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Anatomy of the Brainstem:

A Gaze Into the Stem of Life


M. Ángeles Fernández-Gil, R. Palacios-Bote, M. Leo-Barahona, and J.P. Mora-Encinas

The brainstem has an ectodermal origin and is composed of 4 parts: the diencephalon,
mesencephalon, pons, and medulla oblongata. It serves as the connection between the
cerebral hemispheres with the medulla and the cerebellum and is responsible for basic vital
functions, such as breathing, heartbeat blood pressure, control of consciousness, and
sleep. The brainstem contains both white and gray matter. The gray matter of the brainstem
(neuronal cell bodies) is found in clumps and clusters throughout the brainstem to form the
cranial nerve nuclei, the reticular formation, and pontine nuclei. The white matter consists
of fiber tracts (axons of neuronal cells) passing down from the cerebral cortex—important
for voluntary motor function—and up from peripheral nerves and the spinal cord—where
somatosensory pathways travel—to the highest parts of the brain. The internal structure of
brainstem, although complex, presents a systematical arrangement and is organized in 3
laminae (tectum, tegmentum, and basis), which extend its entire length. The motor pathway
runs down through the basis, which is located at the most anterior part. The cranial nerve
nuclei are settled into the middle layer (the tegmentum), just in front of the 4th ventricle and
are placed, from medial to lateral, on the basis of their function: somatic motor, visceral
motor, visceral sensory, and somatic sensory. All the somatosensory tracts run upward to
the thalamus crossing the tegmentum in front of the cranial nerve nuclei. The tectum,
formed by the quadrigeminal plate and the medullary velum, contains no cranial nuclei, no
tracts and no reticular formation. The knowledge of precise anatomical localization of a
lesion affecting the brainstem is crucial in neurological diagnosis and, on this basis, is
essential to be familiar with the location of the mayor tracts and nuclei appropriately.
Nowadays, current magnetic resonance imaging techniques, although still macroscopic,
allow the fine internal structure of the brainstem to be viewed directly and make it possible
to locate the main intrinsic structures that justify the symptoms of the patient. In this article
we discuss the anatomy of the brainstem and highlight the features and landmarks that are
important in interpreting magnetic resonance imaging.
Semin Ultrasound CT MRI 31:196-219 © 2010 Elsevier Inc. All rights reserved.

T he brainstem is the lower extension of the brain connect-


ing it to the spinal cord and cerebellum. It is made up of
4 sections: diencephalon, midbrain, pons, and medulla ob-
clinically relevant neuroanatomy of the brainstem by provid-
ing the necessary details to approach clinical situations in
neurology.
longata (Fig. 1). Only by knowing the organization of the As we have indicated, the internal structure of brainstem
brainstem can we assist in patient diagnosis. Therefore, to has a systematical arrangement and may be understood by
understand the patient symptoms we need to know the ana- following a logical sequence. We will attempt to clarify some
tomical substrate. Consequently, the knowledge of precise notions and provide some topographic clues and rules that
anatomical localization of the lesion is crucial in neurological may make it easier to remember the location of the most
diagnosis and familiarity with the location of the major tracts important structures. To this, we must begin by explaining
and nuclei is essential. Our aim in this article is to fulfill the some indispensable basic concepts.
need for a succinct but comprehensible overall perspective of
Basic Concepts
Radiology Department, Complejo Hospitalario Universitario de Badajoz, in Brainstem Anatomy
Hospital Infanta Cristina, Madrid, Spain.
Address reprint requests to: Jesús Rincón Jiménez, 82 5°D 06010, Badajoz, The basic functional unit in the central nervous system (CNS) is
Spain. E-mail: mafdezgil@telefonica.net the neuron, which is formed by a cell body, from whose surface

196 0887-2171/10/$-see front matter © 2010 Elsevier Inc. All rights reserved.
doi:10.1053/j.sult.2010.03.006
Anatomy of the brainstem 197

