

INTERACTIVE LINK

View this video to review the two processes that give rise to the bones of the skull and body. What are the two mechanisms by which the bones of the
body are formed and which bones are formed by each mechanism?

HOMEOSTATIC IMBALANCES
Craniosynostosis

The premature closure (fusion) of a suture line is a condition called craniosynostosis. This error in the normal developmental process results in abnormal
growth of the skull and deformity of the head. It is produced either by defects in the ossification process of the skull bones or failure of the brain to
properly enlarge. Genetic factors are involved, but the underlying cause is unknown. It is a relatively common condition, occurring in approximately
1:2000 births, with males being more commonly affected. Primary craniosynostosis involves the early fusion of one cranial suture, whereas complex
craniosynostosis results from the premature fusion of several sutures.

The early fusion of a suture in primary craniosynostosis prevents any additional enlargement of the cranial bones and skull along this line. Continued
growth of the brain and skull is therefore diverted to other areas of the head, causing an abnormal enlargement of these regions. For example, the early
disappearance of the anterior fontanelle and premature closure of the sagittal suture prevents growth across the top of the head. This is compensated
by upward growth by the bones of the lateral skull, resulting in a long, narrow, wedge-shaped head. This condition, known as scaphocephaly, accounts
for approximately 50 percent of craniosynostosis abnormalities. Although the skull is misshapen, the brain still has adequate room to grow and thus
there is no accompanying abnormal neurological development.

In cases of complex craniosynostosis, several sutures close prematurely. The amount and degree of skull deformity is determined by the location and
extent of the sutures involved. This results in more severe constraints on skull growth, which can alter or impede proper brain growth and development.

Cases of craniosynostosis are usually treated with surgery. A team of physicians will open the skull along the fused suture, which will then allow the skull
bones to resume their growth in this area. In some cases, parts of the skull will be removed and replaced with an artificial plate. The earlier after birth
that surgery is performed, the better the outcome. After treatment, most children continue to grow and develop normally and do not exhibit any
neurological problems.
alveolar process of the mandible
upper border of mandibular body that contains the lower teeth
alveolar process of the maxilla
curved, inferior margin of the maxilla that supports and anchors the upper teeth
angle of the mandible
rounded corner located at outside margin of the body and ramus junction
angle of the rib
portion of rib with greatest curvature; together, the rib angles form the most posterior extent of the thoracic cage
anterior (ventral) sacral foramen
 one of the series of paired openings located on the anterior (ventral) side of the sacrum
anterior arch
anterior portion of the ring-like C1 (atlas) vertebra
anterior cranial fossa
shallowest and most anterior cranial fossa of the cranial base that extends from the frontal bone to the lesser wing of the sphenoid bone
anterior longitudinal ligament
ligament that runs the length of the vertebral column, uniting the anterior aspects of the vertebral bodies
anulus fibrosus
tough, fibrous outer portion of an intervertebral disc, which is strongly anchored to the bodies of the adjacent vertebrae
appendicular skeleton
all bones of the upper and lower limbs, plus the girdle bones that attach each limb to the axial skeleton
articular tubercle
smooth ridge located on the inferior skull, immediately anterior to the mandibular fossa
atlas
first cervical (C1) vertebra
axial skeleton
central, vertical axis of the body, including the skull, vertebral column, and thoracic cage
axis
second cervical (C2) vertebra
body of the rib
shaft portion of a rib
brain case
portion of the skull that contains and protects the brain, consisting of the eight bones that form the cranial base and rounded upper skull
calvaria
(also, skullcap) rounded top of the skull
carotid canal
zig-zag tunnel providing passage through the base of the skull for the internal carotid artery to the brain; begins anteromedial to the styloid
process and terminates in the middle cranial cavity, near the posterior-lateral base of the sella turcica
cervical curve
posteriorly concave curvature of the cervical vertebral column region; a secondary curve of the vertebral column
cervical vertebrae
seven vertebrae numbered as C1–C7 that are located in the neck region of the vertebral column
clavicular notch
paired notches located on the superior-lateral sides of the sternal manubrium, for articulation with the clavicle
coccyx
small bone located at inferior end of the adult vertebral column that is formed by the fusion of four coccygeal vertebrae; also referred to as the
“tailbone”
condylar process of the mandible
thickened upward projection from posterior margin of mandibular ramus
condyle
oval-shaped process located at the top of the condylar process of the mandible
coronal suture
joint that unites the frontal bone to the right and left parietal bones across the top of the skull
coronoid process of the mandible
flattened upward projection from the anterior margin of the mandibular ramus
costal cartilage
hyaline cartilage structure attached to the anterior end of each rib that provides for either direct or indirect attachment of most ribs to the
sternum
costal facet
site on the lateral sides of a thoracic vertebra for articulation with the head of a rib
costal groove
shallow groove along the inferior margin of a rib that provides passage for blood vessels and a nerve
cranial cavity
interior space of the skull that houses the brain
cranium
skull
cribriform plate
small, flattened areas with numerous small openings, located to either side of the midline in the floor of the anterior cranial fossa; formed by
the ethmoid bone
crista galli
small upward projection located at the midline in the floor of the anterior cranial fossa; formed by the ethmoid bone
dens
bony projection (odontoid process) that extends upward from the body of the C2 (axis) vertebra
ear ossicles
three small bones located in the middle ear cavity that serve to transmit sound vibrations to the inner ear
ethmoid air cell
one of several small, air-filled spaces located within the lateral sides of the ethmoid bone, between the orbit and upper nasal cavity
ethmoid bone
unpaired bone that forms the roof and upper, lateral walls of the nasal cavity, portions of the floor of the anterior cranial fossa and medial wall
of orbit, and the upper portion of the nasal septum
external acoustic meatus
ear canal opening located on the lateral side of the skull
external occipital protuberance
small bump located at the midline on the posterior skull
facet
small, flattened area on a bone for an articulation (joint) with another bone, or for muscle attachment
facial bones
fourteen bones that support the facial structures and form the upper and lower jaws and the hard palate
false ribs
vertebrochondral ribs 8–12 whose costal cartilage either attaches indirectly to the sternum via the costal cartilage of the next higher rib or does
not attach to the sternum at all
floating ribs
vertebral ribs 11–12 that do not attach to the sternum or to the costal cartilage of another rib
fontanelle
expanded area of fibrous connective tissue that separates the brain case bones of the skull prior to birth and during the first year after birth
foramen lacerum
irregular opening in the base of the skull, located inferior to the exit of carotid canal
foramen magnum
large opening in the occipital bone of the skull through which the spinal cord emerges and the vertebral arteries enter the cranium
foramen ovale of the middle cranial fossa
oval-shaped opening in the floor of the middle cranial fossa
foramen rotundum
round opening in the floor of the middle cranial fossa, located between the superior orbital fissure and foramen ovale
foramen spinosum
small opening in the floor of the middle cranial fossa, located lateral to the foramen ovale
frontal bone
unpaired bone that forms forehead, roof of orbit, and floor of anterior cranial fossa
frontal sinus
air-filled space within the frontal bone; most anterior of the paranasal sinuses
glabella
slight depression of frontal bone, located at the midline between the eyebrows
greater wings of sphenoid bone
lateral projections of the sphenoid bone that form the anterior wall of the middle cranial fossa and an area of the lateral skull
hard palate
bony structure that forms the roof of the mouth and floor of the nasal cavity, formed by the palatine process of the maxillary bones and the
horizontal plate of the palatine bones
head of the rib
posterior end of a rib that articulates with the bodies of thoracic vertebrae
horizontal plate
medial extension from the palatine bone that forms the posterior quarter of the hard palate
hyoid bone
small, U-shaped bone located in upper neck that does not contact any other bone
hypoglossal canal
paired openings that pass anteriorly from the anterior-lateral margins of the foramen magnum deep to the occipital condyles
hypophyseal (pituitary) fossa
shallow depression on top of the sella turcica that houses the pituitary (hypophyseal) gland
inferior articular process
bony process that extends downward from the vertebral arch of a vertebra that articulates with the superior articular process of the next lower
vertebra
inferior nasal concha
one of the paired bones that project from the lateral walls of the nasal cavity to form the largest and most inferior of the nasal conchae
infraorbital foramen
opening located on anterior skull, below the orbit
infratemporal fossa
space on lateral side of skull, below the level of the zygomatic arch and deep (medial) to the ramus of the mandible
internal acoustic meatus
opening into petrous ridge, located on the lateral wall of the posterior cranial fossa
intervertebral disc
structure located between the bodies of adjacent vertebrae that strongly joins the vertebrae; provides padding, weight bearing ability, and
enables vertebral column movements
intervertebral foramen
opening located between adjacent vertebrae for exit of a spinal nerve
jugular (suprasternal) notch
shallow notch located on superior surface of sternal manubrium
jugular foramen
irregularly shaped opening located in the lateral floor of the posterior cranial cavity
kyphosis
(also, humpback or hunchback) excessive posterior curvature of the thoracic vertebral column region
lacrimal bone
paired bones that contribute to the anterior-medial wall of each orbit
lacrimal fossa
shallow depression in the anterior-medial wall of the orbit, formed by the lacrimal bone that gives rise to the nasolacrimal canal
lambdoid suture
inverted V-shaped joint that unites the occipital bone to the right and left parietal bones on the posterior skull
lamina
portion of the vertebral arch on each vertebra that extends between the transverse and spinous process
lateral pterygoid plate
paired, flattened bony projections of the sphenoid bone located on the inferior skull, lateral to the medial pterygoid plate
lateral sacral crest
paired irregular ridges running down the lateral sides of the posterior sacrum that was formed by the fusion of the transverse processes from
the five sacral vertebrae
lesser wings of the sphenoid bone
lateral extensions of the sphenoid bone that form the bony lip separating the anterior and middle cranial fossae
ligamentum flavum
series of short ligaments that unite the lamina of adjacent vertebrae
lingula
 small flap of bone located on the inner (medial) surface of mandibular ramus, next to the mandibular foramen
lordosis
(also, swayback) excessive anterior curvature of the lumbar vertebral column region
lumbar curve
posteriorly concave curvature of the lumbar vertebral column region; a secondary curve of the vertebral column
lumbar vertebrae
five vertebrae numbered as L1–L5 that are located in lumbar region (lower back) of the vertebral column
mandible
unpaired bone that forms the lower jaw bone; the only moveable bone of the skull
mandibular foramen
opening located on the inner (medial) surface of the mandibular ramus
mandibular fossa
oval depression located on the inferior surface of the skull
mandibular notch
large U-shaped notch located between the condylar process and coronoid process of the mandible
manubrium
expanded, superior portion of the sternum
mastoid process
large bony prominence on the inferior, lateral skull, just behind the earlobe
maxillary bone
(also, maxilla) paired bones that form the upper jaw and anterior portion of the hard palate
maxillary sinus
air-filled space located with each maxillary bone; largest of the paranasal sinuses
medial pterygoid plate
paired, flattened bony projections of the sphenoid bone located on the inferior skull medial to the lateral pterygoid plate; form the posterior
portion of the nasal cavity lateral wall
median sacral crest
irregular ridge running down the midline of the posterior sacrum that was formed from the fusion of the spinous processes of the five sacral
vertebrae
mental foramen
opening located on the anterior-lateral side of the mandibular body
mental protuberance
inferior margin of anterior mandible that forms the chin
middle cranial fossa
centrally located cranial fossa that extends from the lesser wings of the sphenoid bone to the petrous ridge
middle nasal concha
nasal concha formed by the ethmoid bone that is located between the superior and inferior conchae
mylohyoid line
bony ridge located along the inner (medial) surface of the mandibular body
nasal bone
paired bones that form the base of the nose
nasal cavity
opening through skull for passage of air
nasal conchae
curved bony plates that project from the lateral walls of the nasal cavity; include the superior and middle nasal conchae, which are parts of the
ethmoid bone, and the independent inferior nasal conchae bone
nasal septum
flat, midline structure that divides the nasal cavity into halves, formed by the perpendicular plate of the ethmoid bone, vomer bone, and septal
cartilage
nasolacrimal canal
passage for drainage of tears that extends downward from the medial-anterior orbit to the nasal cavity, terminating behind the inferior nasal
conchae
neck of the rib
narrowed region of a rib, next to the rib head
notochord
rod-like structure along dorsal side of the early embryo; largely disappears during later development but does contribute to formation of the
intervertebral discs
nuchal ligament
expanded portion of the supraspinous ligament within the posterior neck; interconnects the spinous processes of the cervical vertebrae and
attaches to the base of the skull
nucleus pulposus
gel-like central region of an intervertebral disc; provides for padding, weight-bearing, and movement between adjacent vertebrae
occipital bone
unpaired bone that forms the posterior portions of the brain case and base of the skull
occipital condyle
paired, oval-shaped bony knobs located on the inferior skull, to either side of the foramen magnum
optic canal
opening spanning between middle cranial fossa and posterior orbit
orbit
bony socket that contains the eyeball and associated muscles
palatine bone
paired bones that form the posterior quarter of the hard palate and a small area in floor of the orbit
palatine process
medial projection from the maxilla bone that forms the anterior three quarters of the hard palate
paranasal sinuses
cavities within the skull that are connected to the conchae that serve to warm and humidify incoming air, produce mucus, and lighten the
weight of the skull; consist of frontal, maxillary, sphenoidal, and ethmoidal sinuses
parietal bone
 paired bones that form the upper, lateral sides of the skull
pedicle
portion of the vertebral arch that extends from the vertebral body to the transverse process
perpendicular plate of the ethmoid bone
downward, midline extension of the ethmoid bone that forms the superior portion of the nasal septum
petrous ridge
petrous portion of the temporal bone that forms a large, triangular ridge in the floor of the cranial cavity, separating the middle and posterior
cranial fossae; houses the middle and inner ear structures
posterior (dorsal) sacral foramen
one of the series of paired openings located on the posterior (dorsal) side of the sacrum
posterior arch
posterior portion of the ring-like C1 (atlas) vertebra
posterior cranial fossa
deepest and most posterior cranial fossa; extends from the petrous ridge to the occipital bone
posterior longitudinal ligament
ligament that runs the length of the vertebral column, uniting the posterior sides of the vertebral bodies
primary curve
anteriorly concave curvatures of the thoracic and sacrococcygeal regions that are retained from the original fetal curvature of the vertebral
column
pterion
H-shaped suture junction region that unites the frontal, parietal, temporal, and sphenoid bones on the lateral side of the skull
ramus of the mandible
vertical portion of the mandible
ribs
thin, curved bones of the chest wall
sacral canal
bony tunnel that runs through the sacrum
sacral foramina
series of paired openings for nerve exit located on both the anterior (ventral) and posterior (dorsal) aspects of the sacrum
sacral hiatus
inferior opening and termination of the sacral canal
sacral promontory
anterior lip of the base (superior end) of the sacrum
sacrococcygeal curve
anteriorly concave curvature formed by the sacrum and coccyx; a primary curve of the vertebral column
sacrum
single bone located near the inferior end of the adult vertebral column that is formed by the fusion of five sacral vertebrae; forms the posterior
portion of the pelvis
sagittal suture
joint that unites the right and left parietal bones at the midline along the top of the skull
sclerotome
medial portion of a somite consisting of mesenchyme tissue that will give rise to bone, cartilage, and fibrous connective tissues
scoliosis
abnormal lateral curvature of the vertebral column
secondary curve
posteriorly concave curvatures of the cervical and lumbar regions of the vertebral column that develop after the time of birth
sella turcica
elevated area of sphenoid bone located at midline of the middle cranial fossa
septal cartilage
flat cartilage structure that forms the anterior portion of the nasal septum
skeleton
bones of the body
skull
bony structure that forms the head, face, and jaws, and protects the brain; consists of 22 bones
somite
one of the paired, repeating blocks of tissue located on either side of the notochord in the early embryo
sphenoid bone
unpaired bone that forms the central base of skull
sphenoid sinus
air-filled space located within the sphenoid bone; most posterior of the paranasal sinuses
spinous process
unpaired bony process that extends posteriorly from the vertebral arch of a vertebra
squamous suture
joint that unites the parietal bone to the squamous portion of the temporal bone on the lateral side of the skull
sternal angle
junction line between manubrium and body of the sternum and the site for attachment of the second rib to the sternum
sternum
flattened bone located at the center of the anterior chest
styloid process
downward projecting, elongated bony process located on the inferior aspect of the skull
stylomastoid foramen
opening located on inferior skull, between the styloid process and mastoid process
superior articular process
bony process that extends upward from the vertebral arch of a vertebra that articulates with the inferior articular process of the next higher
vertebra
superior articular process of the sacrum
paired processes that extend upward from the sacrum to articulate (join) with the inferior articular processes from the L5 vertebra
superior nasal concha
 smallest and most superiorly located of the nasal conchae; formed by the ethmoid bone
superior nuchal line
paired bony lines on the posterior skull that extend laterally from the external occipital protuberance
superior orbital fissure
irregularly shaped opening between the middle cranial fossa and the posterior orbit
supraorbital foramen
opening located on anterior skull, at the superior margin of the orbit
supraorbital margin
superior margin of the orbit
supraspinous ligament
ligament that interconnects the spinous processes of the thoracic and lumbar vertebrae
suture
junction line at which adjacent bones of the skull are united by fibrous connective tissue
temporal bone
paired bones that form the lateral, inferior portions of the skull, with squamous, mastoid, and petrous portions
temporal fossa
shallow space on the lateral side of the skull, above the level of the zygomatic arch
temporal process of the zygomatic bone
short extension from the zygomatic bone that forms the anterior portion of the zygomatic arch
thoracic cage
consists of 12 pairs of ribs and sternum
thoracic curve
anteriorly concave curvature of the thoracic vertebral column region; a primary curve of the vertebral column
thoracic vertebrae
twelve vertebrae numbered as T1–T12 that are located in the thoracic region (upper back) of the vertebral column
transverse foramen
opening found only in the transverse processes of cervical vertebrae
transverse process
paired bony processes that extends laterally from the vertebral arch of a vertebra
true ribs
vertebrosternal ribs 1–7 that attach via their costal cartilage directly to the sternum
tubercle of the rib
small bump on the posterior side of a rib for articulation with the transverse process of a thoracic vertebra
vertebra
individual bone in the neck and back regions of the vertebral column
vertebral (spinal) canal
bony passageway within the vertebral column for the spinal cord that is formed by the series of individual vertebral foramina
vertebral arch
bony arch formed by the posterior portion of each vertebra that surrounds and protects the spinal cord
vertebral column
entire sequence of bones that extend from the skull to the tailbone
vertebral foramen
opening associated with each vertebra defined by the vertebral arch that provides passage for the spinal cord
vomer bone
unpaired bone that forms the inferior and posterior portions of the nasal septum
xiphoid process
small process that forms the inferior tip of the sternum
zygomatic arch
elongated, free-standing arch on the lateral skull, formed anteriorly by the temporal process of the zygomatic bone and posteriorly by the
zygomatic process of the temporal bone
zygomatic bone
cheekbone; paired bones that contribute to the lateral orbit and anterior zygomatic arch
zygomatic process of the temporal bone
extension from the temporal bone that forms the posterior portion of the zygomatic arch

CHAPTER 8

The appendicular skeleton includes all of the limb bones, plus the bones that unite each limb with the axial skeleton (Figure 8.2). The bones that attach
each upper limb to the axial skeleton form the pectoral girdle (shoulder girdle). This consists of two bones, the scapula and clavicle (Figure 8.3). The
clavicle (collarbone) is an S-shaped bone located on the anterior side of the shoulder. It is attached on its medial end to the sternum of the thoracic
cage, which is part of the axial skeleton. The lateral end of the clavicle articulates (joins) with the scapula just above the shoulder joint. You can easily
palpate, or feel with your fingers, the entire length of your clavicle.
 Figure 8.3 Pectoral Girdle The pectoral girdle consists of the clavicle and
the scapula, which serve to attach the upper limb to the sternum of the
axial skeleton.

The scapula (shoulder blade) lies on the posterior aspect of the shoulder.

It is supported by the clavicle and articulates with the humerus (arm bone)
to form the shoulder joint. The scapula is a flat, triangular-shaped bone with
a prominent ridge running across its posterior surface. This ridge extends
out laterally, where it forms the bony tip of the shoulder and joins with the
lateral end of the clavicle. By following along the clavicle, you can palpate
out to the bony tip of the shoulder, and from there, you can move back
across your posterior shoulder to follow the ridge of the scapula. Move your
shoulder around and feel how the clavicle and scapula move together as a
unit. Both of these bones serve as important attachment sites for muscles
that aid with movements of the shoulder and arm.

The right and left pectoral girdles are not joined to each other, allowing
each to operate independently. In addition, the clavicle of each pectoral
girdle is anchored to the axial skeleton by a single, highly mobile joint. This
allows for the extensive mobility of the entire pectoral girdle, which in turn
enhances movements of the shoulder and upper limb.

Clavicle

The clavicle is the only long bone that lies in a horizontal position in the
body (see Figure 8.3). The clavicle has several important functions. First,
anchored by muscles from above, it serves as a strut that extends laterally
to support the scapula. This in turn holds the shoulder joint superiorly and
laterally from the body trunk, allowing for maximal freedom of motion for
the upper limb. The clavicle also transmits forces acting on the upper limb
to the sternum and axial skeleton. Finally, it serves to protect the
underlying nerves and blood vessels as they pass between the trunk of the
body and the upper limb.

The clavicle has three regions: the medial end, the lateral end, and the
shaft. The medial end, known as the sternal end of the clavicle, has a triangular shape and articulates with the manubrium portion of the sternum. This
forms the sternoclavicular joint, which is the only bony articulation between the pectoral girdle of the upper limb and the axial skeleton. This joint
allows considerable mobility, enabling the clavicle and scapula to move in upward/downward and anterior/posterior directions during shoulder
movements. The sternoclavicular joint is indirectly supported by the costoclavicular ligament (costo- = “rib”), which spans the sternal end of the
clavicle and the underlying first rib. The lateral or acromial end of the clavicle articulates with the acromion of the scapula, the portion of the scapula
that forms the bony tip of the shoulder. There are some sex differences in the morphology of the clavicle. In women, the clavicle tends to be shorter,
thinner, and less curved. In men, the clavicle is heavier and longer, and has a greater curvature and rougher surfaces where muscles attach, features
that are more pronounced in manual workers.

The clavicle is the most commonly fractured bone in the body. Such breaks often occur because of the force exerted on the clavicle when a person falls
onto his or her outstretched arms, or when the lateral shoulder receives a strong blow. Because the sternoclavicular joint is strong and rarely dislocated,
excessive force results in the breaking of the clavicle, usually between the middle and lateral portions of the bone. If the fracture is complete, the
shoulder and lateral clavicle fragment will drop due to the weight of the upper limb, causing the person to support the sagging limb with their other hand.
Muscles acting across the shoulder will also pull the shoulder and lateral clavicle anteriorly and medially, causing the clavicle fragments to override. The
clavicle overlies many important blood vessels and nerves for the upper limb, but fortunately, due to the anterior displacement of a broken clavicle, these
structures are rarely affected when the clavicle is fractured.

Scapula

The scapula is also part of the pectoral girdle and thus plays an important role in anchoring the upper limb to the body. The scapula is located on the
posterior side of the shoulder. It is surrounded by muscles on both its anterior (deep) and posterior (superficial) sides, and thus does not articulate with
the ribs of the thoracic cage.

The scapula has several important landmarks (Figure 8.4). The three margins or borders of the scapula, named for their positions within the body, are
the superior border of the scapula, the medial border of the scapula, and the lateral border of the scapula. The suprascapular notch is located
lateral to the midpoint of the superior border. The corners of the triangular scapula, at either end of the medial border, are the superior angle of the
scapula, located between the medial and superior borders, and the inferior angle of the scapula, located between the medial and lateral borders. The
inferior angle is the most inferior portion of the scapula, and is particularly important because it serves as the attachment point for several powerful
muscles involved in shoulder and upper limb movements. The remaining corner of the scapula, between the superior and lateral borders, is the location
of the glenoid cavity (glenoid fossa). This shallow depression articulates with the humerus bone of the arm to form the glenohumeral joint (shoulder
joint). The small bony bumps located immediately above and below the glenoid cavity are the supraglenoid tubercle and the infraglenoid tubercle,
respectively. These provide attachments for muscles of the arm.
 Figure 8.4 Scapula The isolated scapula is shown
here from its anterior (deep) side and its posterior
(superficial) side.

The scapula also has two prominent projections.

Toward the lateral end of the superior border, between
the suprascapular notch and glenoid cavity, is the
hook-like coracoid process (coracoid = “shaped like a
crow’s beak”). This process projects anteriorly and
curves laterally. At the shoulder, the coracoid process
is located inferior to the lateral end of the clavicle. It is
anchored to the clavicle by a strong ligament, and
serves as the attachment site for muscles of the
anterior chest and arm. On the posterior aspect,
the spine of the scapula is a long and prominent
ridge that runs across its upper portion. Extending
laterally from the spine is a flattened and expanded
region called the acromion or acromial process. The
acromion forms the bony tip of the superior shoulder
region and articulates with the lateral end of the
clavicle, forming the acromioclavicular
joint (see Figure 8.3). Together, the clavicle, acromion, and spine of the
scapula form a V-shaped bony line that provides for the attachment of neck
and back muscles that act on the shoulder, as well as muscles that pass
across the shoulder joint to act on the arm.

The scapula has three depressions, each of which is called a fossa (plural

= fossae). Two of these are found on the posterior scapula, above and
below the scapular spine. Superior to the spine is the narrow supraspinous
fossa, and inferior to the spine is the broad infraspinous fossa. The
anterior (deep) surface of the scapula forms the broad subscapular fossa.
All of these fossae provide large surface areas for the attachment of
muscles that cross the shoulder joint to act on the humerus.

The acromioclavicular joint transmits forces from the upper limb to the clavicle. The ligaments around this joint are relatively weak. A hard fall onto the
elbow or outstretched hand can stretch or tear the acromioclavicular ligaments, resulting in a moderate injury to the joint. However, the primary support
for the acromioclavicular joint comes from a very strong ligament called the coracoclavicular ligament (see Figure 8.3). This connective tissue band
anchors the coracoid process of the scapula to the inferior surface of the acromial end of the clavicle and thus provides important indirect support for the
acromioclavicular joint. Following a strong blow to the lateral shoulder, such as when a hockey player is driven into the boards, a complete dislocation of
the acromioclavicular joint can result. In this case, the acromion is thrust under the acromial end of the clavicle, resulting in ruptures of both the
acromioclavicular and coracoclavicular ligaments. The scapula then separates from the clavicle, with the weight of the upper limb pulling the shoulder
downward. This dislocation injury of the acromioclavicular joint is known as a “shoulder separation” and is common in contact sports such as hockey,
football, or martial arts.

The upper limb is divided into three regions. These consist of the arm, located between the shoulder and elbow joints; the forearm, which is between
the elbow and wrist joints; and the hand, which is located distal to the wrist. There are 30 bones in each upper limb (see Figure 8.2). The humerus is
the single bone of the upper arm, and the ulna (medially) and the radius (laterally) are the paired bones of the forearm. The base of the hand contains
eight bones, each called a carpal bone, and the palm of the hand is formed by five bones, each called a metacarpal bone. The fingers and thumb
contain a total of 14 bones, each of which is a phalanx bone of the hand.

Humerus

The humerus is the single bone of the upper arm region (Figure 8.5). At its proximal end is the head of the humerus. This is the large, round, smooth
region that faces medially. The head articulates with the glenoid cavity of the scapula to form the glenohumeral (shoulder) joint. The margin of the
smooth area of the head is the anatomical neck of the humerus. Located on the lateral side of the proximal humerus is an expanded bony area called
the greater tubercle. The smaller lesser tubercle of the humerus is found on the anterior aspect of the humerus. Both the greater and lesser tubercles
serve as attachment sites for muscles that act across the shoulder joint. Passing between the greater and lesser tubercles is the narrow intertubercular
groove (sulcus), which is also known as the bicipital groove because it provides passage for a tendon of the biceps brachii muscle. The surgical
neck is located at the base of the expanded, proximal end of the humerus, where it joins the narrow shaft of the humerus. The surgical neck is a
common site of arm fractures. The deltoid tuberosity is a roughened, V-shaped region located on the lateral side in the middle of the humerus shaft. As
its name indicates, it is the site of attachment for the deltoid muscle.

Figure 8.5 Humerus and Elbow Joint The humerus is the single bone of the upper arm region. It articulates with the radius and ulna bones of the
forearm to form the elbow joint.

Distally, the humerus becomes flattened. The prominent bony projection on the medial side is the medial epicondyle of the humerus. The much
smaller lateral epicondyle of the humerus is found on the lateral side of the distal humerus. The roughened ridge of bone above the lateral epicondyle
is the lateral supracondylar ridge. All of these areas are attachment points for muscles that act on the forearm, wrist, and hand. The powerful grasping
muscles of the anterior forearm arise from the medial epicondyle, which is thus larger and more robust than the lateral epicondyle that gives rise to the
weaker posterior forearm muscles.

The distal end of the humerus has two articulation areas, which join the ulna and radius bones of the forearm to form the elbow joint. The more medial
of these areas is the trochlea, a spindle- or pulley-shaped region (trochlea = “pulley”), which articulates with the ulna bone. Immediately lateral to the
trochlea is the capitulum (“small head”), a knob-like structure located on the anterior surface of the distal humerus. The capitulum articulates with the
radius bone of the forearm. Just above these bony areas are two small depressions. These spaces accommodate the forearm bones when the elbow is
fully bent (flexed). Superior to the trochlea is the coronoid fossa, which receives the coronoid process of the ulna, and above the capitulum is the radial
fossa, which receives the head of the radius when the elbow is flexed. Similarly, the posterior humerus has the olecranon fossa, a larger depression
that receives the olecranon process of the ulna when the forearm is fully extended.

Ulna

The ulna is the medial bone of the forearm. It runs parallel to the radius, which is the lateral bone of the forearm (Figure 8.6). The proximal end of the
ulna resembles a crescent wrench with its large, C-shaped trochlear notch. This region articulates with the trochlea of the humerus as part of the elbow
joint. The inferior margin of the trochlear notch is formed by a prominent lip of bone called the coronoid process of the ulna. Just below this on the
anterior ulna is a roughened area called the ulnar tuberosity. To the lateral side and slightly inferior to the trochlear notch is a small, smooth area called
the radial notch of the ulna. This area is the site of articulation between the proximal radius and the ulna, forming the proximal radioulnar joint. The
posterior and superior portions of the proximal ulna make up the olecranon process, which forms the bony tip of the elbow.

Figure 8.6 Ulna and Radius The ulna is located on the medial side of the

forearm, and the radius is on the lateral side. These bones are attached to
each other by an interosseous membrane.

More distal is the shaft of the ulna. The lateral side of the shaft forms a
ridge called the interosseous border of the ulna. This is the line of
attachment for the interosseous membrane of the forearm, a sheet of
dense connective tissue that unites the ulna and radius bones. The small,
rounded area that forms the distal end is the head of the ulna. Projecting
from the posterior side of the ulnar head is the styloid process of the ulna,
a short bony projection. This serves as an attachment point for a connective
tissue structure that unites the distal ends of the ulna and radius.

In the anatomical position, with the elbow fully extended and the palms
facing forward, the arm and forearm do not form a straight line. Instead, the
forearm deviates laterally by 5–15 degrees from the line of the arm. This
deviation is called the carrying angle. It allows the forearm and hand to
swing freely or to carry an object without hitting the hip. The carrying angle
is larger in females to accommodate their wider pelvis.

Radius

The radius runs parallel to the ulna, on the lateral (thumb) side of the
forearm (see Figure 8.6). The head of the radius is a disc-shaped structure
that forms the proximal end. The small depression on the surface of the
head articulates with the capitulum of the humerus as part of the elbow joint,
whereas the smooth, outer margin of the head articulates with the radial
notch of the ulna at the proximal radioulnar joint. The neck of the radius is
the narrowed region immediately below the expanded head. Inferior to this
point on the medial side is the radial tuberosity, an oval-shaped, bony
protuberance that serves as a muscle attachment point. The shaft of the
radius is slightly curved and has a small ridge along its medial side. This
ridge forms the interosseous border of the radius, which, like the similar
border of the ulna, is the line of attachment for the interosseous membrane
that unites the two forearm bones. The distal end of the radius has a smooth surface for articulation with two carpal bones to form the radiocarpal
joint or wrist joint (Figure 8.7 and Figure 8.8). On the medial side of the distal radius is the ulnar notch of the radius. This shallow depression
articulates with the head of the ulna, which together form the distal radioulnar joint. The lateral end of the radius has a pointed projection called
the styloid process of the radius. This provides attachment for ligaments that support the lateral side of the wrist joint. Compared to the styloid process
of the ulna, the styloid process of the radius projects more distally, thereby limiting the range of movement for lateral deviations of the hand at the wrist
joint.

Carpal Bones

The wrist and base of the hand are formed by a series of eight small carpal bones (see Figure 8.7). The carpal bones are arranged in two rows, forming
a proximal row of four carpal bones and a distal row of four carpal bones. The bones in the proximal row, running from the lateral (thumb) side to the
medial side, are the scaphoid (“boat-shaped”), lunate (“moon-shaped”), triquetrum (“three-cornered”), and pisiform (“pea-shaped”) bones. The small,
rounded pisiform bone articulates with the anterior surface of the triquetrum bone. The pisiform thus projects anteriorly, where it forms the bony bump
that can be felt at the medial base of your hand. The distal bones (lateral to medial) are the trapezium (“table”), trapezoid (“resembles a
table”), capitate (“head-shaped”), and hamate (“hooked bone”) bones. The hamate bone is characterized by a prominent bony extension on its anterior
side called the hook of the hamate bone.

A helpful mnemonic for remembering the arrangement of the carpal bones is “So Long To Pinky, Here Comes The Thumb.” This mnemonic starts on the
lateral side and names the proximal bones from lateral to medial (scaphoid, lunate, triquetrum, pisiform), then makes a U-turn to name the distal bones
from medial to lateral (hamate, capitate, trapezoid, trapezium). Thus, it starts and finishes on the lateral side.

Figure 8.7 Bones of the Wrist and Hand The

eight carpal bones form the base of the hand.
These are arranged into proximal and distal
rows of four bones each. The metacarpal bones
form the palm of the hand. The thumb and
fingers consist of the phalanx bones.

The carpal bones form the base of the hand.

This can be seen in the radiograph (X-ray
image) of the hand that shows the relationships
of the hand bones to the skin creases of the
hand (see Figure 8.8). Within the carpal bones,
the four proximal bones are united to each other
by ligaments to form a unit. Only three of these
bones, the scaphoid, lunate, and triquetrum,
contribute to the radiocarpal joint. The scaphoid
and lunate bones articulate directly with the
distal end of the radius, whereas the triquetrum
bone articulates with a fibrocartilaginous pad
that spans the radius and styloid process of the
ulna. The distal end of the ulna thus does not
directly articulate with any of the carpal bones.

The four distal carpal bones are also held

together as a group by ligaments. The proximal
and distal rows of carpal bones articulate with
each other to form the midcarpal
joint (see Figure 8.8). Together, the radiocarpal
and midcarpal joints are responsible for all movements of the hand at the wrist. The distal carpal bones also articulate with the metacarpal bones of the
hand.

