464 UNIT 3 Control and Regulation

Figure 13–2 Gross Anatomy of the Adult Spinal Cord.

Posterior median sulcus
KEY
Spinal cord and vetebral regions Posterior root
Spinal White matter
Cervical ganglion

Central Gray
Thoracic matter
canal

Lumbar
C1 Spinal Anterior
Sacral C2 nerve root
C3 Anterior median fissure
C4 C3
C5
Cervical C6 C1
spinal Cervical
C7 C2
nerves enlargement
C8 C3
T1 Spinal cord
T2 C4
T3 C5 Vertebrae
T4 C6
T5 C7
T6 C8
13 T7 T1
Thoracic T2
T8 Posterior
spinal T3
median sulcus
nerves T9 T4
T10 T5
T11 T6 T3
Lumbosacral
T12 enlargement T7
Conus T8
L1 medullaris
T9
L2
T10
Inferior
Lumbar L3 tip of T11
spinal spinal cord
nerves L4 T12
Cauda equina
L1
L5
L2

S1
Sacral spinal L3 L1
nerves S2
S3 L4
S4
S5 L5
Coccygeal Filum terminale S1
nerve (Co1) (in coccygeal S2
ligament) S3
S4
S5
S5 1
Co S2

a The superficial anatomy and orientation of the adult b Lateral view of adult c Inferior views of cross sections
spinal cord. The numbers to the left identify the vertebrae and spinal cord. through representative
spinal nerves and indicate where the nerve roots Note that the spinal cord segments of the spinal cord,
leave the vertebral canal. The adult spinal cord segments for S1–S5 are level showing the arrangement of
extends from the brain only to the level of vertebrae with the T12–L1 vertebrae. gray matter and white matter.
L1–L 2; the spinal segments found at representative
locations are indicated in the cross sections.
 Chapter 13 The Spinal Cord, Spinal Nerves, and Spinal Reflexes 465

The entire spinal cord is divided into 31 segments based on the

origins of the spinal nerves. A letter and number designation,
Figure 13–3 The Spinal Cord and Spinal Meninges.
the same method used to identify vertebrae, identifies each Anterior rootlets Gray matter
segment. For example, C 3 , the spinal segment in the upper- of spinal nerve White matter
most section of Figure 13–2c, is the third cervical segment. Spinal ganglion
Anterior root
The posterior (dorsal) surface of the spinal cord has a shal-
low longitudinal groove, the posterior median sulcus. The
Spinal nerve
anterior median fissure is a deeper groove along the anterior
Posterior root
(ventral) surface. In addition, the spinal cord has an internal Meninges
passageway called the central canal. As noted in Chapter 12, Posterior rootlets
of spinal nerve Pia mater
this space is filled with cerebrospinal fluid (CSF), which acts
as a shock absorber and a medium for the diffusion of dis- Arachnoid
mater
solved gases, nutrients, chemical messengers, and wastes.
Dura mater
The amount of gray matter is greatest in segments of
the spinal cord dedicated to the sensory and motor con-
trol of the limbs. These segments are expanded, forming two
enlargements of the spinal cord. The cervical enlargement
supplies nerves to the shoulder and upper limbs. The
lumbosacral enlargement innervates structures of the pelvis
and lower limbs. 13
Inferior to the lumbosacral enlargement, the spinal
cord becomes tapered and conical. This region is the conus a A posterior view of
the spinal cord,
medullaris (med-y u -LAR-us). The filum terminale (F I -lum
showing the meningeal
ter-mi-NAH-l a; “terminal thread”), a slender strand of fibrous layers, superficial landmarks,
tissue, extends from the inferior tip of the conus medullaris. and distribution of gray matter
It continues along the length of the vertebral canal as far as and white matter
the second sacral vertebra, where it longitudinally supports the
spinal cord as a part of the coccygeal ligament.
Meninges
Subarachnoid
Spinal Roots and Ganglia space Dura mater
Every spinal segment is associated with a pair of spinal Arachnoid mater
Rami
ANTERIOR Pia mater
ganglia (posterior root ganglia) located near the spinal cord communicantes
(Figure 13–2c). These ganglia contain the cell bodies of sen- Vertebral
body Autonomic
sory neurons. The axons of the neurons form the posterior (sympathetic)
roots or dorsal roots, which bring sensory information into ganglion
the spinal cord. A pair of anterior roots, or ventral roots, Anterior root
contains the axons of motor neurons that extend into the of spinal
nerve
periphery to control somatic and visceral effectors. The roots
of all spinal nerves divide in fanlike fashion, forming rootlets, Anterior
ramus
before entering or leaving the spinal cord. A spinal posterior
Posterior
root may divide into 8–12 rootlets (Figure 13–3a). ramus
Spinal cord
On both sides, the posterior and anterior roots of each seg-
ment pass between the vertebral canal and the periphery at the Adipose tissue Denticulate Spinal
in epidural space ligament ganglion
intervertebral foramen between successive vertebrae. The spinal
ganglion lies between the pedicles of the adjacent vertebrae. POSTERIOR
The spinal cord continues to enlarge and elongate until b A sectional view through the spinal cord and meninges,
approximately 4 years of age. Up to that time, enlargement of showing the relationship of the meninges, spinal cord,
the spinal cord keeps pace with the growth of the vertebral col- and spinal nerves
umn. Throughout this time, the posterior and anterior roots are
very short, and they enter the intervertebral foramina immedi-
What is the area between the arachnoid mater and the
ately adjacent to their spinal segment. After age 4, the vertebral ? pia mater?
column continues to elongate, but the spinal cord does not.
 Chapter 13 The Spinal Cord, Spinal Nerves, and Spinal Reflexes 469

Figure 13–5 The Sectional Organization of the Spinal Cord.

Posterior Posterior Posterior gray

white column median sulcus commissure
Functional Organization
of Gray Matter
From posterior root
The cell bodies of neurons
in the gray matter of the
spinal cord are organized
into functional groups
called nuclei.
Posterior
Somatic
horn
Sensory nuclei
Lateral Visceral
Spinal Lateral horn
white Visceral
ganglion
column Anterior Motor nuclei
horn Somatic

To anterior
root
13
Anterior root
Anterior gray commissure
Anterior white commissure
Anterior white column
Anterior median fissure

a The left half of this sectional view shows important anatomical landmarks, including
the three columns of white matter. The right half indicates the functional organization
of the nuclei in the anterior, lateral, and posterior horns. The red arrows represent
sensory input from the posterior root and motor output to the anterior root.

POSTERIOR Structural Organization

of Gray Matter
Posterior
median sulcus The projections of gray
matter toward the outer
surface of the spinal cord
Posterior gray are called horns.
commissure
Posterior horn
Dura mater

Arachnoid mater Lateral horn

(broken)
Posterior
root
Central canal
Anterior horn
Anterior gray
commissure

Anterior median
fissure Spinal ganglion
Pia mater ANTERIOR

Anterior root

b A micrograph of a transverse section through the spinal

cord, showing major landmarks in and surrounding the cord.

? In which horn are somatic motor nuclei located?

 470 UNIT 3 Control and Regulation

dense network of collagen fibers. The fibers of the perineurium,

Checkpoint
the middle layer, extend inward from the epineurium. These con-
6. Differentiate between sensory nuclei and motor nective tissue partitions divide the nerve into a series of compart-
nuclei. ments, called fascicles, that contain bundles of axons. Delicate
7. A man with polio has lost the use of his leg muscles. connective tissue fibers of the endoneurium, the innermost layer,
In which area of his spinal cord are the virus-infected extend from the perineurium and surround individual axons.
motor neurons expected to be?
Arteries and veins penetrate the epineurium and branch
8. A disease that damages myelin sheaths would affect
within the perineurium. Capillaries leaving the perineurium
which part of the spinal cord?
branch in the endoneurium. They supply the axons and Schwann
See the blue Answers tab at the back of the book. cells of the nerve and the fibroblasts of the connective tissues.
As the spinal nerves extend into the periphery, they branch
and interconnect, forming the peripheral nerves that inner-
13- 4 Spinal nerves extend to form
vate body tissues and organs. The connective tissue sheaths of
peripheral nerves, sometimes forming peripheral nerves are the same as, and are continuous with,
plexuses along the way; these nerves those of spinal nerves.
carry sensory and motor information Recovery from an injury to a peripheral nerve may depend
Learning Outcome Describe the major components of a spinal on the extent of the injury. If a peripheral axon is severed but
nerve, describe a nerve plexus, and relate the distribution pattern not displaced, normal function may eventually return as the
of spinal nerves to the regions they innervate.
cut stump grows across the site of injury, away from the cell
13 In this section we consider the structure and function of spi- body and along its former path. Repairs made after an entire
nal nerves and the interwoven networks of spinal nerves called peripheral nerve has been damaged are generally incom-
nerve plexuses. plete, primarily due to problems with axon alignment and
regrowth. Various technologically sophisticated life-changing
Anatomy of Spinal Nerves and life-saving surgical procedures designed to improve nerve
regeneration and repair are currently under evaluation.
Every segment of the spinal cord is connected to a pair of spinal
nerves. Surrounding each spinal nerve is a series of connective
tissue layers continuous with those of the associated peripheral
Peripheral Distribution and Function
nerves (Figure 13–6). These wrapped layers, best seen in sectional
of Spinal Nerves
view, are comparable to those associated with skeletal muscles. The A dermatome is the specific bilateral region of the skin sur-
epineurium (ep-ih-NU R-e -um), or outermost layer, consists of a face monitored by a single pair of spinal nerves. Each pair of

Figure 13–6 A Peripheral Nerve. A diagrammatic view and an electron micrograph of a typical spinal nerve. Note the connective tissue
layers that are continuous with the associated spinal nerve.

Blood vessels

Connective Tissue
Layers

Epineurium covering
peripheral nerve
Perineurium (around
one fascicle)

Endoneurium

Schwann cell
Myelinated
axon
Fascicle
a A typical peripheral nerve and its b A scanning electron micrograph showing
connective tissue wrappings peripheral nerve fibers (SEM × 340)

> Media resource available for this topic: Practice Anatomy Lab: Histology: Nervous Tissue
 Figure 13–8

SPOTLIGHT Structure, Function, and Peripheral Distribution of

Spinal Nerves (T1 –L 2 )

SENSORY INFORMATION
A spinal nerve collects sensory information
from peripheral structures and delivers it to sensory
nuclei in the thoracic or superior lumbar segments of the spinal
cord. The posterior and anterior rami, and the white rami of the
rami communicantes, also contain sensory fibers.

From interoceptors From exteroceptors, 4

of back proprioceptors of back The posterior root of
each spinal nerve carries
3 sensory information to
The posterior ramus carries the spinal cord.
sensory information from the skin
and skeletal muscles of the back.
Somatic
sensory nuclei
2
The anterior ramus carries sensory
information from the ventrolateral
body surface, structures in the
body wall, and the limbs.

Spinal
ganglion
From exteroceptors,
proprioceptors of
body wall, limbs

From interoceptors
of body wall, limbs Rami
communicantes
Visceral
Anterior sensory nuclei
KEY
root
= Somatic
sensations
= Visceral 1
sensations The sympathetic nerve
carries sensory information From interoceptors
from the visceral organs. of visceral organs

472
 MOTOR COMMANDS
A spinal nerve distributes motor commands
that originate in motor nuclei of the thoracic or
superior lumbar segments of the spinal cord.

To skeletal Postganglionic fibers to

muscles smooth muscles, glands,
of back and adipose tissue of back
2
The spinal nerve forms just
3
lateral to the intervertebral The posterior ramus contains
foramen, where the posterior somatic motor and visceral motor
and anterior roots unite. fibers that innervate the skin and
Spinal skeletal muscles of the back.
Posterior root ganglion
4
The axons in the relatively
1 large anterior ramus supply
The anterior root of each the ventrolateral body surface,
spinal nerve contains the structures in the body wall,
axons of somatic motor and and the limbs.
visceral motor neurons.