in the CNS is called tract, fasciculus, brachium, peduncle, or


lemniscus. When this bundle of axons leaves the CNS (the
segment that connects the CNS with peripheral structures) it
is called a nerve.1
The internal structure of the brainstem is organized into 3
laminae, which extend its entire length. From dorsal to ven-
tral, these laminae are tectum, tegmentum, and basis. Their
composition varies. The tectum contains the quadrigeminal
plate in the midbrain, the superior medullary velum in the
pons, and the inferior medullary velum in the medulla ob-
longata. There are no cranial nerve nuclei, no reticular for-
mation, and no tracts.
The basis is the ventral lamina and contains the pyramidal
pathway (corticobulbar and corticospinal tracts). At the pons
it is largest because of the presence of numerous nuclei (pon-
tine nuclei), which receive numerous fibers that project to
cerebellum, forming the pontocerebellar pathway. In the
midbrain the pes cerebri form the basis and in the medulla
this lamina corresponds to the pyramids.
The tegmentum is placed between the tectum and basis
and is divided in 2 layers: the dorsal contains all the somato-
motor and general sensory cranial nerve nuclei. The ventral
contains the supplementary motor nuclei: the substantia
nigra and red nucleus (midbrain) and the inferior olivary
nucleus (medulla), which are located close to the basis. The
running tracts (sensory tracts or lemniscus) cross the rest of
the tegmentum and they are surrounded by the reticular
formation.3
Thus, the gray matter of the brainstem (nuclei) lies close to
the central canal (Sylvian aqueduct and 4th ventricle),
whereas the white matter is in the middle of the tegmentum
(with the exception of the pyramidal pathway, which runs
Figure 1 (A) Schematic sagittal view of the brainstem showing its 4
parts. Yellow: diencephalon; blue: midbrain; gray: pons; red: me-
anteriorly; Fig. 2).
dulla oblongata. (B) Sagittal gross specimen showing the natural
appearance of the brainstem, which correlate with (C) sagittal mag- Brainstem Embryology
netic resonance T1-weighted image. Note the superior medullary
velum (arrow) and the disposition of the medial lemniscus in the The whole CNS has an ectodermic origin. The brain is devel-
sagittal plane (arrowheads). The medulla oblongata and the spinal oped from the anterior end of the neural tube, which at an early
cord junction is at the level of the foramen magnum (dotted line).
(Color version of figure is available online.)

project one or more processes called neurites. Some neurites,


called dendrites, receive information (electrophysiological im-
pulses), which are conducted toward the cell body from where
the impulses travel far away along other lengthy tubular neurite
called axon. The gray matter is formed by the body neurons, and
the white matter corresponds to the axons. Information is chem-
ically transmitted from 1 neuron to other through a connection
between axons and dendrites, or synapse. A chain of communi-
cating neurons is a pathway.
The gray matter of the brain stem (neuronal cell bodies) is
grouped in clusters, which form the nuclei of the cranial nerves.
The white matter is composed by the axons of multiple path-
ways connecting the neurons of the cerebral cortex with the Figure 2 Laminae of the brainstem. Pink: basis; blue: tegmentum;
spinal cord, the cerebellum, and the cranial nerve nuclei. green: tectum. Note that the cranial nerve nuclei are located in front
It is important to know that the same structure may have of the aqueduct and IV ventricle, inside of the tegmentum. (Color
different names. Therefore, a bundle of pathway axons lying version of figure is available online.)
198 M.Á. Fernández-Gil et al

period becomes expanded into 3 vesicles that demarcate the pouches. The cavities of these diverticula are the rudiments
territory for cerebral hemispheres and brainstem. These consti- of the lateral ventricles. The anterior part of the forebrain,
tute the 3 primary cerebral vesicles and correspond, respec- including the rudiments of the cerebral hemispheres, is
tively, to the future forebrain (prosencephalon), midbrain (mes- called telencephalon, and its posterior portion is termed di-
encephalon), and hindbrain (rhombencephalon). encephalon. The cavity of the diencephalon is the rudiment
The midbrain or mesencephalon is the only one that does not of the 3rd ventricle.
undergo any further segmentation. The first (forebrain) and Simultaneously, the rhombencephalon is segmented into
third (hindbrain) divide once more into a series of 5 vesicles that 2 structures: the anterior one, named metencephalon, con-
become the major portions of the CNS within the skull. sisting of the pons, cerebellum, and the intermediate part of
The forebrain grows forward and the cerebral hemispheres the 4th ventricle; and the myelencephalon, which comprises
originate as diverticula, which rapidly expand to form 2 large the medulla oblongata and the lower part of the 4th ventricle.