Figure 8.8 Bones of the Hand This radiograph

shows the position of the bones within the hand.
Note the carpal bones that form the base of the
hand. (credit: modification of work by Trace
Meek)

In the articulated hand, the carpal bones form a

U-shaped grouping. A strong ligament called
the flexor retinaculum spans the top of this U-
shaped area to maintain this grouping of the
carpal bones. The flexor retinaculum is attached
laterally to the trapezium and scaphoid bones,
and medially to the hamate and pisiform bones.
Together, the carpal bones and the flexor
retinaculum form a passageway called
the carpal tunnel, with the carpal bones forming
the walls and floor, and the flexor retinaculum
forming the roof of this space (Figure 8.9). The
tendons of nine muscles of the anterior forearm
and an important nerve pass through this narrow
tunnel to enter the hand. Overuse of the muscle
tendons or wrist injury can produce inflammation
and swelling within this space. This produces
compression of the nerve, resulting in carpal tunnel syndrome, which is characterized by pain or numbness, and muscle weakness in those areas of the
hand supplied by this nerve.
 Figure 8.9 Carpal Tunnel The carpal tunnel is the passageway by which
nine muscle tendons and a major nerve enter the hand from the anterior
forearm. The walls and floor of the carpal tunnel are formed by the U-
shaped grouping of the carpal bones, and the roof is formed by the flexor
retinaculum, a strong ligament that anteriorly unites the bones.

Metacarpal Bones

The palm of the hand contains five elongated metacarpal bones. These
bones lie between the carpal bones of the wrist and the bones of the fingers
and thumb (see Figure 8.7). The proximal end of each metacarpal bone
articulates with one of the distal carpal bones. Each of these articulations is
a carpometacarpal joint (see Figure 8.8). The expanded distal end of each
metacarpal bone articulates at the metacarpophalangeal joint with the
proximal phalanx bone of the thumb or one of the fingers. The distal end
also forms the knuckles of the hand, at the base of the fingers. The
metacarpal bones are numbered 1–5, beginning at the thumb.

The first metacarpal bone, at the base of the thumb, is separated from the other metacarpal bones. This allows it a freedom of motion that is
independent of the other metacarpal bones, which is very important for thumb mobility. The remaining metacarpal bones are united together to form the
palm of the hand. The second and third metacarpal bones are firmly anchored in place and are immobile. However, the fourth and fifth metacarpal
bones have limited anterior-posterior mobility, a motion that is greater for the fifth bone. This mobility is important during power gripping with the hand
(Figure 8.10). The anterior movement of these bones, particularly the fifth metacarpal bone, increases the strength of contact for the medial hand during
gripping actions.

Figure 8.10 Hand During Gripping During tight gripping—compare (b) to (a)—the fourth and, particularly, the fifth metatarsal bones are pulled
anteriorly. This increases the contact between the object and the medial side of the hand, thus improving the firmness of the grip.

Phalanx Bones

The fingers and thumb contain 14 bones, each of which is called a phalanx bone (plural = phalanges), named after the ancient Greek phalanx (a
rectangular block of soldiers). The thumb (pollex) is digit number 1 and has two phalanges, a proximal phalanx, and a distal phalanx bone (see Figure
8.7). Digits 2 (index finger) through 5 (little finger) have three phalanges each, called the proximal, middle, and distal phalanx bones.
An interphalangeal joint is one of the articulations between adjacent phalanges of the digits (see Figure 8.8).

DISORDERS OF THE...

Appendicular System: Fractures of Upper Limb Bones

Due to our constant use of the hands and the rest of our upper limbs, an injury to any of these areas will cause a significant loss of functional ability.
Many fractures result from a hard fall onto an outstretched hand. The resulting transmission of force up the limb may result in a fracture of the humerus,
radius, or scaphoid bones. These injuries are especially common in elderly people whose bones are weakened due to osteoporosis.

Falls onto the hand or elbow, or direct blows to the arm, can result in fractures of the humerus (Figure 8.11). Following a fall, fractures at the surgical
neck, the region at which the expanded proximal end of the humerus joins with the shaft, can result in an impacted fracture, in which the distal portion of
the humerus is driven into the proximal portion. Falls or blows to the arm can also produce transverse or spiral fractures of the humeral shaft.

In children, a fall onto the tip of the elbow frequently results in a distal humerus fracture. In these, the olecranon of the ulna is driven upward, resulting in
a fracture across the distal humerus, above both epicondyles (supracondylar fracture), or a fracture between the epicondyles, thus separating one or
both of the epicondyles from the body of the humerus (intercondylar fracture). With these injuries, the immediate concern is possible compression of the
artery to the forearm due to swelling of the surrounding tissues. If compression occurs, the resulting ischemia (lack of oxygen) due to reduced blood flow
can quickly produce irreparable damage to the forearm muscles. In addition, four major nerves for shoulder and upper limb muscles are closely
associated with different regions of the humerus, and thus, humeral fractures may also damage these nerves.

Another frequent injury following a fall onto an outstretched hand is a Colles fracture (“col-lees”) of the distal radius (see Figure 8.11). This involves a
complete transverse fracture across the distal radius that drives the separated distal fragment of the radius posteriorly and superiorly. This injury results
in a characteristic “dinner fork” bend of the forearm just above the wrist due to the posterior displacement of the hand. This is the most frequent forearm
fracture and is a common injury in persons over the age of 50, particularly in older women with osteoporosis. It also commonly occurs following a high-
speed fall onto the hand during activities such as snowboarding or skating.

The most commonly fractured carpal bone is the scaphoid, often resulting from a fall onto the hand. Deep pain at the lateral wrist may yield an initial
diagnosis of a wrist sprain, but a radiograph taken several weeks after the injury, after tissue swelling has subsided, will reveal the fracture. Due to the
poor blood supply to the scaphoid bone, healing will be slow and there is the danger of bone necrosis and subsequent degenerative joint disease of the
wrist.
Figure 8.11 Fractures of the Humerus and Radius Falls or direct blows can result in
fractures of the surgical neck or shaft of the humerus. Falls onto the elbow can fracture
the distal humerus. A Colles fracture of the distal radius is the most common forearm
fracture.

INTERACTIVE LINK

Watch this video to learn about a Colles fracture, a break of the distal radius, usually
caused by falling onto an outstretched hand. When would surgery be required and how
would the fracture be repaired in this case?

Learning Objectives

By the end of this section, you will be able to:

 Define the pelvic girdle and describe the bones and ligaments of the pelvis
 Explain the three regions of the hip bone and identify their bony landmarks
 Describe the openings of the pelvis and the boundaries of the greater and
lesser pelvis
The pelvic girdle (hip girdle) is formed by a single bone, the hip bone or coxal bone (coxal = “hip”), which serves as the attachment point for each
lower limb. Each hip bone, in turn, is firmly joined to the axial skeleton via its attachment to the sacrum of the vertebral column. The right and left hip
bones also converge anteriorly to attach to each other. The bony pelvis is the entire structure formed by the two hip bones, the sacrum, and, attached
inferiorly to the sacrum, the coccyx (Figure 8.12).

Unlike the bones of the pectoral girdle, which are highly mobile to enhance the range of upper limb movements, the bones of the pelvis are strongly
united to each other to form a largely immobile, weight-bearing structure. This is important for stability because it enables the weight of the body to be
easily transferred laterally from the vertebral column, through the pelvic girdle and hip joints, and into either lower limb whenever the other limb is not
bearing weight. Thus, the immobility of the pelvis provides a strong foundation for the upper body as it rests on top of the mobile lower limbs.

Figure 8.12 Pelvis The pelvic girdle is formed by a single hip bone.

The hip bone attaches the lower limb to the axial skeleton through
its articulation with the sacrum. The right and left hip bones, plus the
sacrum and the coccyx, together form the pelvis.

Hip Bone

The hip bone, or coxal bone, forms the pelvic girdle portion of the
pelvis. The paired hip bones are the large, curved bones that form
the lateral and anterior aspects of the pelvis. Each adult hip bone is
formed by three separate bones that fuse together during the late
teenage years. These bony components are the ilium, ischium, and
pubis (Figure 8.13). These names are retained and used to define
the three regions of the adult hip bone.

Figure 8.13 The Hip Bone The adult hip bone consists of three

regions. The ilium forms the large, fan-shaped superior portion, the
ischium forms the posteroinferior portion, and the pubis forms the anteromedial portion.

The ilium is the fan-like, superior region that forms the largest part of the hip bone. It is firmly united to the sacrum at the largely immobile sacroiliac
joint (see Figure 8.12). The ischium forms the posteroinferior region of each hip bone. It supports the body when sitting. The pubis forms the anterior
portion of the hip bone. The pubis curves medially, where it joins to the pubis of the opposite hip bone at a specialized joint called the pubic symphysis.

Ilium

When you place your hands on your waist, you can feel the arching, superior margin of the ilium along your waistline (see Figure 8.13). This curved,
superior margin of the ilium is the iliac crest. The rounded, anterior termination of the iliac crest is the anterior superior iliac spine. This important
bony landmark can be felt at your anterolateral hip. Inferior to the anterior superior iliac spine is a rounded protuberance called the anterior inferior iliac
spine. Both of these iliac spines serve as attachment points for muscles of the thigh. Posteriorly, the iliac crest curves downward to terminate as
the posterior superior iliac spine. Muscles and ligaments surround but do not cover this bony landmark, thus sometimes producing a depression seen
as a “dimple” located on the lower back. More inferiorly is the posterior inferior iliac spine. This is located at the inferior end of a large, roughened area
called the auricular surface of the ilium. The auricular surface articulates with the auricular surface of the sacrum to form the sacroiliac joint. Both the
posterior superior and posterior inferior iliac spines serve as attachment points for the muscles and very strong ligaments that support the sacroiliac joint.

The shallow depression located on the anteromedial (internal) surface of the upper ilium is called the iliac fossa. The inferior margin of this space is
formed by the arcuate line of the ilium, the ridge formed by the pronounced change in curvature between the upper and lower portions of the ilium. The
large, inverted U-shaped indentation located on the posterior margin of the lower ilium is called the greater sciatic notch.

Ischium

The ischium forms the posterolateral portion of the hip bone (see Figure 8.13). The large, roughened area of the inferior ischium is the ischial
tuberosity. This serves as the attachment for the posterior thigh muscles and also carries the weight of the body when sitting. You can feel the ischial
tuberosity if you wiggle your pelvis against the seat of a chair. Projecting superiorly and anteriorly from the ischial tuberosity is a narrow segment of bone
called the ischial ramus. The slightly curved posterior margin of the ischium above the ischial tuberosity is the lesser sciatic notch. The bony
projection separating the lesser sciatic notch and greater sciatic notch is the ischial spine.

Pubis

The pubis forms the anterior portion of the hip bone (see Figure 8.13). The enlarged medial portion of the pubis is the pubic body. Located superiorly
on the pubic body is a small bump called the pubic tubercle. The superior pubic ramus is the segment of bone that passes laterally from the pubic
body to join the ilium. The narrow ridge running along the superior margin of the superior pubic ramus is the pectineal line of the pubis.

The pubic body is joined to the pubic body of the opposite hip bone by the pubic symphysis. Extending downward and laterally from the body is
the inferior pubic ramus. The pubic arch is the bony structure formed by the pubic symphysis, and the bodies and inferior pubic rami of the adjacent
pubic bones. The inferior pubic ramus extends downward to join the ischial ramus. Together, these form the single ischiopubic ramus, which extends
from the pubic body to the ischial tuberosity. The inverted V-shape formed as the ischiopubic rami from both sides come together at the pubic symphysis
is called the subpubic angle.

Pelvis

The pelvis consists of four bones: the right and left hip bones, the sacrum, and the coccyx (see Figure 8.12). The pelvis has several important functions.
Its primary role is to support the weight of the upper body when sitting and to transfer this weight to the lower limbs when standing. It serves as an
attachment point for trunk and lower limb muscles, and also protects the internal pelvic organs. When standing in the anatomical position, the pelvis is
tilted anteriorly. In this position, the anterior superior iliac spines and the pubic tubercles lie in the same vertical plane, and the anterior (internal) surface
of the sacrum faces forward and downward.

The three areas of each hip bone, the ilium, pubis, and ischium, converge centrally to form a deep, cup-shaped cavity called the acetabulum. This is
located on the lateral side of the hip bone and is part of the hip joint. The large opening in the anteroinferior hip bone between the ischium and pubis is
the obturator foramen. This space is largely filled in by a layer of connective tissue and serves for the attachment of muscles on both its internal and
external surfaces.

Several ligaments unite the bones of the pelvis (Figure 8.14). The largely immobile sacroiliac joint is supported by a pair of strong ligaments that are
attached between the sacrum and ilium portions of the hip bone. These are the anterior sacroiliac ligament on the anterior side of the joint and
the posterior sacroiliac ligament on the posterior side. Also spanning the sacrum and hip bone are two additional ligaments. The sacrospinous
ligament runs from the sacrum to the ischial spine, and the sacrotuberous ligament runs from the sacrum to the ischial tuberosity. These ligaments
help to support and immobilize the sacrum as it carries the weight of the body.

Figure 8.14 Ligaments of the Pelvis The posterior sacroiliac

ligament supports the sacroiliac joint. The sacrospinous
ligament spans the sacrum to the ischial spine, and the
sacrotuberous ligament spans the sacrum to the ischial
tuberosity. The sacrospinous and sacrotuberous ligaments
contribute to the formation of the greater and lesser sciatic
foramina.

INTERACTIVE LINK

Watch this video for a 3-D view of the pelvis and its associated

ligaments. What is the large opening in the bony pelvis, located
between the ischium and pubic regions, and what two parts of
the pubis contribute to the formation of this opening?

The sacrospinous and sacrotuberous ligaments also help to

define two openings on the posterolateral sides of the pelvis
through which muscles, nerves, and blood vessels for the lower limb exit. The superior opening is the greater sciatic foramen. This large opening is
formed by the greater sciatic notch of the hip bone, the sacrum, and the sacrospinous ligament. The smaller, more inferior lesser sciatic foramen is
formed by the lesser sciatic notch of the hip bone, together with the sacrospinous and sacrotuberous ligaments.

The space enclosed by the bony pelvis is divided into two regions (Figure 8.15). The broad, superior region, defined laterally by the large, fan-like portion
of the upper hip bone, is called the greater pelvis (greater pelvic cavity; false pelvis). This broad area is occupied by portions of the small and large
intestines, and because it is more closely associated with the abdominal cavity, it is sometimes referred to as the false pelvis. More inferiorly, the narrow,
rounded space of the lesser pelvis (lesser pelvic cavity; true pelvis) contains the bladder and other pelvic organs, and thus is also known as the true
pelvis. The pelvic brim (also known as the pelvic inlet) forms the superior margin of the lesser pelvis, separating it from the greater pelvis. The pelvic
brim is defined by a line formed by the upper margin of the pubic symphysis anteriorly, and the pectineal line of the pubis, the arcuate line of the ilium,
and the sacral promontory (the anterior margin of the superior sacrum) posteriorly. The inferior limit of the lesser pelvic cavity is called the pelvic outlet.
This large opening is defined by the inferior margin of the pubic symphysis anteriorly, and the ischiopubic ramus, the ischial tuberosity, the
sacrotuberous ligament, and the inferior tip of the coccyx posteriorly. Because of the anterior tilt of the pelvis, the lesser pelvis is also angled, giving it an
anterosuperior (pelvic inlet) to posteroinferior (pelvic outlet) orientation.
Figure 8.15 Male and Female Pelvis The female pelvis is adapted for childbirth and is broader, with a larger subpubic angle, a rounder pelvic brim, and
a wider and more shallow lesser pelvic cavity than the male pelvis.

Comparison of the Female and Male Pelvis

The differences between the adult female and male pelvis relate to function and body size. In general, the bones of the male pelvis are thicker and
heavier, adapted for support of the male’s heavier physical build and stronger muscles. The greater sciatic notch of the male hip bone is narrower and
deeper than the broader notch of females. Because the female pelvis is adapted for childbirth, it is wider than the male pelvis, as evidenced by the
distance between the anterior superior iliac spines (see Figure 8.15). The ischial tuberosities of females are also farther apart, which increases the size
of the pelvic outlet. Because of this increased pelvic width, the subpubic angle is larger in females (greater than 80 degrees) than it is in males (less than
70 degrees). The female sacrum is wider, shorter, and less curved, and the sacral promontory projects less into the pelvic cavity, thus giving the female
pelvic inlet (pelvic brim) a more rounded or oval shape compared to males. The lesser pelvic cavity of females is also wider and more shallow than the
narrower, deeper, and tapering lesser pelvis of males. Because of the obvious differences between female and male hip bones, this is the one bone of
the body that allows for the most accurate sex determination. Table 8.1 provides an overview of the general differences between the female and male
pelvis.

Overview of Differences between the Female and Male Pelvis

Table8.1

CAREER CONNECTION

Forensic Pathology and Forensic Anthropology

A forensic pathologist (also known as a medical examiner) is a medically trained physician who has been specifically trained in pathology to examine the
bodies of the deceased to determine the cause of death. A forensic pathologist applies his or her understanding of disease as well as toxins, blood and
DNA analysis, firearms and ballistics, and other factors to assess the cause and manner of death. At times, a forensic pathologist will be called to testify
under oath in situations that involve a possible crime. Forensic pathology is a field that has received much media attention on television shows or
following a high-profile death.

While forensic pathologists are responsible for determining whether the cause of someone’s death was natural, a suicide, accidental, or a homicide,
there are times when uncovering the cause of death is more complex, and other skills are needed. Forensic anthropology brings the tools and
knowledge of physical anthropology and human osteology (the study of the skeleton) to the task of investigating a death. A forensic anthropologist
assists medical and legal professionals in identifying human remains. The science behind forensic anthropology involves the study of archaeological
excavation; the examination of hair; an understanding of plants, insects, and footprints; the ability to determine how much time has elapsed since the
person died; the analysis of past medical history and toxicology; the ability to determine whether there are any postmortem injuries or alterations of the
skeleton; and the identification of the decedent (deceased person) using skeletal and dental evidence.

Due to the extensive knowledge and understanding of excavation techniques, a forensic anthropologist is an integral and invaluable team member to
have on-site when investigating a crime scene, especially when the recovery of human skeletal remains is involved. When remains are bought to a
forensic anthropologist for examination, he or she must first determine whether the remains are in fact human. Once the remains have been identified as
belonging to a person and not to an animal, the next step is to approximate the individual’s age, sex, race, and height. The forensic anthropologist does
not determine the cause of death, but rather provides information to the forensic pathologist, who will use all of the data collected to make a final
determination regarding the cause of death.

Like the upper limb, the lower limb is divided into three regions. The thigh is that portion of the lower limb located between the hip joint and knee joint.
The leg is specifically the region between the knee joint and the ankle joint. Distal to the ankle is the foot. The lower limb contains 30 bones. These are
the femur, patella, tibia, fibula, tarsal bones, metatarsal bones, and phalanges (see Figure 8.2). The femur is the single bone of the thigh. The patella is
the kneecap and articulates with the distal femur. The tibia is the larger, weight-bearing bone located on the medial side of the leg, and the fibula is the
thin bone of the lateral leg. The bones of the foot are divided into three groups. The posterior portion of the foot is formed by a group of seven bones,
each of which is known as a tarsal bone, whereas the mid-foot contains five elongated bones, each of which is a metatarsal bone. The toes contain 14
small bones, each of which is a phalanx bone of the foot.
 Femur

The femur, or thigh bone, is the single bone of the thigh region (Figure 8.16).
It is the longest and strongest bone of the body, and accounts for
approximately one-quarter of a person’s total height. The rounded, proximal
end is the head of the femur, which articulates with the acetabulum of the
hip bone to form the hip joint. The fovea capitis is a minor indentation on
the medial side of the femoral head that serves as the site of attachment for
the ligament of the head of the femur. This ligament spans the femur and
acetabulum, but is weak and provides little support for the hip joint. It does,
however, carry an important artery that supplies the head of the femur.

Figure 8.16 Femur and Patella The femur is the single bone of the thigh

region. It articulates superiorly with the hip bone at the hip joint, and inferiorly
with the tibia at the knee joint. The patella only articulates with the distal end
of the femur.

The narrowed region below the head is the neck of the femur. This is a
common area for fractures of the femur. The greater trochanter is the large,
upward, bony projection located above the base of the neck. Multiple muscles
that act across the hip joint attach to the greater trochanter, which, because
of its projection from the femur, gives additional leverage to these muscles.
The greater trochanter can be felt just under the skin on the lateral side of
your upper thigh. The lesser trochanter is a small, bony prominence that lies
on the medial aspect of the femur, just below the neck. A single, powerful
muscle attaches to the lesser trochanter. Running between the greater and
lesser trochanters on the anterior side of the femur is the
roughened intertrochanteric line. The trochanters are also connected on the
posterior side of the femur by the larger intertrochanteric crest.

The elongated shaft of the femur has a slight anterior bowing or curvature.

At its proximal end, the posterior shaft has the gluteal tuberosity, a
roughened area extending inferiorly from the greater trochanter. More
inferiorly, the gluteal tuberosity becomes continuous with the linea
aspera (“rough line”). This is the roughened ridge that passes distally along
the posterior side of the mid-femur. Multiple muscles of the hip and thigh regions make long, thin attachments to the femur along the linea aspera.

The distal end of the femur has medial and lateral bony expansions. On the lateral side, the smooth portion that covers the distal and posterior aspects
of the lateral expansion is the lateral condyle of the femur. The roughened area on the outer, lateral side of the condyle is the lateral epicondyle of
the femur. Similarly, the smooth region of the distal and posterior medial femur is the medial condyle of the femur, and the irregular outer, medial side
of this is the medial epicondyle of the femur. The lateral and medial condyles articulate with the tibia to form the knee joint. The epicondyles provide
attachment for muscles and supporting ligaments of the knee. The adductor tubercle is a small bump located at the superior margin of the medial
epicondyle. Posteriorly, the medial and lateral condyles are separated by a deep depression called the intercondylar fossa. Anteriorly, the smooth
surfaces of the condyles join together to form a wide groove called the patellar surface, which provides for articulation with the patella bone. The
combination of the medial and lateral condyles with the patellar surface gives the distal end of the femur a horseshoe (U) shape.

INTERACTIVE LINK

Watch this video to view how a fracture of the mid-femur is surgically repaired. How are the two portions of the broken femur stabilized during surgical
repair of a fractured femur?

Patella

The patella (kneecap) is largest sesamoid bone of the body (see Figure 8.16). A sesamoid bone is a bone that is incorporated into the tendon of a
muscle where that tendon crosses a joint. The sesamoid bone articulates with the underlying bones to prevent damage to the muscle tendon due to
rubbing against the bones during movements of the joint. The patella is found in the tendon of the quadriceps femoris muscle, the large muscle of the
anterior thigh that passes across the anterior knee to attach to the tibia. The patella articulates with the patellar surface of the femur and thus prevents
rubbing of the muscle tendon against the distal femur. The patella also lifts the tendon away from the knee joint, which increases the leverage power of
the quadriceps femoris muscle as it acts across the knee. The patella does not articulate with the tibia.

INTERACTIVE LINK

Visit this site to perform a virtual knee replacement surgery. The prosthetic knee components must be properly aligned to function properly. How is this
alignment ensured?

HOMEOSTATIC IMBALANCES
Runner’s Knee

Runner’s knee, also known as patellofemoral syndrome, is the most common overuse injury among runners. It is most frequent in adolescents and
young adults, and is more common in females. It often results from excessive running, particularly downhill, but may also occur in athletes who do a lot
of knee bending, such as jumpers, skiers, cyclists, weight lifters, and soccer players. It is felt as a dull, aching pain around the front of the knee and deep
to the patella. The pain may be felt when walking or running, going up or down stairs, kneeling or squatting, or after sitting with the knee bent for an
extended period.

Patellofemoral syndrome may be initiated by a variety of causes, including individual variations in the shape and movement of the patella, a direct blow
to the patella, or flat feet or improper shoes that cause excessive turning in or out of the feet or leg. These factors may cause in an imbalance in the
muscle pull that acts on the patella, resulting in an abnormal tracking of the patella that allows it to deviate too far toward the lateral side of the patellar
surface on the distal femur.

Because the hips are wider than the knee region, the femur has a diagonal orientation within the thigh, in contrast to the vertically oriented tibia of the leg
(Figure 8.17). The Q-angle is a measure of how far the femur is angled laterally away from vertical. The Q-angle is normally 10–15 degrees, with
females typically having a larger Q-angle due to their wider pelvis. During extension of the knee, the quadriceps femoris muscle pulls the patella both
superiorly and laterally, with the lateral pull greater in women due to their large Q-angle. This makes women more vulnerable to developing
patellofemoral syndrome than men. Normally, the large lip on the lateral side of the patellar surface of the femur compensates for the lateral pull on the
patella, and thus helps to maintain its proper tracking.

However, if the pull produced by the medial and lateral sides of the quadriceps femoris muscle is not properly balanced, abnormal tracking of the patella
toward the lateral side may occur. With continued use, this produces pain and could result in damage to the articulating surfaces of the patella and
femur, and the possible future development of arthritis. Treatment generally involves stopping the activity that produces knee pain for a period of time,
followed by a gradual resumption of activity. Proper strengthening of the quadriceps femoris muscle to correct for imbalances is also important to help
prevent reoccurrence.

Figure 8.17 The Q-Angle The Q-angle is a measure of the amount of lateral deviation of the femur from the
vertical line of the tibia. Adult females have a larger Q-angle due to their wider pelvis than adult males.

Tibia

The tibia (shin bone) is the medial bone of the leg and is larger than the fibula, with which it is paired (Figure
8.18). The tibia is the main weight-bearing bone of the lower leg and the second longest bone of the body, after
the femur. The medial side of the tibia is located immediately under the skin, allowing it to be easily palpated
down the entire length of the medial leg.

Figure 8.18 Tibia and Fibula The tibia is

the larger, weight-bearing bone located on
the medial side of the leg. The fibula is the
slender bone of the lateral side of the leg
and does not bear weight.

The proximal end of the tibia is greatly

expanded. The two sides of this expansion
form the medial condyle of the tibia and
the lateral condyle of the tibia. The tibia
does not have epicondyles. The top surface
of each condyle is smooth and flattened.
These areas articulate with the medial and
lateral condyles of the femur to form
the knee joint. Between the articulating
surfaces of the tibial condyles is
the intercondylar eminence, an irregular,
elevated area that serves as the inferior
attachment point for two supporting
ligaments of the knee.

The tibial tuberosity is an elevated area on

the anterior side of the tibia, near its proximal end. It is the final site of
attachment for the muscle tendon associated with the patella. More inferiorly,
the shaft of the tibia becomes triangular in shape.

The anterior apex of this triangle forms the anterior border of the tibia,
which begins at the tibial tuberosity and runs inferiorly along the length of the tibia. Both the anterior border and the medial side of the triangular shaft
are located immediately under the skin and can be easily palpated along the entire length of the tibia. A small ridge running down the lateral side of the
tibial shaft is the interosseous border of the tibia. This is for the attachment of the interosseous membrane of the leg, the sheet of dense connective
tissue that unites the tibia and fibula bones. Located on the posterior side of the tibia is the soleal line, a diagonally running, roughened ridge that
begins below the base of the lateral condyle, and runs down and medially across the proximal third of the posterior tibia. Muscles of the posterior leg
attach to this line.

The large expansion found on the medial side of the distal tibia is the medial malleolus (“little hammer”). This forms the large bony bump found on the
medial side of the ankle region. Both the smooth surface on the inside of the medial malleolus and the smooth area at the distal end of the tibia articulate
with the talus bone of the foot as part of the ankle joint. On the lateral side of the distal tibia is a wide groove called the fibular notch. This area
articulates with the distal end of the fibula, forming the distal tibiofibular joint.

Fibula

The fibula is the slender bone located on the lateral side of the leg (see Figure 8.18). The fibula does not bear weight. It serves primarily for muscle
attachments and thus is largely surrounded by muscles. Only the proximal and distal ends of the fibula can be palpated.

The head of the fibula is the small, knob-like, proximal end of the fibula. It articulates with the inferior aspect of the lateral tibial condyle, forming
the proximal tibiofibular joint. The thin shaft of the fibula has the interosseous border of the fibula, a narrow ridge running down its medial side for
the attachment of the interosseous membrane that spans the fibula and tibia. The distal end of the fibula forms the lateral malleolus, which forms the
easily palpated bony bump on the lateral side of the ankle. The deep (medial) side of the lateral malleolus articulates with the talus bone of the foot as
part of the ankle joint. The distal fibula also articulates with the fibular notch of the tibia.

Tarsal Bones

The posterior half of the foot is formed by seven tarsal bones (Figure 8.19). The most superior bone is the talus. This has a relatively square-shaped,
upper surface that articulates with the tibia and fibula to form the ankle joint. Three areas of articulation form the ankle joint: The superomedial surface
of the talus bone articulates with the medial malleolus of the tibia, the top of the talus articulates with the distal end of the tibia, and the lateral side of the
talus articulates with the lateral malleolus of the fibula. Inferiorly, the talus articulates with the calcaneus (heel bone), the largest bone of the foot, which
forms the heel. Body weight is transferred from the tibia to the talus to the calcaneus, which rests on the ground. The medial calcaneus has a prominent
bony extension called the sustentaculum tali (“support for the talus”) that supports the medial side of the talus bone.

Figure 8.19 Bones of the Foot The bones of

the foot are divided into three groups. The
posterior foot is formed by the seven tarsal
bones. The mid-foot has the five metatarsal
bones. The toes contain the phalanges.

The cuboid bone articulates with the anterior

end of the calcaneus bone. The cuboid has a
deep groove running across its inferior
surface, which provides passage for a muscle
tendon. The talus bone articulates anteriorly
with the navicular bone, which in turn
articulates anteriorly with the three cuneiform
(“wedge-shaped”) bones. These bones are
the medial cuneiform, the intermediate
cuneiform, and the lateral cuneiform. Each
of these bones has a broad superior surface
and a narrow inferior surface, which together
produce the transverse (medial-lateral)
curvature of the foot. The navicular and lateral
cuneiform bones also articulate with the
medial side of the cuboid bone.

INTERACTIVE LINK

Use this tutorial to review the bones of the

foot. Which tarsal bones are in the proximal, intermediate, and distal groups?

Metatarsal Bones

The anterior half of the foot is formed by the five metatarsal bones, which are located between the tarsal bones of the posterior foot and the phalanges
of the toes (see Figure 8.19). These elongated bones are numbered 1–5, starting with the medial side of the foot. The first metatarsal bone is shorter
and thicker than the others. The second metatarsal is the longest. The base of the metatarsal bone is the proximal end of each metatarsal bone.
These articulate with the cuboid or cuneiform bones. The base of the fifth metatarsal has a large, lateral expansion that provides for muscle attachments.
This expanded base of the fifth metatarsal can be felt as a bony bump at the midpoint along the lateral border of the foot. The expanded distal end of
each metatarsal is the head of the metatarsal bone. Each metatarsal bone articulates with the proximal phalanx of a toe to form
a metatarsophalangeal joint. The heads of the metatarsal bones also rest on the ground and form the ball (anterior end) of the foot.
Phalanges

The toes contain a total of 14 phalanx bones (phalanges), arranged in a similar manner as the phalanges of the fingers (see Figure 8.19). The toes are
numbered 1–5, starting with the big toe (hallux). The big toe has two phalanx bones, the proximal and distal phalanges. The remaining toes all have
proximal, middle, and distal phalanges. A joint between adjacent phalanx bones is called an interphalangeal joint.

Arches of the Foot

When the foot comes into contact with the ground during walking, running, or jumping activities, the impact of the body weight puts a tremendous
amount of pressure and force on the foot. During running, the force applied to each foot as it contacts the ground can be up to 2.5 times your body
weight. The bones, joints, ligaments, and muscles of the foot absorb this force, thus greatly reducing the amount of shock that is passed superiorly into
the lower limb and body. The arches of the foot play an important role in this shock-absorbing ability. When weight is applied to the foot, these arches
will flatten somewhat, thus absorbing energy. When the weight is removed, the arch rebounds, giving “spring” to the step. The arches also serve to
distribute body weight side to side and to either end of the foot.

The foot has a transverse arch, a medial longitudinal arch, and a lateral longitudinal arch (see Figure 8.19). The transverse arch forms the medial-lateral
curvature of the mid-foot. It is formed by the wedge shapes of the cuneiform bones and bases (proximal ends) of the first to fourth metatarsal bones.
This arch helps to distribute body weight from side to side within the foot, thus allowing the foot to accommodate uneven terrain.

The longitudinal arches run down the length of the foot. The lateral longitudinal arch is relatively flat, whereas the medial longitudinal arch is larger
(taller). The longitudinal arches are formed by the tarsal bones posteriorly and the metatarsal bones anteriorly. These arches are supported at either
end, where they contact the ground. Posteriorly, this support is provided by the calcaneus bone and anteriorly by the heads (distal ends) of the
metatarsal bones. The talus bone, which receives the weight of the body, is located at the top of the longitudinal arches. Body weight is then conveyed
from the talus to the ground by the anterior and posterior ends of these arches. Strong ligaments unite the adjacent foot bones to prevent disruption of
the arches during weight bearing. On the bottom of the foot, additional ligaments tie together the anterior and posterior ends of the arches. These
ligaments have elasticity, which allows them to stretch somewhat during weight bearing, thus allowing the longitudinal arches to spread. The stretching
of these ligaments stores energy within the foot, rather than passing these forces into the leg. Contraction of the foot muscles also plays an important
role in this energy absorption. When the weight is removed, the elastic ligaments recoil and pull the ends of the arches closer together. This recovery of
the arches releases the stored energy and improves the energy efficiency of walking.

Stretching of the ligaments that support the longitudinal arches can lead to pain. This can occur in overweight individuals, with people who have jobs that
involve standing for long periods of time (such as a waitress), or walking or running long distances. If stretching of the ligaments is prolonged, excessive,
or repeated, it can result in a gradual lengthening of the supporting ligaments, with subsequent depression or collapse of the longitudinal arches,
particularly on the medial side of the foot. This condition is called pes planus (“flat foot” or “fallen arches”).

Embryologically, the appendicular skeleton arises from mesenchyme, a type of embryonic tissue that can differentiate into many types of tissues,
including bone or muscle tissue. Mesenchyme gives rise to the bones of the upper and lower limbs, as well as to the pectoral and pelvic girdles.
Development of the limbs begins near the end of the fourth embryonic week, with the upper limbs appearing first. Thereafter, the development of the
upper and lower limbs follows similar patterns, with the lower limbs lagging behind the upper limbs by a few days.

Each upper and lower limb initially develops as a small bulge called a limb bud, which appears on the lateral side of the early embryo. The upper limb
bud appears near the end of the fourth week of development, with the lower limb bud appearing shortly after (Figure 8.20).

Figure 8.20 Embryo at Seven Weeks Limb buds are visible in an embryo at the end of the seventh week of development (embryo derived from an
ectopic pregnancy). (credit: Ed Uthman/flickr)

Initially, the limb buds consist of a core of mesenchyme covered by a layer of ectoderm. The ectoderm at the end of the limb bud thickens to form a
narrow crest called the apical ectodermal ridge. This ridge stimulates the underlying mesenchyme to rapidly proliferate, producing the outgrowth of the
developing limb. As the limb bud elongates, cells located farther from the apical ectodermal ridge slow their rates of cell division and begin to
differentiate. In this way, the limb develops along a proximal-to-distal axis.

During the sixth week of development, the distal ends of the upper and lower limb buds expand and flatten into a paddle shape. This region will become
the hand or foot. The wrist or ankle areas then appear as a constriction that develops at the base of the paddle. Shortly after this, a second constriction
on the limb bud appears at the future site of the elbow or knee. Within the paddle, areas of tissue undergo cell death, producing separations between the
growing fingers and toes. Also during the sixth week of development, mesenchyme within the limb buds begins to differentiate into hyaline cartilage that
will form models of the future limb bones.

The early outgrowth of the upper and lower limb buds initially has the limbs positioned so that the regions that will become the palm of the hand or the
bottom of the foot are facing medially toward the body, with the future thumb or big toe both oriented toward the head. During the seventh week of
development, the upper limb rotates laterally by 90 degrees, so that the palm of the hand faces anteriorly and the thumb points laterally. In contrast, the
lower limb undergoes a 90-degree medial rotation, thus bringing the big toe to the medial side of the foot.