Visceral motor nuclei To skeletal

muscles of body
Somatic motor nuclei

Postganglionic fibers to
KEY smooth muscles, and
glands of body wall, limbs
= Somatic motor
commands
Sympathetic
= Visceral motor ganglion Rami communicantes
commands 5
The white ramus communicans is the first branch from
the spinal nerve and carries visceral motor fibers to a
Postganglionic fibers nearby sympathetic ganglion. Because these
to smooth muscles, preganglionic axons are myelinated, this branch has a
glands, visceral organs light color and is therefore known as the white ramus.
in thoracic cavity White rami are only found between T1 and L2.
7 6
A sympathetic nerve
The gray ramus communicans contains post-
contains preganglionic
Preganglionic fibers to ganglionic fibers that innervate glands and smooth
and postganglionic fibers
sympathetic ganglia innervating muscles in the body wall or limbs. These fibers are
innervating structures in
abdominopelvic viscera unmyelinated and have a dark gray color. Gray rami
the thoracic cavity.
are associated with each spinal nerve.

Together, the white and gray rami are known as the

rami communicantes (RĀ-mī ko-mū-ni-KAN-tēz),
or communicating branches (singular, ramus
communicans).

473
 474 UNIT 3 Control and Regulation

Figure 13–9 Nerve Plexuses and Peripheral Nerves.

Major Nerve of the

Cervical Plexus
Lesser occipital nerve

Cervical Plexus (C1–C5 ) Great auricular nerve

The cervical plexus C1 Transverse cervical nerve

innervates the muscles of C2
C3
Supraclavicular nerve
the neck and the diaphragm.
C4
C5 Phrenic nerve
C6
Brachial Plexus (C5–T1) C7
C8 Major Nerves of the
The brachial plexus T1
T2
Brachial Plexus
innervates the pectoral
T3
girdles and upper limbs. Axillary nerve
T4
T5
T6 Musculocutaneous nerve
T7
T8 Radial nerve
T9
13 T10 Ulnar nerve
Lumbosacral Plexus
T11
The lumbosacral plexus Median nerve
innervates the pelvic girdle T12
and lower limbs.
L1
Thoracic nerves
Lumbar plexus (T12–L4)
L2
Major nerves of the lumbar Major Nerves of the
plexus are the femoral, L3 Lumbar Plexus
obturator, and saphenous
L4
nerves. Iliohypogastric nerve
L5
Sacral plexus (L4–S4) S1 Ilio-inguinal nerve
Major nerves of the sacral S2 Genitofemoral nerve
plexus are the gluteal and S3
sciatic nerves.
S4 Femoral nerve
S5
Obturator nerve
Co1
Lateral femoral
cutaneous nerve

Major Nerves of the

Sacral Plexus

Superior
Gluteal nerves
Inferior

Pudendal nerve

Sciatic nerve
Saphenous nerve

? The axillary, radial, and ulnar nerves branch from which plexus?
 Chapter 13 The Spinal Cord, Spinal Nerves, and Spinal Reflexes 475

The information, however, is not precise. More exact conclu- producing a series of compound nerve trunks. Such a complex
sions can be drawn if there is loss of motor control, based on interwoven network of nerves is called a nerve plexus (PLEK-
the origin and distribution of the peripheral nerves originating sus; plexus, braid).
at nerve plexuses. Only the anterior rami form plexuses. These four major
plexuses are the (1) cervical plexus, (2) brachial plexus, (3) lumbar
Nerve Plexuses plexus, and (4) sacral plexus (Figure 13–9). The nerves aris-
ing at these plexuses contain sensory as well as motor fibers
The simple distribution pattern of posterior and anterior rami
because these nerves form from the fusion of anterior rami
in Spotlight Figure 13–8 applies to spinal nerves T1 –L 2 . But in
(Spotlight Figure 13–8).
segments controlling the skeletal muscles of the neck, upper
Chapter 11 includes tables listing the nerves that control
limbs, or lower limbs, the situation is more complicated.
the major axial and appendicular muscles. As we proceed, it
During development, small skeletal muscles innervated by
may be helpful to refer to Tables 11–2 through 11–19 in that
different anterior rami typically fuse to form larger muscles
chapter.
with compound origins. The anatomical distinctions between
the component muscles may disappear, but separate anterior
rami continue to provide sensory innervation and motor con- The Cervical Plexus
trol to each part of the compound muscle. As they converge, The cervical plexus consists of the anterior rami of spinal nerves
the anterior rami of adjacent spinal nerves blend their fibers, C 1 –C 5 (Figures 13–9, 13–10). The branches of the cervical

Figure 13–10 The Cervical Plexus.

13

Nerves of the Cervical Plexus

Spinal
Segments Nerve and Distribution

Cranial
Great Auricular
Nerves C2–C3 Skin over the posterior aspect
of the ear and the neck
Accessory
nerve (XI) Lesser Occipital
Hypoglossal C2 Skin of the neck and the
nerve (XII) scalp posterior and superior
to the ear

Transverse Cervical
Nerve Roots of
Cervical Plexus C3–C4 Skin of the anterior triangle of
the neck
C1
C2 Ansa Cervicalis
C3 C1–C4 Five of the extrinsic laryngeal
C4 muscles by way of cranial
nerve XII
C5
Geniohyoid
Thyrohyoid
Omohyoid
Sternothyroid
Sternohyoid
Clavicle
Phrenic

C3–C5 Diaphragm

Supraclaviculars

C3–C4 Skin of the neck and shoulder

 476 UNIT 3 Control and Regulation

Figure 13–11 The Brachial Plexus.

Brachial Plexus: Major Nerves of the Arm and Forearm

Spinal
Segments Nerve and Distribution

Musculocutaneous

C5–T1 Flexor muscles on the arm (biceps brachii, brachialis, and

coracobrachialis); sensory from skin over the lateral surface of
the forearm through the lateral antebrachial cutaneous nerve

Radial

C5–T1 Many extensor muscles on the arm and forearm (triceps

brachii, anconeus, extensor carpi radialis, extensor carpi
ulnaris, and brachioradialis); and abductor pollicis by the
deep branch; sensory from skin over the posterolateral Median nerve
surface of the limb through the posterior brachial cutaneus
nerve, posterior antebrachial cutaneous nerve, and the
superficial branch (radial half of the hand)
Ulnar nerve
Median

C6–T1 Flexor muscles on the forearm (flexor carpi radialis and Lateral antebrachial
13 palmaris longus); pronator quadratus and pronator teres; cutaneous nerve
digital flexors (through the anterior interosseous nerve);
sensory from skin over the anterolateral surface of the hand Superficial branch
of radial nerve

Ulnar

C8–T1 Flexor carpi ulnaris, flexor digitorum profundus, adductor Deep branch of radial nerve
pollicis, and small digital muscles by the deep branch;
sensory from skin over medial surface of the hand Radial nerve
through the superficial branch

a Major nerves of the arm, forearm, and

hand originating at the right brachial
plexus, anterior view

+ Clinical Note Sensory

Innervation in the Hand
Sensory function in the hand is serviced by three
nerves of the brachial plexus. In carpal tunnel
Palmar digital syndrome, compression of one or more nerves at
nerves
the wrist causes changes or loss of sensation in
the areas innervated by the affected nerve(s).
plexus innervate the muscles of the neck and extend into the
thoracic cavity, where they control the diaphragm. The phrenic
Radial
(FREN-ik) nerve is the major nerve of the cervical plexus. The nerve
left and right phrenic nerves supply the diaphragm, a key breath-
Ulnar
ing muscle. Other branches of this nerve plexus are distributed nerve
to the skin of the neck and the superior part of the chest.

Tips Median
nerve
A useful mnemonic for remembering a function of three im-
portant cervical nerves is “ C 3 ,C 4 ,C 5 keep the diaphragm Anterior Posterior
alive.”
 Chapter 13 The Spinal Cord, Spinal Nerves, and Spinal Reflexes 477

Nerves of the Brachial Plexus Cords Divisions Trunks Anterior Rami Roots
The nerves of the brachial plexus arise from one or The anterior The brachial plexus is
more trunks or cords whose names indicate their rami converge formed by the anterior
positions relative to the axillary artery, a large artery to form the rami of spinal nerves
supplying the upper limb. superior, C5–T1.
middle, and
Spinal inferior trunks.
Segments Nerve and Distribution

Dorsal Scapular

C5 Rhomboid and levator scapulae

Subclavian C5
C5–C6 Subclavius

Suprascapular Superior
trunk
C5–C6 Supraspinatus and infraspinatus; sensory C6
from shoulder joint and scapula Lateral cord
Posterior cord
Pectoral (medial and lateral)
Medial cord Middle
C5–T1 Pectoralis muscles
C7 13
trunk

Subscapular nerves

C5–C6 Subscapularis and teres major Inferior trunk C8

Thoracodorsal

C6–C8 Latissimus dorsi

Axillary T1
First
C5–C6 Deltoid and teres minor rib

Medial Antebrachial Cutaneous

C8–T1 Sensory from skin over

anterior and medial surface KEY
of the arm and forearm
= Anterior rami roots
Musculocutaneous (see part a) = Trunks

= Divisions
Median (see part a)
= Cords
Ulnar (see part a) = Peripheral nerves

Radial (see part a)

Long Thoracic b Right brachial plexus, anterior view

C5–C7 Serratus anterior

The Brachial Plexus of spinal nerve axons, which become the smaller branches
The brachial plexus innervates the pectoral girdle and upper called cords. We name both trunks and cords according to
limb, with contributions from the anterior rami (also called their location relative to the axillary artery, a large artery supply-
roots) of spinal nerves C 5 –T1 (Figures 13–9, 13–11). The bra- ing the upper limb. Hence, we have superior, middle, and inferior
chial plexus can also have fibers from C 4 , T 2 , or both. trunks, and lateral, medial, and posterior cords. The lateral cord
The rami of these spinal nerves form trunks and cords forms the musculocutaneous nerve exclusively and, together
before they become peripheral nerves. Trunks are large bundles with the medial cord, contributes to the median nerve.
 478 UNIT 3 Control and Regulation

Figure 13–12 The Lumbar and Sacral Plexuses.

Nerve Roots of
Lumbar Plexus T12 subcostal nerve Lumbar Plexus
Spinal The lumbar
Segments Nerve and Distribution T12 plexus is formed
by the anterior
Iliohypogastric rami of T12 – L4.
T12–L1 External and internal oblique and transverse abdominis; L1
T12
skin over the inferior abdomen and buttocks
L1
Ilio-inguinal
L2
L2
L1 Abdominal muscles (with iliohypogastric nerve); skin over
superior, medial thigh and portions of external genitalia
L3
Genitofemoral L3

L1–L2 Skin over anteromedial thigh and portions of external genitalia

Lateral Femoral Cutaneous L4

L4
L2–L3 Skin over anterior, lateral, and posterior thigh

Femoral
13 L5
L2–L4 Quadriceps femoris, sartorius, pectineus, and iliopsoas; skin of
the anteromedial thigh, and medial surface of the leg and foot

Obturator Lumbosacral trunk

L2–L4 Gracilis, and adductor magnus, brevis and longus; skin from
a Lumbar plexus, anterior view
the medial surface of the thigh

Nerve Roots of
Lumbosacral trunk
Sacral Plexus
Sacral Plexus
L4 The sacral plexus
Spinal is formed by a
Segments Nerve and Distribution branch from L4
and the anterior
Superior Gluteal rami of L5–S4.
L5
L4–S2 Gluteus minimus, gluteus medius, and tensor L5
fasciae latae

Inferior Gluteal S1

L5–S2 Gluteus maximus S2

Posterior Femoral Cutaneous

Sacrum S3
S1–S3 Skin over perineum and posterior thigh and leg
S4
Sciatic

L4–S3 Semimembranosus, semitendinosus, and adductor magnus;

branches into tibial and common fibular nerves S5

Pudendal Co1

S2–S4 Muscles of the perineum; skin of external genitalia,

bulbospongiosus and ischiocavernosus

b Sacral plexus, anterior view

 Chapter 13 The Spinal Cord, Spinal Nerves, and Spinal Reflexes 479

Iliohypogastric nerve
Clinical Note Superior gluteal
+
Ilio-inguinal nerve
Sensory Innervation in the nerve
Genitofemoral nerve
Lateral femoral
Ankle and Foot Inferior gluteal nerve

cutaneous nerve Pudendal nerve

Femoral nerve These illustrations show the normal distribu- Posterior femoral
Obturator nerve tion of sensory nerves innervating the ankle cutaneous nerve
Superior gluteal nerve and foot. Mapping of touch and pain per- Sciatic nerve
ception can be combined with observations
Inferior gluteal nerve
of muscle function to detect nerve damage
Pudendal nerve affecting specific peripheral nerves.