Figure 3 Sequential diagrammatic picture of embryologic development of the central nervous system. During week 2,
the embryo is a bilaminar disk, consisting of ectoderm and endoderm. The cells of the ectoderm migrate to the median
plane of the dorsum of the embryonic disk and form a linear band called the primitive streak. Cells from the primitive
streak proliferate and migrate down toward the endoderm (to form the embryonic mesoderm) and cranially toward
the prechordal plate as part of the notocordal process, which will develop the notocord. The notocord induces the
overlying ectoderm to form the neural plate that will eventually give rise to the CNS. The neural plate grows along the
notochord axis and beyond. Then, it forms a median longitudinal neural groove with neural folds on either side. At
the top of each fold are the cells that will make up the neural crest. The neural folds move inward and fuse to form the
neural tube. As the tube pinches off, the areas of neural crest cells on either side separate and form a flat mass to each
side of the neural tube. These neural crest cells will become ganglia of spinal, autonomic and cranial nerves. Finally, the
neural tube submerges beneath the ectoderm to become a separate structure. The brain is developed from the anterior
end of the neural tube, which at an early period becomes expanded into 3 vesicles that demarcate the territory for
cerebral hemispheres and brainstem. (Color version of figure is available online.)
Anatomy of the brainstem 199

Figure 6 Axial 3-dimensional balanced gradient-recall echo sequences


shows no gray-white matter differentiation but clearly delineate the edges of
the medulla and the cisternal portions of the cranial nerves. Note the cis-
ternal portions of the lower cranial nerves complex (arrows). Anterior me-
dian fissure (open arrow); pyramids (arrowhead); bulging of the olives
(white arrowhead); and preolivary sulcus (thick arrow).

The 3rd and 4th ventricles are connected by the lumen of


the canal that passes through the mesencephalon, which
tightens and turns into the Sylvian aqueduct. Growth and
development go on and, around the 3rd month of develop-
Figure 4 Anterior view of the medulla oblongata in a gross specimen. p,
ment, the telencephalic vesicles can be easily identified as the
pons; Py, pyramid; thick arrow, anterior median fissure; curved arrow,
cerebral hemispheres, which have grown beyond the dien-
decussation area of the pyramids; o, olive; arrowhead, postolivary
groove; thin arrow, preolivary groove; asterisk, meningeal sheets. cephalon (Fig. 3).3,4
(Color version of figure is available online.)
The Medulla Oblongata
The medulla oblongata connects the spinal cord to the pons.
The junction of the medulla and the spinal cord is located,

Figure 7 Posterior view of the medulla oblongata and floor of the 4th
Figure 5 Coronal inversion-recovery (IR) T1-weighted magnetic reso- ventricle in a gross specimen. Posterior median sulcus (arrow); gracile tu-
nance scan shows a clear discrimination between gray-white matters in bercle (white arrowhead); cuneate tubercle (black arrowhead); inferior cer-
cerebral hemispheres and basal ganglia. This differentiation is scarce in the ebellar peduncles (open arrows); hypoglossal eminence (black asterisk);
brainstem but we can identify some superficial structures, which correlates facial colliculus (black dotted circle); and area acustica (green dotted form).
with those viewed at the gross specimen. P, pons; Py, pyramid; O, olive; Median sulcus (thick arrow). Medial eminence (me); area cinerea (white
and thick arrow, anterior median fissure. Note the cisternal portions of 4th, arrow); striae medullare (thin arrow). (Color version of figure is available
5th, 7th, and 8th cranial nerves (arrows from up to down). online.)
200 M.Á. Fernández-Gil et al

dal tract cross the midline and sweep dorsally and laterally
into the lateral funiculus to form the lateral corticospinal tract
in the spinal cord. The fascicles that cross the midline are
called the decussation of the pyramids. A smaller number of
corticospinal axons in the pyramids remain on the same side
and continue into the spinal cord as the medial corticospinal
tract, located in the ventral funiculus (Figs. 4 and 5). Imme-
diately and posteriorly to the pyramids are 2 rounded and
elongated elevations, the olives, which are shaped by the
underlying inferior olivary nuclei (Figs. 4 and 5).
Between the pyramids and the olives there is a groove, the
preolivary groove, where emerge the rootlets of the hypoglos-
sal nerve (12th cranial nerve). Posterior to the olives is the
postolivary groove, where appear the roots of the glossopha-
ryngeal and vagus nerves and the cranial roots of the acces-
sory nerve (9th, 10th, 11th cranial nerves, respectively; Figs.
4 and 6). Posterior to the olives are the inferior cerebellar
peduncles, which connect the medulla to the cerebellum
(Fig. 7).
The posterior surface of the medulla oblongata is di-
Figure 8 3-T high-resolution axial PD-weighted image shows the pyra-
vided in 2 parts: the lower half, whose surface is continu-
midal pathway (arrowhead), medial lemniscus (arrow), olivary bulging
(asterisk), and inferior cerebellar peduncles (open arrow). ous with the posterior aspect of the spinal cord and where
the posterior median sulcus can be found, and the supe-
rior half, which forms the lower part of the floor of the 4th
approximately, at the level of the foramen magnum (Fig. 1). ventricle. On each side of the median sulcus there are and
On its anterior surface is the anterior median fissure, which 2 elongated swellings, the gracile tubercle, produced by
runs continuously inferior to the anterior median fissure of the underlying gracile nucleus, and lateral to this, the
the spinal cord. On each side of the median fissure there is a cuneate tubercle, produced by the underlying cunate nu-
swelling called the pyramids that are composed of axons cleus (Fig. 7).5-7
from the precentral gyrus of the cerebral cortex that are des- Coronal images through the medulla oblongata clearly
tined to innervate the spinal cord gray matter, the corticospi- may depict the anterior structures as pyramids and olives,
nal or pyramidal tracts. The distal regions of the pyramids which can be demonstrated, as well, in axial sections
become thin, and at that point some fascicles of the pyrami- where cisternal portions of the lower cranial nerve com-