Ossification of Appendicular Bones

All of the girdle and limb bones, except for the clavicle, develop by the process of endochondral ossification. This process begins as the mesenchyme
within the limb bud differentiates into hyaline cartilage to form cartilage models for future bones. By the twelfth week, a primary ossification center will
have appeared in the diaphysis (shaft) region of the long bones, initiating the process that converts the cartilage model into bone. A secondary
ossification center will appear in each epiphysis (expanded end) of these bones at a later time, usually after birth. The primary and secondary
ossification centers are separated by the epiphyseal plate, a layer of growing hyaline cartilage. This plate is located between the diaphysis and each
epiphysis. It continues to grow and is responsible for the lengthening of the bone. The epiphyseal plate is retained for many years, until the bone
reaches its final, adult size, at which time the epiphyseal plate disappears and the epiphysis fuses to the diaphysis. (Seek additional content on
ossification in the chapter on bone tissue.)

Small bones, such as the phalanges, will develop only one secondary ossification center and will thus have only a single epiphyseal plate. Large bones,
such as the femur, will develop several secondary ossification centers, with an epiphyseal plate associated with each secondary center. Thus,
ossification of the femur begins at the end of the seventh week with the appearance of the primary ossification center in the diaphysis, which rapidly
expands to ossify the shaft of the bone prior to birth. Secondary ossification centers develop at later times. Ossification of the distal end of the femur, to
form the condyles and epicondyles, begins shortly before birth. Secondary ossification centers also appear in the femoral head late in the first year after
birth, in the greater trochanter during the fourth year, and in the lesser trochanter between the ages of 9 and 10 years. Once these areas have ossified,
their fusion to the diaphysis and the disappearance of each epiphyseal plate follow a reversed sequence. Thus, the lesser trochanter is the first to fuse,
doing so at the onset of puberty (around 11 years of age), followed by the greater trochanter approximately 1 year later. The femoral head fuses
between the ages of 14–17 years, whereas the distal condyles of the femur are the last to fuse, between the ages of 16–19 years. Knowledge of the age
at which different epiphyseal plates disappear is important when interpreting radiographs taken of children. Since the cartilage of an epiphyseal plate is
less dense than bone, the plate will appear dark in a radiograph image. Thus, a normal epiphyseal plate may be mistaken for a bone fracture.

The clavicle is the one appendicular skeleton bone that does not develop via endochondral ossification. Instead, the clavicle develops through the
process of intramembranous ossification. During this process, mesenchymal cells differentiate directly into bone-producing cells, which produce the
clavicle directly, without first making a cartilage model. Because of this early production of bone, the clavicle is the first bone of the body to begin
ossification, with ossification centers appearing during the fifth week of development. However, ossification of the clavicle is not complete until age 25.

DISORDERS OF THE...

Appendicular System: Congenital Clubfoot

Clubfoot, also known as talipes, is a congenital (present at birth) disorder of unknown cause and is the most common deformity of the lower limb. It
affects the foot and ankle, causing the foot to be twisted inward at a sharp angle, like the head of a golf club (Figure 8.21). Clubfoot has a frequency of
about 1 out of every 1,000 births, and is twice as likely to occur in a male child as in a female child. In 50 percent of cases, both feet are affected.

Figure 8.21 Clubfoot Clubfoot is a common deformity of the ankle and foot that is present at birth. Most cases are corrected without surgery, and
affected individuals will grow up to lead normal, active lives. (credit: James W. Hanson)

At birth, children with a clubfoot have the heel turned inward and the anterior foot twisted so that the lateral side of the foot is facing inferiorly, commonly
due to ligaments or leg muscles attached to the foot that are shortened or abnormally tight. These pull the foot into an abnormal position, resulting in
bone deformities. Other symptoms may include bending of the ankle that lifts the heel of the foot and an extremely high foot arch. Due to the limited
range of motion in the affected foot, it is difficult to place the foot into the correct position. Additionally, the affected foot may be shorter than normal, and
the calf muscles are usually underdeveloped on the affected side. Despite the appearance, this is not a painful condition for newborns. However, it must
be treated early to avoid future pain and impaired walking ability.

Although the cause of clubfoot is idiopathic (unknown), evidence indicates that fetal position within the uterus is not a contributing factor. Genetic factors
are involved, because clubfoot tends to run within families. Cigarette smoking during pregnancy has been linked to the development of clubfoot,
particularly in families with a history of clubfoot.

Previously, clubfoot required extensive surgery. Today, 90 percent of cases are successfully treated without surgery using new corrective casting
techniques. The best chance for a full recovery requires that clubfoot treatment begin during the first 2 weeks after birth. Corrective casting gently
stretches the foot, which is followed by the application of a holding cast to keep the foot in the proper position. This stretching and casting is repeated
weekly for several weeks. In severe cases, surgery may also be required, after which the foot typically remains in a cast for 6 to 8 weeks. After the cast
is removed following either surgical or nonsurgical treatment, the child will be required to wear a brace part-time (at night) for up to 4 years. In addition,
special exercises will be prescribed, and the child must also wear special shoes. Close monitoring by the parents and adherence to postoperative
instructions are imperative in minimizing the risk of relapse.

Despite these difficulties, treatment for clubfoot is usually successful, and the child will grow up to lead a normal, active life. Numerous examples of
individuals born with a clubfoot who went on to successful careers include Dudley Moore (comedian and actor), Damon Wayans (comedian and actor),
Troy Aikman (three-time Super Bowl-winning quarterback), Kristi Yamaguchi (Olympic gold medalist in figure skating), Mia Hamm (two-time Olympic
gold medalist in soccer), and Charles Woodson (Heisman trophy and Super Bowl winner).

acetabulum
large, cup-shaped cavity located on the lateral side of the hip bone; formed by the junction of the ilium, pubis, and ischium portions of the hip
bone
acromial end of the clavicle
lateral end of the clavicle that articulates with the acromion of the scapula
acromial process
acromion of the scapula
acromioclavicular joint
articulation between the acromion of the scapula and the acromial end of the clavicle
acromion
flattened bony process that extends laterally from the scapular spine to form the bony tip of the shoulder
adductor tubercle
small, bony bump located on the superior aspect of the medial epicondyle of the femur
anatomical neck
line on the humerus located around the outside margin of the humeral head
ankle joint
joint that separates the leg and foot portions of the lower limb; formed by the articulations between the talus bone of the foot inferiorly, and the
distal end of the tibia, medial malleolus of the tibia, and lateral malleolus of the fibula superiorly
anterior border of the tibia
narrow, anterior margin of the tibia that extends inferiorly from the tibial tuberosity
anterior inferior iliac spine
small, bony projection located on the anterior margin of the ilium, below the anterior superior iliac spine
anterior sacroiliac ligament
strong ligament between the sacrum and the ilium portions of the hip bone that supports the anterior side of the sacroiliac joint
anterior superior iliac spine
rounded, anterior end of the iliac crest
apical ectodermal ridge
enlarged ridge of ectoderm at the distal end of a limb bud that stimulates growth and elongation of the limb
arcuate line of the ilium
smooth ridge located at the inferior margin of the iliac fossa; forms the lateral portion of the pelvic brim
arm
region of the upper limb located between the shoulder and elbow joints; contains the humerus bone
auricular surface of the ilium
roughened area located on the posterior, medial side of the ilium of the hip bone; articulates with the auricular surface of the sacrum to form
the sacroiliac joint
base of the metatarsal bone
expanded, proximal end of each metatarsal bone
bicipital groove
intertubercular groove; narrow groove located between the greater and lesser tubercles of the humerus
calcaneus
heel bone; posterior, inferior tarsal bone that forms the heel of the foot
capitate
from the lateral side, the third of the four distal carpal bones; articulates with the scaphoid and lunate proximally, the trapezoid laterally, the
hamate medially, and primarily with the third metacarpal distally
capitulum
knob-like bony structure located anteriorly on the lateral, distal end of the humerus
carpal bone
one of the eight small bones that form the wrist and base of the hand; these are grouped as a proximal row consisting of (from lateral to
medial) the scaphoid, lunate, triquetrum, and pisiform bones, and a distal row containing (from lateral to medial) the trapezium, trapezoid,
capitate, and hamate bones
carpal tunnel
passageway between the anterior forearm and hand formed by the carpal bones and flexor retinaculum
carpometacarpal joint
articulation between one of the carpal bones in the distal row and a metacarpal bone of the hand
clavicle
collarbone; elongated bone that articulates with the manubrium of the sternum medially and the acromion of the scapula laterally
coracoclavicular ligament
strong band of connective tissue that anchors the coracoid process of the scapula to the lateral clavicle; provides important indirect support for
the acromioclavicular joint
coracoid process
short, hook-like process that projects anteriorly and laterally from the superior margin of the scapula
coronoid fossa
depression on the anterior surface of the humerus above the trochlea; this space receives the coronoid process of the ulna when the elbow is
maximally flexed
coronoid process of the ulna
projecting bony lip located on the anterior, proximal ulna; forms the inferior margin of the trochlear notch
costoclavicular ligament
band of connective tissue that unites the medial clavicle with the first rib
coxal bone
hip bone
cuboid
tarsal bone that articulates posteriorly with the calcaneus bone, medially with the lateral cuneiform bone, and anteriorly with the fourth and fifth
metatarsal bones
deltoid tuberosity
roughened, V-shaped region located laterally on the mid-shaft of the humerus
distal radioulnar joint
articulation between the head of the ulna and the ulnar notch of the radius
distal tibiofibular joint
articulation between the distal fibula and the fibular notch of the tibia
elbow joint
joint located between the upper arm and forearm regions of the upper limb; formed by the articulations between the trochlea of the humerus
and the trochlear notch of the ulna, and the capitulum of the humerus and the head of the radius
femur
thigh bone; the single bone of the thigh
fibula
thin, non-weight-bearing bone found on the lateral side of the leg
fibular notch
wide groove on the lateral side of the distal tibia for articulation with the fibula at the distal tibiofibular joint
flexor retinaculum
 strong band of connective tissue at the anterior wrist that spans the top of the U-shaped grouping of the carpal bones to form the roof of the
carpal tunnel
foot
portion of the lower limb located distal to the ankle joint
forearm
region of the upper limb located between the elbow and wrist joints; contains the radius and ulna bones
fossa
(plural = fossae) shallow depression on the surface of a bone
fovea capitis
minor indentation on the head of the femur that serves as the site of attachment for the ligament to the head of the femur
glenohumeral joint
shoulder joint; formed by the articulation between the glenoid cavity of the scapula and the head of the humerus
glenoid cavity
(also, glenoid fossa) shallow depression located on the lateral scapula, between the superior and lateral borders
gluteal tuberosity
roughened area on the posterior side of the proximal femur, extending inferiorly from the base of the greater trochanter
greater pelvis
(also, greater pelvic cavity or false pelvis) broad space above the pelvic brim defined laterally by the fan-like portion of the upper ilium
greater sciatic foramen
pelvic opening formed by the greater sciatic notch of the hip bone, the sacrum, and the sacrospinous ligament
greater sciatic notch
large, U-shaped indentation located on the posterior margin of the ilium, superior to the ischial spine
greater trochanter
large, bony expansion of the femur that projects superiorly from the base of the femoral neck
greater tubercle
enlarged prominence located on the lateral side of the proximal humerus
hallux
big toe; digit 1 of the foot
hamate
from the lateral side, the fourth of the four distal carpal bones; articulates with the lunate and triquetrum proximally, the fourth and fifth
metacarpals distally, and the capitate laterally
hand
region of the upper limb distal to the wrist joint
head of the femur
rounded, proximal end of the femur that articulates with the acetabulum of the hip bone to form the hip joint
head of the fibula
small, knob-like, proximal end of the fibula; articulates with the inferior aspect of the lateral condyle of the tibia
head of the humerus
smooth, rounded region on the medial side of the proximal humerus; articulates with the glenoid fossa of the scapula to form the glenohumeral
(shoulder) joint
head of the metatarsal bone
expanded, distal end of each metatarsal bone
head of the radius
disc-shaped structure that forms the proximal end of the radius; articulates with the capitulum of the humerus as part of the elbow joint, and
with the radial notch of the ulna as part of the proximal radioulnar joint
head of the ulna
small, rounded distal end of the ulna; articulates with the ulnar notch of the distal radius, forming the distal radioulnar joint
hip bone
coxal bone; single bone that forms the pelvic girdle; consists of three areas, the ilium, ischium, and pubis
hip joint
joint located at the proximal end of the lower limb; formed by the articulation between the acetabulum of the hip bone and the head of the
femur
hook of the hamate bone
bony extension located on the anterior side of the hamate carpal bone
humerus
single bone of the upper arm
iliac crest
curved, superior margin of the ilium
iliac fossa
shallow depression found on the anterior and medial surfaces of the upper ilium
ilium
superior portion of the hip bone
inferior angle of the scapula
inferior corner of the scapula located where the medial and lateral borders meet
inferior pubic ramus
narrow segment of bone that passes inferiorly and laterally from the pubic body; joins with the ischial ramus to form the ischiopubic ramus
infraglenoid tubercle
small bump or roughened area located on the lateral border of the scapula, near the inferior margin of the glenoid cavity
infraspinous fossa
broad depression located on the posterior scapula, inferior to the spine
intercondylar eminence
irregular elevation on the superior end of the tibia, between the articulating surfaces of the medial and lateral condyles
intercondylar fossa
deep depression on the posterior side of the distal femur that separates the medial and lateral condyles
intermediate cuneiform
middle of the three cuneiform tarsal bones; articulates posteriorly with the navicular bone, medially with the medial cuneiform bone, laterally
with the lateral cuneiform bone, and anteriorly with the second metatarsal bone
interosseous border of the fibula
 small ridge running down the medial side of the fibular shaft; for attachment of the interosseous membrane between the fibula and tibia
interosseous border of the radius
narrow ridge located on the medial side of the radial shaft; for attachment of the interosseous membrane between the ulna and radius bones
interosseous border of the tibia
small ridge running down the lateral side of the tibial shaft; for attachment of the interosseous membrane between the tibia and fibula
interosseous border of the ulna
narrow ridge located on the lateral side of the ulnar shaft; for attachment of the interosseous membrane between the ulna and radius
interosseous membrane of the forearm
sheet of dense connective tissue that unites the radius and ulna bones
interosseous membrane of the leg
sheet of dense connective tissue that unites the shafts of the tibia and fibula bones
interphalangeal joint
articulation between adjacent phalanx bones of the hand or foot digits
intertrochanteric crest
short, prominent ridge running between the greater and lesser trochanters on the posterior side of the proximal femur
intertrochanteric line
small ridge running between the greater and lesser trochanters on the anterior side of the proximal femur
intertubercular groove (sulcus)
bicipital groove; narrow groove located between the greater and lesser tubercles of the humerus
ischial ramus
bony extension projecting anteriorly and superiorly from the ischial tuberosity; joins with the inferior pubic ramus to form the ischiopubic ramus
ischial spine
pointed, bony projection from the posterior margin of the ischium that separates the greater sciatic notch and lesser sciatic notch
ischial tuberosity
large, roughened protuberance that forms the posteroinferior portion of the hip bone; weight-bearing region of the pelvis when sitting
ischiopubic ramus
narrow extension of bone that connects the ischial tuberosity to the pubic body; formed by the junction of the ischial ramus and inferior pubic
ramus
ischium
posteroinferior portion of the hip bone
knee joint
joint that separates the thigh and leg portions of the lower limb; formed by the articulations between the medial and lateral condyles of the
femur, and the medial and lateral condyles of the tibia
lateral border of the scapula
diagonally oriented lateral margin of the scapula
lateral condyle of the femur
smooth, articulating surface that forms the distal and posterior sides of the lateral expansion of the distal femur
lateral condyle of the tibia
lateral, expanded region of the proximal tibia that includes the smooth surface that articulates with the lateral condyle of the femur as part of
the knee joint
lateral cuneiform
most lateral of the three cuneiform tarsal bones; articulates posteriorly with the navicular bone, medially with the intermediate cuneiform bone,
laterally with the cuboid bone, and anteriorly with the third metatarsal bone
lateral epicondyle of the femur
roughened area of the femur located on the lateral side of the lateral condyle
lateral epicondyle of the humerus
small projection located on the lateral side of the distal humerus
lateral malleolus
expanded distal end of the fibula
lateral supracondylar ridge
narrow, bony ridge located along the lateral side of the distal humerus, superior to the lateral epicondyle
leg
portion of the lower limb located between the knee and ankle joints
lesser pelvis
(also, lesser pelvic cavity or true pelvis) narrow space located within the pelvis, defined superiorly by the pelvic brim (pelvic inlet) and inferiorly
by the pelvic outlet
lesser sciatic foramen
pelvic opening formed by the lesser sciatic notch of the hip bone, the sacrospinous ligament, and the sacrotuberous ligament
lesser sciatic notch
shallow indentation along the posterior margin of the ischium, inferior to the ischial spine
lesser trochanter
small, bony projection on the medial side of the proximal femur, at the base of the femoral neck
lesser tubercle
small, bony prominence located on anterior side of the proximal humerus
ligament of the head of the femur
ligament that spans the acetabulum of the hip bone and the fovea capitis of the femoral head
limb bud
small elevation that appears on the lateral side of the embryo during the fourth or fifth week of development, which gives rise to an upper or
lower limb
linea aspera
longitudinally running bony ridge located in the middle third of the posterior femur
lunate
from the lateral side, the second of the four proximal carpal bones; articulates with the radius proximally, the capitate and hamate distally, the
scaphoid laterally, and the triquetrum medially
medial border of the scapula
elongated, medial margin of the scapula
medial condyle of the femur
smooth, articulating surface that forms the distal and posterior sides of the medial expansion of the distal femur
medial condyle of the tibia
medial, expanded region of the proximal tibia that includes the smooth surface that articulates with the medial condyle of the femur as part of
the knee joint
medial cuneiform
most medial of the three cuneiform tarsal bones; articulates posteriorly with the navicular bone, laterally with the intermediate cuneiform bone,
and anteriorly with the first and second metatarsal bones
medial epicondyle of the femur
roughened area of the distal femur located on the medial side of the medial condyle
medial epicondyle of the humerus
enlarged projection located on the medial side of the distal humerus
medial malleolus
bony expansion located on the medial side of the distal tibia
metacarpal bone
one of the five long bones that form the palm of the hand; numbered 1–5, starting on the lateral (thumb) side of the hand
metacarpophalangeal joint
articulation between the distal end of a metacarpal bone of the hand and a proximal phalanx bone of the thumb or a finger
metatarsal bone
one of the five elongated bones that forms the anterior half of the foot; numbered 1–5, starting on the medial side of the foot
metatarsophalangeal joint
articulation between a metatarsal bone of the foot and the proximal phalanx bone of a toe
midcarpal joint
articulation between the proximal and distal rows of the carpal bones; contributes to movements of the hand at the wrist
navicular
tarsal bone that articulates posteriorly with the talus bone, laterally with the cuboid bone, and anteriorly with the medial, intermediate, and
lateral cuneiform bones
neck of the femur
narrowed region located inferior to the head of the femur
neck of the radius
narrowed region immediately distal to the head of the radius
obturator foramen
large opening located in the anterior hip bone, between the pubis and ischium regions
olecranon fossa
large depression located on the posterior side of the distal humerus; this space receives the olecranon process of the ulna when the elbow is
fully extended
olecranon process
expanded posterior and superior portions of the proximal ulna; forms the bony tip of the elbow
patella
kneecap; the largest sesamoid bone of the body; articulates with the distal femur
patellar surface
smooth groove located on the anterior side of the distal femur, between the medial and lateral condyles; site of articulation for the patella
pectineal line
narrow ridge located on the superior surface of the superior pubic ramus
pectoral girdle
shoulder girdle; the set of bones, consisting of the scapula and clavicle, which attaches each upper limb to the axial skeleton
pelvic brim
pelvic inlet; the dividing line between the greater and lesser pelvic regions; formed by the superior margin of the pubic symphysis, the
pectineal lines of each pubis, the arcuate lines of each ilium, and the sacral promontory
pelvic girdle
hip girdle; consists of a single hip bone, which attaches a lower limb to the sacrum of the axial skeleton
pelvic inlet
pelvic brim
pelvic outlet
inferior opening of the lesser pelvis; formed by the inferior margin of the pubic symphysis, right and left ischiopubic rami and sacrotuberous
ligaments, and the tip of the coccyx
pelvis
ring of bone consisting of the right and left hip bones, the sacrum, and the coccyx
phalanx bone of the foot
(plural = phalanges) one of the 14 bones that form the toes; these include the proximal and distal phalanges of the big toe, and the proximal,
middle, and distal phalanx bones of toes two through five
phalanx bone of the hand
(plural = phalanges) one of the 14 bones that form the thumb and fingers; these include the proximal and distal phalanges of the thumb, and
the proximal, middle, and distal phalanx bones of the fingers two through five
pisiform
from the lateral side, the fourth of the four proximal carpal bones; articulates with the anterior surface of the triquetrum
pollex
(also, thumb) digit 1 of the hand
posterior inferior iliac spine
small, bony projection located at the inferior margin of the auricular surface on the posterior ilium
posterior sacroiliac ligament
strong ligament spanning the sacrum and ilium of the hip bone that supports the posterior side of the sacroiliac joint
posterior superior iliac spine
rounded, posterior end of the iliac crest
proximal radioulnar joint
articulation formed by the radial notch of the ulna and the head of the radius
proximal tibiofibular joint
articulation between the head of the fibula and the inferior aspect of the lateral condyle of the tibia
pubic arch
bony structure formed by the pubic symphysis, and the bodies and inferior pubic rami of the right and left pubic bones
pubic body
enlarged, medial portion of the pubis region of the hip bone
pubic symphysis
joint formed by the articulation between the pubic bodies of the right and left hip bones
pubic tubercle
small bump located on the superior aspect of the pubic body
pubis
anterior portion of the hip bone
radial fossa
small depression located on the anterior humerus above the capitulum; this space receives the head of the radius when the elbow is
maximally flexed
radial notch of the ulna
small, smooth area on the lateral side of the proximal ulna; articulates with the head of the radius as part of the proximal radioulnar joint
radial tuberosity
oval-shaped, roughened protuberance located on the medial side of the proximal radius
radiocarpal joint
wrist joint, located between the forearm and hand regions of the upper limb; articulation formed proximally by the distal end of the radius and
the fibrocartilaginous pad that unites the distal radius and ulna bone, and distally by the scaphoid, lunate, and triquetrum carpal bones
radius
bone located on the lateral side of the forearm
sacroiliac joint
joint formed by the articulation between the auricular surfaces of the sacrum and ilium
sacrospinous ligament
ligament that spans the sacrum to the ischial spine of the hip bone
sacrotuberous ligament
ligament that spans the sacrum to the ischial tuberosity of the hip bone
scaphoid
from the lateral side, the first of the four proximal carpal bones; articulates with the radius proximally, the trapezoid, trapezium, and capitate
distally, and the lunate medially
scapula
shoulder blade bone located on the posterior side of the shoulder
shaft of the femur
cylindrically shaped region that forms the central portion of the femur
shaft of the fibula
elongated, slender portion located between the expanded ends of the fibula
shaft of the humerus
narrow, elongated, central region of the humerus
shaft of the radius
narrow, elongated, central region of the radius
shaft of the tibia
triangular-shaped, central portion of the tibia
shaft of the ulna
narrow, elongated, central region of the ulna
soleal line
small, diagonally running ridge located on the posterior side of the proximal tibia
spine of the scapula
prominent ridge passing mediolaterally across the upper portion of the posterior scapular surface
sternal end of the clavicle
medial end of the clavicle that articulates with the manubrium of the sternum
sternoclavicular joint
articulation between the manubrium of the sternum and the sternal end of the clavicle; forms the only bony attachment between the pectoral
girdle of the upper limb and the axial skeleton
styloid process of the radius
pointed projection located on the lateral end of the distal radius
styloid process of the ulna
short, bony projection located on the medial end of the distal ulna
subpubic angle
inverted V-shape formed by the convergence of the right and left ischiopubic rami; this angle is greater than 80 degrees in females and less
than 70 degrees in males
subscapular fossa
broad depression located on the anterior (deep) surface of the scapula
superior angle of the scapula
corner of the scapula between the superior and medial borders of the scapula
superior border of the scapula
superior margin of the scapula
superior pubic ramus
narrow segment of bone that passes laterally from the pubic body to join the ilium
supraglenoid tubercle
small bump located at the superior margin of the glenoid cavity
suprascapular notch
small notch located along the superior border of the scapula, medial to the coracoid process
supraspinous fossa
narrow depression located on the posterior scapula, superior to the spine
surgical neck
region of the humerus where the expanded, proximal end joins with the narrower shaft
sustentaculum tali
bony ledge extending from the medial side of the calcaneus bone
talus
 tarsal bone that articulates superiorly with the tibia and fibula at the ankle joint; also articulates inferiorly with the calcaneus bone and anteriorly
with the navicular bone
tarsal bone
one of the seven bones that make up the posterior foot; includes the calcaneus, talus, navicular, cuboid, medial cuneiform, intermediate
cuneiform, and lateral cuneiform bones
thigh
portion of the lower limb located between the hip and knee joints
tibia
shin bone; the large, weight-bearing bone located on the medial side of the leg
tibial tuberosity
elevated area on the anterior surface of the proximal tibia
trapezium
from the lateral side, the first of the four distal carpal bones; articulates with the scaphoid proximally, the first and second metacarpals distally,
and the trapezoid medially
trapezoid
from the lateral side, the second of the four distal carpal bones; articulates with the scaphoid proximally, the second metacarpal distally, the
trapezium laterally, and the capitate medially
triquetrum
from the lateral side, the third of the four proximal carpal bones; articulates with the lunate laterally, the hamate distally, and has a facet for the
pisiform
trochlea - pulley-shaped region located medially at the distal end of the humerus; articulates at the elbow with the trochlear notch of the ulna
trochlear notch - large, C-shaped depression located on the anterior side of the proximal ulna; articulates at the elbow with the trochlea of the humerus
ulna - bone located on the medial side of the forearm
ulnar notch of the radius - shallow, smooth area located on the medial side of the distal radius; articulates with the head of the ulna at the distal
radioulnar joint
ulnar tuberosity -roughened area located on the anterior, proximal ulna inferior to the coronoid process

CHAPTER 9

A joint, also called an articulation, is any place where adjacent bones or bone and cartilage come together (articulate with each other) to form a
connection. Joints are classified both structurally and functionally. Structural classifications of joints take into account whether the adjacent bones are
strongly anchored to each other by fibrous connective tissue or cartilage, or whether the adjacent bones articulate with each other within a fluid-filled
space called a joint cavity. Functional classifications describe the degree of movement available between the bones, ranging from immobile, to slightly
mobile, to freely moveable joints. The amount of movement available at a particular joint of the body is related to the functional requirements for that
joint. Thus immobile or slightly moveable joints serve to protect internal organs, give stability to the body, and allow for limited body movement. In
contrast, freely moveable joints allow for much more extensive movements of the body and limbs.

Structural Classification of Joints

The structural classification of joints is based on whether the articulating surfaces of the adjacent bones are directly connected by fibrous connective tissue or
cartilage, or whether the articulating surfaces contact each other within a fluid-filled joint cavity. These differences serve to divide the joints of the body into three
structural classifications. A fibrous joint is where the adjacent bones are united by fibrous connective tissue. At a cartilaginous joint, the bones are joined by hyaline
cartilage or fibrocartilage. At a synovial joint, the articulating surfaces of the bones are not directly connected, but instead come into contact with each other within a
joint cavity that is filled with a lubricating fluid. Synovial joints allow for free movement between the bones and are the most common joints of the body.

Functional Classification of Joints

The functional classification of joints is determined by the amount of mobility found between the adjacent bones. Joints are thus functionally classified as a
synarthrosis or immobile joint, an amphiarthrosis or slightly moveable joint, or as a diarthrosis, which is a freely moveable joint (arthroun = “to fasten by a joint”).
Depending on their location, fibrous joints may be functionally classified as a synarthrosis (immobile joint) or an amphiarthrosis (slightly mobile joint). Cartilaginous
joints are also functionally classified as either a synarthrosis or an amphiarthrosis joint. All synovial joints are functionally classified as a diarthrosis joint.

Synarthrosis

An immobile or nearly immobile joint is called a synarthrosis. The immobile nature of these joints provide for a strong union between the articulating bones. This is
important at locations where the bones provide protection for internal organs. Examples include sutures, the fibrous joints between the bones of the skull that
surround and protect the brain (Figure 9.2), and the manubriosternal joint, the cartilaginous joint that unites the manubrium and body of the sternum for protection
of the heart.

Figure 9.2 Suture Joints of Skull The suture joints of the skull are an example of a synarthrosis, an immobile or
essentially immobile joint.

Amphiarthrosis

An amphiarthrosis is a joint that has limited mobility. An example of this type of joint is the cartilaginous joint
that unites the bodies of adjacent vertebrae. Filling the gap between the vertebrae is a thick pad of
fibrocartilage called an intervertebral disc (Figure 9.3). Each intervertebral disc strongly unites the vertebrae but
still allows for a limited amount of movement between them. However, the small movements available between adjacent vertebrae can sum together along the
length of the vertebral column to provide for large ranges of body movements.

Another example of an amphiarthrosis is the pubic symphysis of the pelvis. This is a cartilaginous joint in which the pubic regions of the right and left hip bones are
strongly anchored to each other by fibrocartilage. This joint normally has very little mobility. The strength of the pubic symphysis is important in conferring weight-
bearing stability to the pelvis.

Figure 9.3 Intervertebral Disc An intervertebral disc unites the bodies of adjacent vertebrae within the vertebral column. Each disc allows for limited movement
between the vertebrae and thus functionally forms an amphiarthrosis type of joint. Intervertebral discs are made of fibrocartilage and thereby structurally form a
symphysis type of cartilaginous joint.

Diarthrosis

A freely mobile joint is classified as a diarthrosis. These types of joints include all synovial joints of the body, which provide the majority of body movements. Most
diarthrotic joints are found in the appendicular skeleton and thus give the limbs a wide range of motion. These joints are divided into three categories, based on the
number of axes of motion provided by each. An axis in anatomy is described as the movements in reference to the three anatomical planes: transverse, frontal, and
sagittal. Thus, diarthroses are classified as uniaxial (for movement in one plane), biaxial (for movement in two planes), or multiaxial joints (for movement in all three
anatomical planes).

A uniaxial joint only allows for a motion in a single plane (around a single axis). The elbow joint, which only allows for bending or straightening, is an example of a
uniaxial joint. A biaxial joint allows for motions within two planes. An example of a biaxial joint is a metacarpophalangeal joint (knuckle joint) of the hand. The joint
allows for movement along one axis to produce bending or straightening of the finger, and movement along a second axis, which allows for spreading of the fingers
away from each other and bringing them together. A joint that allows for the several directions of movement is called a multiaxial joint (polyaxial or triaxial joint).
This type of diarthrotic joint allows for movement along three axes (Figure 9.4). The shoulder and hip joints are multiaxial joints. They allow the upper or lower limb
to move in an anterior-posterior direction and a medial-lateral direction. In addition, the limb can also be rotated around its long axis. This third movement results in
rotation of the limb so that its anterior surface is moved either toward or away from the midline of the body.

Figure 9.4 Multiaxial Joint A multiaxial joint, such as the hip joint, allows for three types of
movement: anterior-posterior, medial-lateral, and rotational.

At a fibrous joint, the adjacent bones are directly connected to each other by fibrous
connective tissue, and thus the bones do not have a joint cavity between them (Figure 9.5).
The gap between the bones may be narrow or wide. There are three types of fibrous joints.
A suture is the narrow fibrous joint found between most bones of the skull. At a syndesmosis
joint, the bones are more widely separated but are held together by a narrow band of fibrous
connective tissue called a ligament or a wide sheet of connective tissue called an
interosseous membrane. This type of fibrous joint is found between the shaft regions of the
long bones in the forearm and in the leg. Lastly, a gomphosis is the narrow fibrous joint
between the roots of a tooth and the bony socket in the jaw into which the tooth fits.

Figure 9.5 Fibrous Joints Fibrous joints form strong

connections between bones. (a) Sutures join most bones of the
skull. (b) An interosseous membrane forms a syndesmosis
between the radius and ulna bones of the forearm. (c) A
gomphosis is a specialized fibrous joint that anchors a tooth to its
socket in the jaw.

Suture

All the bones of the skull, except for the mandible, are joined to
each other by a fibrous joint called a suture. The fibrous
connective tissue found at a suture (“to bind or sew”) strongly
unites the adjacent skull bones and thus helps to protect the brain and form the face. In adults, the skull bones are closely opposed and fibrous
connective tissue fills the narrow gap between the bones. The suture is frequently convoluted, forming a tight union that prevents most movement
between the bones. (See Figure 9.5a.) Thus, skull sutures are functionally classified as a synarthrosis, although some sutures may allow for slight
movements between the cranial bones.

In newborns and infants, the areas of connective tissue between the bones are much wider, especially in those areas on the top and sides of the skull
that will become the sagittal, coronal, squamous, and lambdoid sutures. These broad areas of connective tissue are called fontanelles (Figure 9.6).
During birth, the fontanelles provide flexibility to the skull, allowing the bones to push closer together or to overlap slightly, thus aiding movement of the
infant’s head through the birth canal. After birth, these expanded regions of connective tissue allow for rapid growth of the skull and enlargement of the
brain. The fontanelles greatly decrease in width during the first year after birth as the skull bones enlarge. When the connective tissue between the
adjacent bones is reduced to a narrow layer, these fibrous joints are now called sutures. At some sutures, the connective tissue will ossify and be
converted into bone, causing the adjacent bones to fuse to each other. This fusion between bones is called a synostosis (“joined by bone”). Examples
of synostosis fusions between cranial bones are found both early and late in life. At the time of birth, the frontal and maxillary bones consist of right and
left halves joined together by sutures, which disappear by the eighth year as the halves fuse together to form a single bone. Late in life, the sagittal,
coronal, and lambdoid sutures of the skull will begin to ossify and fuse, causing the suture line to gradually disappear.

Figure 9.6 The Newborn Skull The fontanelles of a newborn’s skull are broad areas of

fibrous connective tissue that form fibrous joints between the bones of the skull.

Syndesmosis

A syndesmosis (“fastened with a band”) is a type of fibrous joint in which two parallel

bones are united to each other by fibrous connective tissue. The gap between the bones
may be narrow, with the bones joined by ligaments, or the gap may be wide and filled in by
a broad sheet of connective tissue called an interosseous membrane.

In the forearm, the wide gap between the shaft portions of the radius and ulna bones are
strongly united by an interosseous membrane (see Figure 9.5b). Similarly, in the leg, the
shafts of the tibia and fibula are also united by an interosseous membrane. In addition, at the distal tibiofibular joint, the articulating surfaces of the bones
lack cartilage and the narrow gap between the bones is anchored by fibrous connective tissue and ligaments on both the anterior and posterior aspects
of the joint. Together, the interosseous membrane and these ligaments form the tibiofibular syndesmosis.

The syndesmoses found in the forearm and leg serve to unite parallel bones and prevent their separation. However, a syndesmosis does not prevent all
movement between the bones, and thus this type of fibrous joint is functionally classified as an amphiarthrosis. In the leg, the syndesmosis between the
tibia and fibula strongly unites the bones, allows for little movement, and firmly locks the talus bone in place between the tibia and fibula at the ankle
joint. This provides strength and stability to the leg and ankle, which are important during weight bearing. In the forearm, the interosseous membrane is
flexible enough to allow for rotation of the radius bone during forearm movements. Thus in contrast to the stability provided by the tibiofibular
syndesmosis, the flexibility of the antebrachial interosseous membrane allows for the much greater mobility of the forearm.