Posterior femoral
cutaneous nerve (cut) Saphenous
Sciatic nerve nerve

Sural
nerve Tibial nerve
Saphenous nerve Common
fibular Common fibular
nerve nerve 13

Common fibular
nerve
Tibial
Superficial fibular nerve
nerve Saphenous Sural nerve
Sural nerve
Deep fibular nerve
nerve

Saphenous Sural
nerve nerve
c Nerves of the lumbar d Nerves of the sacral
and sacral plexuses, plexus, posterior view
anterior view Fibular
Tibial
nerve
nerve

The ulnar nerve is the other major nerve of the medial cord. sciatic (s ı -AT-ik) nerve and the pudendal nerve. The sciatic
The posterior cord gives rise to the axillary nerve and the nerve passes posterior to the femur, deep to the long head of
radial nerve. the biceps femoris. As it approaches the knee, the sciatic nerve
divides into two branches: the common fibular nerve (or
The Lumbar and Sacral Plexuses common peroneal nerve) and the tibial nerve. The sural nerve,
The lumbar plexus and the sacral plexus arise from the lum- formed by branches of the tibial nerve, is a sensory nerve inner-
bar and sacral segments of the spinal cord, respectively. The vating the lateral portion of the foot. A section of this nerve is
nerves arising at these plexuses innervate the pelvic girdle and often removed for use in nerve grafts.
lower limbs (Figures 13–9, 13–12). Discussions of motor performance usually make a dis-
The lumbar plexus contains axons from the anterior rami tinction between the conscious ability to control motor
of spinal nerves T12 –L 4 . The major nerves of this plexus are the function—something that requires communication and feed-
genitofemoral nerve, the lateral femoral cutaneous nerve, back between the brain and spinal cord—and automatic motor
and the femoral nerve. responses coordinated entirely within the spinal cord. These
The sacral plexus contains axons from the anterior rami of automatic responses, called reflexes, are motor responses to
spinal nerves L 4 –S 4 . Two major nerves arise at this plexus: the specific stimuli. In the rest of this chapter, we look at how
 SPOTLIGHT
Figure 13–14

Spinal Reflexes

Reflexes are rapid, automatic responses to specific stimuli. They preserve homeostasis
by making rapid adjustments in the function of organs or organ systems.

a EVENTS IN A REFLEX ARC

The “wiring” of a single
1 2 Posterior Sensory Sensation
reflex is called a reflex Arrival of Activation of a root neuron relayed to the
arc. It begins at a stimulus and sensory (stimulated) brain by axon
receptor and ends at a activation of neuron collaterals
peripheral effector such receptor
as a muscle fiber or a Spinal cord
gland cell. A simple 3
reflex arc, such as this
REFLEX Information
withdrawal reflex, processing
ARC
consists of five steps. Receptor in the CNS
Stimulus

5
Response by a Excitatory interneuron
Effector
peripheral Anterior root
effector 4 Motor neuron (stimulated)
Activation of a
motor neuron

MONOSYNAPTIC REFLEXES
In the simplest reflex arc, a sensory neuron synapses directly on a motor neuron.
Because there is only one synapse, it is called a monosynaptic reflex. Transmis-
sion across a chemical synapse always involves a synaptic delay, but with only
one synapse, the delay between the stimulus and the response is minimized.

b Stretch Reflex
Sensory neuron (stimulated)
The stretch reflex is an Receptor
example of a monosynaptic Stretch (muscle
spindle) Spinal cord
reflex. Because there is only
one synapse, there is little
delay between sensory input Stimulus REFLEX
and motor output. These ARC
reflexes control the most
rapid motor responses of the
nervous system. A stretch
reflex provides automatic Effector
Contraction
regulation of skeletal muscle
length. Motor neuron (stimulated)

The patellar reflex is an example of a stretch reflex.

The stimulus is a tap on the patellar tendon that
Response stretches receptors within the quadriceps femoris.
The response is a brief contraction of those muscles,
which produces a noticeable kick.

482
POLYSYNAPTIC REFLEXES
Polysynaptic reflexes can produce far more complicated responses
than monosynaptic reflexes, because the interneurons can control
motor neurons that activate several muscle groups simultaneously.
c Withdrawal Reflex
A withdrawal reflex moves affected parts of the body away Distribution within gray matter horns to
other segments of the spinal cord
from a stimulus. A flexor reflex is an example of a withdrawal
reflex that affects the muscles of a limb. In this example, the Sensory neuron
stimulus of a hot frying pan causes the contraction of the (stimulated)
flexor muscles of the arm, yanking the forearm
and hand away from the pan.
This response occurs while
Excitatory interneuron
pain sensations are
simultaneously Motor neuron (stimulated)
Painful
ascending to stimulus
the brain. Flexors
stimulated
Motor neuron Inhibitory
(inhibited) interneuron
Extensors
inhibited

d Crossed Extensor Reflex

A crossed extensor reflex involves a contralateral reflex To motor neurons in other
segments of the spinal cord
arc (contra, opposite). In other words, a motor response
to the stimulus also occurs on a side opposite the
stimulus. The crossed extensor reflex complements the
Excitatory
flexor reflex, and the two occur simultaneously. When interneuron
you step on a tack, the flexor reflex pulls the affected
foot away from the ground, while the crossed extensor Sensory
reflex straightens the other leg to support your body Inhibitory
neuron
interneuron
weight. In the crossed extensor reflex, interneurons (stimulated)
responding to the pain have axons that cross to the
other side of the spinal cord. There they stimulate motor
neurons that control the extensor muscles of the Motor neuron
uninjured leg. As a result, your opposite leg straightens (inhibited)
to support the shifting weight. Reverberating circuits use
Motor neuron
positive feedback to ensure that the movement lasts (stimulated)
long enough to be effective. All of this happens without Extensors
motor commands from higher centers of the brain. inhibited
Flexors Extensors stimulated
stimulated
Flexors inhibited
KEY
Sensory neuron Motor neuron
(stimulated) (inhibited)
Excitatory Inhibitory
interneuron interneuron
Motor neuron
(stimulated) Painful
stimulus

483
 Chapter 14 The Brain and Cranial Nerves 495

Figure 14–1 An Introduction to Brain Structures and Functions.

Left cerebral
hemisphere
Gyri

a Cerebrum Sulci

• Conscious thought processes,

intellectual functions Fissures
• Memory storage and processing
• Conscious and subconscious regulation
of skeletal muscle contractions

c Diencephalon
Thalamus
• Relays and processes
sensory information

Hypothalamus
• Controls emotions, 14
autonomic functions,
and hormone
b Cerebellum
production • Coordinates complex
Spinal somatic motor
cord patterns
Brainstem • Adjusts output of
d Midbrain other somatic motor
• Processes visual centers in brain and
and auditory data spinal cord
• Generates reflexive
somatic motor
responses
• Maintains
consciousness

e Pons
• Relays sensory f Medulla oblongata
information to
cerebellum and • Relays sensory information to thalamus and
thalamus to other portions of the brainstem
• Subconscious • Autonomic centers for regulation of visceral
somatic and visceral function (cardiovascular, respiratory, and
motor centers digestive system activities)

? What structures make up the brainstem?

It contains centers involved with emotions, autonomic function, a variety of important processing centers and nuclei (clearly distin-
and hormone production. The infundibulum, a narrow stalk, con- guishable masses of brain neurons) that relay information headed
nects the hypothalamus to the pituitary gland, a part of the to or from the cerebrum or cerebellum. The brainstem includes
endocrine system. This connection integrates the nervous and the midbrain, pons, and medulla oblongata (Figure 14–1d,e,f).
endocrine systems.
■ The midbrain contains nuclei that process visual and
auditory information and control reflexes triggered by these
The Brainstem stimuli. For example, immediate, reflexive responses to a
The diencephalon is a structural and functional link between the loud, unexpected noise (eye movements and head turning)
cerebral hemispheres and the brainstem. The brainstem contains are directed by nuclei in the midbrain. This region also
 496 UNIT 3 Control and Regulation

contains centers (groups of nerve cells governing a specific “forebrain”; the mesencephalon, or “midbrain”; and the rhomben-
function) that help maintain consciousness (Figure 14–1d). cephalon (rom-ben-SEF-ah-lon), or “hindbrain.”
■ The pons of the brain connects the cerebellum to the Table 14–1 summarizes the fates of the primary brain
brainstem (pons is Latin for “bridge”). In addition to tracts vesicles. The prosencephalon and rhombencephalon are sub-
(collections of CNS axons) and relay centers, the pons divided further, forming secondary brain vesicles, while the
contains nuclei involved with somatic and visceral motor mesencephalon develops but does not divide:
control (Figure 14–1e). ■ The prosencephalon forms the telencephalon (tel-en-SEF-
■ The medulla oblongata connects the brain to the spinal a-lon; telos, end) and the diencephalon. The telencephalon
cord. Near the pons, the posterior wall of the medulla ultimately forms the cerebrum, and the diencephalon
oblongata is thin and membranous. The inferior portion forms the thalamus.
of the medulla oblongata resembles the spinal cord in ■ The walls of the mesencephalon thicken, and the neural
that it has a narrow central canal. The medulla oblongata tube becomes a relatively narrow passageway, much like
relays sensory information to the thalamus and to centers the central canal of the spinal cord.
in other portions of the brainstem. The medulla oblongata
■ The portion of the rhombencephalon adjacent to the
also contains major centers that regulate autonomic
mesencephalon forms the metencephalon (met-en-
function, such as heart rate, blood pressure, and digestion
SEF-ah-lon; meta, after). The posterior portion of the
(Figure 14–1f).
metencephalon later becomes the cerebellum, and the
In considering the regions of the brain in this chapter, we anterior portion develops into the pons. The portion of
the rhombencephalon closer to the spinal cord forms
14 begin at the inferior portion of the medulla oblongata because
this region has the simplest organization. We then move up to the myelencephalon (m ı -el-en-SEF-ah-lon; myelon,
regions of increasing structural and functional complexity. spinal cord), which becomes the medulla oblongata.

Embryology of the Brain Ventricles of the Brain

The embryonic origin of the brain can help us understand its During development, the neural tube within the cerebral hemi-
internal organization. The central nervous system (CNS) begins spheres, diencephalon, metencephalon, and medulla oblongata
as a hollow cylinder known as the neural tube. At the cephalic expands to form chambers called ventricles (VEN-trih-klz).
portion of the neural tube, three swellings called the primary The ventricular system is composed of four ventricles (two lateral
brain vesicles develop. They are named for their relative posi- ventricles, then the third and fourth ventricles) and their pas-
tions: the prosencephalon (proz-en-SEF-ah-lon; proso, forward), or sageways (interventricular foramen and cerebral aqueduct).

Table 14–1 Development of the Brain

Primary Brain Vesicles (3 weeks) Secondary Brain Vesicles (6 weeks) Brain Regions at Birth Ventricular System

Prosencephalon Telencephalon Cerebrum Lateral ventricles

(Forebrain)
Diencephalon Diencephalon Third ventricle

Mesencephalon Midbrain Cerebral aqueduct

Mesencephalon
(Midbrain)
Cerebellum Fourth ventricle
Metencephalon and Pons

Rhombencephalon
(Hindbrain)
Medulla Fourth ventricle
Myelencephalon oblongata
 Chapter 14 The Brain and Cranial Nerves 497

Figure 14–2 Ventricular System. The orientation and extent of the ventricles as they would appear if the brain
were transparent.