The interosseous membranes of the leg and forearm also provide areas for muscle attachment. Damage to a syndesmotic joint, which usually results
from a fracture of the bone with an accompanying tear of the interosseous membrane, will produce pain, loss of stability of the bones, and may damage
the muscles attached to the interosseous membrane. If the fracture site is not properly immobilized with a cast or splint, contractile activity by these
muscles can cause improper alignment of the broken bones during healing.

Gomphosis

A gomphosis (“fastened with bolts”) is the specialized fibrous joint that anchors the root of a tooth into its bony socket within the maxillary bone (upper
jaw) or mandible bone (lower jaw) of the skull. A gomphosis is also known as a peg-and-socket joint. Spanning between the bony walls of the socket and
the root of the tooth are numerous short bands of dense connective tissue, each of which is called a periodontal ligament (see Figure 9.5c). Due to the
immobility of a gomphosis, this type of joint is functionally classified as a synarthrosis.

As the name indicates, at a cartilaginous joint, the adjacent bones are united by cartilage, a tough but flexible type of connective tissue. These types of
joints lack a joint cavity and involve bones that are joined together by either hyaline cartilage or fibrocartilage (Figure 9.7). There are two types of
cartilaginous joints. A synchondrosis is a cartilaginous joint where the bones are joined by hyaline cartilage. Also classified as a synchondrosis are
places where bone is united to a cartilage structure, such as between the anterior end of a rib and the costal cartilage of the thoracic cage. The second
type of cartilaginous joint is a symphysis, where the bones are joined by fibrocartilage.

Figure 9.7 Cartilaginous Joints At cartilaginous joints, bones are

united by hyaline cartilage to form a synchondrosis or by fibrocartilage
to form a symphysis. (a) The hyaline cartilage of the epiphyseal plate
(growth plate) forms a synchondrosis that unites the shaft (diaphysis)
and end (epiphysis) of a long bone and allows the bone to grow in
length. (b) The pubic portions of the right and left hip bones of the pelvis
are joined together by fibrocartilage, forming the pubic symphysis.

Synchondrosis

A synchondrosis (“joined by cartilage”) is a cartilaginous joint where

bones are joined together by hyaline cartilage, or where bone is united
to hyaline cartilage. A synchondrosis may be temporary or permanent. A temporary synchondrosis is the epiphyseal plate (growth plate) of a growing
long bone. The epiphyseal plate is the region of growing hyaline cartilage that unites the diaphysis (shaft) of the bone to the epiphysis (end of the bone).
Bone lengthening involves growth of the epiphyseal plate cartilage and its replacement by bone, which adds to the diaphysis. For many years during
childhood growth, the rates of cartilage growth and bone formation are equal and thus the epiphyseal plate does not change in overall thickness as the
bone lengthens. During the late teens and early 20s, growth of the cartilage slows and eventually stops. The epiphyseal plate is then completely
replaced by bone, and the diaphysis and epiphysis portions of the bone fuse together to form a single adult bone. This fusion of the diaphysis and
epiphysis is a synostosis. Once this occurs, bone lengthening ceases. For this reason, the epiphyseal plate is considered to be a temporary
synchondrosis. Because cartilage is softer than bone tissue, injury to a growing long bone can damage the epiphyseal plate cartilage, thus stopping
bone growth and preventing additional bone lengthening.
Growing layers of cartilage also form synchondroses that join together the ilium, ischium, and pubic portions of the hip bone during childhood and
adolescence. When body growth stops, the cartilage disappears and is replaced by bone, forming synostoses and fusing the bony components together
into the single hip bone of the adult. Similarly, synostoses unite the sacral vertebrae that fuse together to form the adult sacrum.

Examples of permanent synchondroses are found in the thoracic cage. One example is the first sternocostal joint, where the first rib is anchored to the
manubrium by its costal cartilage. (The articulations of the remaining costal cartilages to the sternum are all synovial joints.) Additional synchondroses
are formed where the anterior end of the other 11 ribs is joined to its costal cartilage. Unlike the temporary synchondroses of the epiphyseal plate, these
permanent synchondroses retain their hyaline cartilage and thus do not ossify with age. Due to the lack of movement between the bone and cartilage,
both temporary and permanent synchondroses are functionally classified as a synarthrosis.

Symphysis

A cartilaginous joint where the bones are joined by fibrocartilage is called a symphysis (“growing together”). Fibrocartilage is very strong because it
contains numerous bundles of thick collagen fibers, thus giving it a much greater ability to resist pulling and bending forces when compared with hyaline
cartilage. This gives symphyses the ability to strongly unite the adjacent bones, but can still allow for limited movement to occur. Thus, a symphysis is
functionally classified as an amphiarthrosis.

The gap separating the bones at a symphysis may be narrow or wide. Examples in which the gap between the bones is narrow include the pubic
symphysis and the manubriosternal joint. At the pubic symphysis, the pubic portions of the right and left hip bones of the pelvis are joined together by
fibrocartilage across a narrow gap. Similarly, at the manubriosternal joint, fibrocartilage unites the manubrium and body portions of the sternum.

The intervertebral symphysis is a wide symphysis located between the bodies of adjacent vertebrae of the vertebral column. Here a thick pad of
fibrocartilage called an intervertebral disc strongly unites the adjacent vertebrae by filling the gap between them. The width of the intervertebral
symphysis is important because it allows for small movements between the adjacent vertebrae. In addition, the thick intervertebral disc provides
cushioning between the vertebrae, which is important when carrying heavy objects or during high-impact activities such as running or jumping.

Synovial joints are the most common type of joint in the body (Figure 9.8). A key structural characteristic for a synovial joint that is not seen at fibrous or
cartilaginous joints is the presence of a joint cavity. This fluid-filled space is the site at which the articulating surfaces of the bones contact each other.
Also unlike fibrous or cartilaginous joints, the articulating bone surfaces at a synovial joint are not directly connected to each other with fibrous
connective tissue or cartilage. This gives the bones of a synovial joint the ability to move smoothly against each other, allowing for increased joint
mobility.

Figure 9.8 Synovial Joints Synovial joints allow for smooth movements between the

adjacent bones. The joint is surrounded by an articular capsule that defines a joint cavity
filled with synovial fluid. The articulating surfaces of the bones are covered by a thin layer of
articular cartilage. Ligaments support the joint by holding the bones together and resisting
excess or abnormal joint motions.

Structural Features of Synovial Joints

Synovial joints are characterized by the presence of a joint cavity. The walls of this space
are formed by the articular capsule, a fibrous connective tissue structure that is attached to
each bone just outside the area of the bone’s articulating surface. The bones of the joint
articulate with each other within the joint cavity.

Friction between the bones at a synovial joint is prevented by the presence of the articular
cartilage, a thin layer of hyaline cartilage that covers the entire articulating surface of each
bone. However, unlike at a cartilaginous joint, the articular cartilages of each bone are not
continuous with each other. Instead, the articular cartilage acts like a Teflon® coating over
the bone surface, allowing the articulating bones to move smoothly against each other
without damaging the underlying bone tissue. Lining the inner surface of the articular
capsule is a thin synovial membrane. The cells of this membrane secrete synovial
fluid (synovia = “a thick fluid”), a thick, slimy fluid that provides lubrication to further reduce
friction between the bones of the joint. This fluid also provides nourishment to the articular cartilage, which does not contain blood vessels. The ability of
the bones to move smoothly against each other within the joint cavity, and the freedom of joint movement this provides, means that each synovial joint is
functionally classified as a diarthrosis.

Outside of their articulating surfaces, the bones are connected together by ligaments, which are strong bands of fibrous connective tissue. These
strengthen and support the joint by anchoring the bones together and preventing their separation. Ligaments allow for normal movements at a joint, but
limit the range of these motions, thus preventing excessive or abnormal joint movements. Ligaments are classified based on their relationship to the
fibrous articular capsule. An extrinsic ligament is located outside of the articular capsule, an intrinsic ligament is fused to or incorporated into the wall
of the articular capsule, and an intracapsular ligament is located inside of the articular capsule.

At many synovial joints, additional support is provided by the muscles and their tendons that act across the joint. A tendon is the dense connective
tissue structure that attaches a muscle to bone. As forces acting on a joint increase, the body will automatically increase the overall strength of
contraction of the muscles crossing that joint, thus allowing the muscle and its tendon to serve as a “dynamic ligament” to resist forces and support the
joint. This type of indirect support by muscles is very important at the shoulder joint, for example, where the ligaments are relatively weak.
Additional Structures Associated with Synovial Joints

A few synovial joints of the body have a fibrocartilage structure located between the articulating bones. This is called an articular disc, which is
generally small and oval-shaped, or a meniscus, which is larger and C-shaped. These structures can serve several functions, depending on the specific
joint. In some places, an articular disc may act to strongly unite the bones of the joint to each other. Examples of this include the articular discs found at
the sternoclavicular joint or between the distal ends of the radius and ulna bones. At other synovial joints, the disc can provide shock absorption and
cushioning between the bones, which is the function of each meniscus within the knee joint. Finally, an articular disc can serve to smooth the
movements between the articulating bones, as seen at the temporomandibular joint. Some synovial joints also have a fat pad, which can serve as a
cushion between the bones.

Additional structures located outside of a synovial joint serve to prevent friction between the bones of the joint and the overlying muscle tendons or skin.
A bursa (plural = bursae) is a thin connective tissue sac filled with lubricating liquid. They are located in regions where skin, ligaments, muscles, or
muscle tendons can rub against each other, usually near a body joint (Figure 9.9). Bursae reduce friction by separating the adjacent structures,
preventing them from rubbing directly against each other. Bursae are classified by their location. A subcutaneous bursa is located between the skin
and an underlying bone. It allows skin to move smoothly over the bone. Examples include the prepatellar bursa located over the kneecap and the
olecranon bursa at the tip of the elbow. A submuscular bursa is found between a muscle and an underlying bone, or between adjacent muscles. These
prevent rubbing of the muscle during movements. A large submuscular bursa, the trochanteric bursa, is found at the lateral hip, between the greater
trochanter of the femur and the overlying gluteus maximus muscle. A subtendinous bursa is found between a tendon and a bone. Examples include
the subacromial bursa that protects the tendon of shoulder muscle as it passes under the acromion of the scapula, and the suprapatellar bursa that
separates the tendon of the large anterior thigh muscle from the distal femur just above the knee.

Figure 9.9 Bursae Bursae are fluid-filled sacs that serve to prevent friction

between skin, muscle, or tendon and an underlying bone. Three major bursae and
a fat pad are part of the complex joint that unites the femur and tibia of the leg.

A tendon sheath is similar in structure to a bursa, but smaller. It is a connective

tissue sac that surrounds a muscle tendon at places where the tendon crosses a
joint. It contains a lubricating fluid that allows for smooth motions of the tendon
during muscle contraction and joint movements.

HOMEOSTATIC IMBALANCES

Bursitis

Bursitis is the inflammation of a bursa near a joint. This will cause pain, swelling,
or tenderness of the bursa and surrounding area, and may also result in joint
stiffness. Bursitis is most commonly associated with the bursae found at or near
the shoulder, hip, knee, or elbow joints. At the shoulder, subacromial bursitis may
occur in the bursa that separates the acromion of the scapula from the tendon of a shoulder muscle as it passes deep to the acromion. In the hip region,
trochanteric bursitis can occur in the bursa that overlies the greater trochanter of the femur, just below the lateral side of the hip. Ischial bursitis occurs in
the bursa that separates the skin from the ischial tuberosity of the pelvis, the bony structure that is weight bearing when sitting. At the knee, inflammation
and swelling of the bursa located between the skin and patella bone is prepatellar bursitis (“housemaid’s knee”), a condition more commonly seen today
in roofers or floor and carpet installers who do not use knee pads. At the elbow, olecranon bursitis is inflammation of the bursa between the skin and
olecranon process of the ulna. The olecranon forms the bony tip of the elbow, and bursitis here is also known as “student’s elbow.”

Bursitis can be either acute (lasting only a few days) or chronic. It can arise from muscle overuse, trauma, excessive or prolonged pressure on the skin,
rheumatoid arthritis, gout, or infection of the joint. Repeated acute episodes of bursitis can result in a chronic condition. Treatments for the disorder
include antibiotics if the bursitis is caused by an infection, or anti-inflammatory agents, such as nonsteroidal anti-inflammatory drugs (NSAIDs) or
corticosteroids if the bursitis is due to trauma or overuse. Chronic bursitis may require that fluid be drained, but additional surgery is usually not required.

Types of Synovial Joints

Synovial joints are subdivided based on the shapes of the articulating surfaces of the bones that form each joint. The six types of synovial joints are
pivot, hinge, condyloid, saddle, plane, and ball-and socket-joints (Figure 9.10).
 Figure 9.10 Types of Synovial Joints The six types of synovial joints
allow the body to move in a variety of ways. (a) Pivot joints allow for
rotation around an axis, such as between the first and second cervical
vertebrae, which allows for side-to-side rotation of the head. (b) The
hinge joint of the elbow works like a door hinge. (c) The articulation
between the trapezium carpal bone and the first metacarpal bone at the
base of the thumb is a saddle joint. (d) Plane joints, such as those
between the tarsal bones of the foot, allow for limited gliding movements
between bones. (e) The radiocarpal joint of the wrist is a condyloid joint.
(f) The hip and shoulder joints are the only ball-and-socket joints of the
body.

Pivot Joint

At a pivot joint, a rounded portion of a bone is enclosed within a ring

formed partially by the articulation with another bone and partially by a
ligament (see Figure 9.10a). The bone rotates within this ring. Since the
rotation is around a single axis, pivot joints are functionally classified as a
uniaxial diarthrosis type of joint. An example of a pivot joint is the
atlantoaxial joint, found between the C1 (atlas) and C2 (axis) vertebrae.
Here, the upward projecting dens of the axis articulates with the inner
aspect of the atlas, where it is held in place by a ligament. Rotation at this
joint allows you to turn your head from side to side. A second pivot joint is
found at the proximal radioulnar joint. Here, the head of the radius is
largely encircled by a ligament that holds it in place as it articulates with
the radial notch of the ulna. Rotation of the radius allows for forearm
movements.

Hinge Joint

In a hinge joint, the convex end of one bone articulates with the concave
end of the adjoining bone (see Figure 9.10b). This type of joint allows only for bending and straightening motions along a single axis, and thus hinge
joints are functionally classified as uniaxial joints. A good example is the elbow joint, with the articulation between the trochlea of the humerus and the
trochlear notch of the ulna. Other hinge joints of the body include the knee, ankle, and interphalangeal joints between the phalanx bones of the fingers
and toes.

Condyloid Joint

At a condyloid joint (ellipsoid joint), the shallow depression at the end of one bone articulates with a rounded structure from an adjacent bone or bones
(see Figure 9.10e). The knuckle (metacarpophalangeal) joints of the hand between the distal end of a metacarpal bone and the proximal phalanx bone
are condyloid joints. Another example is the radiocarpal joint of the wrist, between the shallow depression at the distal end of the radius bone and the
rounded scaphoid, lunate, and triquetrum carpal bones. In this case, the articulation area has a more oval (elliptical) shape. Functionally, condyloid joints
are biaxial joints that allow for two planes of movement. One movement involves the bending and straightening of the fingers or the anterior-posterior
movements of the hand. The second movement is a side-to-side movement, which allows you to spread your fingers apart and bring them together, or to
move your hand in a medial-going or lateral-going direction.

Saddle Joint

At a saddle joint, both of the articulating surfaces for the bones have a saddle shape, which is concave in one direction and convex in the other
(see Figure 9.10c). This allows the two bones to fit together like a rider sitting on a saddle. Saddle joints are functionally classified as biaxial joints. The
primary example is the first carpometacarpal joint, between the trapezium (a carpal bone) and the first metacarpal bone at the base of the thumb. This
joint provides the thumb the ability to move away from the palm of the hand along two planes. Thus, the thumb can move within the same plane as the
palm of the hand, or it can jut out anteriorly, perpendicular to the palm. This movement of the first carpometacarpal joint is what gives humans their
distinctive “opposable” thumbs. The sternoclavicular joint is also classified as a saddle joint.

Plane Joint

At a plane joint (gliding joint), the articulating surfaces of the bones are flat or slightly curved and of approximately the same size, which allows the
bones to slide against each other (see Figure 9.10d). The motion at this type of joint is usually small and tightly constrained by surrounding ligaments.
Based only on their shape, plane joints can allow multiple movements, including rotation. Thus plane joints can be functionally classified as a multiaxial
joint. However, not all of these movements are available to every plane joint due to limitations placed on it by ligaments or neighboring bones. Thus,
depending upon the specific joint of the body, a plane joint may exhibit only a single type of movement or several movements. Plane joints are found
between the carpal bones (intercarpal joints) of the wrist or tarsal bones (intertarsal joints) of the foot, between the clavicle and acromion of the scapula
(acromioclavicular joint), and between the superior and inferior articular processes of adjacent vertebrae (zygapophysial joints).
Ball-and-Socket Joint

The joint with the greatest range of motion is the ball-and-socket joint. At these joints, the rounded head of one bone (the ball) fits into the concave
articulation (the socket) of the adjacent bone (see Figure 9.10f). The hip joint and the glenohumeral (shoulder) joint are the only ball-and-socket joints of
the body. At the hip joint, the head of the femur articulates with the acetabulum of the hip bone, and at the shoulder joint, the head of the humerus
articulates with the glenoid cavity of the scapula.

Ball-and-socket joints are classified functionally as multiaxial joints. The femur and the humerus are able to move in both anterior-posterior and medial-
lateral directions and they can also rotate around their long axis. The shallow socket formed by the glenoid cavity allows the shoulder joint an extensive
range of motion. In contrast, the deep socket of the acetabulum and the strong supporting ligaments of the hip joint serve to constrain movements of the
femur, reflecting the need for stability and weight-bearing ability at the hip.

AGING AND THE...

Joints

Arthritis is a common disorder of synovial joints that involves inflammation of the joint. This often results in significant joint pain, along with swelling,
stiffness, and reduced joint mobility. There are more than 100 different forms of arthritis. Arthritis may arise from aging, damage to the articular cartilage,
autoimmune diseases, bacterial or viral infections, or unknown (probably genetic) causes.

The most common type of arthritis is osteoarthritis, which is associated with aging and “wear and tear” of the articular cartilage (Figure 9.11). Risk
factors that may lead to osteoarthritis later in life include injury to a joint; jobs that involve physical labor; sports with running, twisting, or throwing
actions; and being overweight. These factors put stress on the articular cartilage that covers the surfaces of bones at synovial joints, causing the
cartilage to gradually become thinner. As the articular cartilage layer wears down, more pressure is placed on the bones. The joint responds by
increasing production of the lubricating synovial fluid, but this can lead to swelling of the joint cavity, causing pain and joint stiffness as the articular
capsule is stretched. The bone tissue underlying the damaged articular cartilage also responds by thickening, producing irregularities and causing the
articulating surface of the bone to become rough or bumpy. Joint movement then results in pain and inflammation. In its early stages, symptoms of
osteoarthritis may be reduced by mild activity that “warms up” the joint, but the symptoms may worsen following exercise. In individuals with more
advanced osteoarthritis, the affected joints can become more painful and therefore are difficult to use effectively, resulting in increased immobility. There
is no cure for osteoarthritis, but several treatments can help alleviate the pain. Treatments may include lifestyle changes, such as weight loss and low-
impact exercise, and over-the-counter or prescription medications that help to alleviate the pain and inflammation. For severe cases, joint replacement
surgery (arthroplasty) may be required.

Joint replacement is a very invasive procedure, so other treatments are always tried before surgery. However arthroplasty can provide relief from chronic
pain and can enhance mobility within a few months following the surgery. This type of surgery involves replacing the articular surfaces of the bones with
prosthesis (artificial components). For example, in hip arthroplasty, the worn or damaged parts of the hip joint, including the head and neck of the femur
and the acetabulum of the pelvis, are removed and replaced with artificial joint components. The replacement head for the femur consists of a rounded
ball attached to the end of a shaft that is inserted inside the diaphysis of the femur. The acetabulum of the pelvis is reshaped and a replacement socket
is fitted into its place. The parts, which are always built in advance of the surgery, are sometimes custom made to produce the best possible fit for a
patient.

Gout is a form of arthritis that results from the deposition of uric acid crystals within a body joint. Usually only one or a few joints are affected, such as the
big toe, knee, or ankle. The attack may only last a few days, but may return to the same or another joint. Gout occurs when the body makes too much
uric acid or the kidneys do not properly excrete it. A diet with excessive fructose has been implicated in raising the chances of a susceptible individual
developing gout.

Other forms of arthritis are associated with various autoimmune diseases, bacterial infections of the joint, or unknown genetic causes. Autoimmune
diseases, including rheumatoid arthritis, scleroderma, or systemic lupus erythematosus, produce arthritis because the immune system of the body
attacks the body joints. In rheumatoid arthritis, the joint capsule and synovial membrane become inflamed. As the disease progresses, the articular
cartilage is severely damaged or destroyed, resulting in joint deformation, loss of movement, and severe disability. The most commonly involved joints
are the hands, feet, and cervical spine, with corresponding joints on both sides of the body usually affected, though not always to the same extent.
Rheumatoid arthritis is also associated with lung fibrosis, vasculitis (inflammation of blood vessels), coronary heart disease, and premature mortality.
With no known cure, treatments are aimed at alleviating symptoms. Exercise, anti-inflammatory and pain medications, various specific disease-
modifying anti-rheumatic drugs, or surgery are used to treat rheumatoid arthritis.
 Figure 9.11 Osteoarthritis Osteoarthritis of a synovial joint results from aging or
prolonged joint wear and tear. These cause erosion and loss of the articular
cartilage covering the surfaces of the bones, resulting in inflammation that causes
joint stiffness and pain.

Synovial
joints allow the
body a

tremendous range of movements. Each movement at a synovial joint results

from the contraction or relaxation of the muscles that are attached to the bones on
either side of the articulation. The type of movement that can be produced at a
synovial joint is determined by its structural type. While the ball-and-socket joint
gives the greatest range of movement at an individual joint, in other regions of the
body, several joints may work together to produce a particular movement.
Overall, each type of synovial joint is necessary to provide the body with its
great flexibility and mobility. There are many types of movement that can occur at
synovial joints (Table 9.1). Movement types are generally paired, with one being the
opposite of the other. Body movements are always described in relation to the
anatomical position of the body: upright stance, with upper limbs to the side of
body and palms facing forward. Refer to Figure 9.12 as you go through this
section.

Figure 9.12 Movements of the Body, Part 1 Synovial joints give the body many ways in which to move. (a)–(b) Flexion and extension motions are in
the sagittal (anterior–posterior) plane of motion. These movements take place at the shoulder, hip, elbow, knee, wrist, metacarpophalangeal,
metatarsophalangeal, and interphalangeal joints. (c)–(d) Anterior bending of the head or vertebral column is flexion, while any posterior-going movement
is extension. (e) Abduction and adduction are motions of the limbs, hand, fingers, or toes in the coronal (medial–lateral) plane of movement. Moving the
limb or hand laterally away from the body, or spreading the fingers or toes, is abduction. Adduction brings the limb or hand toward or across the midline
of the body, or brings the fingers or toes together. Circumduction is the movement of the limb, hand, or fingers in a circular pattern, using the sequential
combination of flexion, adduction, extension, and abduction motions. Adduction/abduction and circumduction take place at the shoulder, hip, wrist,
metacarpophalangeal, and metatarsophalangeal joints. (f) Turning of the head side to side or twisting of the body is rotation. Medial and lateral rotation
of the upper limb at the shoulder or lower limb at the hip involves turning the anterior surface of the limb toward the midline of the body (medial or
internal rotation) or away from the midline (lateral or external rotation).

Figure 9.13 Movements of the Body, Part 2 (g) Supination of the forearm turns the hand to the palm forward position in which the radius and ulna are
parallel, while forearm pronation turns the hand to the palm backward position in which the radius crosses over the ulna to form an "X." (h) Dorsiflexion
of the foot at the ankle joint moves the top of the foot toward the leg, while plantar flexion lifts the heel and points the toes. (i) Eversion of the foot moves
the bottom (sole) of the foot away from the midline of the body, while foot inversion faces the sole toward the midline. (j) Protraction of the mandible
pushes the chin forward, and retraction pulls the chin back. (k) Depression of the mandible opens the mouth, while elevation closes it. (l) Opposition of
the thumb brings the tip of the thumb into contact with the tip of the fingers of the same hand and reposition brings the thumb back next to the index
finger.

Flexion and Extension

Flexion and extension are typically movements that take place within the sagittal plane and involve anterior or posterior movements of the neck, trunk,
or limbs. For the vertebral column, flexion (anterior flexion) is an anterior (forward) bending of the neck or trunk, while extension involves a posterior-
directed motion, such as straightening from a flexed position or bending backward. Lateral flexion of the vertebral column occurs in the coronal plane
and is defined as the bending of the neck or trunk toward the right or left side.. These movements of the vertebral column involve both the symphysis
joint formed by each intervertebral disc, as well as the plane type of synovial joint formed between the inferior articular processes of one vertebra and
the superior articular processes of the next lower vertebra.

In the limbs, flexion decreases the angle between the bones (bending of the joint), while extension increases the angle and straightens the joint. For the
upper limb, all anterior-going motions are flexion and all posterior-going motions are extension. These include anterior-posterior movements of the arm
at the shoulder, the forearm at the elbow, the hand at the wrist, and the fingers at the metacarpophalangeal and interphalangeal joints. For the thumb,
extension moves the thumb away from the palm of the hand, within the same plane as the palm, while flexion brings the thumb back against the index
finger or into the palm. These motions take place at the first carpometacarpal joint. In the lower limb, bringing the thigh forward and upward is flexion at
the hip joint, while any posterior-going motion of the thigh is extension. Note that extension of the thigh beyond the anatomical (standing) position is
greatly limited by the ligaments that support the hip joint. Knee flexion is the bending of the knee to bring the foot toward the posterior thigh, and
extension is the straightening of the knee. Flexion and extension movements are seen at the hinge, condyloid, saddle, and ball-and-socket joints of the
limbs (see Figure 9.12a-d).
Hyperextension is the abnormal or excessive extension of a joint beyond its normal range of motion, thus resulting in injury. Similarly, hyperflexion is
excessive flexion at a joint. Hyperextension injuries are common at hinge joints such as the knee or elbow. In cases of “whiplash” in which the head is
suddenly moved backward and then forward, a patient may experience both hyperextension and hyperflexion of the cervical region.

Abduction and Adduction

Abduction and adduction motions occur within the coronal plane and involve medial-lateral motions of the limbs, fingers, toes, or thumb. Abduction
moves the limb laterally away from the midline of the body, while adduction is the opposing movement that brings the limb toward the body or across the
midline. For example, abduction is raising the arm at the shoulder joint, moving it laterally away from the body, while adduction brings the arm down to
the side of the body. Similarly, abduction and adduction at the wrist moves the hand away from or toward the midline of the body. Spreading the fingers
or toes apart is also abduction, while bringing the fingers or toes together is adduction. For the thumb, abduction is the anterior movement that brings the
thumb to a 90° perpendicular position, pointing straight out from the palm. Adduction moves the thumb back to the anatomical position, next to the index
finger. Abduction and adduction movements are seen at condyloid, saddle, and ball-and-socket joints (see Figure 9.12e).

Circumduction

Circumduction is the movement of a body region in a circular manner, in which one end of the body region being moved stays relatively stationary
while the other end describes a circle. It involves the sequential combination of flexion, adduction, extension, and abduction at a joint. This type of
motion is found at biaxial condyloid and saddle joints, and at multiaxial ball-and-sockets joints (see Figure 9.12e).

Rotation

Rotation can occur within the vertebral column, at a pivot joint, or at a ball-and-socket joint. Rotation of the neck or body is the twisting movement
produced by the summation of the small rotational movements available between adjacent vertebrae. At a pivot joint, one bone rotates in relation to
another bone. This is a uniaxial joint, and thus rotation is the only motion allowed at a pivot joint. For example, at the atlantoaxial joint, the first cervical
(C1) vertebra (atlas) rotates around the dens, the upward projection from the second cervical (C2) vertebra (axis). This allows the head to rotate from
side to side as when shaking the head “no.” The proximal radioulnar joint is a pivot joint formed by the head of the radius and its articulation with the
ulna. This joint allows for the radius to rotate along its length during pronation and supination movements of the forearm.

Rotation can also occur at the ball-and-socket joints of the shoulder and hip. Here, the humerus and femur rotate around their long axis, which moves
the anterior surface of the arm or thigh either toward or away from the midline of the body. Movement that brings the anterior surface of the limb toward
the midline of the body is called medial (internal) rotation. Conversely, rotation of the limb so that the anterior surface moves away from the midline
is lateral (external) rotation (see Figure 9.12f). Be sure to distinguish medial and lateral rotation, which can only occur at the multiaxial shoulder and
hip joints, from circumduction, which can occur at either biaxial or multiaxial joints.

Supination and Pronation

Supination and pronation are movements of the forearm. In the anatomical position, the upper limb is held next to the body with the palm facing forward.
This is the supinated position of the forearm. In this position, the radius and ulna are parallel to each other. When the palm of the hand faces
backward, the forearm is in the pronated position, and the radius and ulna form an X-shape.

Supination and pronation are the movements of the forearm that go between these two positions. Pronation is the motion that moves the forearm from
the supinated (anatomical) position to the pronated (palm backward) position. This motion is produced by rotation of the radius at the proximal radioulnar
joint, accompanied by movement of the radius at the distal radioulnar joint. The proximal radioulnar joint is a pivot joint that allows for rotation of the head
of the radius. Because of the slight curvature of the shaft of the radius, this rotation causes the distal end of the radius to cross over the distal ulna at the
distal radioulnar joint. This crossing over brings the radius and ulna into an X-shape position. Supination is the opposite motion, in which rotation of the
radius returns the bones to their parallel positions and moves the palm to the anterior facing (supinated) position. It helps to remember that supination is
the motion you use when scooping up soup with a spoon (see Figure 9.13g).

Dorsiflexion and Plantar Flexion

Dorsiflexion and plantar flexion are movements at the ankle joint, which is a hinge joint. Lifting the front of the foot, so that the top of the foot moves
toward the anterior leg is dorsiflexion, while lifting the heel of the foot from the ground or pointing the toes downward is plantar flexion. These are the
only movements available at the ankle joint (see Figure 9.13h).

Inversion and Eversion

Inversion and eversion are complex movements that involve the multiple plane joints among the tarsal bones of the posterior foot (intertarsal joints) and
thus are not motions that take place at the ankle joint. Inversion is the turning of the foot to angle the bottom of the foot toward the midline,
while eversion turns the bottom of the foot away from the midline. The foot has a greater range of inversion than eversion motion. These are important
motions that help to stabilize the foot when walking or running on an uneven surface and aid in the quick side-to-side changes in direction used during
active sports such as basketball, racquetball, or soccer (see Figure 9.13i).

Protraction and Retraction

Protraction and retraction are anterior-posterior movements of the scapula or mandible. Protraction of the scapula occurs when the shoulder is moved
forward, as when pushing against something or throwing a ball. Retraction is the opposite motion, with the scapula being pulled posteriorly and medially,
toward the vertebral column. For the mandible, protraction occurs when the lower jaw is pushed forward, to stick out the chin, while retraction pulls the
lower jaw backward. (See Figure 9.13j.)
Depression and Elevation

Depression and elevation are downward and upward movements of the scapula or mandible. The upward movement of the scapula and shoulder is
elevation, while a downward movement is depression. These movements are used to shrug your shoulders. Similarly, elevation of the mandible is the
upward movement of the lower jaw used to close the mouth or bite on something, and depression is the downward movement that produces opening of
the mouth (see Figure 9.13k).

Excursion

Excursion is the side to side movement of the mandible. Lateral excursion moves the mandible away from the midline, toward either the right or left
side. Medial excursion returns the mandible to its resting position at the midline.

Superior Rotation and Inferior Rotation

Superior and inferior rotation are movements of the scapula and are defined by the direction of movement of the glenoid cavity. These motions involve
rotation of the scapula around a point inferior to the scapular spine and are produced by combinations of muscles acting on the scapula.
During superior rotation, the glenoid cavity moves upward as the medial end of the scapular spine moves downward. This is a very important motion
that contributes to upper limb abduction. Without superior rotation of the scapula, the greater tubercle of the humerus would hit the acromion of the
scapula, thus preventing any abduction of the arm above shoulder height. Superior rotation of the scapula is thus required for full abduction of the upper
limb. Superior rotation is also used without arm abduction when carrying a heavy load with your hand or on your shoulder. You can feel this rotation
when you pick up a load, such as a heavy book bag and carry it on only one shoulder. To increase its weight-bearing support for the bag, the shoulder
lifts as the scapula superiorly rotates. Inferior rotation occurs during limb adduction and involves the downward motion of the glenoid cavity with
upward movement of the medial end of the scapular spine.

Opposition and Reposition

Opposition is the thumb movement that brings the tip of the thumb in contact with the tip of a finger. This movement is produced at the first
carpometacarpal joint, which is a saddle joint formed between the trapezium carpal bone and the first metacarpal bone. Thumb opposition is produced
by a combination of flexion and abduction of the thumb at this joint. Returning the thumb to its anatomical position next to the index finger is
called reposition (see Figure 9.13l).

Each synovial joint of the body is specialized to perform certain movements. The movements that are allowed are determined by the structural
classification for each joint. For example, a multiaxial ball-and-socket joint has much more mobility than a uniaxial hinge joint. However, the ligaments
and muscles that support a joint may place restrictions on the total range of motion available. Thus, the ball-and-socket joint of the shoulder has little in
the way of ligament support, which gives the shoulder a very large range of motion. In contrast, movements at the hip joint are restricted by strong
ligaments, which reduce its range of motion but confer stability during standing and weight bearing.

This section will examine the anatomy of selected synovial joints of the body. Anatomical names for most joints are derived from the names of the bones
that articulate at that joint, although some joints, such as the elbow, hip, and knee joints are exceptions to this general naming scheme.

Articulations of the Vertebral Column

In addition to being held together by the intervertebral discs, adjacent vertebrae also articulate with each other at synovial joints formed between the
superior and inferior articular processes called zygapophysial joints (facet joints) (see Figure 9.3). These are plane joints that provide for only limited
motions between the vertebrae. The orientation of the articular processes at these joints varies in different regions of the vertebral column and serves to
determine the types of motions available in each vertebral region. The cervical and lumbar regions have the greatest ranges of motions.

In the neck, the articular processes of cervical vertebrae are flattened and generally face upward or downward. This orientation provides the cervical
vertebral column with extensive ranges of motion for flexion, extension, lateral flexion, and rotation. In the thoracic region, the downward projecting and
overlapping spinous processes, along with the attached thoracic cage, greatly limit flexion, extension, and lateral flexion. However, the flattened and
vertically positioned thoracic articular processes allow for the greatest range of rotation within the vertebral column. The lumbar region allows for
considerable extension, flexion, and lateral flexion, but the orientation of the articular processes largely prohibits rotation.

The articulations formed between the skull, the atlas (C1 vertebra), and the axis (C2 vertebra) differ from the articulations in other vertebral areas and
play important roles in movement of the head. The atlanto-occipital joint is formed by the articulations between the superior articular processes of the
atlas and the occipital condyles on the base of the skull. This articulation has a pronounced U-shaped curvature, oriented along the anterior-posterior
axis. This allows the skull to rock forward and backward, producing flexion and extension of the head. This moves the head up and down, as when
shaking your head “yes.”

The atlantoaxial joint, between the atlas and axis, consists of three articulations. The paired superior articular processes of the axis articulate with the
inferior articular processes of the atlas. These articulating surfaces are relatively flat and oriented
horizontally. The third articulation is the pivot joint formed between the dens, which projects
upward from the body of the axis, and the inner aspect of the anterior arch of the atlas (Figure
9.14). A strong ligament passes posterior to the dens to hold it in position against the anterior
arch. These articulations allow the atlas to rotate on top of the axis, moving the head toward the
right or left, as when shaking your head “no.”
 Figure 9.14 Atlantoaxial Joint The atlantoaxial joint is a pivot type of joint between the dens
portion of the axis (C2 vertebra) and the anterior arch of the atlas (C1 vertebra), with the dens
held in place by a ligament.