Cerebral hemispheres
Cerebral Ventricular System
hemispheres of the Brain

Lateral ventricles

Interventricular
foramen

Third ventricle

Cerebral aqueduct

Fourth ventricle
Pons

Medulla oblongata
Spinal cord Central canal Central canal Cerebellum

a Ventricular system, lateral view b Ventricular system, anterior view

14

Each cerebral hemisphere contains a large lateral ventricle 14-2 The brain is protected and
(Figure 14–2). The septum pellucidum, a thin plate of brain tis-
supported by the cranial meninges,
sue, separates the two lateral ventricles. Because there are two
lateral ventricles, the ventricle in the diencephalon is called the
cerebrospinal fluid, and the blood-brain
third ventricle. The two lateral ventricles are not directly con- barrier
nected, but each communicates with the third ventricle of the Learning Outcome Explain how the brain is protected and
diencephalon through an interventricular foramen. supported, and discuss the formation, circulation, and function of
cerebrospinal fluid.
The midbrain has a slender canal known as the cerebral
aqueduct. This passageway connects the third ventricle with The delicate tissues of the brain are protected from physical
the fourth ventricle. The superior portion of the fourth ven- forces by the bones of the cranium, membranes called the cra-
tricle lies between the posterior surface of the pons and the nial meninges, and cerebrospinal fluid. In addition, the nervous
anterior surface of the cerebellum. The fourth ventricle extends tissue of the brain is biochemically isolated from the general
into the superior portion of the medulla oblongata. There this circulation by the blood-brain barrier.
ventricle narrows and becomes continuous with the central
canal of the spinal cord. The Cranial Meninges
Ependymal cells, a type of neuroglia, line the ventricles.
The layers that make up the cranial meninges—the cranial
These cells produce cerebrospinal fluid (CSF) that fills the ven-
dura mater, arachnoid mater, and pia mater—are continuous
tricles and continuously circulates in the CNS. The CSF passes
with the spinal meninges. However, the cranial meninges
between the interior and exterior of the CNS through three
have distinctive anatomical and functional characteristics
foramina in the roof of the fourth ventricle. We describe these
(Figure 14–3a).
foramina in Section 14-7.

Dura Mater and Dural Folds

Checkpoint
The dura mater consists of outer and inner fibrous lay-
1. Name the four major regions of the brain. ers. The outer layer is fused to the periosteum of the cranial
2. What brain regions make up the brainstem? bones. As a result, there is no epidural space superficial to the
3. Which primary brain vesicle is destined to form the dura mater, as occurs along the spinal cord. The outer perios-
cerebellum, pons, and medulla oblongata? teal cranial dura and inner meningeal cranial dura are typically
See the blue Answers tab at the back of the book. fused together.
 498 UNIT 3 Control and Regulation

Figure 14–3 The Relationships among the Brain, Cranium, and Cranial Meninges.

Cranium Periosteal cranial dura

Dura mater
Dural sinus

Subarachnoid space Meningeal cranial dura

Pia mater Subdural space

Cerebral cortex Arachnoid mater

Dura mater Superior sagittal Cranium Inferior sagittal

sinus sinus

Dural folds
Cerebrum
Falx cerebri
Tentorium
cerebelli
14 Falx cerebelli

Cerebellum

Medulla
oblongata Transverse
Spinal cord sinus

a A lateral view of the brain, showing its position in b A diagrammatic view, showing the orientation of the three largest
the cranium and the organization of the meninges dural folds: the falx cerebri, tentorium cerebelli, and falx cerebelli

? List the maters surrounding the brain from the deepest layer to the most
superficial layer.

In several locations, the meningeal cranial dura extends and posteriorly to the internal occipital crest, a ridge along the
into the cranial cavity, forming a sheet that dips inward and inner surface of the occipital bone. The superior sagittal
then returns. These inward projections, known as dural folds, sinus and the inferior sagittal sinus, two large dural
provide additional stabilization and support to the brain. venous sinuses, lie within this dural fold. The posterior
Dural venous sinuses are large collecting veins located within margin of the falx cerebri intersects the tentorium cerebelli.
the dural folds. The veins of the brain open into these sinuses, ■ The tentorium cerebelli (ten-TO-re -um ser-e-BEL-e ; tentorium,
which deliver the venous blood to the veins of the neck. The tent) protects the cerebellum and separates the cerebral
three largest dural folds are called the falx cerebri, the tentorium hemispheres from the cerebellum. It extends across the cranium
cerebelli, and the falx cerebelli (Figure 14–3b): at right angles to the falx cerebri. The transverse sinus is a
■ The falx cerebri (FALKS SER-e-br ı ; falx, sickle shaped) paired dural venous sinus that runs along the occipital bone.
is a fold of dura mater that projects between the cerebral It allows blood to drain from the back of the head.
hemispheres in the longitudinal cerebral fissure. Its inferior ■ The falx cerebelli divides the two cerebellar hemispheres
portions attach anteriorly to the crista galli of the ethmoid along the midsagittal line inferior to the tentorium cerebelli.
 SPOTLIGHT
Figure 14–4

Formation and Circulation of Cerebrospinal Fluid

Nutrients,
Cerebrospinal fluid (CSF) completely O2
Interstitial fluid
surrounds and bathes the exposed surfaces in thalamus
Capillaries
of the CNS. The CSF has three important
functions: Wastes, Neuron
CO2
• Supporting the brain.
• Cushioning the brain and spinal cord Astrocyte
against physical trauma.
• Transporting nutrients, chemical
messengers, and wastes.
Choroid plexus
1 ependymal cells Ependymal cells
The choroid plexus produces and maintains CSF. Two
extensive folds of the choroid plexus originate in the roof Removal
of waste
of the third ventricle and extend through the interventric-
ular foramina. These folds cover the floors of the lateral
Production Cerebrospinal fluid
ventricles. In the inferior brainstem, a region of the
of CSF in third ventricle
choroid plexus in the roof of the fourth ventricle projects
between the cerebellum and the pons.
Tight junction

Choroid plexus
of third ventricle Choroid plexus

Dura mater Arachnoid Arachnoid

(periosteal layer) granulation trabecula

Cranium CSF fluid

Superior movement
sagittal sinus
Dura mater
(meningeal
layer)
Subdural
space

Arachnoid
mater
Cerebral
Choroid cortex Subarachnoid
2 plexus space
The CSF circulates from the of fourth Pia mater
choroid plexus through the ventricle
ventricles and fills the central 5
canal of the spinal cord. As it Fingerlike extensions of the arachnoid membrane,
3
circulates, materials diffuse called the arachnoid villi, penetrate the meningeal layer
The CSF reaches the
between the CSF and the of the dura mater and extend into the superior sagittal
subarachnoid space
interstitial fluid of the CNS sinus. In adults, these extensions form large arachnoid
through two lateral
across the ependymal cells. granulations. CSF is absorbed into the venous
apertures and a single
circulation at the arachnoid granulations.
median aperture in the roof
Spinal cord of the fourth ventricle.

Central canal 4
Dura mater
Cerebrospinal fluid then The choroid plexus produces CSF at a rate of
flows through the about 500 mL, or 2.1 cups, per day. The total
Conus medullaris subarachnoid space volume of CSF at any moment is approximately
surrounding the brain,
150 mL. The entire volume of CSF is replaced
Arachnoid mater spinal cord, and cauda
about every 8 hours. Despite this rapid
equina.
Cauda equina turnover, the composition of CSF is closely
regulated, and the rate of removal normally
Filum terminale
keeps pace with the rate of production.
500
 Chapter 14 The Brain and Cranial Nerves 503

Figure 14–5 The Diencephalon and Brainstem. The brainstem is made up of the medulla oblongata,
pons, and midbrain.

Optic chiasm Infundibulum

Diencephalon
Cerebral peduncle
Thalamus Lateral geniculate nucleus
Optic tract Cranial
Medial geniculate nucleus Thalamus
Nerves
Cranial
Nerves Midbrain II
II Superior colliculus
Inferior colliculus Optic tract III
III
IV Cerebral
Cerebellar Peduncles IV
peduncle
Superior cerebellar Mammillary V
V peduncle Pons
body
Pons VI
Middle cerebellar
VI peduncle VII
VII Inferior cerebellar VIII
VIII peduncle IX
IX X
X
XI Medulla XI
oblongata
XII 14
XII Medulla
Spinal oblongata
nerve C1 Ventral roots
of spinal
Spinal Spinal
nerves C1
nerve C2 cord
and C2
Spinal cord
a Lateral view b Anterior view

Choroid plexus Third ventricle

in third and
Thalamus
fourth ventricles
Pineal gland

Corpora Quadrigemina
Superior colliculi

Inferior colliculi
CN IV

Cerebral peduncle

Trochlear
Cerebellar Peduncles
nerve (IV)
Superior
Middle
Choroid plexus
in roof of Inferior
fourth ventricle Fourth
ventricle

Posterior roots
of spinal nerves
C1 and C2

c Posterior view d Posterior view, cadaver dissection

 504 UNIT 3 Control and Regulation

information to the thalamus. Tracts leaving these brainstem The inferior olivary complex consists of three nuclei
nuclei cross to the opposite side of the brain before that collectively form the inferior olivary nucleus. They relay
reaching their destinations. This crossing over is called a information to the cerebellar cortex about somatic motor
decussation (d e -kuh-S A -shun; decussatio, crossing over), and commands as they are issued by motor centers at higher lev-
the site is the decussation of pyramids. els. The bulk of the olivary nuclei creates the olives, promi-
nent olive-shaped bulges along the ventrolateral surface of the
In addition, a few important paired nuclei are found in the
medulla oblongata.
medulla oblongata. The solitary nuclei (nuclei of solitary tract) are
Note that Figure 14–6 also contains a table that summa-
the visceral sensory nuclei, receiving information from the spinal
rizes the major components of the medulla oblongata and
and cranial nerves. This information is integrated and forwarded to
their functions.
other autonomic centers in the medulla oblongata and elsewhere.

Figure 14–6 The Medulla Oblongata.