Temporomandibular Joint

The temporomandibular joint (TMJ) is the joint that allows for opening (mandibular
depression) and closing (mandibular elevation) of the mouth, as well as side-to-side and
protraction/retraction motions of the lower jaw. This joint involves the articulation between the
mandibular fossa and articular tubercle of the temporal bone, with the condyle (head) of the
mandible. Located between these bony structures, filling the gap between the skull and
mandible, is a flexible articular disc (Figure 9.15). This disc serves to smooth the movements
between the temporal bone and mandibular condyle.

Movement at the TMJ during opening and closing of the mouth involves both gliding and
hinge motions of the mandible. With the mouth closed, the mandibular condyle and articular disc are located within the mandibular fossa of the temporal
bone. During opening of the mouth, the mandible hinges downward and at the same time is pulled anteriorly, causing both the condyle and the articular
disc to glide forward from the mandibular fossa onto the downward projecting articular tubercle. The net result is a forward and downward motion of the
condyle and mandibular depression. The temporomandibular joint is supported by an extrinsic ligament that anchors the mandible to the skull. This
ligament spans the distance between the base of the skull and the lingula on the medial side of the mandibular ramus.

Dislocation of the TMJ may occur when opening the mouth too wide (such as when taking a large bite) or following a blow to the jaw, resulting in the
mandibular condyle moving beyond (anterior to) the articular tubercle. In this case, the individual would not be able to close his or her mouth.
Temporomandibular joint disorder is a painful condition that may arise due to arthritis, wearing of the articular cartilage covering the bony surfaces of the
joint, muscle fatigue from overuse or grinding of the teeth, damage to the articular disc within the joint, or jaw injury. Temporomandibular joint disorders
can also cause headache, difficulty chewing, or even the inability to move the jaw (lock jaw). Pharmacologic agents for pain or other therapies, including
bite guards, are used as treatments.

Figure 9.15 Temporomandibular Joint The temporomandibular joint is the articulation between the temporal bone of the skull and the condyle of the
mandible, with an articular disc located between these bones. During depression of the mandible (opening of the mouth), the mandibular condyle moves
both forward and hinges downward as it travels from the mandibular fossa onto the articular tubercle.

INTERACTIVE LINK

Watch this video to learn about TMJ. Opening of the mouth requires the combination of two motions at the temporomandibular joint, an anterior gliding
motion of the articular disc and mandible and the downward hinging of the mandible. What is the initial movement of the mandible during opening and
how much mouth opening does this produce?

Shoulder Joint

The shoulder joint is called the glenohumeral joint. This is a ball-and-socket joint formed by the articulation between the head of the humerus and the
glenoid cavity of the scapula (Figure 9.16). This joint has the largest range of motion of any joint in the body. However, this freedom of movement is due
to the lack of structural support and thus the enhanced mobility is offset by a loss of stability.

Figure 9.16 Glenohumeral Joint The glenohumeral (shoulder)

joint is a ball-and-socket joint that provides the widest range of
motions. It has a loose articular capsule and is supported by
ligaments and the rotator cuff muscles.

The large range of motions at the shoulder joint is provided by the

articulation of the large, rounded humeral head with the small and
shallow glenoid cavity, which is only about one third of the size of
the humeral head. The socket formed by the glenoid cavity is
deepened slightly by a small lip of fibrocartilage called the glenoid
labrum, which extends around the outer margin of the cavity. The
articular capsule that surrounds the glenohumeral joint is relatively
thin and loose to allow for large motions of the upper limb. Some
structural support for the joint is provided by thickenings of the
articular capsule wall that form weak intrinsic ligaments. These
include the coracohumeral ligament, running from the coracoid
process of the scapula to the anterior humerus, and three
ligaments, each called a glenohumeral ligament, located on the
anterior side of the articular capsule. These ligaments help to
strengthen the superior and anterior capsule walls.
However, the primary support for the shoulder joint is provided by muscles crossing the joint, particularly the four rotator cuff muscles. These muscles
(supraspinatus, infraspinatus, teres minor, and subscapularis) arise from the scapula and attach to the greater or lesser tubercles of the humerus. As
these muscles cross the shoulder joint, their tendons encircle the head of the humerus and become fused to the anterior, superior, and posterior walls of
the articular capsule. The thickening of the capsule formed by the fusion of these four muscle tendons is called the rotator cuff. Two bursae,
the subacromial bursa and the subscapular bursa, help to prevent friction between the rotator cuff muscle tendons and the scapula as these tendons
cross the glenohumeral joint. In addition to their individual actions of moving the upper limb, the rotator cuff muscles also serve to hold the head of the
humerus in position within the glenoid cavity. By constantly adjusting their strength of contraction to resist forces acting on the shoulder, these muscles
serve as “dynamic ligaments” and thus provide the primary structural support for the glenohumeral joint.

Injuries to the shoulder joint are common. Repetitive use of the upper limb, particularly in abduction such as during throwing, swimming, or racquet
sports, may lead to acute or chronic inflammation of the bursa or muscle tendons, a tear of the glenoid labrum, or degeneration or tears of the rotator
cuff. Because the humeral head is strongly supported by muscles and ligaments around its anterior, superior, and posterior aspects, most dislocations of
the humerus occur in an inferior direction. This can occur when force is applied to the humerus when the upper limb is fully abducted, as when diving to
catch a baseball and landing on your hand or elbow. Inflammatory responses to any shoulder injury can lead to the formation of scar tissue between the
articular capsule and surrounding structures, thus reducing shoulder mobility, a condition called adhesive capsulitis (“frozen shoulder”).

Elbow Joint

The elbow joint is a uniaxial hinge joint formed by the humeroulnar joint, the articulation between the trochlea of the humerus and the trochlear notch
of the ulna. Also associated with the elbow are the humeroradial joint and the proximal radioulnar joint. All three of these joints are enclosed within a
single articular capsule (Figure 9.17).

The articular capsule of the elbow is thin on its anterior and posterior aspects, but is thickened along its outside margins by strong intrinsic ligaments.
These ligaments prevent side-to-side movements and hyperextension. On the medial side is the triangular ulnar collateral ligament. This arises from
the medial epicondyle of the humerus and attaches to the medial side of the proximal ulna. The strongest part of this ligament is the anterior portion,
which resists hyperextension of the elbow. The ulnar collateral ligament may be injured by frequent, forceful extensions of the forearm, as is seen in
baseball pitchers. Reconstructive surgical repair of this ligament is referred to as Tommy John surgery, named for the former major league pitcher who
was the first person to have this treatment.

The lateral side of the elbow is supported by the radial collateral ligament. This arises from the lateral epicondyle of the humerus and then blends into
the lateral side of the annular ligament. The annular ligament encircles the head of the radius. This ligament supports the head of the radius as it
articulates with the radial notch of the ulna at the proximal radioulnar joint. This is a pivot joint that allows for rotation of the radius during supination and
pronation of the forearm.

Figure 9.17 Elbow Joint (a) The elbow is a hinge joint that

allows only for flexion and extension of the forearm. (b) It is
supported by the ulnar and radial collateral ligaments. (c) The
annular ligament supports the head of the radius at the
proximal radioulnar joint, the pivot joint that allows for rotation
of the radius.

Hip Joint

The hip joint is a multiaxial ball-and-socket joint between the

head of the femur and the acetabulum of the hip bone (Figure
9.18). The hip carries the weight of the body and thus requires
strength and stability during standing and walking. For these
reasons, its range of motion is more limited than at the
shoulder joint.

The acetabulum is the socket portion of the hip joint. This

space is deep and has a large articulation area for the femoral
head, thus giving stability and weight bearing ability to the
joint. The acetabulum is further deepened by the acetabular
labrum, a fibrocartilage lip attached to the outer margin of the
acetabulum. The surrounding articular capsule is strong, with
several thickened areas forming intrinsic ligaments. These
ligaments arise from the hip bone, at the margins of the
acetabulum, and attach to the femur at the base of the neck. The ligaments are the iliofemoral ligament, pubofemoral ligament, and ischiofemoral
ligament, all of which spiral around the head and neck of the femur. The ligaments are tightened by extension at the hip, thus pulling the head of the
femur tightly into the acetabulum when in the upright, standing position. Very little additional extension of the thigh is permitted beyond this vertical
position. These ligaments thus stabilize the hip joint and allow you to maintain an upright standing position with only minimal muscle contraction. Inside
of the articular capsule, the ligament of the head of the femur (ligamentum teres) spans between the acetabulum and femoral head. This intracapsular
ligament is normally slack and does not provide any significant joint support, but it does provide a pathway for an important artery that supplies the head
of the femur.
 The hip is prone to osteoarthritis, and thus was the first joint for which a replacement prosthesis was
developed. A common injury in elderly individuals, particularly those with weakened bones due to
osteoporosis, is a “broken hip,” which is actually a fracture of the femoral neck. This may result from a
fall, or it may cause the fall. This can happen as one lower limb is taking a step and all of the body weight
is placed on the other limb, causing the femoral neck to break and producing a fall. Any accompanying
disruption of the blood supply to the femoral neck or head can lead to necrosis of these areas, resulting in
bone and cartilage death. Femoral fractures usually require surgical treatment, after which the patient will
need mobility assistance for a prolonged period, either from family members or in a long-term care
facility. Consequentially, the associated health care costs of “broken hips” are substantial. In addition, hip
fractures are associated with increased rates of morbidity (incidences of disease) and mortality (death).
Surgery for a hip fracture followed by prolonged bed rest may lead to life-threatening complications,
including pneumonia, infection of pressure ulcers (bedsores), and thrombophlebitis (deep vein
thrombosis; blood clot formation) that can result in a pulmonary embolism (blood clot within the lung).

Figure 9.18 Hip Joint (a) The ball-and-socket joint of the hip is a multiaxial joint that provides both
stability and a wide range of motion. (b–c) When standing, the supporting ligaments are tight, pulling the
head of the femur into the acetabulum.

Knee Joint

The knee joint is the largest joint of the body (Figure 9.19). It actually consists of three articulations.
The femoropatellar joint is found between the patella and the distal femur. The medial tibiofemoral
joint and lateral tibiofemoral joint are located between the medial and lateral condyles of the femur and
the medial and lateral condyles of the tibia. All of these articulations are enclosed within a single articular
capsule. The knee functions as a hinge joint, allowing flexion and extension of the leg. This action is generated by both rolling and gliding motions of the
femur on the tibia. In addition, some rotation of the leg is available when the knee is flexed, but not when extended. The knee is well constructed for
weight bearing in its extended position, but is vulnerable to injuries associated with hyperextension, twisting, or blows to the medial or lateral side of the
joint, particularly while weight bearing.

At the femoropatellar joint, the patella slides vertically within a groove on the distal femur. The patella is a sesamoid bone incorporated into the tendon of
the quadriceps femoris muscle, the large muscle of the anterior thigh. The patella serves to protect the quadriceps tendon from friction against the distal
femur. Continuing from the patella to the anterior tibia just below the knee is the patellar ligament. Acting via the patella and patellar ligament, the
quadriceps femoris is a powerful muscle that acts to extend the leg at the knee. It also serves as a “dynamic ligament” to provide very important support
and stabilization for the knee joint.

The medial and lateral tibiofemoral joints are the articulations between the rounded condyles of the femur and the relatively flat condyles of the tibia.
During flexion and extension motions, the condyles of the femur both roll and glide over the surfaces of the tibia. The rolling action produces flexion or
extension, while the gliding action serves to maintain the femoral condyles centered over the tibial condyles, thus ensuring maximal bony, weight-
bearing support for the femur in all knee positions. As the knee comes into full extension, the femur undergoes a slight medial rotation in relation to tibia.
The rotation results because the lateral condyle of the femur is slightly smaller than the medial condyle. Thus, the lateral condyle finishes its rolling
motion first, followed by the medial condyle. The resulting small medial rotation of the femur serves to “lock” the knee into its fully extended and most
stable position. Flexion of the knee is initiated by a slight lateral rotation of the femur on the tibia, which “unlocks” the knee. This lateral rotation motion is
produced by the popliteus muscle of the posterior leg.

Located between the articulating surfaces of the femur and tibia are two articular discs, the medial meniscus and lateral meniscus (see Figure 9.19b).
Each is a C-shaped fibrocartilage structure that is thin along its inside margin and thick along the outer margin. They are attached to their tibial condyles,
but do not attach to the femur. While both menisci are free to move during knee motions, the medial meniscus shows less movement because it is
anchored at its outer margin to the articular capsule and tibial collateral ligament. The menisci provide padding between the bones and help to fill the
gap between the round femoral condyles and flattened tibial condyles. Some areas of each meniscus lack an arterial blood supply and thus these areas
heal poorly if damaged.

The knee joint has multiple ligaments that provide support, particularly in the extended position (see Figure 9.19c). Outside of the articular capsule,
located at the sides of the knee, are two extrinsic ligaments. The fibular collateral ligament (lateral collateral ligament) is on the lateral side and spans
from the lateral epicondyle of the femur to the head of the fibula. The tibial collateral ligament (medial collateral ligament) of the medial knee runs from
the medial epicondyle of the femur to the medial tibia. As it crosses the knee, the tibial collateral ligament is firmly attached on its deep side to the
articular capsule and to the medial meniscus, an important factor when considering knee injuries. In the fully extended knee position, both collateral
ligaments are taut (tight), thus serving to stabilize and support the extended knee and preventing side-to-side or rotational motions between the femur
and tibia.

The articular capsule of the posterior knee is thickened by intrinsic ligaments that help to resist knee hyperextension. Inside the knee are two
intracapsular ligaments, the anterior cruciate ligament and posterior cruciate ligament. These ligaments are anchored inferiorly to the tibia at the
intercondylar eminence, the roughened area between the tibial condyles. The cruciate ligaments are named for whether they are attached anteriorly or
posteriorly to this tibial region. Each ligament runs diagonally upward to attach to the inner aspect of a femoral condyle. The cruciate ligaments are
named for the X-shape formed as they pass each other (cruciate means “cross”). The posterior cruciate ligament is the stronger ligament. It serves to
support the knee when it is flexed and weight bearing, as when walking downhill. In this position, the posterior cruciate ligament prevents the femur from
sliding anteriorly off the top of the tibia. The anterior cruciate ligament becomes tight when the knee is extended, and thus resists hyperextension.
 Figure 9.19 Knee Joint (a) The knee joint is the
largest joint of the body. (b)–(c) It is supported by the
tibial and fibular collateral ligaments located on the
sides of the knee outside of the articular capsule,
and the anterior and posterior cruciate ligaments
found inside the capsule. The medial and lateral
menisci provide padding and support between the
femoral condyles and tibial condyles.

DISORDERS OF THE...

Joints

Injuries to the knee are common. Since this joint is

primarily supported by muscles and ligaments,
injuries to any of these structures will result in pain
or knee instability. Injury to the posterior cruciate
ligament occurs when the knee is flexed and the
tibia is driven posteriorly, such as falling and landing
on the tibial tuberosity or hitting the tibia on the
dashboard when not wearing a seatbelt during an
automobile accident. More commonly, injuries occur
when forces are applied to the extended knee,
particularly when the foot is planted and unable to
move. Anterior cruciate ligament injuries can result
with a forceful blow to the anterior knee, producing hyperextension, or when a runner makes a quick change of direction that produces both twisting and
hyperextension of the knee.

A worse combination of injuries can occur with a hit to the lateral side of the extended knee (Figure 9.20). A moderate blow to the lateral knee will cause
the medial side of the joint to open, resulting in stretching or damage to the tibial collateral ligament. Because the medial meniscus is attached to the
tibial collateral ligament, a stronger blow can tear the ligament and also damage the medial meniscus. This is one reason that the medial meniscus is 20
times more likely to be injured than the lateral meniscus. A powerful blow to the lateral knee produces a “terrible triad” injury, in which there is a
sequential injury to the tibial collateral ligament, medial meniscus, and anterior cruciate ligament.

Arthroscopic surgery has greatly improved the surgical treatment of knee injuries and reduced subsequent recovery times. This procedure involves a
small incision and the insertion into the joint of an arthroscope, a pencil-thin instrument that allows for visualization of the joint interior. Small surgical
instruments are also inserted via additional incisions. These tools allow a surgeon to remove or repair a torn meniscus or to reconstruct a ruptured
cruciate ligament. The current method for anterior cruciate ligament replacement involves using a portion of the patellar ligament. Holes are drilled into
the cruciate ligament attachment points on the tibia and femur, and the patellar ligament graft, with small areas of attached bone still intact at each end,
is inserted into these holes. The bone-to-bone sites at each end of the graft heal rapidly and strongly, thus enabling a rapid recovery.

Figure 9.20 Knee Injury A strong blow to the lateral side of the extended

knee will cause three injuries, in sequence: tearing of the tibial collateral
ligament, damage to the medial meniscus, and rupture of the anterior
cruciate ligament.

Ankle and Foot Joints

The ankle is formed by the talocrural joint (Figure 9.21). It consists of the

articulations between the talus bone of the foot and the distal ends of the
tibia and fibula of the leg (crural = “leg”). The superior aspect of the talus
bone is square-shaped and has three areas of articulation. The top of the
talus articulates with the inferior tibia. This is the portion of the ankle joint
that carries the body weight between the leg and foot. The sides of the
talus are firmly held in position by the articulations with the medial
malleolus of the tibia and the lateral malleolus of the fibula, which prevent
any side-to-side motion of the talus. The ankle is thus a uniaxial hinge joint that allows only for dorsiflexion and plantar flexion of the foot.

Additional joints between the tarsal bones of the posterior foot allow for the movements of foot inversion and eversion. Most important for these
movements is the subtalar joint, located between the talus and calcaneus bones. The joints between the talus and navicular bones and the calcaneus
and cuboid bones are also important contributors to these movements. All of the joints between tarsal bones are plane joints. Together, the small
motions that take place at these joints all contribute to the production of inversion and eversion foot motions.

Like the hinge joints of the elbow and knee, the talocrural joint of the ankle is supported by several strong ligaments located on the sides of the joint.
These ligaments extend from the medial malleolus of the tibia or lateral malleolus of the fibula and anchor to the talus and calcaneus bones. Since they
are located on the sides of the ankle joint, they allow for dorsiflexion and plantar flexion of the foot. They also prevent abnormal side-to-side and twisting
movements of the talus and calcaneus bones during eversion and inversion of the foot. On the medial side is the broad deltoid ligament. The deltoid
ligament supports the ankle joint and also resists excessive eversion of the foot. The lateral side of the ankle has several smaller ligaments. These
include the anterior talofibular ligament and the posterior talofibular ligament, both of which span between the talus bone and the lateral malleolus
of the fibula, and the calcaneofibular ligament, located between the calcaneus bone and fibula. These ligaments support the ankle and also resist
excess inversion of the foot.

Figure 9.21 Ankle Joint The talocrural (ankle) joint is a uniaxial hinge joint that only allows
for dorsiflexion or plantar flexion of the foot. Movements at the subtalar joint, between the
talus and calcaneus bones, combined with motions at other intertarsal joints, enables
eversion/inversion movements of the foot. Ligaments that unite the medial or lateral
malleolus with the talus and calcaneus bones serve to support the talocrural joint and to
resist excess eversion or inversion of the foot.

DISORDERS OF THE...

Joints

The ankle is the most frequently injured joint in the body, with the most common injury being
an inversion ankle sprain. A sprain is the stretching or tearing of the supporting ligaments.
Excess inversion causes the talus bone to tilt laterally, thus damaging the ligaments on the
lateral side of the ankle. The anterior talofibular ligament is most commonly injured, followed
by the calcaneofibular ligament. In severe inversion injuries, the forceful lateral movement of
the talus not only ruptures the lateral ankle ligaments, but also fractures the distal fibula.

Less common are eversion sprains of the ankle, which involve stretching of the deltoid
ligament on the medial side of the ankle. Forcible eversion of the foot, for example, with an
awkward landing from a jump or when a football player has a foot planted and is hit on the
lateral ankle, can result in a Pott’s fracture and dislocation of the ankle joint. In this injury, the very strong deltoid ligament does not tear, but instead
shears off the medial malleolus of the tibia. This frees the talus, which moves laterally and fractures the distal fibula. In extreme cases, the posterior
margin of the tibia may also be sheared off.

Above the ankle, the distal ends of the tibia and fibula are united by a strong syndesmosis formed by the interosseous membrane and ligaments at the
distal tibiofibular joint. These connections prevent separation between the distal ends of the tibia and fibula and maintain the talus locked into position
between the medial malleolus and lateral malleolus. Injuries that produce a lateral twisting of the leg on top of the planted foot can result in stretching or
tearing of the tibiofibular ligaments, producing a syndesmotic ankle sprain or “high ankle sprain.”

Most ankle sprains can be treated using the RICE technique: Rest, Ice, Compression, and Elevation. Reducing joint mobility using a brace or cast may
be required for a period of time. More severe injuries involving ligament tears or bone fractures may require surgery.

Joints form during embryonic development in conjunction with the formation and growth of the associated bones. The embryonic tissue that gives rise to
all bones, cartilages, and connective tissues of the body is called mesenchyme. In the head, mesenchyme will accumulate at those areas that will
become the bones that form the top and sides of the skull. The mesenchyme in these areas will develop directly into bone through the process of
intramembranous ossification, in which mesenchymal cells differentiate into bone-producing cells that then generate bone tissue. The mesenchyme
between the areas of bone production will become the fibrous connective tissue that fills the spaces between the developing bones. Initially, the
connective tissue-filled gaps between the bones are wide, and are called fontanelles. After birth, as the skull bones grow and enlarge, the gaps between
them decrease in width and the fontanelles are reduced to suture joints in which the bones are united by a narrow layer of fibrous connective tissue.

The bones that form the base and facial regions of the skull develop through the process of endochondral ossification. In this process, mesenchyme
accumulates and differentiates into hyaline cartilage, which forms a model of the future bone. The hyaline cartilage model is then gradually, over a
period of many years, displaced by bone. The mesenchyme between these developing bones becomes the fibrous connective tissue of the suture joints
between the bones in these regions of the skull.

A similar process of endochondral ossification gives rises to the bones and joints of the limbs. The limbs initially develop as small limb buds that appear
on the sides of the embryo around the end of the fourth week of development. Starting during the sixth week, as each limb bud continues to grow and
elongate, areas of mesenchyme within the bud begin to differentiate into the hyaline cartilage that will form models for of each of the future bones. The
synovial joints will form between the adjacent cartilage models, in an area called the joint interzone. Cells at the center of this interzone region undergo
cell death to form the joint cavity, while surrounding mesenchyme cells will form the articular capsule and supporting ligaments. The process of
endochondral ossification, which converts the cartilage models into bone, begins by the twelfth week of embryonic development. At birth, ossification of
much of the bone has occurred, but the hyaline cartilage of the epiphyseal plate will remain throughout childhood and adolescence to allow for bone
lengthening. Hyaline cartilage is also retained as the articular cartilage that covers the surfaces of the bones at synovial joints.
CHAPTER 10

Muscle is one of the four primary tissue types of the body, and the body contains three
types of muscle tissue: skeletal muscle, cardiac muscle, and smooth muscle (Figure
10.2). All three muscle tissues have some properties in common; they all exhibit a
quality called excitability as their plasma membranes can change their electrical
states (from polarized to depolarized) and send an electrical wave called an action
potential along the entire length of the membrane. While the nervous system can
influence the excitability of cardiac and smooth muscle to some degree, skeletal
muscle completely depends on signaling from the nervous system to work properly.
On the other hand, both cardiac muscle and smooth muscle can respond to other
stimuli, such as hormones and local stimuli.

Figure 10.2 The Three Types of Muscle Tissue The body contains three types of

muscle tissue: (a) skeletal muscle, (b) smooth muscle, and (c) cardiac muscle.
(Micrographs provided by the Regents of University of Michigan Medical School ©
2012)

The muscles all begin the actual process of contracting (shortening) when a protein
called actin is pulled by a protein called myosin. This occurs in striated muscle
(skeletal and cardiac) after specific binding sites on the actin have been exposed in
response to the interaction between calcium ions (Ca++) and proteins (troponin and
tropomyosin) that “shield” the actin-binding sites. Ca++ also is required for the
contraction of smooth muscle, although its role is different: here Ca++ activates
enzymes, which in turn activate myosin heads. All muscles require adenosine
triphosphate (ATP) to continue the process of contracting, and they all relax when the
Ca++ is removed and the actin-binding sites are re-shielded.

A muscle can return to its original length when relaxed due to a quality of muscle
tissue called elasticity. It can recoil back to its original length due to elastic fibers.
Muscle tissue also has the quality of extensibility; it can stretch or
extend. Contractility allows muscle tissue to pull on its attachment points and shorten
with force.

Differences among the three muscle types include the microscopic organization of
their contractile proteins—actin and myosin. The actin and myosin proteins are
arranged very regularly in the cytoplasm of individual muscle cells (referred to as
fibers) in both skeletal muscle and cardiac muscle, which creates a pattern, or stripes,
called striations. The striations are visible with a light microscope under high
magnification (see Figure 10.2). Skeletal muscle fibers are multinucleated structures
that compose the skeletal muscle. Cardiac muscle fibers each have one to two nuclei
and are physically and electrically connected to each other so that the entire heart
contracts as one unit (called a syncytium).

Because the actin and myosin are not arranged in such regular fashion in smooth muscle, the cytoplasm of a smooth muscle fiber (which has only a
single nucleus) has a uniform, nonstriated appearance (resulting in the name smooth muscle). However, the less organized appearance of smooth
muscle should not be interpreted as less efficient. Smooth muscle in the walls of arteries is a critical component that regulates blood pressure necessary
to push blood through the circulatory system; and smooth muscle in the skin, visceral organs, and internal passageways is essential for moving all
materials through the body.

The best-known feature of skeletal muscle is its ability to contract and cause movement. Skeletal muscles act not only to produce movement but also to
stop movement, such as resisting gravity to maintain posture. Small, constant adjustments of the skeletal muscles are needed to hold a body upright or
balanced in any position. Muscles also prevent excess movement of the bones and joints, maintaining skeletal stability and preventing skeletal structure
damage or deformation. Joints can become misaligned or dislocated entirely by pulling on the associated bones; muscles work to keep joints stable.
Skeletal muscles are located throughout the body at the openings of internal tracts to control the movement of various substances. These muscles allow
functions, such as swallowing, urination, and defecation, to be under voluntary control. Skeletal muscles also protect internal organs (particularly
abdominal and pelvic organs) by acting as an external barrier or shield to external trauma and by supporting the weight of the organs.

Skeletal muscles contribute to the maintenance of homeostasis in the body by generating heat. Muscle contraction requires energy, and when ATP is
broken down, heat is produced. This heat is very noticeable during exercise, when sustained muscle movement causes body temperature to rise, and in
cases of extreme cold, when shivering produces random skeletal muscle contractions to generate heat.

Each skeletal muscle is an organ that consists of various integrated tissues. These tissues include the skeletal muscle fibers, blood vessels, nerve
fibers, and connective tissue. Each skeletal muscle has three layers of connective tissue (called “mysia”) that enclose it and provide structure to the
muscle as a whole, and also compartmentalize the muscle fibers within the muscle (Figure 10.3). Each muscle is wrapped in a sheath of dense, irregular
connective tissue called the epimysium, which allows a muscle to contract and move powerfully while maintaining its structural integrity. The epimysium
also separates muscle from other tissues and organs in the area, allowing the muscle to move independently.
 Figure 10.3 The Three Connective Tissue Layers Bundles of muscle fibers,
called fascicles, are covered by the perimysium. Muscle fibers are covered by
the endomysium.

Inside each skeletal muscle, muscle fibers are organized into individual
bundles, each called a fascicle, by a middle layer of connective tissue called
the perimysium. This fascicular organization is common in muscles of the
limbs; it allows the nervous system to trigger a specific movement of a muscle
by activating a subset of muscle fibers within a bundle, or fascicle of the
muscle. Inside each fascicle, each muscle fiber is encased in a thin connective
tissue layer of collagen and reticular fibers called the endomysium. The
endomysium contains the extracellular fluid and nutrients to support the
muscle fiber. These nutrients are supplied via blood to the muscle tissue.

In skeletal muscles that work with tendons to pull on bones, the collagen in the
three tissue layers (the mysia) intertwines with the collagen of a tendon. At the
other end of the tendon, it fuses with the periosteum coating the bone. The
tension created by contraction of the muscle fibers is then transferred though
the mysia, to the tendon, and then to the periosteum to pull on the bone for
movement of the skeleton. In other places, the mysia may fuse with a broad,
tendon-like sheet called an aponeurosis, or to fascia, the connective tissue
between skin and bones. The broad sheet of connective tissue in the lower
back that the latissimus dorsi muscles (the “lats”) fuse into is an example of an
aponeurosis.

Every skeletal muscle is also richly supplied by blood vessels for nourishment,
oxygen delivery, and waste removal. In addition, every muscle fiber in a skeletal muscle is supplied by the axon branch of a somatic motor neuron,
which signals the fiber to contract. Unlike cardiac and smooth muscle, the only way to functionally contract a skeletal muscle is through signaling from
the nervous system.

Skeletal Muscle Fibers

Because skeletal muscle cells are long and cylindrical, they are commonly referred to as muscle fibers. Skeletal muscle fibers can be quite large for
human cells, with diameters up to 100 μm and lengths up to 30 cm (11.8 in) in the Sartorius of the upper leg. During early development, embryonic
myoblasts, each with its own nucleus, fuse with up to hundreds of other myoblasts to form the multinucleated skeletal muscle fibers. Multiple nuclei
mean multiple copies of genes, permitting the production of the large amounts of proteins and enzymes needed for muscle contraction.

Some other terminology associated with muscle fibers is rooted in the Greek sarco, which means “flesh.” The plasma membrane of muscle fibers is
called the sarcolemma, the cytoplasm is referred to as sarcoplasm, and the specialized smooth endoplasmic reticulum, which stores, releases, and
retrieves calcium ions (Ca++) is called the sarcoplasmic reticulum (SR) (Figure 10.4). As will soon be described, the functional unit of a skeletal muscle
fiber is the sarcomere, a highly organized arrangement of the contractile myofilaments actin (thin filament) and myosin (thick filament), along with other
support proteins.

Figure 10.4 Muscle Fiber A skeletal muscle fiber is

surrounded by a plasma membrane called the sarcolemma,
which contains sarcoplasm, the cytoplasm of muscle cells. A
muscle fiber is composed of many fibrils, which give the cell
its striated appearance.

The Sarcomere

The striated appearance of skeletal muscle fibers is due to

the arrangement of the myofilaments of actin and myosin in
sequential order from one end of the muscle fiber to the
other. Each packet of these microfilaments and their
regulatory proteins, troponin and tropomyosin (along with
other proteins) is called a sarcomere.

The sarcomere is the functional unit of the muscle fiber. The

sarcomere itself is bundled within the myofibril that runs the
entire length of the muscle fiber and attaches to the
sarcolemma at its end. As myofibrils contract, the entire
muscle cell contracts. Because myofibrils are only
approximately 1.2 μm in diameter, hundreds to thousands
(each with thousands of sarcomeres) can be found inside
one muscle fiber. Each sarcomere is approximately 2 μm in
length with a three-dimensional cylinder-like arrangement
and is bordered by structures called Z-discs (also called Z-
lines, because pictures are two-dimensional), to which the actin myofilaments are anchored (Figure 10.5). Because the actin and its troponin-
tropomyosin complex (projecting from the Z-discs toward the center of the sarcomere) form strands that are thinner than the myosin, it is called the thin
filament of the sarcomere. Likewise, because the myosin strands and their multiple heads (projecting from the center of the sarcomere, toward but not
 all to way to, the Z-discs) have more mass and are thicker, they are called the thick
filament of the sarcomere.

Figure 10.5 The Sarcomere The sarcomere, the region from one Z-line to

the next Z-line, is the functional unit of a skeletal muscle fiber.

The Neuromuscular Junction

Another specialization of the skeletal muscle is the site where a motor

neuron’s terminal meets the muscle fiber—called the neuromuscular
junction (NMJ). This is where the muscle fiber first responds to signaling by
the motor neuron. Every skeletal muscle fiber in every skeletal muscle is
innervated by a motor neuron at the NMJ. Excitation signals from the neuron
are the only way to functionally activate the fiber to contract.

Excitation-Contraction Coupling

All living cells have membrane potentials, or electrical gradients across their
membranes. The inside of the membrane is usually around -60 to -90 mV,
relative to the outside. This is referred to as a cell’s membrane potential.
Neurons and muscle cells can use their membrane potentials to generate
electrical signals. They do this by controlling the movement of charged
particles, called ions, across their membranes to create electrical currents.
This is achieved by opening and closing specialized proteins in the
membrane called ion channels. Although the currents generated by ions
moving through these channel proteins are very small, they form the basis of
both neural signaling and muscle contraction.

Both neurons and skeletal muscle cells are electrically excitable, meaning that they
are able to generate action potentials. An action potential is a special type of
electrical signal that can travel along a cell membrane as a wave. This allows a signal
to be transmitted quickly and faithfully over long distances.

Although the term excitation-contraction coupling confuses or scares some

students, it comes down to this: for a skeletal muscle fiber to contract, its membrane
must first be “excited”—in other words, it must be stimulated to fire an action potential. The muscle fiber action potential, which sweeps along the
sarcolemma as a wave, is “coupled” to the actual contraction through the release of calcium ions (Ca++) from the SR. Once released, the Ca++ interacts
with the shielding proteins, forcing them to move aside so that the actin-binding sites are available for attachment by myosin heads. The myosin then
pulls the actin filaments toward the center, shortening the muscle fiber.

In skeletal muscle, this sequence begins with signals from the somatic motor division of the nervous system. In other words, the “excitation” step in
skeletal muscles is always triggered by signaling from the nervous system (Figure 10.6).

Figure 10.6 Motor End-Plate and Innervation At the NMJ, the axon terminal releases ACh. The motor end-plate is the location of the ACh-receptors in
the muscle fiber sarcolemma. When ACh molecules are released, they diffuse across a minute space called the synaptic cleft and bind to the receptors.

The motor neurons that tell the skeletal muscle fibers to contract originate in the spinal cord, with a smaller number located in the brainstem for
activation of skeletal muscles of the face, head, and neck. These neurons have long processes, called axons, which are specialized to transmit action
potentials long distances— in this case, all the way from the spinal cord to the muscle itself (which may be up to three feet away). The axons of multiple
neurons bundle together to form nerves, like wires bundled together in a cable.

Signaling begins when a neuronal action potential travels along the axon of a motor neuron, and then along the individual branches to terminate at the
NMJ. At the NMJ, the axon terminal releases a chemical messenger, or neurotransmitter, called acetylcholine (ACh). The ACh molecules diffuse
across a minute space called the synaptic cleft and bind to ACh receptors located within the motor end-plate of the sarcolemma on the other side of
the synapse. Once ACh binds, a channel in the ACh receptor opens and positively charged ions can pass through into the muscle fiber, causing it
to depolarize, meaning that the membrane potential of the muscle fiber becomes less negative (closer to zero.)

As the membrane depolarizes, another set of ion channels called voltage-gated sodium channels are triggered to open. Sodium ions enter the muscle
fiber, and an action potential rapidly spreads (or “fires”) along the entire membrane to initiate excitation-contraction coupling.

Things happen very quickly in the world of excitable membranes (just think about how quickly you can snap your fingers as soon as you decide to do it).
Immediately following depolarization of the membrane, it repolarizes, re-establishing the negative membrane potential. Meanwhile, the ACh in the
synaptic cleft is degraded by the enzyme acetylcholinesterase (AChE) so that the ACh cannot rebind to a receptor and reopen its channel, which would
cause unwanted extended muscle excitation and contraction.

Propagation of an action potential along the sarcolemma is the excitation portion of excitation-contraction coupling. Recall that this excitation actually
triggers the release of calcium ions (Ca++) from its storage in the cell’s SR. For the action potential to reach the membrane of the SR, there are periodic
invaginations in the sarcolemma, called T-tubules (“T” stands for “transverse”). You will recall that the diameter of a muscle fiber can be up to 100 μm,
 so these T-tubules ensure that the membrane can get close to the SR in the
sarcoplasm. The arrangement of a T-tubule with the membranes of SR on either side
is called a triad (Figure 10.7). The triad surrounds the cylindrical structure called
a myofibril, which contains actin and myosin.

Figure 10.7 The T-tubule Narrow T-tubules permit the conduction of

electrical impulses. The SR functions to regulate intracellular levels of
calcium. Two terminal cisternae (where enlarged SR connects to the T-
tubule) and one T-tubule comprise a triad—a “threesome” of membranes,
with those of SR on two sides and the T-tubule sandwiched between them.

The T-tubules carry the action potential into the interior of the cell, which
triggers the opening of calcium channels in the membrane of the adjacent
SR, causing Ca++ to diffuse out of the SR and into the sarcoplasm. It is the
arrival of Ca++ in the sarcoplasm that initiates contraction of the muscle fiber
by its contractile units, or sarcomeres.

The sequence of events that result in the contraction of an individual muscle

fiber begins with a signal—the neurotransmitter, ACh—from the motor
neuron innervating that fiber. The local membrane of the fiber will depolarize as
positively charged sodium ions (Na+) enter, triggering an action potential that spreads
to the rest of the membrane which will depolarize, including the T-tubules. This
triggers the release of calcium ions (Ca++) from storage in the sarcoplasmic reticulum
(SR). The Ca++ then initiates contraction, which is sustained by ATP (Figure 10.8). As
long as Ca++ ions remain in the sarcoplasm to bind to troponin, which keeps the actin-
binding sites “unshielded,” and as long as ATP is available to drive the cross-bridge
cycling and the pulling of actin strands by myosin, the muscle fiber will continue to shorten to an anatomical limit.

Figure 10.8 Contraction of a Muscle Fiber A cross-bridge forms between actin and the myosin heads triggering contraction. As long as Ca++ ions
remain in the sarcoplasm to bind to troponin, and as long as ATP is available, the muscle fiber will continue to shorten.

Muscle contraction usually stops when signaling from the motor neuron ends, which repolarizes the sarcolemma and T-tubules, and closes the voltage-
gated calcium channels in the SR. Ca++ ions are then pumped back into the SR, which causes the tropomyosin to reshield (or re-cover) the binding sites
on the actin strands. A muscle also can stop contracting when it runs out of ATP and becomes fatigued (Figure 10.9).

Figure 10.9 Relaxation of a Muscle Fiber Ca++ ions are pumped back into the

SR, which causes the tropomyosin to reshield the binding sites on the actin
strands. A muscle may also stop contracting when it runs out of ATP and
becomes fatigued.

The molecular events of muscle fiber shortening occur within the fiber’s
sarcomeres (see Figure 10.10). The contraction of a striated muscle fiber occurs
as the sarcomeres, linearly arranged within myofibrils, shorten as myosin heads
pull on the actin filaments.

The region where thick and thin filaments overlap has a dense appearance, as
there is little space between the filaments. This zone where thin and thick
filaments overlap is very important to muscle contraction, as it is the site where
filament movement starts. Thin filaments, anchored at their ends by the Z-discs,
do not extend completely into the central region that only contains thick
filaments, anchored at their bases at a spot called the M-line. A myofibril is
composed of many sarcomeres running along its length; thus, myofibrils and
muscle cells contract as the sarcomeres contract.

The Sliding Filament Model of Contraction

When signaled by a motor neuron, a skeletal muscle fiber contracts as the thin
filaments are pulled and then slide past the thick filaments within the fiber’s
sarcomeres. This process is known as the sliding filament model of muscle
contraction (Figure 10.10). The sliding can only occur when myosin-binding
sites on the actin filaments are exposed by a series of steps that begins with
Ca++ entry into the sarcoplasm.

Figure 10.10 The Sliding Filament Model of Muscle Contraction When a

sarcomere contracts, the Z lines move closer together, and the I band becomes
 smaller. The A band stays the same width. At full contraction, the thin and
thick filaments overlap completely.

Tropomyosin is a protein that winds around the chains of the actin filament
and covers the myosin-binding sites to prevent actin from binding to myosin.
Tropomyosin binds to troponin to form a troponin-tropomyosin complex. The
troponin-tropomyosin complex prevents the myosin “heads” from binding to
the active sites on the actin microfilaments. Troponin also has a binding site
for Ca++ ions.

To initiate muscle contraction, tropomyosin has to expose the myosin-

binding site on an actin filament to allow cross-bridge formation between the
actin and myosin microfilaments. The first step in the process of contraction
is for Ca++ to bind to troponin so that tropomyosin can slide away from the
binding sites on the actin strands. This allows the myosin heads to bind to
these exposed binding sites and form cross-bridges. The thin filaments are
then pulled by the myosin heads to slide past the thick filaments toward the
center of the sarcomere. But each head can only pull a very short distance
before it has reached its limit and must be “re-cocked” before it can pull
again, a step that requires ATP.

ATP and Muscle Contraction

For thin filaments to continue to slide past thick filaments during muscle
contraction, myosin heads must pull the actin at the binding sites, detach,
re-cock, attach to more binding sites, pull, detach, re-cock, etc. This
repeated movement is known as the cross-bridge cycle. This motion of the
myosin heads is similar to the oars when an individual rows a boat: The
paddle of the oars (the myosin heads) pull, are lifted from the water
(detach), repositioned (re-cocked) and then immersed again to pull (Figure
10.11). Each cycle requires energy, and the action of the myosin heads in
the sarcomeres repetitively pulling on the thin filaments also requires
energy, which is provided by ATP.

Figure 10.11 Skeletal Muscle Contraction (a) The active site on actin is

exposed as calcium binds to troponin. (b) The myosin head is attracted to actin, and myosin binds actin at its actin-binding site, forming the cross-bridge.
(c) During the power stroke, the phosphate generated in the previous contraction cycle is released. This results in the myosin head pivoting toward the
center of the sarcomere, after which the attached ADP and phosphate group are released. (d) A new molecule of ATP attaches to the myosin head,
causing the cross-bridge to detach. (e) The myosin head hydrolyzes ATP to ADP and phosphate, which returns the myosin to the cocked position.

Cross-bridge formation occurs when the myosin head attaches to the actin while adenosine diphosphate (ADP) and inorganic phosphate (P i) are still
bound to myosin (Figure 10.11a,b). Pi is then released, causing myosin to form a stronger attachment to the actin, after which the myosin head moves
toward the M-line, pulling the actin along with it. As actin is pulled, the filaments move approximately 10 nm toward the M-line. This movement is called
the power stroke, as movement of the thin filament occurs at this step (Figure 10.11c). In the absence of ATP, the myosin head will not detach from
actin.

One part of the myosin head attaches to the binding site on the actin, but the head has another binding site for ATP. ATP binding causes the myosin
head to detach from the actin (Figure 10.11d). After this occurs, ATP is converted to ADP and Pi by the intrinsic ATPase activity of myosin. The energy
released during ATP hydrolysis changes the angle of the myosin head into a cocked position (Figure 10.11e). The myosin head is now in position for
further movement.

When the myosin head is cocked, myosin is in a high-energy configuration. This energy is expended as the myosin head moves through the power
stroke, and at the end of the power stroke, the myosin head is in a low-energy position. After the power stroke, ADP is released; however, the formed
cross-bridge is still in place, and actin and myosin are bound together. As long as ATP is available, it readily attaches to myosin, the cross-bridge cycle
can recur, and muscle contraction can continue.

Note that each thick filament of roughly 300 myosin molecules has multiple myosin heads, and many cross-bridges form and break continuously during
muscle contraction. Multiply this by all of the sarcomeres in one myofibril, all the myofibrils in one muscle fiber, and all of the muscle fibers in one
skeletal muscle, and you can understand why so much energy (ATP) is needed to keep skeletal muscles working. In fact, it is the loss of ATP that
results in the rigor mortis observed soon after someone dies. With no further ATP production possible, there is no ATP available for myosin heads to
detach from the actin-binding sites, so the cross-bridges stay in place, causing the rigidity in the skeletal muscles.

Sources of ATP

ATP supplies the energy for muscle contraction to take place. In addition to its direct role in the cross-bridge cycle, ATP also provides the energy for the
active-transport Ca++ pumps in the SR. Muscle contraction does not occur without sufficient amounts of ATP. The amount of ATP stored in muscle is
very low, only sufficient to power a few seconds worth of contractions. As it is broken down, ATP must therefore be regenerated and replaced quickly to
allow for sustained contraction. There are three mechanisms by which ATP can be regenerated: creatine phosphate metabolism, anaerobic glycolysis,
and fermentation and aerobic respiration.

Creatine phosphate is a molecule that can store energy in its phosphate bonds. In a resting muscle, excess ATP transfers its energy to creatine,
producing ADP and creatine phosphate. This acts as an energy reserve that can be used to quickly create more ATP. When the muscle starts to
contract and needs energy, creatine phosphate transfers its phosphate back to ADP to form ATP and creatine. This reaction is catalyzed by the enzyme
creatine kinase and occurs very quickly; thus, creatine phosphate-derived ATP powers the first few seconds of muscle contraction. However, creatine
phosphate can only provide approximately 15 seconds worth of energy, at which point another energy source has to be used (Figure 10.12).

Figure 10.12 Muscle Metabolism (a) Some ATP is stored in a resting muscle. As

contraction starts, it is used up in seconds. More ATP is generated from creatine
phosphate for about 15 seconds. (b) Each glucose molecule produces two ATP and
two molecules of pyruvic acid, which can be used in aerobic respiration or converted to
lactic acid. If oxygen is not available, pyruvic acid is converted to lactic acid, which may
contribute to muscle fatigue. This occurs during strenuous exercise when high amounts
of energy are needed but oxygen cannot be sufficiently delivered to muscle. (c) Aerobic
respiration is the breakdown of glucose in the presence of oxygen (O2) to produce
carbon dioxide, water, and ATP. Approximately 95 percent of the ATP required for
resting or moderately active muscles is provided by aerobic respiration, which takes
place in mitochondria.

As the ATP produced by creatine phosphate is depleted, muscles turn to glycolysis as

an ATP source. Glycolysis is an anaerobic (non-oxygen-dependent) process that
breaks down glucose (sugar) to produce ATP; however, glycolysis cannot generate
ATP as quickly as creatine phosphate. Thus, the switch to glycolysis results in a slower
rate of ATP availability to the muscle. The sugar used in glycolysis can be provided by
blood glucose or by metabolizing glycogen that is stored in the muscle. The breakdown
of one glucose molecule produces two ATP and two molecules of pyruvic acid, which
can be used in aerobic respiration or when oxygen levels are low, converted to lactic
acid (Figure 10.12b).

If oxygen is available, pyruvic acid is used in aerobic respiration. However, if oxygen is

not available, pyruvic acid is converted to lactic acid, which may contribute to muscle
fatigue. This conversion allows the recycling of the enzyme NAD+ from NADH, which is needed for glycolysis to continue. This occurs during strenuous
exercise when high amounts of energy are needed but oxygen cannot be sufficiently delivered to muscle. Glycolysis itself cannot be sustained for very
long (approximately 1 minute of muscle activity), but it is useful in facilitating short bursts of high-intensity output. This is because glycolysis does not
utilize glucose very efficiently, producing a net gain of two ATPs per molecule of glucose, and the end product of lactic acid, which may contribute to
muscle fatigue as it accumulates.

Aerobic respiration is the breakdown of glucose or other nutrients in the presence of oxygen (O2) to produce carbon dioxide, water, and ATP.
Approximately 95 percent of the ATP required for resting or moderately active muscles is provided by aerobic respiration, which takes place in
mitochondria. The inputs for aerobic respiration include glucose circulating in the bloodstream, pyruvic acid, and fatty acids. Aerobic respiration is much
more efficient than anaerobic glycolysis, producing approximately 36 ATPs per molecule of glucose versus four from glycolysis. However, aerobic
respiration cannot be sustained without a steady supply of O2 to the skeletal muscle and is much slower (Figure 10.12c). To compensate, muscles store
small amount of excess oxygen in proteins call myoglobin, allowing for more efficient muscle contractions and less fatigue. Aerobic training also
increases the efficiency of the circulatory system so that O2 can be supplied to the muscles for longer periods of time.

Muscle fatigue occurs when a muscle can no longer contract in response to signals from the nervous system. The exact causes of muscle fatigue are
not fully known, although certain factors have been correlated with the decreased muscle contraction that occurs during fatigue. ATP is needed for
normal muscle contraction, and as ATP reserves are reduced, muscle function may decline. This may be more of a factor in brief, intense muscle output
rather than sustained, lower intensity efforts. Lactic acid buildup may lower intracellular pH, affecting enzyme and protein activity. Imbalances in Na + and
K+ levels as a result of membrane depolarization may disrupt Ca++ flow out of the SR. Long periods of sustained exercise may damage the SR and the
sarcolemma, resulting in impaired Ca++ regulation.

Intense muscle activity results in an oxygen debt, which is the amount of oxygen needed to compensate for ATP produced without oxygen during
muscle contraction. Oxygen is required to restore ATP and creatine phosphate levels, convert lactic acid to pyruvic acid, and, in the liver, to convert
lactic acid into glucose or glycogen. Other systems used during exercise also require oxygen, and all of these combined processes result in the
increased breathing rate that occurs after exercise. Until the oxygen debt has been met, oxygen intake is elevated, even after exercise has stopped.

Relaxation of a Skeletal Muscle

Relaxing skeletal muscle fibers, and ultimately, the skeletal muscle, begins with the motor neuron, which stops releasing its chemical signal, ACh, into
the synapse at the NMJ. The muscle fiber will repolarize, which closes the gates in the SR where Ca++ was being released. ATP-driven pumps will move
Ca++ out of the sarcoplasm back into the SR. This results in the “reshielding” of the actin-binding sites on the thin filaments. Without the ability to form
cross-bridges between the thin and thick filaments, the muscle fiber loses its tension and relaxes.

Muscle Strength
The number of skeletal muscle fibers in a given muscle is genetically determined and does not change. Muscle strength is directly related to the amount
of myofibrils and sarcomeres within each fiber. Factors, such as hormones and stress (and artificial anabolic steroids), acting on the muscle can
increase the production of sarcomeres and myofibrils within the muscle fibers, a change called hypertrophy, which results in the increased mass and
bulk in a skeletal muscle. Likewise, decreased use of a skeletal muscle results in atrophy, where the number of sarcomeres and myofibrils disappear
(but not the number of muscle fibers). It is common for a limb in a cast to show atrophied muscles when the cast is removed, and certain diseases, such
as polio, show atrophied muscles.

DISORDERS OF THE...

Muscular System

Duchenne muscular dystrophy (DMD) is a progressive weakening of the skeletal muscles. It is one of several diseases collectively referred to as
“muscular dystrophy.” DMD is caused by a lack of the protein dystrophin, which helps the thin filaments of myofibrils bind to the sarcolemma. Without
sufficient dystrophin, muscle contractions cause the sarcolemma to tear, causing an influx of Ca++, leading to cellular damage and muscle fiber
degradation. Over time, as muscle damage accumulates, muscle mass is lost, and greater functional impairments develop.

DMD is an inherited disorder caused by an abnormal X chromosome. It primarily affects males, and it is usually diagnosed in early childhood. DMD
usually first appears as difficulty with balance and motion, and then progresses to an inability to walk. It continues progressing upward in the body from
the lower extremities to the upper body, where it affects the muscles responsible for breathing and circulation. It ultimately causes death due to
respiratory failure, and those afflicted do not usually live past their 20s.

Because DMD is caused by a mutation in the gene that codes for dystrophin, it was thought that introducing healthy myoblasts into patients might be an
effective treatment. Myoblasts are the embryonic cells responsible for muscle development, and ideally, they would carry healthy genes that could
produce the dystrophin needed for normal muscle contraction. This approach has been largely unsuccessful in humans. A recent approach has involved
attempting to boost the muscle’s production of utrophin, a protein similar to dystrophin that may be able to assume the role of dystrophin and prevent
cellular damage from occurring.

To move an object, referred to as load, the sarcomeres in the muscle fibers of the skeletal muscle must shorten. The force generated by the contraction
of the muscle (or shortening of the sarcomeres) is called muscle tension. However, muscle tension also is generated when the muscle is contracting
against a load that does not move, resulting in two main types of skeletal muscle contractions: isotonic contractions and isometric contractions.

In isotonic contractions, where the tension in the muscle stays constant, a load is moved as the length of the muscle changes (shortens). There are
two types of isotonic contractions: concentric and eccentric. A concentric contraction involves the muscle shortening to move a load. An example of
this is the biceps brachii muscle contracting when a hand weight is brought upward with increasing muscle tension. As the biceps brachii contract, the
angle of the elbow joint decreases as the forearm is brought toward the body. Here, the biceps brachii contracts as sarcomeres in its muscle fibers are
shortening and cross-bridges form; the myosin heads pull the actin. An eccentric contraction occurs as the muscle tension diminishes and the muscle
lengthens. In this case, the hand weight is lowered in a slow and controlled manner as the amount of cross-bridges being activated by nervous system
stimulation decreases. In this case, as tension is released from the biceps brachii, the angle of the elbow joint increases. Eccentric contractions are also
used for movement and balance of the body.

An isometric contraction occurs as the muscle produces tension without changing the angle of a skeletal joint. Isometric contractions involve
sarcomere shortening and increasing muscle tension, but do not move a load, as the force produced cannot overcome the resistance provided by the
load. For example, if one attempts to lift a hand weight that is too heavy, there will be sarcomere activation and shortening to a point, and ever-
increasing muscle tension, but no change in the angle of the elbow joint. In everyday living, isometric contractions are active in maintaining posture and
maintaining bone and joint stability. However, holding your head in an upright position occurs not because the muscles cannot move the head, but
because the goal is to remain stationary and not produce movement. Most actions of the body are the result of a combination of isotonic and isometric
contractions working together to produce a wide range of outcomes (Figure 10.13).

Figure 10.13 Types of Muscle Contractions During isotonic contractions, muscle length

changes to move a load. During isometric contractions, muscle length does not change because
the load exceeds the tension the muscle can generate.

All of these muscle activities are under the exquisite control of the nervous system. Neural
control regulates concentric, eccentric and isometric contractions, muscle fiber recruitment, and
muscle tone. A crucial aspect of nervous system control of skeletal muscles is the role of motor
units.

Motor Units

As you have learned, every skeletal muscle fiber must be innervated by the axon terminal of a
motor neuron in order to contract. Each muscle fiber is innervated by only one motor neuron.
The actual group of muscle fibers in a muscle innervated by a single motor neuron is called
a motor unit. The size of a motor unit is variable depending on the nature of the muscle.

A small motor unit is an arrangement where a single motor neuron supplies a small number of
muscle fibers in a muscle. Small motor units permit very fine motor control of the muscle. The
best example in humans is the small motor units of the extraocular eye muscles that move the
eyeballs. There are thousands of muscle fibers in each muscle, but every six or so fibers are
supplied by a single motor neuron, as the axons branch to form synaptic connections at their individual NMJs. This allows for exquisite control of eye
movements so that both eyes can quickly focus on the same object. Small motor units are also involved in the many fine movements of the fingers and
thumb of the hand for grasping, texting, etc.

A large motor unit is an arrangement where a single motor neuron supplies a large number of muscle fibers in a muscle. Large motor units are
concerned with simple, or “gross,” movements, such as powerfully extending the knee joint. The best example is the large motor units of the thigh
muscles or back muscles, where a single motor neuron will supply thousands of muscle fibers in a muscle, as its axon splits into thousands of branches.

There is a wide range of motor units within many skeletal muscles, which gives the nervous system a wide range of control over the muscle. The small
motor units in the muscle will have smaller, lower-threshold motor neurons that are more excitable, firing first to their skeletal muscle fibers, which also
tend to be the smallest. Activation of these smaller motor units, results in a relatively small degree of contractile strength (tension) generated in the
muscle. As more strength is needed, larger motor units, with bigger, higher-threshold motor neurons are enlisted to activate larger muscle fibers. This
increasing activation of motor units produces an increase in muscle contraction known as recruitment. As more motor units are recruited, the muscle
contraction grows progressively stronger. In some muscles, the largest motor units may generate a contractile force of 50 times more than the smallest
motor units in the muscle. This allows a feather to be picked up using the biceps brachii arm muscle with minimal force, and a heavy weight to be lifted
by the same muscle by recruiting the largest motor units.

When necessary, the maximal number of motor units in a muscle can be recruited simultaneously, producing the maximum force of contraction for that
muscle, but this cannot last for very long because of the energy requirements to sustain the contraction. To prevent complete muscle fatigue, motor units
are generally not all simultaneously active, but instead some motor units rest while others are active, which allows for longer muscle contractions. The
nervous system uses recruitment as a mechanism to efficiently utilize a skeletal muscle.

The Length-Tension Range of a Sarcomere

When a skeletal muscle fiber contracts, myosin heads attach to actin to form cross-bridges followed by the thin filaments sliding over the thick filaments
as the heads pull the actin, and this results in sarcomere shortening, creating the tension of the muscle contraction. The cross-bridges can only form
where thin and thick filaments already overlap, so that the length of the sarcomere has a direct influence on the force generated when the sarcomere
shortens. This is called the length-tension relationship.

The ideal length of a sarcomere to produce maximal tension occurs at 80 percent to 120 percent of its resting length, with 100 percent being the state
where the medial edges of the thin filaments are just at the most-medial myosin heads of the thick filaments (Figure 10.14). This length maximizes the
overlap of actin-binding sites and myosin heads. If a sarcomere is stretched past this ideal length (beyond 120 percent), thick and thin filaments do not
overlap sufficiently, which results in less tension produced. If a sarcomere is shortened beyond 80 percent, the zone of overlap is reduced with the thin
filaments jutting beyond the last of the myosin heads and shrinks the H zone, which is normally composed of myosin tails. Eventually, there is nowhere
else for the thin filaments to go and the amount of tension is diminished. If the muscle is stretched to the point where thick and thin filaments do not
overlap at all, no cross-bridges can be formed, and no tension is produced in that sarcomere. This amount of stretching does not usually occur, as
accessory proteins and connective tissue oppose extreme stretching.

Figure 10.14 The Ideal Length of a Sarcomere Sarcomeres produce

maximal tension when thick and thin filaments overlap between about 80
percent to 120 percent.

The Frequency of Motor Neuron Stimulation

A single action potential from a motor neuron will produce a single contraction
in the muscle fibers of its motor unit. This isolated contraction is called
a twitch. A twitch can last for a few milliseconds or 100 milliseconds,
depending on the muscle type. The tension produced by a single twitch can
be measured by a myogram, an instrument that measures the amount of
tension produced over time (Figure 10.15). Each twitch undergoes three
phases. The first phase is the latent period, during which the action potential
is being propagated along the sarcolemma and Ca++ ions are released from
the SR. This is the phase during which excitation and contraction are being
coupled but contraction has yet to occur. The contraction phase occurs
next. The Ca++ ions in the sarcoplasm have bound to troponin, tropomyosin
has shifted away from actin-binding sites, cross-bridges formed, and
sarcomeres are actively shortening to the point of peak tension. The
last phase is the relaxation phase, when tension decreases as
contraction stops. Ca++ ions are pumped out of the sarcoplasm into
the SR, and cross-bridge cycling stops, returning the muscle fibers to
their resting state.

Figure 10.15 A Myogram of a Muscle Twitch A single muscle

twitch has a latent period, a contraction phase when tension
increases, and a relaxation phase when tension decreases. During
the latent period, the action potential is being propagated along the
sarcolemma. During the contraction phase, Ca++ ions in the
sarcoplasm bind to troponin, tropomyosin moves from actin-binding
sites, cross-bridges form, and sarcomeres shorten. During the
relaxation phase, tension decreases as Ca++ ions are pumped out of
the sarcoplasm and cross-bridge cycling stops.
Although a person can experience a muscle “twitch,” a single twitch does not produce any significant muscle activity in a living body. A series of action
potentials to the muscle fibers is necessary to produce a muscle contraction that can produce work. Normal muscle contraction is more sustained, and it
can be modified by input from the nervous system to produce varying amounts of force; this is called a graded muscle response. The frequency of
action potentials (nerve impulses) from a motor neuron and the number of motor neurons transmitting action potentials both affect the tension produced
in skeletal muscle.

The rate at which a motor neuron fires action potentials affects the tension produced in the skeletal muscle. If the fibers are stimulated while a previous
twitch is still occurring, the second twitch will be stronger. This response is called wave summation, because the excitation-contraction coupling effects
of successive motor neuron signaling is summed, or added together (Figure 10.16a). At the molecular level, summation occurs because the second
stimulus triggers the release of more Ca++ ions, which become available to activate additional sarcomeres while the muscle is still contracting from the
first stimulus. Summation results in greater contraction of the motor unit.

Figure 10.16 Wave Summation and Tetanus (a) The excitation-

contraction coupling effects of successive motor neuron signaling is
added together which is referred to as wave summation. The bottom
of each wave, the end of the relaxation phase, represents the point of
stimulus. (b) When the stimulus frequency is so high that the
relaxation phase disappears completely, the contractions become
continuous; this is called tetanus.

If the frequency of motor neuron signaling increases, summation and

subsequent muscle tension in the motor unit continues to rise until it
reaches a peak point. The tension at this point is about three to four
times greater than the tension of a single twitch, a state referred to as incomplete tetanus. During incomplete tetanus, the muscle goes through quick
cycles of contraction with a short relaxation phase for each. If the stimulus frequency is so high that the relaxation phase disappears completely,
contractions become continuous in a process called complete tetanus (Figure 10.16b).

During tetanus, the concentration of Ca++ ions in the sarcoplasm allows virtually all of the sarcomeres to form cross-bridges and shorten, so that a
contraction can continue uninterrupted (until the muscle fatigues and can no longer produce tension).

Treppe

When a skeletal muscle has been dormant for an extended period and then activated to contract, with all other things being equal, the initial contractions
generate about one-half the force of later contractions. The muscle tension increases in a graded manner that to some looks like a set of stairs. This
tension increase is called treppe, a condition where muscle contractions become more efficient. It’s also known as the “staircase effect” (Figure 10.17).

Figure 10.17 Treppe When muscle tension increases in a graded manner that looks like a set of stairs, it is called treppe. The bottom of each wave
represents the point of stimulus.

It is believed that treppe results from a higher concentration of Ca++ in the sarcoplasm resulting from the
steady stream of signals from the motor neuron. It can only be maintained with adequate ATP.

Muscle Tone

Skeletal muscles are rarely completely relaxed, or flaccid. Even if a muscle is not producing movement,
it is contracted a small amount to maintain its contractile proteins and produce muscle tone. The
tension produced by muscle tone allows muscles to continually stabilize joints and maintain posture.

Muscle tone is accomplished by a complex interaction between the nervous system and skeletal
muscles that results in the activation of a few motor units at a time, most likely in a cyclical manner. In
this manner, muscles never fatigue completely, as some motor units can recover while others are
active.

The absence of the low-level contractions that lead to muscle tone is referred to as hypotonia, and can
result from damage to parts of the central nervous system (CNS), such as the cerebellum, or from loss of innervations to a skeletal muscle, as in
poliomyelitis. Hypotonic muscles have a flaccid appearance and display functional impairments, such as weak reflexes. Conversely, excessive muscle
tone is referred to as hypertonia, accompanied by hyperreflexia (excessive reflex responses), often the result of damage to upper motor neurons in the
CNS. Hypertonia can present with muscle rigidity (as seen in Parkinson’s disease) or spasticity, a phasic change in muscle tone, where a limb will “snap”
back from passive stretching (as seen in some strokes).

Two criteria to consider when classifying the types of muscle fibers are how fast some fibers contract relative to others, and how fibers produce ATP.
Using these criteria, there are three main types of skeletal muscle fibers. Slow oxidative (SO) fibers contract relatively slowly and use aerobic
respiration (oxygen and glucose) to produce ATP. Fast oxidative (FO) fibers have fast contractions and primarily use aerobic respiration, but because
they may switch to anaerobic respiration (glycolysis), can fatigue more quickly than SO fibers. Lastly, fast glycolytic (FG) fibers have fast contractions
and primarily use anaerobic glycolysis. The FG fibers fatigue more quickly than the others. Most skeletal muscles in a human contain(s) all three types,
although in varying proportions.
The speed of contraction is dependent on how quickly myosin’s ATPase hydrolyzes ATP to produce cross-bridge action. Fast fibers hydrolyze ATP
approximately twice as quickly as slow fibers, resulting in much quicker cross-bridge cycling (which pulls the thin filaments toward the center of the
sarcomeres at a faster rate). The primary metabolic pathway used by a muscle fiber determines whether the fiber is classified as oxidative or glycolytic.
If a fiber primarily produces ATP through aerobic pathways it is oxidative. More ATP can be produced during each metabolic cycle, making the fiber
more resistant to fatigue. Glycolytic fibers primarily create ATP through anaerobic glycolysis, which produces less ATP per cycle. As a result, glycolytic
fibers fatigue at a quicker rate.

The oxidative fibers contain many more mitochondria than the glycolytic fibers, because aerobic metabolism, which uses oxygen (O 2) in the metabolic
pathway, occurs in the mitochondria. The SO fibers possess a large number of mitochondria and are capable of contracting for longer periods because
of the large amount of ATP they can produce, but they have a relatively small diameter and do not produce a large amount of tension. SO fibers are
extensively supplied with blood capillaries to supply O2 from the red blood cells in the bloodstream. The SO fibers also possess myoglobin, an O2-
carrying molecule similar to O2-carrying hemoglobin in the red blood cells. The myoglobin stores some of the needed O 2 within the fibers themselves
(and gives SO fibers their red color). All of these features allow SO fibers to produce large quantities of ATP, which can sustain muscle activity without
fatiguing for long periods of time.

The fact that SO fibers can function for long periods without fatiguing makes them useful in maintaining posture, producing isometric contractions,
stabilizing bones and joints, and making small movements that happen often but do not require large amounts of energy. They do not produce high
tension, and thus they are not used for powerful, fast movements that require high amounts of energy and rapid cross-bridge cycling.

FO fibers are sometimes called intermediate fibers because they possess characteristics that are intermediate between fast fibers and slow fibers. They
produce ATP relatively quickly, more quickly than SO fibers, and thus can produce relatively high amounts of tension. They are oxidative because they
produce ATP aerobically, possess high amounts of mitochondria, and do not fatigue quickly. However, FO fibers do not possess significant myoglobin,
giving them a lighter color than the red SO fibers. FO fibers are used primarily for movements, such as walking, that require more energy than postural
control but less energy than an explosive movement, such as sprinting. FO fibers are useful for this type of movement because they produce more
tension than SO fibers but they are more fatigue-resistant than FG fibers.

FG fibers primarily use anaerobic glycolysis as their ATP source. They have a large diameter and possess high amounts of glycogen, which is used in
glycolysis to generate ATP quickly to produce high levels of tension. Because they do not primarily use aerobic metabolism, they do not possess
substantial numbers of mitochondria or significant amounts of myoglobin and therefore have a white color. FG fibers are used to produce rapid, forceful
contractions to make quick, powerful movements. These fibers fatigue quickly, permitting them to only be used for short periods. Most muscles possess
a mixture of each fiber type. The predominant fiber type in a muscle is determined by the primary function of the muscle.

Physical training alters the appearance of skeletal muscles and can produce changes in muscle performance. Conversely, a lack of use can result in
decreased performance and muscle appearance. Although muscle cells can change in size, new cells are not formed when muscles grow. Instead,
structural proteins are added to muscle fibers in a process called hypertrophy, so cell diameter increases. The reverse, when structural proteins are lost
and muscle mass decreases, is called atrophy. Age-related muscle atrophy is called sarcopenia. Cellular components of muscles can also undergo
changes in response to changes in muscle use.

Endurance Exercise

Slow fibers are predominantly used in endurance exercises that require little force but involve numerous repetitions. The aerobic metabolism used by
slow-twitch fibers allows them to maintain contractions over long periods. Endurance training modifies these slow fibers to make them even more
efficient by producing more mitochondria to enable more aerobic metabolism and more ATP production. Endurance exercise can also increase the
amount of myoglobin in a cell, as increased aerobic respiration increases the need for oxygen. Myoglobin is found in the sarcoplasm and acts as an
oxygen storage supply for the mitochondria.

The training can trigger the formation of more extensive capillary networks around the fiber, a process called angiogenesis, to supply oxygen and
remove metabolic waste. To allow these capillary networks to supply the deep portions of the muscle, muscle mass does not greatly increase in order to
maintain a smaller area for the diffusion of nutrients and gases. All of these cellular changes result in the ability to sustain low levels of muscle
contractions for greater periods without fatiguing.

The proportion of SO muscle fibers in muscle determines the suitability of that muscle for endurance, and may benefit those participating in endurance
activities. Postural muscles have a large number of SO fibers and relatively few FO and FG fibers, to keep the back straight (Figure 10.18). Endurance
athletes, like marathon-runners also would benefit from a larger proportion of SO fibers, but it is unclear if the most-successful marathoners are those
with naturally high numbers of SO fibers, or whether the most successful marathon runners develop high numbers of SO fibers with repetitive training.
Endurance training can result in overuse injuries such as stress fractures and joint and tendon inflammation.

Figure 10.18 Marathoners Long-distance runners have a large number of SO fibers and relatively few FO and FG fibers. (credit: “Tseo2”/Wikimedia
Commons)

Resistance Exercise

Resistance exercises, as opposed to endurance exercise, require large amounts of FG fibers to produce short, powerful movements that are not
repeated over long periods. The high rates of ATP hydrolysis and cross-bridge formation in FG fibers result in powerful muscle contractions. Muscles
used for power have a higher ratio of FG to SO/FO fibers, and trained athletes possess even higher levels of FG fibers in their muscles. Resistance
exercise affects muscles by increasing the formation of myofibrils, thereby increasing the thickness of muscle fibers. This added structure causes
hypertrophy, or the enlargement of muscles, exemplified by the large skeletal muscles seen in body builders and other athletes (Figure 10.19). Because
this muscular enlargement is achieved by the addition of structural proteins, athletes trying to build muscle mass often ingest large amounts of protein.
Figure 10.19 Hypertrophy Body builders have a large number of FG fibers and relatively few FO and SO fibers. (credit: Lin Mei/flickr)

Except for the hypertrophy that follows an increase in the number of sarcomeres and myofibrils in a skeletal muscle, the cellular changes observed
during endurance training do not usually occur with resistance training. There is usually no significant increase in mitochondria or capillary density.
However, resistance training does increase the development of connective tissue, which adds to the overall mass of the muscle and helps to contain
muscles as they produce increasingly powerful contractions. Tendons also become stronger to prevent tendon damage, as the force produced by
muscles is transferred to tendons that attach the muscle to bone.

For effective strength training, the intensity of the exercise must continually be increased. For instance, continued weight lifting without increasing the
weight of the load does not increase muscle size. To produce ever-greater results, the weights lifted must become increasingly heavier, making it more
difficult for muscles to move the load. The muscle then adapts to this heavier load, and an even heavier load must be used if even greater muscle mass
is desired.

If done improperly, resistance training can lead to overuse injuries of the muscle, tendon, or bone. These injuries can occur if the load is too heavy or if
the muscles are not given sufficient time between workouts to recover or if joints are not aligned properly during the exercises. Cellular damage to
muscle fibers that occurs after intense exercise includes damage to the sarcolemma and myofibrils. This muscle damage contributes to the feeling of
soreness after strenuous exercise, but muscles gain mass as this damage is repaired, and additional structural proteins are added to replace the
damaged ones. Overworking skeletal muscles can also lead to tendon damage and even skeletal damage if the load is too great for the muscles to bear.