Autonomic Centers
Reticular formation

Medulla Cardiovascular centers

14 oblongata
Solitary nuclei
Pons

Relay Stations

Inferior olive Olivary nucleus

Pyramids Cuneate nucleus

Decussation of Spinal cord Gracile nucleus

pyramids
Lateral white columns
a Anterior view

The Medulla Oblongata

Region/Nucleus Function
GRAY MATTER
Inferior olivary complex Relays information from the red nucleus, other
midbrain centers, and the cerebral cortex to the
Attachments to
cerebellum
membranous
roof of fourth Reflex centers
ventricle Cardiovascular centers Regulate heart rate and force of contraction
Respiratory rhythmicity centers Set the basic pace of respiratory movements

Gracile nucleus
Cuneate nucleus } Relay somatic information to the thalamus

Other nuclei/centers: Sensory and motor nuclei of five cranial nerves

Cranial nerves VIII (in part), Nuclei relay ascending information from the spinal cord
IX, X, XI (in part), and XII to higher centers
Posterior median sulcus
Reticular formation Contains nuclei and centers that regulate vital
Lateral funiculus autonomic functions; extends into the pons and
midbrain
Posterior funiculi
WHITE MATTER
Ascending and descending Link the brain with the spinal cord
b Posterolateral view tracts within funiculi
 Chapter 14 The Brain and Cranial Nerves 505

■ Nuclei Involved with the Control of Respiration. Research on

Checkpoint
laboratory animals has identified regions in the medulla
8. Identify the components of the medulla oblongata and pons, known as centers, that appear to modify
that are responsible for relaying somatic sensory breathing activities. Two pontine centers, the apneustic
information to the thalamus.
(ap-N U S-tik) center located in the middle or lower pons
9. The medulla oblongata is one of the smallest sections and the pneumotaxic (n u -mo-TAK-sik) center located in the
of the brain, yet damage there can cause death, whereas
pons, process information originating in the respiratory
similar damage in the cerebrum might go unnoticed. Why?
rhythmicity centers of the medulla oblongata. These centers
See the blue Answers tab at the back of the book. are further discussed in Chapter 23.
Nuclei and Tracts That Process and Relay Information Sent to or
14- 4 Brainstem: The pons contains
■

from the Cerebellum. The pons links the cerebellum with the
nuclei that process and tracts that relay brainstem, cerebrum, and spinal cord.
sensory and motor information ■ Ascending, Descending, and Transverse Pontine Fibers.
Learning Outcome List the main components of the pons and Longitudinal tracts interconnect other portions of the CNS.
specify the functions of each. Tracts of the cerebellum (middle cerebellar peduncles) are
The pons links the cerebellum with the midbrain, diencepha- connected to the transverse pontine fibers, which cross
lon, cerebrum, and spinal cord. Figure 14–5 shows the spatial the anterior surface of the pons. These fibers are axons that
relationship of the pons with these structures. link nuclei of the pons (pontine nuclei) with the cerebellum
Important features and regions of the pons are shown in of the opposite side.
Figure 14–7, along with a summary of its major components and 14
their functions. The pons contains four groups of components:
Checkpoint
■ Sensory and Motor Nuclei of Cranial Nerves. These cranial
nerves (V, VI, VII, and VIII) innervate the jaw muscles, the 10. Name the four groups of components found in the pons.
anterior surface of the face, one of the extrinsic eye muscles 11. If the respiratory centers of the pons were damaged,
(the lateral rectus), and the sense organs of the internal ear what respiratory controls might be lost?
(the vestibular and cochlear nuclei). See the blue Answers tab at the back of the book.

Figure 14–7 The Pons. The Pons

Region/Nucleus Function
WHITE MATTER
Descending tracts Carry motor commands from
higher centers to nuclei of cranial
or spinal nerves
Ascending tracts Carry sensory information from
brainstem nuclei to the thalamus
Transverse Interconnect cerebellar
pontine fibers hemispheres
Pons
Cerebellum GRAY MATTER

Apneustic and Adjust activities of the respiratory

Fourth pneumotaxic centers rhythmicity centers in the medulla
ventricle oblongata

Medulla oblongata Reticular formation Automatic processing of incoming

sensations and outgoing motor
Spinal cord Inferior olivary commands
nucleus
Nuclei of cranial Relay sensory information and
nerves V, VI, VII, issue somatic motor commands
and VIII (in part)
Other nuclei/relay Relay sensory and motor
centers information to the cerebellum
 Chapter 14 The Brain and Cranial Nerves 507

Figure 14–8 The Midbrain.

The Midbrain
Region/Nuclei Function
GRAY MATTER
Pineal gland Tectum (roof) Integrate visual information with other sensory input;
Superior colliculi initiate reflex responses to visual stimuli

Thalamus
Inferior colliculi Relay auditory information to medial geniculate nuclei;
initiate reflex responses to auditory stimuli
Walls and floor

Substantia nigra Regulates activity in the basal nuclei

Red nucleus Subconscious control of upper limb position and
background muscle tone
Reticular formation Automatic processing of incoming sensations and
outgoing motor commands; can initiate involuntary
motor responses to stimuli; helps maintain
consciousness
Other nuclei/centers Nuclei associated with cranial nerves III and IV

WHITE MATTER
a A posterior view. The
Cerebral peduncles Connect primary motor cortex with motor neurons
underlying nuclei are in brain and spinal cord; carry ascending sensory
14
colored only on the right. information to thalamus

Thalamus

Corpora
Quadrigemina ANTERIOR
Cerebral
Superior colliculi peduncle

Inferior colliculi Substantia

nigra
Cerebral peduncle

Trochlear nerve (IV) Red nucleus Tegmentum

Cerebral aqueduct
Superior
cerebellar Superior colliculus
peduncle Cerebellum
Tectum (roof)
Fourth ventricle
POSTERIOR

b Posterior view of a cadaver dissection of c A superior view of a transverse section

the diencephalon and brainstem. at the level of the midbrain.

■ Programming and Fine-Tuning Movements Controlled at the The cerebellum receives proprioceptive information from the
Conscious and Subconscious Levels. The cerebellum refines spinal cord and monitors all proprioceptive, visual, tactile, balance,
learned movement patterns, such as riding a bicycle or and auditory sensations received by the brain. Most axons that carry
playing the piano. It performs this function indirectly by sensory information do not synapse in the cerebellar nuclei but
regulating activity along motor pathways at the cerebral pass through the deeper layers of the cerebellum on their way to the
cortex, basal nuclei, and motor centers in the brainstem. Purkinje cells of the cerebellar cortex. Information about the motor
The cerebellum compares the motor commands with commands issued at the conscious and subconscious levels reaches
proprioceptive information (position sense) and stimulates the Purkinje cells indirectly, after being relayed by nuclei in the pons
any adjustments needed to make the movement smooth. or by the cerebellar nuclei embedded within the arbor vitae.
 508 UNIT 3 Control and Regulation

Figure 14–9 The Cerebellum.

Vermis Vermis

Anterior lobe

Primary fissure

Folia

Posterior
lobe

Left Hemisphere Right Hemisphere Left Hemisphere Right Hemisphere

of Cerebellum of Cerebellum of Cerebellum of Cerebellum

a The posterior, superior surface of the cerebellum, showing major anatomical landmarks and regions
14

Dendrites

Cell body of
Purkinje cell

Purkinje cells LM × 100 The Cerebellum

Region/Nuclei Function
GRAY MATTER
Anterior lobe
Cerebellar cortex Involuntary coordination and control of
Midbrain ongoing body movements
Cerebellar nuclei Involuntary coordination and control of
ongoing body movements
WHITE MATTER
Choroid plexus Pons
of the fourth
Arbor vitae Connects cerebellar cortex and nuclei
ventricle
with cerebellar peduncles
Flocculonodular
lobe
Cerebellar peduncles
Superior Link cerebellum with midbrain,
Medulla oblongata diencephalon, and cerebrum
Middle Carry communications between the
Posterior lobe
cerebellum and pons
Inferior Link the cerebellum with the medulla
oblongata and spinal cord

Transverse pontine Interconnect pontine nuclei with the

b A sectional view of the cerebellum, showing the fibers opposite cerebellar hemisphere
arrangement of gray matter and white matter

? What makes up the gray matter in the cerebellum?

 510 UNIT 3 Control and Regulation

Figure 14–10 The Thalamus. Table 14–2 The Thalamus

Nuclei/Body Function
Frontal lobe Parietal lobe
Anterior nuclei Part of the limbic system
Medial nuclei Integrate sensory information for
Occipital projection to the frontal lobes
lobe
Ventral nuclei Project sensory information to the primary
sensory cortex; relay information from
cerebellum and basal nuclei to motor area
of cerebral cortex
Dorsal nuclei
Lateral dorsal nucleus Projects information to parietal,
occipitoparietal, and temporal cortex; may
play a role in memory
Pulvinar nuclei Integrate sensory information for
a A lateral projection to association areas of cerebral
cortex
view of the brain, color coded
to indicate the regions that Lateral geniculate body Projects visual information to the visual
receive input from the thalamic cortex; integrates sensory information and
influences emotional states
nuclei shown in part b
Medial geniculate body Projects auditory information to the
auditory cortex; integrates sensory
Limbic Frontal Parietal lobe Association areas information and influences emotional
system lobes and cingulate of cerebral cortex states
14
gyrus

Medial nuclei
Anterior
nuclei
Lateral nuclei components of the brainstem. It also controls a variety of auto-
Pulvinar nomic functions and forms the link between the nervous and
endocrine systems.
V e n t r a l
nucl ei Auditory
input Structure of the Hypothalamus
Medial
The hypothalamus extends from the area superior to the
geniculate optic chiasm (the location where axons within the optic nerves
Basal
body arrive at the brain and cross over) to the posterior margins of
nuclei
General Lateral the mammillary (mammilla, nipple) bodies (Figure 14–11a).
sensory Visual geniculate Immediately posterior to the optic chiasm, a narrow stalk
Cerebellum input input body
called the infundibulum (in-fun-DIB- u -lum; infundibulum,
b An enlarged view of the thalamic nuclei of the left side funnel) extends inferiorly, connecting the floor of the hypo-
thalamus to the pituitary gland (Figure 14–11b). The floor of
the hypothalamus between the infundibulum and the mam-
■ The lateral geniculate (je-NIK-y u -l a t) body receives millary bodies is the tuber cinereum (T U -ber si-nir-E -um;
visual information over the optic tract, which originates at tuber, swelling).
the eyes. The output of the lateral geniculate body goes
to the occipital lobes of the cerebral hemispheres and to the Functions of the Hypothalamus
midbrain. The medial geniculate body relays auditory The hypothalamus contains important nuclei that function
information to the appropriate area of the cerebral cortex as control and integrative centers in addition to those asso-
from specialized receptors of the internal ear. This body ciated with the limbic system. These centers are shown in
forms feedback loops with the limbic system and the Figure 14–11b, along with a summary of their functions.
parietal lobes of the cerebral hemispheres. Hypothalamic centers may be stimulated by (1) sensory
information from the cerebrum, brainstem, and spinal
cord; (2) changes in the compositions of the CSF and inter-
The Hypothalamus stitial fluid; or (3) chemical stimuli in the circulating blood
The hypothalamus contains centers involved with emotions that move rapidly across highly permeable capillaries to enter
and visceral processes that affect the cerebrum as well as the hypothalamus (where there is no blood-brain barrier).
 Chapter 14 The Brain and Cranial Nerves 511

Figure 14–11 The Hypothalamus in Sagittal Section.

Parietal
lobe
Corpus
callosum

Septum
pellucidum Choroid plexus

Thalamus
(surrounds
Fornix
third ventricle)
Anterior Pineal gland
cerebral artery

Frontal lobe Hypothalamus

Cerebral
Anterior aqueduct 14
commissure
Cerebellum
Optic chiasm
Optic nerve Fourth
ventricle
a The hypothalamus and adjacent Infundibulum (cut) Tuber cinereum Mammillary body
portions of the brain

Interthalamic adhesion

The Hypothalamus
Thalamus Region/Nuclei Function
Paraventricular nucleus Secretes oxytocin; stimulates smooth muscle
contractions in uterus and mammary glands

Pre-optic area Regulates body temperature by control of

autonomic centers in the medulla oblongata
Hypothalamus
Tuber
cinereum Autonomic centers Control heart rate and blood pressure by
regulation of autonomic centers in the medulla
Sympathetic
oblongata
Optic Parasympathetic
chiasm
Lateral tuberal nuclei Produce inhibitory and releasing hormones that
Mid- control endocrine cells of the anterior lobe of the
Infundibulum brain pituitary gland (adenohypophysis)

Mammillary bodies Control feeding reflexes (licking, swallowing)

Suprachiasmatic nucleus Regulates daily (circadian) rhythms

Pituitary Pons
gland
Supra-optic nucleus Secretes antidiuretic hormone; restricts water loss
by the kidneys

b A diagram of the hypothalamus, showing the locations

and functions of major nuclei and centers
 Chapter 14 The Brain and Cranial Nerves 513

the limbic lobe. The cingulate (SIN-gy u -l a t) gyrus (cingulum, Another cerebral nucleus is the amygdaloid (ah-MIG-
belt) sits superior to the corpus callosum. The dentate gyrus duh-loyd; amygdale, almond) body, commonly referred to as
and the parahippocampal (pah-rah-hip-o-KAM-pal) gyrus the amygdala (Figure 14–12b). It appears to act as an interface
form the posterior and inferior portions of the limbic lobe between the limbic system, the cerebrum, and various sensory
(Figure 14–12b). systems. It plays a role in regulating heart rate, in responding
These gyri conceal the hippocampus, a nucleus inferior to to fear and anxiety as well as controlling the “fight or flight”
the floor of the lateral ventricle. To early anatomists, this struc- response by the sympathetic division of the autonomic nervous
ture resembled a sea horse (hippocampus) when removed from system (ANS), and in linking emotions with specific memories.
the brain. This nucleus is important in learning, especially in The fornix (FOR-niks, arch) is a tract of cerebral white
the storage and retrieval of new long-term memories. matter that connects the hippocampus with the hypothalamus

Figure 14–12 The Limbic System.