Performance-Enhancing Substances

Some athletes attempt to boost their performance by using various agents that may enhance muscle performance. Anabolic steroids are one of the
more widely known agents used to boost muscle mass and increase power output. Anabolic steroids are a form of testosterone, a male sex hormone
that stimulates muscle formation, leading to increased muscle mass.

Endurance athletes may also try to boost the availability of oxygen to muscles to increase aerobic respiration by using substances such as erythropoietin
(EPO), a hormone normally produced in the kidneys, which triggers the production of red blood cells. The extra oxygen carried by these blood cells can
then be used by muscles for aerobic respiration. Human growth hormone (hGH) is another supplement, and although it can facilitate building muscle
mass, its main role is to promote the healing of muscle and other tissues after strenuous exercise. Increased hGH may allow for faster recovery after
muscle damage, reducing the rest required after exercise, and allowing for more sustained high-level performance.

Although performance-enhancing substances often do improve performance, most are banned by governing bodies in sports and are illegal for
nonmedical purposes. Their use to enhance performance raises ethical issues of cheating because they give users an unfair advantage over nonusers.
A greater concern, however, is that their use carries serious health risks. The side effects of these substances are often significant, nonreversible, and in
some cases fatal. The physiological strain caused by these substances is often greater than what the body can handle, leading to effects that are
unpredictable and dangerous. Anabolic steroid use has been linked to infertility, aggressive behavior, cardiovascular disease, and brain cancer.

Similarly, some athletes have used creatine to increase power output. Creatine phosphate provides quick bursts of ATP to muscles in the initial stages of
contraction. Increasing the amount of creatine available to cells is thought to produce more ATP and therefore increase explosive power output,
although its effectiveness as a supplement has been questioned.

EVERYDAY CONNECTION

Aging and Muscle Tissue

Although atrophy due to disuse can often be reversed with exercise, muscle atrophy with age, referred to as sarcopenia, is irreversible. This is a primary
reason why even highly trained athletes succumb to declining performance with age. This decline is noticeable in athletes whose sports require strength
and powerful movements, such as sprinting, whereas the effects of age are less noticeable in endurance athletes such as marathon runners or long-
distance cyclists. As muscles age, muscle fibers die, and they are replaced by connective tissue and adipose tissue (Figure 10.20). Because those
tissues cannot contract and generate force as muscle can, muscles lose the ability to produce powerful contractions. The decline in muscle mass causes
a loss of strength, including the strength required for posture and mobility. This may be caused by a reduction in FG fibers that hydrolyze ATP quickly to
produce short, powerful contractions. Muscles in older people sometimes possess greater numbers of SO fibers, which are responsible for longer
contractions and do not produce powerful movements. There may also be a reduction in the size of motor units, resulting in fewer fibers being stimulated
and less muscle tension being produced.

Sarcopenia can be delayed to some extent by exercise, as training adds structural proteins and causes cellular changes that can offset the effects of
atrophy. Increased exercise can produce greater numbers of cellular mitochondria, increase capillary density, and increase the mass and strength of
connective tissue. The effects of age-related atrophy are especially pronounced in people who are sedentary, as the loss of muscle cells is displayed as
functional impairments such as trouble with locomotion, balance, and posture. This can lead to a decrease in quality of life and medical problems, such
as joint problems because the muscles that stabilize bones and joints are weakened. Problems with locomotion and balance can also cause various
injuries due to falls.
Cardiac muscle tissue is only found in the heart. Highly
coordinated contractions of cardiac muscle pump blood
into the vessels of the circulatory system. Similar to
skeletal muscle, cardiac muscle is striated and organized
into sarcomeres, possessing the same banding
organization as skeletal muscle (Figure 10.21). However,
cardiac muscle fibers are shorter than skeletal muscle
fibers and usually contain only one nucleus, which is
located in the central region of the cell. Cardiac muscle
fibers also possess many mitochondria and myoglobin,
as ATP is produced primarily through aerobic
metabolism. Cardiac muscle fibers cells also are
extensively branched and are connected to one another
at their ends by intercalated discs. An intercalated
disc allows the cardiac muscle cells to contract in a
wave-like pattern so that the heart can work as a pump.

Intercalated discs are part of the sarcolemma and contain two structures important
in cardiac muscle contraction: gap junctions and desmosomes. A gap junction forms
channels between adjacent cardiac muscle fibers that allow the depolarizing current
produced by cations to flow from one cardiac muscle cell to the next. This joining is
called electric coupling, and in cardiac muscle it allows the quick transmission of
action potentials and the coordinated contraction of the entire heart. This network of
electrically connected cardiac muscle cells creates a functional unit of contraction
called a syncytium. The remainder of the intercalated disc is composed of
desmosomes. A desmosome is a cell structure that anchors the ends of cardiac
muscle fibers together so the cells do not pull apart during the stress of individual
fibers contracting (Figure 10.22).

Figure 10.22 Cardiac Muscle Intercalated discs are part of the cardiac muscle

sarcolemma and they contain gap junctions and desmosomes.

Contractions of the heart (heartbeats) are controlled by specialized cardiac muscle cells called pacemaker cells that directly control heart rate. Although
cardiac muscle cannot be consciously controlled, the pacemaker cells respond to signals from the autonomic nervous system (ANS) to speed up or slow
down the heart rate. The pacemaker cells can also respond to various hormones that modulate heart rate to control blood pressure.

The wave of contraction that allows the heart to work as a unit, called a functional syncytium, begins with the pacemaker cells. This group of cells is self-
excitable and able to depolarize to threshold and fire action potentials on their own, a feature called autorhythmicity; they do this at set intervals which
determine heart rate. Because they are connected with gap junctions to surrounding muscle fibers and the specialized fibers of the heart’s conduction
system, the pacemaker cells are able to transfer the depolarization to the other cardiac muscle fibers in a manner that allows the heart to contract in a
coordinated manner.

Another feature of cardiac muscle is its relatively long action potentials in its fibers, having a sustained depolarization “plateau.” The plateau is produced
by Ca++ entry though voltage-gated calcium channels in the sarcolemma of cardiac muscle fibers. This sustained depolarization (and Ca ++ entry)
provides for a longer contraction than is produced by an action potential in skeletal muscle. Unlike skeletal muscle, a large percentage of the Ca ++ that
initiates contraction in cardiac muscles comes from outside the cell rather than from the SR.

Smooth muscle (so-named because the cells do not have striations) is present
in the walls of hollow organs like the urinary bladder, uterus, stomach,
intestines, and in the walls of passageways, such as the arteries and veins of
the circulatory system, and the tracts of the respiratory, urinary, and
reproductive systems (Figure 10.23ab). Smooth muscle is also present in the
eyes, where it functions to change the size of the iris and alter the shape of the
lens; and in the skin where it causes hair to stand erect in response to cold
temperature or fear.

Smooth muscle fibers are spindle-shaped (wide in the middle and tapered at
both ends, somewhat like a football) and have a single nucleus; they range from
about 30 to 200 μm (thousands of times shorter than skeletal muscle fibers),
and they produce their own connective tissue, endomysium. Although they do
not have striations and sarcomeres, smooth muscle fibers do have actin and
myosin contractile proteins, and thick and thin filaments. These thin filaments
are anchored by dense bodies. A dense body is analogous to the Z-discs of
skeletal and cardiac muscle fibers and is fastened to the sarcolemma. Calcium
ions are supplied by the SR in the fibers and by sequestration from the
extracellular fluid through membrane indentations called calveoli.

Because smooth muscle cells do not contain troponin, cross-bridge formation is

not regulated by the troponin-tropomyosin complex but instead by the
regulatory protein calmodulin. In a smooth muscle fiber, external Ca++ ions passing through opened calcium channels in the sarcolemma, and additional
Ca++ released from SR, bind to calmodulin. The Ca++-calmodulin complex then activates an enzyme called myosin (light chain) kinase, which, in turn,
activates the myosin heads by phosphorylating them (converting ATP to ADP and Pi, with the Pi attaching to the head). The heads can then attach to
actin-binding sites and pull on the thin filaments. The thin filaments also are anchored to the dense bodies; the structures invested in the inner
membrane of the sarcolemma (at adherens junctions) that also have cord-like intermediate filaments attached to them. When the thin filaments slide
past the thick filaments, they pull on the dense bodies, structures tethered to the sarcolemma, which then pull on the intermediate filaments networks
throughout the sarcoplasm. This arrangement causes the entire muscle fiber to contract in a manner whereby the ends are pulled toward the center,
causing the midsection to bulge in a corkscrew motion (Figure 10.24).

Figure 10.24 Muscle Contraction The dense bodies and

intermediate filaments are networked through the
sarcoplasm, which cause the muscle fiber to contract.

Although smooth muscle contraction relies on the

presence of Ca++ ions, smooth muscle fibers have a much
smaller diameter than skeletal muscle cells. T-tubules are not required to reach the interior of the cell and therefore not necessary to transmit an action
potential deep into the fiber. Smooth muscle fibers have a limited calcium-storing SR but have calcium channels in the sarcolemma (similar to cardiac
muscle fibers) that open during the action potential along the sarcolemma. The influx of extracellular Ca++ ions, which diffuse into the sarcoplasm to
reach the calmodulin, accounts for most of the Ca++ that triggers contraction of a smooth muscle cell.

Muscle contraction continues until ATP-dependent calcium pumps actively transport Ca++ ions back into the SR and out of the cell. However, a low
concentration of calcium remains in the sarcoplasm to maintain muscle tone. This remaining calcium keeps the muscle slightly contracted, which is
important in certain tracts and around blood vessels.

Because most smooth muscles must function for long periods without rest, their power output is relatively low, but contractions can continue without
using large amounts of energy. Some smooth muscle can also maintain contractions even as Ca++ is removed and myosin kinase is
inactivated/dephosphorylated. This can happen as a subset of cross-bridges between myosin heads and actin, called latch-bridges, keep the thick and
thin filaments linked together for a prolonged period, and without the need for ATP. This allows for the maintaining of muscle “tone” in smooth muscle
that lines arterioles and other visceral organs with very little energy expenditure.

Smooth muscle is not under voluntary control; thus, it is called involuntary muscle. The triggers for smooth muscle contraction include hormones, neural
stimulation by the ANS, and local factors. In certain locations, such as the walls of visceral organs, stretching the muscle can trigger its contraction (the
stress-relaxation response).

Axons of neurons in the ANS do not form the highly organized NMJs with smooth muscle, as seen between motor neurons and skeletal muscle fibers.
Instead, there is a series of neurotransmitter-filled bulges called varicosities as an axon courses through smooth muscle, loosely forming motor units
(Figure 10.25). A varicosity releases neurotransmitters into the synaptic cleft. Also, visceral muscle in the walls of the hollow organs (except the heart)
contains pacesetter cells. A pacesetter cell can spontaneously trigger action potentials and contractions in the muscle.

Figure 10.25 Motor Units A series of axon-like swelling, called

varicosities or “boutons,” from autonomic neurons form motor
units through the smooth muscle.

Smooth muscle is organized in two ways: as single-unit smooth

muscle, which is much more common; and as multiunit smooth
muscle. The two types have different locations in the body and
have different characteristics. Single-unit muscle has its muscle
fibers joined by gap junctions so that the muscle contracts as a
single unit. This type of smooth muscle is found in the walls of
all visceral organs except the heart (which has cardiac muscle
in its walls), and so it is commonly called visceral muscle.
Because the muscle fibers are not constrained by the
organization and stretchability limits of sarcomeres, visceral smooth muscle has a stress-relaxation response. This means that as the muscle of a
hollow organ is stretched when it fills, the mechanical stress of the stretching will trigger contraction, but this is immediately followed by relaxation so that
the organ does not empty its contents prematurely. This is important for hollow organs, such as the stomach or urinary bladder, which continuously
expand as they fill. The smooth muscle around these organs also can maintain a muscle tone when the organ empties and shrinks, a feature that
prevents “flabbiness” in the empty organ. In general, visceral smooth muscle produces slow, steady contractions that allow substances, such as food in
the digestive tract, to move through the body.

Multiunit smooth muscle cells rarely possess gap junctions, and thus are not electrically coupled. As a result, contraction does not spread from one cell
to the next, but is instead confined to the cell that was originally stimulated. Stimuli for multiunit smooth muscles come from autonomic nerves or
hormones but not from stretching. This type of tissue is found around large blood vessels, in the respiratory airways, and in the eyes.

Hyperplasia in Smooth Muscle

Similar to skeletal and cardiac muscle cells, smooth muscle can undergo hypertrophy to increase in size. Unlike other muscle, smooth muscle can also
divide to produce more cells, a process called hyperplasia. This can most evidently be observed in the uterus at puberty, which responds to increased
estrogen levels by producing more uterine smooth muscle fibers, and greatly increases the size of the myometrium.
Most muscle tissue of the body arises from embryonic mesoderm. Paraxial mesodermal cells adjacent to the neural tube form blocks of cells
called somites. Skeletal muscles, excluding those of the head and limbs, develop from mesodermal somites, whereas skeletal muscle in the head and
limbs develop from general mesoderm. Somites give rise to myoblasts. A myoblast is a muscle-forming stem cell that migrates to different regions in the
body and then fuse(s) to form a syncytium, or myotube. As a myotube is formed from many different myoblast cells, it contains many nuclei, but has a
continuous cytoplasm. This is why skeletal muscle cells are multinucleate, as the nucleus of each contributing myoblast remains intact in the mature
skeletal muscle cell. However, cardiac and smooth muscle cells are not multinucleate because the myoblasts that form their cells do not fuse.

Gap junctions develop in the cardiac and single-unit smooth muscle in the early stages of development. In skeletal muscles, ACh receptors are initially
present along most of the surface of the myoblasts, but spinal nerve innervation causes the release of growth factors that stimulate the formation of
motor end-plates and NMJs. As neurons become active, electrical signals that are sent through the muscle influence the distribution of slow and fast
fibers in the muscle.

Although the number of muscle cells is set during development, satellite cells help to repair skeletal muscle cells. A satellite cell is similar to a myoblast
because it is a type of stem cell; however, satellite cells are incorporated into muscle cells and facilitate the protein synthesis required for repair and
growth. These cells are located outside the sarcolemma and are stimulated to grow and fuse with muscle cells by growth factors that are released by
muscle fibers under certain forms of stress. Satellite cells can regenerate muscle fibers to a very limited extent, but they primarily help to repair damage
in living cells. If a cell is damaged to a greater extent than can be repaired by satellite cells, the muscle fibers are replaced by scar tissue in a process
called fibrosis. Because scar tissue cannot contract, muscle that has sustained significant damage loses strength and cannot produce the same
amount of power or endurance as it could before being damaged.

Smooth muscle tissue can regenerate from a type of stem cell called a pericyte, which is found in some small blood vessels. Pericytes allow smooth
muscle cells to regenerate and repair much more readily than skeletal and cardiac muscle tissue. Similar to skeletal muscle tissue, cardiac muscle does
not regenerate to a great extent. Dead cardiac muscle tissue is replaced by scar tissue, which cannot contract. As scar tissue accumulates, the heart
loses its ability to pump because of the loss of contractile power. However, some minor regeneration may occur due to stem cells found in the blood that
occasionally enter cardiac tissue.

acetylcholine (ACh)
neurotransmitter that binds at a motor end-plate to trigger depolarization
actin
protein that makes up most of the thin myofilaments in a sarcomere muscle fiber
action potential
change in voltage of a cell membrane in response to a stimulus that results in transmission of an electrical signal; unique to neurons and
muscle fibers
aerobic respiration
production of ATP in the presence of oxygen
angiogenesis
formation of blood capillary networks
aponeurosis
broad, tendon-like sheet of connective tissue that attaches a skeletal muscle to another skeletal muscle or to a bone
ATPase
enzyme that hydrolyzes ATP to ADP
atrophy
loss of structural proteins from muscle fibers
autorhythmicity
heart’s ability to control its own contractions
calmodulin
regulatory protein that facilitates contraction in smooth muscles
cardiac muscle
striated muscle found in the heart; joined to one another at intercalated discs and under the regulation of pacemaker cells, which contract as
one unit to pump blood through the circulatory system. Cardiac muscle is under involuntary control.
concentric contraction
muscle contraction that shortens the muscle to move a load
contractility
ability to shorten (contract) forcibly
contraction phase
twitch contraction phase when tension increases
creatine phosphate
phosphagen used to store energy from ATP and transfer it to muscle
dense body
sarcoplasmic structure that attaches to the sarcolemma and shortens the muscle as thin filaments slide past thick filaments
depolarize
to reduce the voltage difference between the inside and outside of a cell’s plasma membrane (the sarcolemma for a muscle fiber), making the
inside less negative than at rest
desmosome
cell structure that anchors the ends of cardiac muscle fibers to allow contraction to occur
eccentric contraction
muscle contraction that lengthens the muscle as the tension is diminished
elasticity
ability to stretch and rebound
endomysium
loose, and well-hydrated connective tissue covering each muscle fiber in a skeletal muscle
epimysium
outer layer of connective tissue around a skeletal muscle
excitability
 ability to undergo neural stimulation
excitation-contraction coupling
sequence of events from motor neuron signaling to a skeletal muscle fiber to contraction of the fiber’s sarcomeres
extensibility
ability to lengthen (extend)
fascicle
bundle of muscle fibers within a skeletal muscle
fast glycolytic (FG)
muscle fiber that primarily uses anaerobic glycolysis
fast oxidative (FO)
intermediate muscle fiber that is between slow oxidative and fast glycolytic fibers
fibrosis
replacement of muscle fibers by scar tissue
glycolysis
anaerobic breakdown of glucose to ATP
graded muscle response
modification of contraction strength
hyperplasia
process in which one cell splits to produce new cells
hypertonia
abnormally high muscle tone
hypertrophy
addition of structural proteins to muscle fibers
hypotonia
abnormally low muscle tone caused by the absence of low-level contractions
intercalated disc
part of the sarcolemma that connects cardiac tissue, and contains gap junctions and desmosomes
isometric contraction
muscle contraction that occurs with no change in muscle length
isotonic contraction
muscle contraction that involves changes in muscle length
lactic acid
product of anaerobic glycolysis
latch-bridges
subset of a cross-bridge in which actin and myosin remain locked together
latent period
the time when a twitch does not produce contraction
motor end-plate
sarcolemma of muscle fiber at the neuromuscular junction, with receptors for the neurotransmitter acetylcholine
motor unit
motor neuron and the group of muscle fibers it innervates
muscle tension
force generated by the contraction of the muscle; tension generated during isotonic contractions and isometric contractions
muscle tone
low levels of muscle contraction that occur when a muscle is not producing movement
myoblast
muscle-forming stem cell
myofibril
long, cylindrical organelle that runs parallel within the muscle fiber and contains the sarcomeres
myogram
instrument used to measure twitch tension
myosin
protein that makes up most of the thick cylindrical myofilament within a sarcomere muscle fiber
myotube
fusion of many myoblast cells
neuromuscular junction (NMJ)
synapse between the axon terminal of a motor neuron and the section of the membrane of a muscle fiber with receptors for the acetylcholine
released by the terminal
neurotransmitter
signaling chemical released by nerve terminals that bind to and activate receptors on target cells
oxygen debt
amount of oxygen needed to compensate for ATP produced without oxygen during muscle contraction
pacesetter cell
cell that triggers action potentials in smooth muscle
pericyte
stem cell that regenerates smooth muscle cells
perimysium
connective tissue that bundles skeletal muscle fibers into fascicles within a skeletal muscle
power stroke
action of myosin pulling actin inward (toward the M line)
pyruvic acid
product of glycolysis that can be used in aerobic respiration or converted to lactic acid
recruitment
increase in the number of motor units involved in contraction
relaxation phase
period after twitch contraction when tension decreases
sarcolemma
plasma membrane of a skeletal muscle fiber
sarcomere
longitudinally, repeating functional unit of skeletal muscle, with all of the contractile and associated proteins involved in contraction
sarcopenia
age-related muscle atrophy
sarcoplasm
cytoplasm of a muscle cell
sarcoplasmic reticulum (SR)
specialized smooth endoplasmic reticulum, which stores, releases, and retrieves Ca++
satellite cell
stem cell that helps to repair muscle cells
skeletal muscle
striated, multinucleated muscle that requires signaling from the nervous system to trigger contraction; most skeletal muscles are referred to as
voluntary muscles that move bones and produce movement
slow oxidative (SO)
muscle fiber that primarily uses aerobic respiration
smooth muscle
nonstriated, mononucleated muscle in the skin that is associated with hair follicles; assists in moving materials in the walls of internal organs,
blood vessels, and internal passageways
somites
blocks of paraxial mesoderm cells
stress-relaxation response
relaxation of smooth muscle tissue after being stretched
synaptic cleft
space between a nerve (axon) terminal and a motor end-plate
T-tubule
projection of the sarcolemma into the interior of the cell
tetanus
a continuous fused contraction
thick filament
the thick myosin strands and their multiple heads projecting from the center of the sarcomere toward, but not all to way to, the Z-discs
thin filament
thin strands of actin and its troponin-tropomyosin complex projecting from the Z-discs toward the center of the sarcomere
treppe
stepwise increase in contraction tension
triad
the grouping of one T-tubule and two terminal cisternae
tropomyosin
regulatory protein that covers myosin-binding sites to prevent actin from binding to myosin
troponin
regulatory protein that binds to actin, tropomyosin, and calcium
twitch
single contraction produced by one action potential
varicosity
enlargement of neurons that release neurotransmitters into synaptic clefts
visceral muscle
smooth muscle found in the walls of visceral organs
voltage-gated sodium channels
membrane proteins that open sodium channels in response to a sufficient voltage change, and initiate and transmit the action potential as
Na+ enters through the channel
wave summation
addition of successive neural stimuli to produce greater contraction

To move the skeleton, the tension created by the contraction of the fibers in most skeletal muscles is transferred to the tendons. The tendons are strong
bands of dense, regular connective tissue that connect muscles to bones. The bone connection is why this muscle tissue is called skeletal muscle.

Interactions of Skeletal Muscles in the Body

To pull on a bone, that is, to change the angle at its synovial joint, which essentially moves the skeleton, a skeletal muscle must also be attached to a fixed part of the
skeleton. The moveable end of the muscle that attaches to the bone being pulled is called the muscle’s insertion, and the end of the muscle attached to a fixed
(stabilized) bone is called the origin. During forearm flexion—bending the elbow—the
brachioradialis assists the brachialis.

Although a number of muscles may be involved in an action, the principal muscle involved is
called the prime mover, or agonist. To lift a cup, a muscle called the biceps brachii is actually
the prime mover; however, because it can be assisted by the brachialis, the brachialis is called
a synergist in this action (Figure 11.2). A synergist can also be a fixator that stabilizes the
bone that is the attachment for the prime mover’s origin.

Figure 11.2 Prime Movers and Synergists The biceps brachii flex the lower arm. The
brachoradialis, in the forearm, and brachialis, located deep to the biceps in the upper arm,
are both synergists that aid in this motion.
A muscle with the opposite action of the prime mover is called an antagonist. Antagonists play two important roles in muscle function: (1) they maintain body or limb
position, such as holding the arm out or standing erect; and (2) they control rapid movement, as in shadow boxing without landing a punch or the ability to check the
motion of a limb.

For example, to extend the knee, a group of four muscles called the quadriceps femoris in the anterior compartment of the thigh are activated (and would be called
the agonists of knee extension). However, to flex the knee joint, an opposite or antagonistic set of muscles called the hamstrings is activated.

As you can see, these terms would also be reversed for the opposing action. If you consider the first action as the knee bending, the hamstrings would be called the
agonists and the quadriceps femoris would then be called the antagonists. See Table 11.1 for a list of some agonists and antagonists.

Agonist and Antagonist Skeletal Muscle Pairs

Agonist Antagonist Movement

Triceps brachii: in the posterior compartment of

Biceps brachii: in the anterior compartment of the arm The biceps brachii flexes the forearm, whereas
the arm

Hamstrings: group of three muscles in the posterior Quadriceps femoris: group of four muscles in the
The hamstrings flex the leg, whereas the quad
compartment of the thigh anterior compartment of the thigh

Flexor digitorum superficialis and flexor digitorum profundus: Extensor digitorum: in the posterior compartment The flexor digitorum superficialis and flexor dig
in the anterior compartment of the forearm of the forearm wrist, whereas the extensor digitorum extends

Table11.1

There are also skeletal muscles that do not pull against the skeleton for movements. For example, there are the muscles that produce facial expressions. The
insertions and origins of facial muscles are in the skin, so that certain individual muscles contract to form a smile or frown, form sounds or words, and raise the
eyebrows. There also are skeletal muscles in the tongue, and the external urinary and anal sphincters that allow for voluntary regulation of urination and defecation,
respectively. In addition, the diaphragm contracts and relaxes to change the volume of the pleural cavities but it does not move the skeleton to do this.

Patterns of Fascicle Organization

Skeletal muscle is enclosed in connective tissue scaffolding at three levels. Each muscle fiber (cell) is covered by endomysium and the entire muscle is covered by
epimysium. When a group of muscle fibers is “bundled” as a unit within the whole muscle by an additional covering of a connective tissue called perimysium, that
bundled group of muscle fibers is called a fascicle. Fascicle arrangement by perimysia is correlated to the force generated by a muscle; it also affects the range of
motion of the muscle. Based on the patterns of fascicle arrangement, skeletal muscles can be classified in several ways. What follows are the most common fascicle
arrangements.

Parallel muscles have fascicles that are arranged in the same

direction as the long axis of the muscle (Figure 11.3). The majority
of skeletal muscles in the body have this type of organization. Some
parallel muscles are flat sheets that expand at the ends to make
broad attachments. Other parallel muscles are rotund with tendons
at one or both ends. Muscles that seem to be plump have a large
mass of tissue located in the middle of the muscle, between the
insertion and the origin, which is known as the central body. A more
common name for this muscle is belly. When a muscle contracts,
the contractile fibers shorten it to an even larger bulge. For
example, extend and then flex your biceps brachii muscle; the large,
middle section is the belly (Figure 11.4). When a parallel muscle has
a central, large belly that is spindle-shaped, meaning it tapers as it
extends to its origin and insertion, it sometimes is called fusiform.

Figure 11.3 Muscle Shapes and Fiber Alignment The skeletal

muscles of the body typically come in seven different general
shapes.

Circular muscles are also called sphincters (see Figure 11.3). When

they relax, the sphincters’ concentrically arranged bundles of
muscle fibers increase the size of the opening, and when they
contract, the size of the opening shrinks to the point of closure. The
orbicularis oris muscle is a circular muscle that goes around the
mouth. When it contracts, the oral opening becomes smaller, as when puckering the lips for whistling. Another example is the orbicularis oculi, one of which
surrounds each eye. Consider, for example, the names of the two orbicularis muscles (orbicularis oris and oribicularis oculi), where part of the first name of both
muscles is the same. The first part of orbicularis, orb (orb = “circular”), is a reference to a round or circular structure; it may also make one think of orbit, such as the
moon’s path around the earth. The word oris (oris = “oral”) refers to the oral cavity, or the mouth. The word oculi (ocular = “eye”) refers to the eye.

There are other muscles throughout the body named by their shape or location. The deltoid is a large, triangular-shaped muscle that covers the shoulder. It is so-
named because the Greek letter delta looks like a triangle. The rectus abdominis (rector = “straight”) is the straight muscle in the anterior wall of the abdomen, while
the rectus femoris is the straight muscle in the anterior compartment of the thigh.

When a muscle has a widespread expansion over a sizable area, but then the fascicles come to a single, common attachment point, the muscle is called convergent.
The attachment point for a convergent muscle could be a tendon, an aponeurosis (a flat, broad tendon), or a raphe (a very slender tendon). The large muscle on the
chest, the pectoralis major, is an example of a convergent muscle because it converges on the greater tubercle of the humerus via a tendon. The temporalis muscle of
the cranium is another.

Pennate muscles (penna = “feathers”) blend into a tendon that runs through the central region of the muscle for its whole length, somewhat like the quill of a feather
with the muscle arranged similar to the feathers. Due to this design, the muscle fibers in a pennate muscle can only pull at an angle, and as a result, contracting
pennate muscles do not move their tendons very far. However, because a pennate muscle generally can hold more muscle fibers within it, it can produce relatively
more tension for its size. There are three subtypes of pennate muscles.

In a unipennate muscle, the fascicles are located on one side of the tendon. The extensor digitorum of the forearm is an example of a unipennate muscle.
A bipennate muscle has fascicles on both sides of the tendon. In some pennate muscles, the muscle fibers wrap around the tendon, sometimes forming individual
fascicles in the process. This arrangement is referred to as multipennate. A common example is the deltoid muscle of the shoulder, which covers the shoulder but has
a single tendon that inserts on the deltoid tuberosity of the humerus.

Because of fascicles, a portion of a multipennate muscle like the deltoid can be stimulated by the nervous system to change the direction of the pull. For example,
when the deltoid muscle contracts, the arm abducts (moves away from midline in the sagittal plane), but when only the anterior fascicle is stimulated, the arm
will abduct and flex (move anteriorly at the shoulder joint).

The Lever System of Muscle and Bone Interactions

Skeletal muscles do not work by themselves. Muscles are arranged in pairs based on their functions. For muscles attached to the bones of the skeleton, the
connection determines the force, speed, and range of movement. These characteristics depend on each other and can explain the general organization of the
muscular and skeletal systems.

The skeleton and muscles act together to move the body. Have you ever used the back of a hammer to remove a nail from wood? The handle acts as a lever and the
head of the hammer acts as a fulcrum, the fixed point that the force is applied to when you pull back or push down on the handle. The effort applied to this system is
the pulling or pushing on the handle to remove the nail, which is the load, or “resistance” to the movement of the handle in the system. Our musculoskeletal system
works in a similar manner, with bones being stiff levers and the articular endings of the bones—encased in synovial joints—acting as fulcrums. The load would be an
object being lifted or any resistance to a movement (your head is a load when you are lifting it), and the effort, or applied force, comes from contracting skeletal
muscle.

The Greeks and Romans conducted the first studies done on the human body in Western culture. The educated class of subsequent societies studied
Latin and Greek, and therefore the early pioneers of anatomy continued to apply Latin and Greek terminology or roots when they named the skeletal
muscles. The large number of muscles in the body and unfamiliar words can make learning the names of the muscles in the body seem daunting, but
understanding the etymology can help. Etymology is the study of how the root of a particular word entered a language and how the use of the word
evolved over time. Taking the time to learn the root of the words is crucial to understanding the vocabulary of anatomy and physiology. When you
understand the names of muscles it will help you remember where the muscles are located and what they do (Figure 11.5, Figure 11.6, and Table 11.2).
Pronunciation of words and terms will take a bit of time to master, but after you have some basic information; the correct names and pronunciations will
become easier.

Figure 11.5 Overview of the Muscular System On the anterior and posterior views of the muscular system above, superficial muscles (those at the
surface) are shown on the right side of the body while deep muscles (those underneath the superficial muscles) are shown on the left half of the body.
For the legs, superficial muscles are shown in the anterior view while the posterior view shows both superficial and deep muscles.

Figure 11.6 Understanding a Muscle Name from the Latin

Mnemonic Device for Latin Roots

Example Latin or Greek Translation Mnemonic Device

ad to; toward ADvance toward your goal

ab away from n/a

sub under SUBmarines move under water.

 Example Latin or Greek Translation Mnemonic Device

ductor something that moves A conDUCTOR makes a train move.

anti against If you are antisocial, you are against engaging in social activities.

epi on top of n/a

apo to the side of n/a

longissimus longest “Longissimus” is longer than the word “long.”

longus long long

brevis short brief

maximus large max

medius medium “Medius” and “medium” both begin with “med.”

minimus tiny; little mini

rectus straight To RECTify a situation is to straighten it out.

multi many If something is MULTIcolored, it has many colors.

uni one A UNIcorn has one horn.

bi/di two If a ring is DIcast, it is made of two metals.

tri three TRIple the amount of money is three times as much.

quad four QUADruplets are four children born at one birth.

externus outside EXternal

internus inside INternal

Table11.2

Anatomists name the skeletal muscles according to a number of criteria, each of which describes the muscle in some way. These include naming the
muscle after its shape, its size compared to other muscles in the area, its location in the body or the location of its attachments to the skeleton, how
many origins it has, or its action.

The skeletal muscle’s anatomical location or its relationship to a particular bone often determines its name. For example, the frontalis muscle is located
on top of the frontal bone of the skull. Similarly, the shapes of some muscles are very distinctive and the names, such as orbicularis, reflect the shape.
For the buttocks, the size of the muscles influences the names: gluteus maximus (largest), gluteus medius (medium), and the
gluteus minimus (smallest). Names were given to indicate length—brevis (short), longus (long)—and to identify position relative to the
midline: lateralis (to the outside away from the midline), and medialis (toward the midline). The direction of the muscle fibers and fascicles are used to
describe muscles relative to the midline, such as the rectus (straight) abdominis, or the oblique (at an angle) muscles of the abdomen.

Some muscle names indicate the number of muscles in a group. One example of this is the quadriceps, a group of four muscles located on the anterior
(front) thigh. Other muscle names can provide information as to how many origins a particular muscle has, such as the biceps brachii. The
prefix bi indicates that the muscle has two origins and tri indicates three origins.

The location of a muscle’s attachment can also appear in its name. When the name of a muscle is based on the attachments, the origin is always named
first. For instance, the sternocleidomastoid muscle of the neck has a dual origin on the sternum (sterno) and clavicle (cleido), and it inserts on the
mastoid process of the temporal bone. The last feature by which to name a muscle is its action. When muscles are named for the movement they
produce, one can find action words in their name. Some examples are flexor (decreases the angle at the joint), extensor (increases the angle at the
joint), abductor (moves the bone away from the midline), or adductor (moves the bone toward the midline).

The skeletal muscles are divided into axial (muscles of the trunk and head) and appendicular (muscles of the arms and legs) categories. This system
reflects the bones of the skeleton system, which are also arranged in this manner. The axial muscles are grouped based on location, function, or both.
Some of the axial muscles may seem to blur the boundaries because they cross over to the appendicular skeleton. The first grouping of the axial
muscles you will review includes the muscles of the head and neck, then you will review the muscles of the vertebral column, and finally you will review
the oblique and rectus muscles.
 Muscles That Create Facial Expression

The origins of the muscles of facial expression are on the surface of the
skull (remember, the origin of a muscle does not move). The insertions
of these muscles have fibers intertwined with connective tissue and the
dermis of the skin. Because the muscles insert in the skin rather than
on bone, when they contract, the skin moves to create facial
expression (Figure 11.7).

Figure 11.7 Muscles of Facial Expression Many of the muscles of

facial expression insert into the skin surrounding the eyelids, nose and
mouth, producing facial expressions by moving the skin rather than
bones.

The orbicularis oris is a circular muscle that moves the lips, and

the orbicularis oculi is a circular muscle that closes the eye.
The occipitofrontalis muscle moves up the scalp and eyebrows. The
muscle has a frontal belly and an occipital (near the occipital bone on the posterior part of the skull) belly. In other words, there is a muscle on the
forehead (frontalis) and one on the back of the head (occipitalis), but there is no muscle across the top of the head. Instead, the two bellies are
connected by a broad tendon called the epicranial aponeurosis, or galea aponeurosis (galea = “helmet”). The physicians originally studying human
anatomy thought the skull looked like an helmet.

A large portion of the face is composed of the buccinator muscle, which compresses the cheek. This muscle allows you to whistle, blow, and suck; and
it contributes to the action of chewing. There are several small facial muscles, one of which is the corrugator supercilii, which is the prime mover of the
eyebrows. Place your finger on your eyebrows at the point of the bridge of the nose. Raise your eyebrows as if you were surprised and lower your
eyebrows as if you were frowning. With
these movements, you can feel the action of
the corrugator supercilii. Additional muscles
of facial

expression are presented in Figure 11.8.