Corpus Pineal
callosum Fornix gland

Cingulate gyrus 14
(superior portion of
limbic lobe)

Anterior
thalamic nuclei Parahippocampal
gyrus (inferior portion
Hypothalamus of limbic lobe)
Mammillary body
Hippocampus (within
Temporal lobe dentate gyrus, the posterior
of cerebrum portion of limbic lobe)
a A diagrammatic sagittal section through the cerebrum,
showing the cortical areas associated with the limbic
system. The parahippocampal gyrus is shown as though
transparent to make deeper limbic components visible.

The Limbic System

Cerebral Components
The Limbic System Limbic lobe cortical areas
Diencephalic Components Corpus callosum Cingulate gyrus (superior
Thalamus portion of limbic lobe)
Anterior nuclei Parahippocampal gyrus
(inferior portion of limbic lobe)
Hypothalamus Dentate gyrus (posterior por-
Hypothalamic nuclei tion of the limbic lobe)
Mammillary body Tracts
Other Components Fornix
Reticular formation Nuclei
(not shown)
Olfactory Amygdaloid body
tract
Hippocampus (within
b A three-dimensional reconstruction of dentate gyrus)
the limbic system, showing the relationships
among the major components.
 Chapter 14 The Brain and Cranial Nerves 515

Figure 14–13 The Brain in Lateral View.

Central sulcus

Frontal
lobe
Parietal lobe

Occipital lobe

Lateral
sulcus Precentral Central Postcentral
Temporal Cerebellum gyrus sulcus gyrus
lobe
Pons
Frontal lobe
Medulla oblongata Parietal lobe
a Lateral view, cadaver brain

14

Occipital
lobe

Temporal
Retractor lobe
Lateral sulcus

Cerebellum

Pons

Medulla oblongata
b Lateral view

Precentral gyrus Central sulcus Postcentral gyrus

Cingulate
Insula gyrus
Parietal lobe
c Retractors along the lateral
sulcus showing the insula Frontal lobe
Parieto-
occipital
sulcus

Occipital
lobe

Pons
Temporal lobe Cerebellum

Medulla oblongata
d Midsagittal section

? The central sulcus divides which two cerebral lobes?

 516 UNIT 3 Control and Regulation

Figure 14–14 Fibers of the White Matter of the Cerebrum.

Fibers of the White Matter of the Cerebrum

Fibers and Commissures Function
Association fibers Interconnect cortical areas within the
same hemisphere
Arcuate fibers Interconnect gyri within a lobe

a Lateral view Longitudinal fasciculi Interconnect the frontal lobe with other
cerebral lobes

Commissures Interconnect and permit communication

between the cerebral hemispheres
Corpus callosum
Longitudinal
cerebral fissure Anterior commissure
14

Projection fibers Connect cerebral cortex to diencephalon,

brainstem, cerebellum, and spinal cord
Internal capsule

b Anterior view

■ Projection fibers link the cerebral cortex to the (Figure 14–15a). They are embedded in the white matter of
diencephalon, brainstem, cerebellum, and spinal cord. All the cerebrum. The radiating projection fibers and commissural
projection fibers pass through the diencephalon. There, fibers travel around or between these nuclei.
axons heading to sensory areas of the cerebral cortex pass Historically, the basal nuclei have been considered part
among the axons descending from motor areas of the cortex. of a larger functional group known as the basal ganglia.
In gross dissection, the ascending fibers and descending This group included the basal nuclei of the cerebrum and
fibers look alike. The entire collection of projection fibers is the associated motor nuclei in the diencephalon and mid-
known as the internal capsule (Figure 14–14b). brain. We will consider the functional interactions among
these components in Chapter 15, but we will avoid the term
“basal ganglia” because ganglia are otherwise restricted to
The Basal Nuclei
the PNS.
While the cerebral cortex is consciously directing a complex The caudate (KOW-d a t) nucleus has a large head and a
movement or solving some intellectual puzzle, other centers slender, curving tail that follows the curve of the lateral ven-
of the cerebrum, diencephalon, and brainstem are processing tricle. The head of the caudate nucleus lies anterior to the len-
sensory information and issuing motor commands outside tiform nucleus. The lentiform nucleus consists of a lateral
conscious awareness. Many of these activities, which occur at putamen (p u -T A -men; Figure 14–15b) and a medial glo-
the subconscious level, are directed by the basal nuclei. bus pallidus (GLO-bus PAL-ih-dus; pale globe). The term
corpus striatum (striated body) has been used to refer to the
Structure of the Basal Nuclei caudate and lentiform nuclei, or to the caudate nucleus and
The basal nuclei are masses of gray matter that lie within putamen. The name refers to the striated (striped) appearance
each hemisphere deep to the floor of the lateral ventricle of the internal capsule as its fibers pass among these nuclei.
 Chapter 14 The Brain and Cranial Nerves 517

Figure 14–15 The Basal Nuclei.

Corpus callosum

Lateral ventricle
(anterior horn)
Head of caudate nucleus
Septum pellucidum
Internal capsule
Fornix (cut edge)
Putamen

Thalamus
Third ventricle
Choroid plexus
Pineal gland Fornix

Lateral ventricle
Head of caudate (posterior horn) 14
nucleus Lentiform
nucleus

Tail of caudate
nucleus

Amygdaloid Thalamus
body b A superior view of two transverse
sections at different levels
a The relative
positions of the
basal nuclei in the
intact brain, lateral view

The Basal Nuclei

Corpus callosum Nuclei Function
Caudate nucleus Subconscious
Lateral ventricle adjustment and
modification of voluntary
Septum motor commands
pellucidum
Internal Lentiform nucleus Subconscious
capsule adjustment and
Putamen
modification of voluntary
Lateral sulcus Globus pallidus motor commands
Insula

Anterior
commissure Claustrum

Tip of inferior
horn of lateral
ventricle Amygdaloid body

c Frontal section

? Which portion of the lentiform nucleus is more medial and which portion is more lateral?
 Chapter 14 The Brain and Cranial Nerves 519

Figure 14–16 Motor, Sensory, and Association Areas of the Cerebral Cortex.

Central sulcus
Parietal Lobe
Frontal Lobe
Primary
Primary motor cortex somatosensory cortex

Premotor cortex
Somatosensory
Retractor association cortex

Occipital Lobe

Visual association area

Prefrontal cortex

Visual cortex

Gustatory cortex

Insula
Temporal Lobe
Lateral sulcus
Auditory association area
Auditory cortex 14
a Major anatomical landmarks on the surface
of the left cerebral hemisphere. The lateral Olfactory cortex
sulcus has been pulled apart to expose the insula.

Frontal eye field

Broca’s Wernicke’s
area area

Prefrontal
cortex

b The left hemisphere generally contains the specialized

language areas. The prefrontal cortex of each hemisphere
is involved with conscious intellectual functions.

Sensory Areas of the occipital lobe receives visual information. The auditory
Like the monitoring gauges in the dashboard of a car, the sen- cortex and olfactory cortex of the temporal lobe receive infor-
sory areas of the cerebral cortex report key information. At mation about hearing and smell, respectively. The gustatory
each location, sensory information is reported in the pattern cortex, which receives information from taste receptors of the
of neuron activity in the cortex. The surface of the postcentral tongue and pharynx, lies in the anterior portion of the insula
gyrus contains the primary somatosensory cortex. Neurons and adjacent portions of the frontal lobe.
in this region receive general somatic sensory information
from receptors for touch, pressure, pain, vibration, or temper- Association Areas
ature. A person is aware of these sensations only when nuclei The sensory and motor regions of the cortex are connected to
in the thalamus relay the information to the primary somato- nearby association areas, regions of the cortex that interpret
sensory cortex. incoming data or coordinate a motor response (Figure 14–16a).
Sight, sound, smell, and taste sensations arrive at other Like the information provided by the gauges in a car, the arriv-
areas of the cerebral cortex (Figure 14–16a). The visual cortex ing data must be noticed and interpreted before the driver
 FOCUS Cranial Nerves

Cranial nerves are PNS components that connect directly to the to other nuclei or to processing centers in the cerebral or cer-
brain. The 12 pairs of cranial nerves are visible on the ventral ebellar cortex. In a similar way, the motor nuclei receive con-
surface of the brain (Figure 14–18). Each has a name related to vergent inputs from higher centers or from other nuclei along
its distribution or its function. the brainstem.
The number assigned to a cranial nerve corresponds to the In this section, we classify cranial nerves as primarily sen-
nerve’s position along the longitudinal axis of the brain, be- sory, special sensory, motor, or mixed (sensory and motor). In
ginning at the cerebrum. In scientific writing, the abbreviation this classification, sensory nerves carry somatic sensory infor-
for cranial nerve, CN, is followed by a Roman numeral. For mation, including touch, pressure, vibration, temperature, or
example, CN II refers to cranial nerve II or the optic nerve. If pain. Special sensory nerves carry the sensations of smell, sight,
the full name of the cranial nerve is given, then only the Roman hearing, or balance. Motor nerves are dominated by the axons
numeral is necessary, such as optic nerve (II). of somatic motor neurons. Mixed nerves have a mixture of sen-
Each cranial nerve attaches to the brain near the associated sory and motor fibers. This classification scheme is useful, but
sensory or motor nuclei. The sensory nuclei act as switching it is based on the primary function, and a cranial nerve can
centers, with the postsynaptic neurons relaying the information have important secondary functions. Here are three examples:

Figure 14–18 Origins of the Cranial Nerves. An inferior view of the brain.

Cranial Nerves

Olfactory bulb: termination

of olfactory nerve (I)

Olfactory tract
Optic nerve (II)
Optic chiasm
Oculomotor nerve (III)
Infundibulum

Optic tract Trochlear nerve (IV)

Mammillary body
Trigeminal nerve (V)

Basilar artery Abducens nerve (VI)

Pons Facial nerve (VII)

Vertebral arteries
Vestibulocochlear
nerve (VIII)
Medulla oblongata
Glossopharyngeal
nerve (IX)
Cerebellum
Vagus nerve (X)

Hypoglossal nerve (XII)

Spinal cord Accessory nerve (XI)

523
■ The olfactory receptors, the visual receptors, and the Figure 14–19 The Olfactory Nerve.
receptors of the internal ear are innervated by cranial
nerves that are dedicated almost entirely to carrying special Olfactory tract Left olfactory
(to olfactory cortex bulb (termination
sensory information. In contrast, the sensation of taste, of olfactory nerve)
of cerebrum)
considered to be one of the special senses, is carried by
axons that form only a small part of large cranial nerves
that have other primary functions.
■ As elsewhere in the PNS, a nerve containing tens of Olfactory
thousands of motor fibers that lead to a skeletal muscle nerve (I)
also contains sensory fibers from muscle spindles and
Golgi tendon organs (proprioceptive sensory nerve Olfactory
endings) in that muscle. We assume that these sensory nerve fibers
fibers are present but ignore them in classifying the
nerve.
■ Regardless of their other functions, several cranial
nerves (III, VII, IX, and X) distribute autonomic fibers to
peripheral ganglia, just as spinal nerves deliver them to
ganglia along the spinal cord. We note the presence of
small numbers of autonomic fibers (discussed further in
Chapter 16) but ignore them in the classification of the
Cribriform plate Olfactory
nerve. of ethmoid epithelium
Cranial nerves are clinically important, in part because they
can provide clues to underlying CNS problems. Clinicians as-
sess cranial nerve function using various standardized tests.
conchae, and the superior parts of the nasal septum. Axons
from these sensory neurons collect to form 20 or more
Tips
bundles that penetrate the cribriform plate of the ethmoid.
Two useful mnemonics for remembering the names of the These bundles are components of the olfactory nerves (I).
cranial nerves in numerical order are “Oh Oh Oh, To Touch These bundles enter the olfactory bulbs, neural masses on
And Feel Very Green Vegetables, Ah Heaven!” and “Oh, either side of the crista galli. The olfactory afferents synapse
Once One Takes The Anatomy Final, Very Good Vacations within the olfactory bulbs. The axons of the postsynaptic
Are Heavenly!”
neurons proceed to the cerebrum along the slender olfac-
tory tracts (Figures 14–19, 14–20).
The Olfactory Nerves (I) Because the olfactory tracts look like typical peripheral
nerves, anatomists about a century ago misidentified these
■ Primary function: Special sensory (smell) tracts as the first cranial nerve. Later studies demonstrated that
■ Origin: Receptors of olfactory epithelium the olfactory tracts and bulbs are part of the cerebrum, but by
then the numbering system was already firmly established.
■ Pass through: Olfactory foramina in cribriform plate of
Anatomists were left with a forest of tiny olfactory nerve bun-
ethmoid
dles lumped together as cranial nerve I.
■ Destination: Olfactory bulbs
The olfactory nerves are the only cranial nerves attached
The first pair of cranial nerves carries special sensory infor- directly to the cerebrum. The rest originate or terminate within
mation responsible for the sense of smell (Figure 14–19). nuclei of the diencephalon or brainstem, and the ascending
The olfactory receptors are specialized neurons in the epithe- sensory information synapses in the thalamus before reaching
lium covering the roof of the nasal cavity, the superior nasal the cerebrum.

524
 optic chiasm (chiasma, a crossing). At the optic chiasm, fibers
Figure 14–20 The Optic Nerve.
from the medial half of each retina cross over to the opposite
side of the brain.
The reorganized axons continue toward the lat-
Eye eral geniculate body of the thalamus as the optic tracts
Pituitary
Olfactory bulb (Figures 14–19, 14–21). After synapsing in the lateral
gland
Olfactory tract geniculate body, projection fibers deliver the information to
Optic nerve (II) the visual cortex of the occipital lobes. With this arrangement,
each cerebral hemisphere receives visual information from the
Optic chiasm lateral half of the retina of the eye on that side and from the
medial half of the retina of the eye of the opposite side. A few
Optic tract axons in the optic tracts bypass the lateral geniculate body and
synapse in the superior colliculi of the midbrain. That pathway
is discussed in Chapter 17, The Special Senses.

The Oculomotor Nerves (III)

■ Primary function: Motor (eye movements)

Lateral
geniculate ■ Origin: Midbrain
body (in ■ Pass through: Superior orbital fissures of sphenoid
thalamus)
■ Destination: Somatic motor: superior, inferior, and
medial rectus muscles; inferior oblique; levator palpebrae
Midbrain Visual cortex Optic projection
(cut) (in occipital lobes) fibers superioris. Visceral motor: intrinsic eye muscles
The midbrain contains the motor nuclei controlling the third
? Which structure receives visual information before it reaches
the visual cortex?
and fourth cranial nerves. Each oculomotor nerve (III) inner-
vates four of the six extrinsic muscles that move the eye and
the levator palpebrae superioris, which raises the upper eyelid
The Optic Nerves (II) (Figure 14–21). On each side of the brain, CN III emerges from
the ventral surface of the midbrain and penetrates the poste-
■ Primary function: Special sensory (vision) rior wall of the orbit at the superior orbital fissure. Individuals
■ Origin: Retina of eye with damage to this nerve often complain of pain over the eye,
droopy eyelids, and double vision, because the movements of
■ Pass through: Optic canals of sphenoid
the left and right eyes cannot be coordinated properly.
■ Destination: Diencephalon by way of the optic chiasm The oculomotor nerve also delivers preganglionic auto-
The optic nerves (II) carry visual information from special nomic fibers to neurons of the ciliary ganglion. These neu-
sensory ganglia in the eyes (Figure 14–20). These nerves con- rons control intrinsic eye muscles. These muscles change the
tain about 1 million sensory nerve fibers. The optic nerves pass diameter of the pupil, adjusting the amount of light entering
through the optic canals of the sphenoid. Then they converge the eye. They also change the shape of the lens to focus images
at the ventral, anterior margin of the diencephalon, at the on the retina.

525
Figure 14–21 Cranial Nerves Controlling the Extrinsic Eye Muscles.

Optic nerve Optic Oculomotor Trochlear nerve

Superior rectus (II) chiasm nerve (III) (IV)
Superior
oblique

Trochlea

Levator
palpebrae
superioris

Trigeminal
nerve (V), cut

Inferior Vestibulocochlear
oblique nerve (VIII), cut

Facial nerve
(VII), cut
Inferior Ciliary Medial Lateral rectus Abducens
rectus ganglion rectus (cut) nerve (VI)

The Trochlear Nerves (IV) The Abducens Nerves (VI)

■ Primary function: Motor (eye movements) ■ Primary function: Motor (eye movements)
■ Origin: Midbrain ■ Origin: Pons
■ Pass through: Superior orbital fissures of sphenoid ■ Pass through: Superior orbital fissures of sphenoid
■ Destination: Superior oblique ■ Destination: Lateral rectus
A trochlear (TROK-l e -ar; trochlea, a pulley) nerve (IV), the The abducens (ab-DU -senz) nerves (VI) innervate the lateral
smallest cranial nerve, innervates the superior oblique of each rectus, the sixth pair of extrinsic eye muscles (Figure 14–21).
eye (Figure 14–21). The trochlea is a pulley-shaped, ligamen- Contraction of the lateral rectus makes the eye look to the side. In
tous sling. Each superior oblique passes through a trochlea on essence, the abducens causes abduction of the eye. Each abducens
its way to its insertion on the surface of the eye. An individual nerve emerges from the inferior surface of the brainstem at the bor-
with damage to cranial nerve IV or to its nucleus has difficulty der between the pons and the medulla oblongata (Figure 14–22).
looking down and to the side. Along with the oculomotor and trochlear nerves from that side, it
reaches the orbit through the superior orbital fissure.

526
Figure 14–22 The Trigeminal Nerve.

Superior orbital fissure

Ophthalmic
Supra-orbital nerve (V1)
nerves
Semilunar
ganglion
Ciliary ganglion
Pons
Trigeminal
Foramen nerve (V)
rotundum
Maxillary nerve (V2)
Infra-orbital
nerve

Foramen ovale

Otic ganglion
Lingual nerve

Submandibular ganglion Mandibular nerve (V3)

Mental nerve Pterygopalatine ganglion

? What are the three divisions of the trigeminal nerve?

The Trigeminal Nerves (V) The pons contains nuclei associated with three cranial nerves
(V, VI, and VII) and contributes to a fourth (VIII). The trigeminal
■ Primary function: Mixed (sensory and motor) to face
(tr ı -JEM-i-nal) nerves (V), the largest cranial nerves, are
■ Origin: Ophthalmic nerve (V1) (sensory): orbital structures, mixed nerves. Each provides both somatic sensory informa-
cornea, nasal cavity, skin of forehead, upper eyelid, tion from the head and face and motor control over muscles
eyebrow, nose (part). Maxillary nerve (V2) (sensory): lower of mastication. Sensory (posterior) and motor (anterior) roots
eyelid, upper lip, gums, and teeth; cheek; nose, palate, and originate on the lateral surface of the pons (Figure 14–22).
pharynx (part). Mandibular nerve (V3) (mixed): sensory The sensory branch is larger, and the enormous semilunar
from lower gums, teeth, and lips; palate and tongue (part); ganglion contains the cell bodies of the sensory neurons.
motor from motor nuclei of pons As the name implies, the trigeminal has three major divi-
■ Pass through (on each side): Ophthalmic nerve sions. The small motor root contributes to only one of the three.
through superior orbital fissure, maxillary nerve through Tic douloureux (du -luh-ROO; douloureux, painful), or trigeminal
foramen rotundum, mandibular nerve through foramen neuralgia, is a painful condition affecting the area innervated by
ovale the maxillary and mandibular divisions of the trigeminal nerve.
■ Destination: Ophthalmic, maxillary, and mandibular People with the condition often report debilitating pain triggered
nerves to sensory nuclei in pons; mandibular nerve also by contact with the lips, tongue, or gums. The cause of the condi-
innervates muscles of mastication tion is unknown.

527
 The trigeminal nerve divisions are associated with the ciliary, ■ Destination: Sensory: sensory nuclei of pons. Somatic
sphenopalatine, submandibular, and otic ganglia. These are autonomic motor: muscles of facial expression. Visceral motor: lacrimal
(parasympathetic) ganglia whose neurons innervate structures of (tear) gland and nasal mucous glands by way of the
the face. Although its nerve fibers may pass around or through these pterygopalatine ganglion; submandibular and sublingual
ganglia, the trigeminal nerve does not contain visceral motor fibers. glands by way of the submandibular ganglion
We discussed the ciliary ganglion in the previous Oculomotor
The facial nerves (VII) are mixed nerves. The cell bodies of
Nerves (III) section and describe the other ganglia next, with the
the sensory neurons are found in the geniculate ganglia, and
branches of the facial nerves (VII) and the glossopharyngeal nerves (IX).
the motor nuclei are in the pons. On each side, the sensory
and motor roots emerge from the pons and enter the internal
Tips acoustic meatus of the temporal bone. Each facial nerve then
To learn the sensory distribution of the trigeminal nerve, place passes through the facial canal to reach the face by way of the
the first three fingers across the face with the index finger just stylomastoid foramen. The nerve then splits to form the tem-
below the eye (ophthalmic), the middle finger on the upper lip poral, zygomatic, buccal, marginal mandibular, and cervical
(maxillary), and the third finger (mandibular) on the chin. branches (Figure 14–23).
The sensory neurons monitor proprioceptors in the facial
The Facial Nerves (VII) muscles and provide deep pressure sensations over the face.
They also receive taste information from receptors along the
■ Primary function: Mixed (sensory and motor) to face
anterior two-thirds of the tongue. Somatic motor fibers control
■ Origin: Sensory: taste receptors on anterior two-thirds of the superficial muscles of the scalp and face and deep muscles
tongue. Motor: motor nuclei of pons near the ear.
■ Pass through: Internal acoustic meatus to canals leading The facial nerves carry preganglionic autonomic fibers to
to the stylomastoid foramina the pterygopalatine and submandibular ganglia. Postganglionic

Figure 14–23 The Facial Nerve.