Figure 11.8 Muscles in Facial Expression

Muscles That Move the Eyes

The movement of the eyeball is under the control of

the extrinsic eye muscles, which originate outside
the eye and insert onto the outer surface of the white
of the eye. These muscles are located inside the eye
socket and cannot be seen on any part of the visible
eyeball (Figure 11.9 and Table 11.3). If you have
ever been to a doctor who held up a finger and asked
you to follow it up, down, and to both sides, he or she
is checking to make sure your eye muscles are acting
in a coordinated pattern.

Figure 11.9 Muscles of the Eyes (a) The extrinsic

eye muscles originate outside of the eye on the skull.
(b) Each muscle inserts onto the eyeball.

Muscles of the Eyes

 Movement Target Target motion direction Prime mover Origin

Moves eyes up and toward nose; rotates eyes from Superior (elevates); medial Common tendinous ring (ring
Eyeballs Superior rectus
1 o’clock to 3 o’clock (adducts) attaches to optic foramen)

Moves eyes down and toward nose; rotates eyes Inferior (depresses); medial Common tendinous ring (ring
Eyeballs Inferior rectus
from 6 o’clock to 3 o’clock (adducts) attaches to optic foramen)

Common tendinous ring (ring

Moves eyes away from nose Eyeballs Lateral (abducts) Lateral rectus
attaches to optic foramen)

Common tendinous ring (ring

Moves eyes toward nose Eyeballs Medial (adducts) Medial rectus
attaches to optic foramen)

Moves eyes up and away from nose; rotates eyeball Superior (elevates); lateral
Eyeballs Inferior oblique Floor of orbit (maxilla)
from 12 o’clock to 9 o’clock (abducts)

Moves eyes down and away from nose; rotates Inferior (depress); lateral
Eyeballs Superior oblique Sphenoid bone
eyeball from 6 o’clock to 9 o’clock (abducts)

Upper Levator palpabrae

Opens eyes Superior (elevates) Roof of orbit (sphenoid bone)
eyelid superioris

Compression along superior–

Closes eyelids Eyelid skin Orbicularis oculi Medial bones composing the o
inferior axis

Table11.3

Muscles That Move the Lower Jaw

In anatomical terminology, chewing is called mastication. Muscles involved in chewing must be able to exert enough pressure to bite through and then
chew food before it is swallowed (Figure 11.10 and Table 11.4). The masseter muscle is the main muscle used for chewing because it elevates the
mandible (lower jaw) to close the mouth, and it is assisted by the temporalis muscle, which retracts the mandible. You can feel the temporalis move by
putting your fingers to your temple as you chew.

Figure 11.10 Muscles That Move the Lower Jaw The muscles that move the lower jaw are typically located within the cheek and originate from
processes in the skull. This provides the jaw muscles with the large amount of leverage needed for chewing.

Muscles of the Lower Jaw

Movement Target Target motion direction Prime mover Origin

Closes mouth; aids chewing Mandible Superior (elevates) Masseter Maxilla ar

 Movement Target Target motion direction Prime mover Origin

(for mass

Closes mouth; pulls lower jaw in under upper jaw Mandible Superior (elevates); posterior (retracts) Temporalis Temporal

Opens mouth; pushes lower jaw out under upper jaw; Inferior (depresses); posterior (protracts); lateral Lateral Pterygoid
Mandible
moves lower jaw side-to-side (abducts); medial (adducts) pterygoid bone

Closes mouth; pushes lower jaw out under upper jaw; Superior (elevates); posterior (protracts); lateral Medial
Mandible Sphenoid
moves lower jaw side-to-side (abducts); medial (adducts) pterygoid

Table11.4

Although the masseter and temporalis are responsible for elevating and closing the jaw to break food into digestible pieces, the medial
pterygoid and lateral pterygoid muscles provide assistance in chewing and moving food within the mouth.

Muscles That Move the Tongue

Although the tongue is obviously

important for tasting food, it is also
necessary for
mastication, deglutition (swallowing),
and speech (Figure 11.11 and Figure
11.12). Because it is so moveable, the
tongue facilitates complex speech
patterns and sounds.

Figure 11.11 Muscles that Move the

Tongue

Figure 11.12 Muscles for Tongue

Movement, Swallowing, and Speech

Tongue muscles can be extrinsic or

intrinsic. Extrinsic tongue muscles insert
into the tongue from outside origins,
and the intrinsic tongue muscles insert
into the tongue from origins within it.
The extrinsic muscles move the whole
tongue in different directions, whereas
the intrinsic muscles allow the tongue to
change its shape (such as, curling the
tongue in a loop or flattening it).

The extrinsic muscles all include the

word root glossus (glossus = “tongue”),
and the muscle names are derived from
where the muscle originates.
The genioglossus (genio = “chin”)
originates on the mandible and allows
the tongue to move downward and
forward. The styloglossus originates
on the styloid bone, and allows upward
and backward motion.
The palatoglossus originates on the
soft palate to elevate the back of the
tongue, and the hyoglossus originates
on the hyoid bone to move the tongue
downward and flatten it.
Muscles of the Anterior Neck

The muscles of the anterior neck assist in deglutition (swallowing) and speech by controlling the positions of the larynx (voice box), and the hyoid bone,
a horseshoe-shaped bone that functions as a solid foundation on which the tongue can move. The muscles of the neck are categorized according to
their position relative to the hyoid bone (Figure 11.13). Suprahyoid muscles are superior to it, and the infrahyoid muscles are located inferiorly.

Figure 11.13 Muscles of the Anterior Neck The anterior

muscles of the neck facilitate swallowing and speech. The
suprahyoid muscles originate from above the hyoid bone in
the chin region. The infrahyoid muscles originate below the
hyoid bone in the lower neck.

The suprahyoid muscles raise the hyoid bone, the floor of the
mouth, and the larynx during deglutition. These include
the digastric muscle, which has anterior and posterior
bellies that work to elevate the hyoid bone and larynx when
one swallows; it also depresses the mandible.
The stylohyoid muscle moves the hyoid bone posteriorly,
elevating the larynx, and the mylohyoid muscle lifts it and
helps press the tongue to the top of the mouth.
The geniohyoid depresses the mandible in addition to
raising and pulling the hyoid bone anteriorly.

The strap-like infrahyoid muscles generally depress the hyoid

bone and control the position of the larynx.
The omohyoid muscle, which has superior and inferior
bellies, depresses the hyoid bone in conjunction with
the sternohyoid and thyrohyoid muscles. The thyrohyoid
muscle also elevates the larynx’s thyroid cartilage, whereas
the sternothyroid depresses it to create different tones of
voice.

Muscles That Move the Head

The head, attached to the top of the vertebral column, is balanced, moved, and rotated by the neck muscles (Table 11.5). When these muscles act
unilaterally, the head rotates. When they contract bilaterally, the head flexes or extends. The major muscle that laterally flexes and rotates the head is
the sternocleidomastoid. In addition, both muscles working together are the flexors of the head. Place your fingers on both sides of the neck and turn
your head to the left and to the right. You will feel the movement originate there. This muscle divides the neck into anterior and posterior triangles when
viewed from the side (Figure 11.14).

Figure 11.
14 Posteri
or and
Lateral
Views of
the
Neck The
superficial
and deep
muscles of
the neck
are

responsible for moving the head, cervical vertebrae, and scapulas.

Muscles That Move the Head

Movement Target Target motion direction Prime mover Origin

Rotates and tilts head to the side; Skull; Individually: rotates head to opposite side;
Sternocleidomastoid Sternum; clavicle
tilts head forward vertebrae bilaterally: flexion
 Movement Target Target motion direction Prime mover Origin

Skull; Individually: laterally flexes and rotates head to Transverse and articular p
Rotates and tilts head backward Semispinalis capitis
vertebrae same side; bilaterally: extension cervical and thoracic vert

Rotates and tilts head to the side; Skull; Individually: laterally flexes and rotates head to Spinous processes of cerv
Splenius capitis
tilts head backward vertebrae same side; bilaterally: extension vertebra

Rotates and tilts head to the side; Skull; Individually: laterally flexes and rotates head to Transverse and articular p
Longissimus capitis
tilts head backward vertebrae same side; bilaterally: extension cervical and thoracic vert

Table11.5

Muscles of the Posterior Neck and the Back

The posterior muscles of the neck are primarily concerned with head movements, like extension. The back muscles stabilize and move the vertebral
column, and are grouped according to the lengths and direction of the fascicles.

The splenius muscles originate at the midline and run laterally and superiorly to their insertions. From the sides and the back of the neck, the splenius
capitis inserts onto the head region, and the splenius cervicis extends onto the cervical region. These muscles can extend the head, laterally flex it,
and rotate it (Figure 11.15).

Figure 11.15 Muscles of the Neck

and Back The large, complex muscles
of the neck and back move the head,
shoulders, and vertebral column.

The erector spinae group forms the

majority of the muscle mass of the back
and it is the primary extensor of the
vertebral column. It controls flexion,
lateral flexion, and rotation of the
vertebral column, and maintains the
lumbar curve. The erector spinae
comprises the iliocostalis (laterally
placed) group, the longissimus
(intermediately placed) group, and the
spinalis (medially placed) group.

The iliocostalis group includes
the iliocostalis cervicis, associated
with the cervical region; the iliocostalis
thoracis, associated with the thoracic
region; and the iliocostalis lumborum,
associated with the lumbar region. The
three muscles of the longissimus
group are the longissimus capitis,
associated with the head region;
the longissimus cervicis, associated
with the cervical region; and
the longissimus thoracis, associated
with the thoracic region. The third
group, the spinalis group, comprises
the spinalis capitis (head region),
the spinalis cervicis (cervical region),
and the spinalis thoracis (thoracic
region).

The transversospinales muscles run
from the transverse processes to the
spinous processes of the vertebrae.
Similar to the erector spinae muscles,
the semispinalis muscles in this group
are named for the areas of the body
with which they are associated. The semispinalis muscles include the semispinalis capitis, the semispinalis cervicis, and the semispinalis thoracis.
The multifidus muscle of the lumbar region helps extend and laterally flex the vertebral column.

Important in the stabilization of the vertebral column is the segmental muscle group, which includes the interspinales and intertransversarii muscles.
These muscles bring together the spinous and transverse processes of each consecutive vertebra. Finally, the scalene muscles work together to flex,
laterally flex, and rotate the head. They also contribute to deep inhalation. The scalene muscles include the anterior scalene muscle (anterior to the
middle scalene), the middle scalene muscle (the longest, intermediate between the anterior and posterior scalenes), and the posterior scalene muscle
(the smallest, posterior to the middle scalene).

It is a complex job to balance the body on two feet and walk upright. The muscles of the vertebral column, thorax, and abdominal wall extend, flex, and
stabilize different parts of the body’s trunk. The deep muscles of the core of the body help maintain posture as well as carry out other functions. The
brain sends out electrical impulses to these various muscle groups to control posture by alternate contraction and relaxation. This is necessary so that
no single muscle group becomes fatigued too quickly. If any one group fails to function, body posture will be compromised.

Muscles of the Abdomen

There are four pairs of abdominal muscles that cover the anterior and lateral abdominal region and meet at the anterior midline. These muscles of the
anterolateral abdominal wall can be divided into four groups: the external obliques, the internal obliques, the transversus abdominis, and the rectus
abdominis (Figure 11.16 and Table 11.6).

Figure 11.16 Muscles of the
Abdomen (a) The anterior
abdominal muscles include the
medially located rectus abdominis,
which is covered by a sheet of
connective tissue called the rectus
sheath. On the flanks of the body,
medial to the rectus abdominis, the
abdominal wall is composed of three
layers. The external oblique muscles
form the superficial layer, while the
internal oblique muscles form the
middle layer, and the transversus
abdominis forms the deepest layer.
(b) The muscles of the lower back
move the lumbar spine but also
assist in femur movements.

Muscles of the Abdomen

Movement

Twisting at waist; also bending to the side

Squeezing abdomen during forceful exhalations

childbirth

Sitting up

Bending to the side

Table11.6

There are three flat skeletal muscles in the antero-lateral wall of the abdomen. The external oblique, closest to the surface, extend inferiorly and
medially, in the direction of sliding one’s four fingers into pants pockets. Perpendicular to it is the intermediate internal oblique, extending superiorly and
medially, the direction the thumbs usually go when the other fingers are in the pants pocket. The deep muscle, the transversus abdominis, is arranged
transversely around the abdomen, similar to the front of a belt on a pair of pants. This arrangement of three bands of muscles in different orientations
allows various movements and rotations of the trunk. The three layers of muscle also help to protect the internal abdominal organs in an area where
there is no bone.
The linea alba is a white, fibrous band that is made of the bilateral rectus sheaths that join at the anterior midline of the body. These enclose
the rectus abdominis muscles (a pair of long, linear muscles, commonly called the “sit-up” muscles) that originate at the pubic crest and symphysis,
and extend the length of the body’s trunk. Each muscle is segmented by three transverse bands of collagen fibers called the tendinous intersections.
This results in the look of “six-pack abs,” as each segment hypertrophies on individuals at the gym who do many sit-ups.

The posterior abdominal wall is formed by the lumbar vertebrae, parts of the ilia of the hip bones, psoas major and iliacus muscles, and quadratus
lumborum muscle. This part of the core plays a key role in stabilizing the rest of the body and maintaining posture.

Muscles of the Thorax

The muscles of the chest serve to facilitate breathing by changing the size of the thoracic cavity (Table 11.7). When you inhale, your chest rises because
the cavity expands. Alternately, when you exhale, your chest falls because the thoracic cavity decreases in size.

Muscles of the Thorax

Movement Target Target motion direction Prime mover Origin

Inhalation; Thoracic Sternum; ribs 6–12; lumb

Compression; expansion Diaphragm
exhalation cavity vertebrae

External Rib superior to each inte

Inhalation;exhalation Ribs Elevation (expands thoracic cavity)
intercostals muscle

Movement along superior/inferior axis to bring ribs closer Internal Rib inferior to each inter
Forced exhalation Ribs
together intercostals muscle

Table11.7

The Diaphragm

The change in volume of the thoracic cavity during breathing is due to the alternate contraction and relaxation of the diaphragm (Figure 11.17). It
separates the thoracic and abdominal cavities, and is dome-shaped at rest. The superior surface of the diaphragm is convex, creating the elevated floor
of the thoracic cavity. The inferior surface is concave, creating the curved roof of the abdominal cavity.

Figure 11.17 Muscles of the Diaphragm The diaphragm separates

the thoracic and abdominal cavities.

Defecating, urination, and even childbirth involve cooperation

between the diaphragm and abdominal muscles (this cooperation is
referred to as the “Valsalva maneuver”). You hold your breath by a
steady contraction of the diaphragm; this stabilizes the volume and
pressure of the peritoneal cavity. When the abdominal muscles
contract, the pressure cannot push the diaphragm up, so it increases
pressure on the intestinal tract (defecation), urinary tract (urination),
or reproductive tract (childbirth).

The inferior surface of the pericardial sac and the inferior surfaces of
the pleural membranes (parietal pleura) fuse onto the central tendon
of the diaphragm. To the sides of the tendon are the skeletal muscle
portions of the diaphragm, which insert into the tendon while having
a number of origins including the xiphoid process of the sternum
anteriorly, the inferior six ribs and their cartilages laterally, and the
lumbar vertebrae and 12th ribs posteriorly.

The diaphragm also includes three openings for the passage of

structures between the thorax and the abdomen. The inferior vena cava passes through the caval opening, and the esophagus and attached nerves
pass through the esophageal hiatus. The aorta, thoracic duct, and azygous vein pass through the aortic hiatus of the posterior diaphragm.

The Intercostal Muscles

There are three sets of muscles, called intercostal muscles, which span each of the intercostal spaces. The principal role of the intercostal muscles is
to assist in breathing by changing the dimensions of the rib cage (Figure 11.18).
 Figure 11.18 Intercostal Muscles The external
intercostals are located laterally on the sides of the body.
The internal intercostals are located medially near the
sternum. The innermost intercostals are located deep to
both the internal and external intercostals.

The 11 pairs of superficial external intercostal muscles

aid in inspiration of air during breathing because when
they contract, they raise the rib cage, which expands it.
The 11 pairs of internal intercostal muscles, just under
the externals, are used for expiration because they draw
the ribs together to constrict the rib cage. The innermost
intercostal muscles are the deepest, and they act as
synergists for the action of the internal intercostals.

Muscles of the Pelvic Floor and Perineum

The pelvic floor is a muscular sheet that defines the

inferior portion of the pelvic cavity. The pelvic diaphragm, spanning
anteriorly to posteriorly from the pubis to the coccyx, comprises the
levator ani and the ischiococcygeus. Its openings include the anal
canal and urethra, and the vagina in women.

The large levator ani consists of two skeletal muscles,

the pubococcygeus and the iliococcygeus (Figure 11.19). The
levator ani is considered the most important muscle of the pelvic floor
because it supports the pelvic viscera. It resists the pressure produced
by contraction of the abdominal muscles so that the pressure is applied
to the colon to aid in defecation and to the uterus to aid in childbirth
(assisted by the ischiococcygeus, which pulls the coccyx anteriorly).
This muscle also creates skeletal muscle sphincters at the urethra and
anus.

Figure 11.19 Muscles of the Pelvic Floor The pelvic floor muscles

support the pelvic organs, resist intra-abdominal pressure, and work as
sphincters for the urethra, rectum, and vagina.

The perineum is the diamond-shaped space between the pubic

symphysis (anteriorly), the coccyx (posteriorly), and the ischial tuberosities (laterally), lying just inferior to the pelvic diaphragm (levator ani and
coccygeus). Divided transversely into triangles, the anterior is the urogenital triangle, which includes the external genitals. The posterior is the anal
triangle, which contains the anus (Figure 11.20). The perineum is also divided into superficial and deep layers with some of the muscles common to
men and women (Figure 11.21). Women also have the compressor urethrae and the sphincter urethrovaginalis, which function to close the vagina.
In men, there is the deep transverse perineal muscle that plays a role in ejaculation.

Figure 11.20 Muscles of the Perineum The perineum muscles play roles in urination in both sexes, ejaculation in men, and vaginal contraction in
women.

Muscles of the shoulder and upper limb can be divided into four groups:
muscles that stabilize and position the pectoral girdle, muscles that move the
arm, muscles that move the forearm, and muscles that move the wrists, hands,
and fingers. The pectoral girdle, or shoulder girdle, consists of the lateral ends
of the clavicle and scapula, along with the proximal end of the humerus, and the
muscles covering these three bones to stabilize the shoulder joint. The girdle
creates a base from which the head of the humerus, in its ball-and-socket joint
with the glenoid fossa of the scapula, can move the arm in multiple directions.

Muscles That Position the Pectoral Girdle

Muscles that position the pectoral girdle are located either on the anterior thorax
or on the posterior thorax (Figure 11.22 and Table 11.8). The anterior muscles
include the subclavius, pectoralis minor, and serratus anterior. The posterior muscles include the trapezius, rhomboid major, and rhomboid
minor. When the rhomboids are contracted, your scapula moves medially, which can pull the shoulder and upper limb posteriorly.
 Figure 11.22 Muscles That Position
the Pectoral Girdle The muscles that
stabilize the pectoral girdle make it a
steady base on which other muscles
can move the arm. Note that the
pectoralis major and deltoid, which
move the humerus, are cut here to
show the deeper positioning muscles.

Muscles that Position the Pectoral

Girdle

Position in the
thorax Movement Target Target motion direction Prime mover Orig

Anterior Stabilizes clavicle during movement by

Clavicle Depression Subclavius First
thorax depressing it

Anterior Rotates shoulder anteriorly (throwing Pectoralis Ante

Scapula; ribs Scapula: depresses; ribs: elevates
thorax motion); assists with inhalation minor certa

Anterior Moves arm from side of body to front of Serratus Mus

Scapula; ribs Scapula: protracts; ribs: elevates
thorax body; assists with inhalation anterior ribs

Posterior Elevates shoulders (shrugging); pulls shoulder Scapula; Scapula: rotests inferiorly, retracts,
Trapezius Skull
thorax blades together; tilts head backwards cervical spine elevates, and depresses; spine: extends

Posterior Stabilizes scapula during pectoral girdle Rhomboid Thor

Scapula Retracts; rotates inferiorly
thorax movement major T5)

Posterior Stabilizes scapula during pectoral girdle Rhomboid Cerv

Scapula Retracts; rotates inferiorly
thorax movement minor verte
 Table11.8

Muscles That Move the

Humerus

Similar to the muscles that position

the pectoral girdle, muscles that
cross the shoulder joint and move
the humerus bone of the arm
include both axial and scapular
muscles (Figure 11.23 and Figure
11.24). The two axial muscles are
the pectoralis major and the
latissimus dorsi. The pectoralis
major is thick and fan-shaped,
covering much of the superior portion of the anterior thorax. The broad, triangular latissimus dorsi is located on the inferior part of the back, where it
inserts into a thick connective tissue shealth called an aponeurosis.

Figure 11.23 Muscles That Move the Humerus (a, c) The muscles that move the humerus anteriorly are generally located on the anterior side of the
body and originate from the sternum (e.g., pectoralis major) or the anterior side of the scapula (e.g., subscapularis). (b) The muscles that move the
humerus superiorly generally originate from the superior surfaces of the scapula and/or the clavicle (e.g., deltoids). The muscles that move the humerus
inferiorly generally originate from middle or lower back (e.g., latissiumus dorsi). (d) The muscles that move the humerus posteriorly are generally located
on the posterior side of the body and insert into the scapula (e.g., infraspinatus).

Figure 11.24 Muscles That Move the

Humerus

The rest of the shoulder muscles

originate on the scapula. The
anatomical and ligamental structure of
the shoulder joint and the arrangements
of the muscles covering it, allows the
arm to carry out different types of
movements. The deltoid, the thick
muscle that creates the rounded lines of
the shoulder is the major abductor of
the arm, but it also facilitates flexing and
medial rotation, as well as extension
and lateral rotation.
The subscapularis originates on the
anterior scapula and medially rotates
the arm. Named for their locations,
the supraspinatus (superior to the
spine of the scapula) and
the infraspinatus (inferior to the spine
of the scapula) abduct the arm, and
laterally rotate the arm, respectively.
The thick and flat teres major is inferior
to the teres minor and extends the arm,
and assists in adduction and medial
rotation of it. The long teres
minor laterally rotates and extends the
arm. Finally,
the coracobrachialis flexes and
adducts the arm.
The tendons of the deep subscapularis, supraspinatus, infraspinatus, and teres minor connect the scapula to the humerus, forming the rotator
cuff (musculotendinous cuff), the circle of tendons around the shoulder joint. When baseball pitchers undergo shoulder surgery it is usually on the
rotator cuff, which becomes pinched and inflamed, and may tear away from the bone due to the repetitive motion of bring the arm overhead to throw a
fast pitch.

Muscles That Move the Forearm

The forearm, made of the radius and ulna bones, has four main types of action at the hinge of the elbow joint: flexion, extension, pronation, and
supination. The forearm flexors include the biceps brachii,
brachialis, and brachioradialis. The extensors are the triceps
brachii and anconeus. The pronators are the pronator teres and
the pronator quadratus, and the supinator is the only one that
turns the forearm anteriorly. When the forearm faces anteriorly, it is
supinated. When the forearm faces posteriorly, it is pronated.

The biceps brachii, brachialis, and brachioradialis flex the forearm. The
two-headed biceps brachii crosses the shoulder and elbow joints to
flex the forearm, also taking part in supinating the forearm at the
radioulnar joints and flexing the arm at the shoulder joint. Deep to the
biceps brachii, the brachialis provides additional power in flexing the
forearm. Finally, the brachioradialis can flex the forearm quickly or help
lift a load slowly. These muscles and their associated blood vessels and
nerves form the anterior compartment of the arm (anterior flexor
compartment of the arm) (Figure 11.25 and Figure 11.26).

Figure 11.25 Muscles That Move the Forearm The muscles

originating in the upper arm flex, extend, pronate, and supinate the
forearm. The muscles originating in the forearm move the wrists,
hands, and fingers.

Muscles That Move the Wrist, Hand,

and Fingers

Wrist, hand, and finger movements are

facilitated by two groups of muscles.
The forearm is the origin of
the extrinsic muscles of the hand.
The palm is the origin of the intrinsic
muscles of the hand.
Muscles of the Arm That Move the Wrists, Hands, and Fingers

The muscles in the anterior compartment of the forearm (anterior flexor compartment of the forearm) originate on the humerus and insert onto
different parts of the hand. These make up the bulk of the forearm. From lateral to medial, the superficial anterior compartment of the
forearm includes the flexor carpi radialis, palmaris longus, flexor carpi ulnaris, and flexor digitorum superficialis. The flexor digitorum
superficialis flexes the hand as well as the digits at the knuckles, which allows for rapid finger movements, as in typing or playing a musical instrument
(see Figure 11.27 and Table 11.9). However, poor ergonomics can irritate the tendons of these muscles as they slide back and forth with the carpal
tunnel of the anterior wrist and pinch the median nerve, which also travels through the tunnel, causing Carpal Tunnel Syndrome. The deep anterior
compartment produces flexion and bends fingers to make a fist. These are the flexor pollicis longus and the flexor digitorum profundus.

The muscles in the superficial posterior compartment of the forearm (superficial posterior extensor compartment of the forearm) originate on the
humerus. These are the extensor radialis longus, extensor carpi radialis brevis, extensor digitorum, extensor digiti minimi, and the extensor
carpi ulnaris.

The muscles of the deep posterior compartment of the forearm (deep posterior extensor compartment of the forearm) originate on the radius and
ulna. These include the abductor pollicis longus, extensor pollicis brevis, extensor pollicis longus, and extensor indicis (see Figure 11.27).

Figure 11.27 Muscles That Move

the Wrist, Hands, and Forearm

The tendons of the forearm

muscles attach to the wrist and

extend into the hand. Fibrous

bands called retinacula sheath the
tendons at the wrist. The flexor
retinaculum extends over the
palmar surface of the hand while
the extensor retinaculum extends
over the dorsal surface of the hand.
Intrinsic Muscles of the Hand

The intrinsic muscles of the hand both originate and insert within it (Figure 11.28). These muscles allow your fingers to also make precise movements
for actions, such as typing or writing. These muscles are divided into three groups. The thenar muscles are on the radial aspect of the palm.
The hypothenar muscles are on the medial aspect of the palm, and the intermediate muscles are midpalmar.

The thenar muscles include the abductor pollicis brevis, opponens pollicis, flexor pollicis brevis, and the adductor pollicis. These muscles form
the thenar eminence, the rounded contour of the base of the thumb, and all act on the thumb. The movements of the thumb play an integral role in
most precise movements of the hand.

The hypothenar muscles include the abductor digiti minimi, flexor digiti minimi brevis, and the opponens digiti minimi. These muscles form
the hypothenar eminence, the rounded contour of the little finger, and as such, they all act on the little finger. Finally, the intermediate muscles act on
all the fingers and include the lumbrical, the palmar interossei, and the dorsal interossei.

Figure 11.28 Intrinsic Muscles of the Hand The intrinsic muscles of the hand both originate and insert within the hand. These muscles provide the fine
motor control of the fingers by flexing, extending, abducting, and adducting the more distal finger and thumb segments.

Intrinsic Muscles of the Hand

Muscle Movement

Thenar muscles Moves thumb toward body

Thenar muscles Moves thumb across palm to touch other fingers

Thenar muscles Flexes thumb

Thenar muscles Moves thumb away from body

Hypothenar muscles Moves little finger toward body

Flexes little
Hypothenar muscles Little finger
finger

Hypothenar muscles Moves little finger across palm to touch thumb

Flexes each finger at metacarpo-phalangeal

Intermediate muscles joints; extends each finger at interphalangeal
joints

Adducts and flexes each finger at metacarpo-

Intermediate muscles phalangeal joints; extends each finger at
interphalangeal joints

Abducts and flexes the three middle fingers at

Intermediate muscles metacarpo-phalangeal joints; extends the three
middle fingers at interphalangeal joints
The appendicular muscles of the lower body position and stabilize the pelvic girdle, which serves as a foundation for the lower limbs. Comparatively,
there is much more movement at the pectoral girdle than
at the pelvic girdle. There is very little movement of the
pelvic girdle because of its connection with the sacrum at
the base of the axial skeleton. The pelvic girdle is less
range of motion because it was designed to stabilize and
support the body.

Muscles of the Thigh

What would happen if the pelvic girdle, which attaches

the lower limbs to the torso, were capable of the same
range of motion as the pectoral girdle? For one thing,
walking would expend more energy if the heads of the
femurs were not secured in the acetabula of the pelvis.
The body’s center of gravity is in the area of the pelvis. If
the center of gravity were not to remain fixed, standing
up would be difficult as well. Therefore, what the leg

muscles lack in range of motion and versatility, they make up for in

size and power, facilitating the body’s stabilization, posture, and
movement.

Gluteal Region Muscles That Move the Femur

Most muscles that insert on the femur (the thigh bone) and move it,
originate on the pelvic girdle. The psoas major and iliacus make
up the iliopsoas group. Some of the largest and most powerful
muscles in the body are the gluteal muscles or gluteal group.
The gluteus maximus is the largest; deep to the gluteus maximus is
the gluteus medius, and deep to the gluteus medius is
the gluteus minimus, the smallest of the trio (Figure
11.29 and Figure 11.30).

Figure 11.29 Hip and Thigh Muscles The large and powerful

muscles of the hip that move the femur generally originate on the
pelvic girdle and insert into the femur. The muscles that move the
lower leg typically originate on the femur and insert into the bones of
the knee joint. The anterior muscles of the femur extend the lower
leg but also aid in flexing the thigh. The posterior muscles of the
femur flex the lower leg but also aid in extending the thigh. A
combination of gluteal and thigh muscles also adduct, abduct, and
rotate the thigh and lower leg.
 Figure 11.30 Gluteal Region
Muscles That Move the Femur

The tensor fascia latae is a

thick, squarish muscle in the
superior aspect of the lateral
thigh. It acts as a synergist of the
gluteus medius and iliopsoas in
flexing and abducting the thigh. It
also helps stabilize the lateral
aspect of the knee by pulling on
the iliotibial tract (band), making
it taut. Deep to the gluteus
maximus,
the piriformis, obturator
internus, obturator
externus, superior
gemellus, inferior gemellus,
and quadratus femoris laterally
rotate the femur at the hip.

The adductor longus, adductor
brevis, and adductor
magnus can both medially and
laterally rotate the thigh
depending on the placement of
the foot. The adductor longus
flexes the thigh, whereas the
adductor magnus extends it.
The pectineus adducts and
flexes the femur at the hip as
well. The pectineus is located in
the femoral triangle, which is
formed at the junction between
the hip and the leg and also
includes the femoral nerve, the
femoral artery, the femoral vein,
and the deep inguinal lymph
nodes.

Thigh Muscles That Move the

Femur, Tibia, and Fibula

Deep fascia in the thigh

separates it into medial, anterior,
and posterior compartments
(see Figure 11.29 and Figure
11.31). The muscles in
the medial compartment of the
thigh are responsible for
adducting the femur at the hip.
Along with the adductor longus,
adductor brevis, adductor
magnus, and pectineus, the
strap-like gracilis adducts the
thigh in addition to flexing the leg
at the knee.

Figure 11.31 Thigh Muscles That

Move the Femur, Tibia, and Fibula

The muscles of the anterior

compartment of the thigh flex the
thigh and extend the leg. This
compartment contains the quadriceps
femoris group, which actually
comprises four muscles that extend and
stabilize the knee. The rectus femoris is on the anterior aspect of the thigh, the vastus lateralis is on the lateral aspect of the thigh, the vastus
medialis is on the medial aspect of the thigh, and the vastus intermedius is between the vastus lateralis and vastus medialis and deep to the rectus
femoris. The tendon common to all four is the quadriceps tendon (patellar tendon), which inserts into the patella and continues below it as the patellar
ligament. The patellar ligament attaches to the tibial tuberosity. In addition to the quadriceps femoris, the sartorius is a band-like muscle that extends
from the anterior superior iliac spine to the medial side of the proximal tibia. This versatile muscle flexes the leg at the knee and flexes, abducts, and
laterally rotates the leg at the hip. This muscle allows us to sit cross-legged.

The posterior compartment of the thigh includes muscles that flex the leg and extend the thigh. The three long muscles on the back of the knee are
the hamstring group, which flexes the knee. These are the biceps femoris, semitendinosus, and semimembranosus. The tendons of these
muscles form the popliteal fossa, the diamond-shaped space at the back of the knee.

Muscles That Move the Feet and Toes

Similar to the thigh muscles, the muscles of the leg are divided by deep fascia into compartments, although the leg has three: anterior, lateral, and
posterior (Figure 11.32 and Figure 11.33).

Figure 11.32 Muscles of the
Lower Leg The muscles of the
anterior compartment of the lower
leg are generally responsible for
dorsiflexion, and the muscles of
the posterior compartment of the
lower leg are generally
responsible for plantar flexion.
The lateral and medial muscles in
both compartments invert, evert,
and rotate the foot.

Figure 11.33 Muscles That
Move the Feet and Toes

The muscles in the anterior

compartment of the leg:
the tibialis anterior, a long and
thick muscle on the lateral
surface of the tibia, the extensor
hallucis longus, deep under it,
and the extensor digitorum
longus, lateral to it, all contribute
to raising the front of the foot
when they contract. The fibularis
tertius, a small muscle that
originates on the anterior surface
of the fibula, is associated with
the extensor digitorum longus
and sometimes fused to it, but is
not present in all people. Thick
bands of connective tissue called
the superior extensor
retinaculum (transverse
ligament of the ankle) and
the inferior extensor
retinaculum, hold the tendons of
these muscles in place during
dorsiflexion.

The lateral compartment of the

leg includes two muscles:
the fibularis longus (peroneus
longus) and the fibularis
brevis (peroneus brevis). The
superficial muscles in
the posterior compartment of
the leg all insert onto
the calcaneal tendon (Achilles
tendon), a strong tendon that
inserts into the calcaneal bone of
the ankle. The muscles in this compartment are large and strong and keep humans upright. The most superficial and visible muscle of the calf is
the gastrocnemius. Deep to the gastrocnemius is the wide, flat soleus. The plantaris runs obliquely between the two; some people may have two of
these muscles, whereas no plantaris is observed in about seven percent of other cadaver dissections. The plantaris tendon is a desirable substitute for
the fascia lata in hernia repair, tendon transplants, and repair of ligaments. There are four deep muscles in the posterior compartment of the leg as well:
the popliteus, flexor digitorum longus, flexor hallucis longus, and tibialis posterior.

The foot also has intrinsic muscles, which originate and insert within it (similar to the intrinsic muscles of the hand). These muscles primarily provide
support for the foot and its arch, and contribute to movements of the toes (Figure 11.34 and Figure 11.35). The principal support for the longitudinal arch
of the foot is a deep fascia called plantar aponeurosis, which runs from the calcaneus bone to the toes (inflammation of this tissue is the cause of
“plantar fasciitis,” which can affect runners. The intrinsic muscles of the foot consist of two groups. The dorsal group includes only one muscle,
the extensor digitorum brevis. The second group is the plantar group, which consists of four layers, starting with the most superficial.

Figure 11.34 Intrinsic Muscles of the Foot The muscles along the dorsal side of the foot (a) generally extend the toes while the muscles of the plantar
side of the foot (b, c, d) generally flex the toes. The plantar muscles exist in three layers, providing the foot the strength to counterbalance the weight of
the body. In this diagram, these three layers are shown from a plantar view beginning with the bottom-most layer just under the plantar skin of the foot
(b) and ending with the top-most layer (d) located just inferior to the foot and toe bones.