Pterygopalatine Greater petrosal Geniculate Facial nerve

ganglion nerve ganglion (VII)
Temporal
branches
Zygomatic
branches
Buccal
Temporal branches
branch
Marginal
Pons mandibular
branch
Cervical branch

b The superficial distribution

Zygomatic Posterior of the five major branches
branches auricular of the facial nerve
nerve
Buccal
branch
Stylomastoid
foramen
Marginal Chorda tympani with
mandibular mandibular nerve
branch
Lingual branch
Cervical with lingual nerve
branch
Submandibular ganglion
a The origin and branches of the facial nerve

528
fibers from the pterygopalatine ganglia innervate the lacrimal We use vestibulocochlear because this term indicates the names
glands and small glands of the nasal cavity and pharynx. The of its two major nerve branches: the vestibular nerve and the
submandibular ganglia innervate the submandibular and sub- cochlear nerve. Each vestibulocochlear nerve lies posterior
lingual (sub-, under + lingual, pertaining to the tongue) glands. to the origin of the facial nerve, straddling the boundary
Bell’s palsy is a cranial nerve disorder that results from an between the pons and the medulla oblongata (Figure 14–24).
inflammation of a facial nerve. The condition is probably due This nerve reaches the sensory receptors of the internal ear by
to a viral infection. Signs and symptoms include paralysis of entering the internal acoustic meatus in company with the
facial muscles on the affected side and loss of taste sensations facial nerve.
from the anterior two-thirds of the tongue. The condition is Each vestibulocochlear nerve has two distinct bundles of
usually painless. In most cases the symptoms fade after a few sensory fibers. The vestibular (vestibulum, cavity) nerve origi-
weeks or months. nates at the receptors of the vestibule, the portion of the internal
ear concerned with balance sensations. The sensory neurons are
The Vestibulocochlear Nerves (VIII) in an adjacent sensory ganglion, and their axons target the ves-
tibular nuclei of the pons and medulla oblongata. These affer-
■ Primary function: Special sensory: balance and ents convey information about the orientation and movement
equilibrium (vestibular nerve) and hearing (cochlear nerve) of the head. The cochlear (cochlea, snail shell) nerve monitors
■ Origin: Monitor receptors of the internal ear (vestibule the receptors in the cochlea, the portion of the internal ear that
and cochlea) provides the sense of hearing. The cell bodies of the sensory
■ Pass through: Internal acoustic meatus of temporal bones neurons are located within a peripheral ganglion (the spiral gan-
glion), and their axons synapse within the cochlear nuclei of
■ Destination: Vestibular and cochlear nuclei of pons and
the pons and medulla oblongata. Axons leaving the vestibular
medulla oblongata
and cochlear nuclei relay the sensory information to other cen-
The vestibulocochlear nerves (VIII) are also known as the ters or initiate reflexive motor responses. Balance and the sense
acoustic nerves, the auditory nerves, and the stato-acoustic nerves. of hearing are discussed in Chapter 17, The Special Senses.

Figure 14–24 The Vestibulocochlear Nerve.

Tympanic cavity Semicircular Vestibular Facial nerve Internal Vestibulocochlear

(middle ear) canals nerve (VII), cut acoustic meatus nerve (VIII)

CN V

Pons

CN VI
CN VII

CN IX
CN XII
CN X
Medulla
oblongata
CN XI

Tympanic membrane Auditory tube Cochlea Cochlear nerve

529
The Glossopharyngeal Nerves (IX) pharynx. Each glossopharyngeal nerve penetrates the cranium
within the jugular foramen, with cranial nerves X and XI.
■ Primary function: Mixed (sensory and motor) to head The glossopharyngeal nerves are mixed nerves, but sen-
and neck sory fibers are most abundant. The sensory neurons on each
■ Origin: Sensory: posterior one-third of the tongue, part of side are in the superior (jugular) ganglion and inferior
the pharynx and palate, carotid arteries of the neck. Motor: (petrosal) ganglion (Figure 14–25). The sensory fibers carry
motor nuclei of medulla oblongata general sensory information from the lining of the pharynx
■ Pass through: Jugular foramina between the occipital and the soft palate to a nucleus in the medulla oblongata.
bone and the temporal bones These nerves also provide taste sensations from the pos-
terior third of the tongue. Additionally, they have special
■ Destination: Sensory: sensory nuclei of medulla oblongata.
receptors that monitor the blood pressure and dissolved gas
Somatic motor: pharyngeal muscles involved in swallowing.
concentrations in the carotid arteries, major blood vessels
Visceral motor: parotid gland by way of the otic ganglion
in the neck.
The medulla oblongata contains the sensory and motor nuclei The somatic motor fibers control the pharyngeal muscles
of cranial nerves IX, X, XI, and XII, in addition to the vestibular involved in swallowing. Visceral motor fibers synapse in the
nucleus of nerve VIII. The glossopharyngeal (glos-o-fah-RIN- otic ganglion. Postganglionic fibers innervate the parotid
j e -al; glossum, tongue) nerves (IX) innervate the tongue and gland of the cheek.

The Vagus Nerves (X)

Figure 14–25 The Glossopharyngeal Nerve.
■ Primary function: Mixed (sensory and motor), widely
distributed in the thorax and abdomen
■ Origin: Sensory: pharynx (part), auricle and external
acoustic meatus (a portion of the external ear), diaphragm,
and visceral organs in thoracic and abdominopelvic
cavities. Motor: motor nuclei in medulla oblongata

Pons
CN V
CN VII
CN VIII
CN VI
Glossopharyngeal
Otic ganglion nerve (IX)
Medulla oblongata
Inferior (petrosal)
ganglion Superior (jugular) ganglion

Pharyngeal branches
Lingual branch
Parotid gland

Carotid branch

Carotid body

Carotid sinus

Common carotid artery

530
 Pharyngeal Vagus nerve
branch (X)
Pons

Medulla oblongata
Auricular branch
to external ear
Superior ganglion
of vagus nerve
Inferior ganglion
of vagus nerve
Pharyngeal branch

Superior laryngeal nerve

Internal branch
Superior Figure 14–26 The Vagus Nerve.
laryngeal nerve External branch

Recurrent Cardiac branches ■ Pass through: Jugular foramina between

laryngeal nerve the occipital bone and the temporal bones
Cardiac plexus ■ Destination: Sensory: sensory nuclei
and autonomic centers of medulla
Right lung Left lung
oblongata. Visceral motor: muscles of the
palate, pharynx, digestive, respiratory, and
cardiovascular systems in the thoracic and
abdominal cavities
The vagus (V A -gus) nerves (X) arise imme-
diately posterior to the attachment of the
glossopharyngeal nerves. Many small rootlets
contribute to their formation. Developmental
studies indicate that these nerves probably
represent the fusion of several smaller cranial
nerves during our evolutionary history. Vagus
means wandering, and the vagus nerves branch
and radiate extensively. Figure 14–26 shows
Liver only the general pattern of distribution.
Anterior Sensory neurons are in the superior
vagal trunk
(jugular) ganglion and the inferior(nodose;
NO-dos) ganglion. Each vagus nerve provides
Spleen
somatic sensory information about the exter-
Celiac plexus
nal acoustic meatus and the diaphragm, and
special sensory information from pharyngeal
Stomach taste receptors. But most of the vagal afferents
carry visceral sensory information from recep-
Pancreas
tors along the esophagus, respiratory tract, and
Colon abdominal viscera as distant as the last por-
Small intestine tions of the large intestine. This visceral sensory
information is vital to the autonomic control of
Hypogastric
plexus
visceral function.

531
 The motor components of the vagus are equally diverse. The accessory nerves (XI) are also known as the spinal acces-
Each vagus nerve carries preganglionic autonomic (parasympa- sory nerves or the spino-accessory nerves. Unlike other cranial
thetic) fibers that affect the heart and control smooth muscles nerves, each accessory nerve has some motor fibers that origi-
and glands within the areas monitored by its sensory fibers, nate in the lateral part of the anterior gray horns of the first
including the stomach, intestines, and gallbladder. Difficulty five cervical segments of the spinal cord (Figure 14–27). These
in swallowing is one of the most common signs of damage to somatic motor fibers form the spinal root of nerve XI. They
either cranial nerve IX or X. Damage to either one prevents the enter the cranium through the foramen magnum. They then
coordination of the swallowing reflex. join the motor fibers of the cranial root, which originates at a
nucleus in the medulla oblongata. The composite nerve leaves
The Accessory Nerves (XI) the cranium through the jugular foramen and divides into two
branches.
■ Primary function: Motor to muscles of the neck and The internal branch of nerve XI joins the vagus nerve
upper back and innervates the voluntary swallowing muscles of the
■ Origin: Motor nuclei of spinal cord and medulla oblongata soft palate and pharynx and the intrinsic muscles that
■ Pass through: Jugular foramina between the occipital control the vocal cords. The external branch of nerve XI
bone and the temporal bones controls the sternocleidomastoid and trapezius of the neck
and back. The motor fibers of this branch originate in the
■ Destination: Internal branch innervates voluntary muscles
lateral gray part of the anterior horns of cervical spinal
of palate, pharynx, and larynx; external branch controls
nerves C 1 to C5.
sternocleidomastoid and trapezius

Figure 14–27 The Accessory and Hypoglossal Nerves.

Hypoglossal nerve (XII)

Trigeminal nerve (V)

Accessory nerve (XI)

Medulla oblongata

Cranial root of CN XI
Internal branch: to palatal,
pharyngeal, and laryngeal
muscles with vagus nerve

Spinal
Intrinsic muscles
root of CN XI
of tongue
Styloglossus
External
Genioglossus branch of CN XI

Geniohyoid Spinal cord

Hyoglossus

Hyoid bone
Trapezius
Thyrohyoid Sternocleidomastoid

Sternohyoid
Ansa cervicalis
Sternothyroid (cervical plexus)

Omohyoid

532
The Hypoglossal Nerves (XII) to reach the skeletal muscles of the tongue (Figure 14–27). This
cranial nerve provides voluntary motor control over move-
■ Primary function: Motor (tongue movements) ments of the tongue. Its condition is checked by having a per-
■ Origin: Motor nuclei of medulla oblongata son stick out their tongue. Damage to one hypoglossal nerve
■ Pass through: Hypoglossal canals of occipital bone or to its associated nuclei causes the tongue to veer toward the
affected side.
■ Destination: Muscles of the tongue
Table 14–4 summarizes the basic distribution and func-
Each hypoglossal (h ı -po-GLOS-ul) nerve (XII) leaves the tion of each cranial nerve.
cranium through the hypoglossal canal. The nerve then curves

Table 14–4 Cranial Nerve Branches and Functions

Cranial Nerve Sensory Primary
(Number) Ganglion Branch Function Foramen Innervation
Olfactory (I) Special Olfactory foramina of Olfactory epithelium
sensory ethmoid
Optic (II) Special Optic canal Retina of eye
sensory
Oculomotor (III) Motor Superior orbital fissure Superior, inferior, and medial rectus muscles;
inferior oblique; levator palpebrae superioris;
intrinsic eye muscles
Trochlear (IV) Motor Superior orbital fissure Superior oblique
Trigeminal (V) Semilunar Mixed Superior orbital fissure Areas associated with the jaws
Ophthalmic Sensory Superior orbital fissure Orbital structures, nasal cavity, skin of forehead,
nerve (V1) upper eyelid, eyebrows, nose (part)
Maxillary Sensory Foramen rotundum Lower eyelid; superior lip, gums, and teeth; cheek,
nerve (V2) nose (part), palate, and pharynx (part)
Mandibular Mixed Foramen ovale Sensory: inferior gums, teeth, lips, palate (part),
nerve (V3) and tongue (part)
Motor: muscles of mastication
Abducens (VI) Motor Superior orbital fissure Lateral rectus
Facial (VII) Geniculate Mixed Internal acoustic meatus Sensory: taste receptors on anterior 2/3 of tongue
to facial canal; exits at Motor: muscles of facial expression, lacrimal gland,
stylomastoid foramen submandibular gland, sublingual glands
Vestibulocochlear Cochlear nerve Special Internal acoustic meatus Cochlea (receptors for hearing) Vestibule (receptors
(VIII) Vestibular sensory for motion and balance)
nerve
Glossopharyngeal Superior Mixed Jugular foramen Sensory: posterior 1/3 of tongue; pharynx and
(IX) (jugular) and palate (part); receptors for blood pressure, pH,
inferior (petrosal) oxygen, and carbon dioxide concentrations
Motor: pharyngeal muscles and parotid gland
Vagus (X) Superior Mixed Jugular foramen Sensory: pharynx; auricle and external acoustic
(jugular) and meatus; diaphragm; visceral organs in thoracic and
inferior (nodose) abdominopelvic cavities
Motor: palatal and pharyngeal muscles and visceral
organs in thoracic and abdominopelvic cavities
Accessory (XI) Internal Motor Jugular foramen Skeletal muscles of palate, pharynx, and larynx
(with vagus nerve)
External Motor Jugular foramen Sternocleidomastoid and trapezius
Hypoglossal (XII) Motor Hypoglossal canal Tongue musculature

533