Table 1 Major Nuclei of the Brainstem


Direction: Medial to Lateral
Somatic Visceral Motor Visceral
Motor (Parasympathetic) Sensory Somatic Sensory
Pure sensory
8th vestibulocochlear
Pure motor
3rd oculomotor Edinger-Westphal
4th trochlear
6th abducens
12th Hypoglossal
Mixed
5th trigeminal Motor Nucl. 5 Spinal/mesencephalic nucleus
7th facial Facial Superior. Saliv. Nucl. N. Solitarius. Sens. N. 5
9th glossopharyngeal Ambiguus Inferior. Saliv. Nucl. N. Solitarius. Sens. N. 5
10th vagus Ambiguus Dorsal Motor N. Solitarius. Sens. N. 5
The functions of the major nuclei of the brainstem have a topographical position from medial to lateral, somatic motor, visceral motor, visceral
sensory and somatic sensory. The distribution of the cranial nerves may be studied systematically. There is only one nucleus that is entirely
sensory (8th). There are 5 motor nuclei (3, 4, 6, 12, which divide equally into 12 and which are located in the midline). The 11th cranial nerve
nucleus is Motor but it is not really in the medulla. It is, in fact, a spinal nucleus.
The rest of the nuclei are mixed (sensory plus motor). All of these have 4 components except the 5th, which has 2 components. Solitarius is
Sensory and ambiguus is Motor.
The cranial nerve autonomic fibers are all parasympathetic (3rd, 7th, 9th, and 10th).
Anatomy of the brainstem 201

Figure 9 Cranial nerve nuclei. (A) Scheme of the cranial nerve nuclei within the brainstem. Note the location and the direction of
their axons. (B) dorsal view scheme of the brainstem and cranial nerve nuclei location. Quadrigeminal plate (asterisks); lateral and
medial geniculate bodies (arrows). The nuclei of the cranial nerve 3rd to 12th lie in the brainstem, just in front of the floor of the 4th
ventricle and the sylvian aqueduct. Four are located in the medulla oblongata (9th, 10th, 11th, 12th); 4 in the pons (5th, 6th, 7th,
8th) and 4 above the pons (3rd, 4th lay in the midbrain). The 1st (olfactory) and the 2nd (optic) are above the brainstem. The 4
motor nuclei that are in the midline of the brainstem are those that divides equally into 12 that is 3, 4, 6, and 12. They emerge
through the anterior midline surface, except the 4th, which exits from the dorsal part of the midbrain after its decussation in the
superior medullary velum. Although 5, 7, and 9 cranial nerve nuclei have motor function, they also have sensory components and
they do not divide evenly into 12. Thus, they are not pure motor and, on this basis, they are not located medially. The 8th cranial
nerve nucleus (entirely sensory) and 11th (pure motor, which does not divide equally into 12), are located laterally.20 Salivatory and
lacrimal nuclei (s). (Color version of figure is available online.)

Figure 11 Axial anatomic section at pontomedullary junction stained with


Mulligan’s method, which shows the gray substance tinged in blue. P,
pons; Py, pyramids; icp, inferior cerebellar peduncle; ML, medial lemnis-
cus; arrow, medial longitudinal fasciculus; dotted line, tegmental trajectory
of the fibers of 9th and 10th cranial nerve complex. White arrows, cisternal
portions of the 5th cranial nerve; IV, 4th ventricle; curved arrow, inferior
Figure 10 Axial drawing of the medulla oblongata showing the tegmentum medullary velum; CP, choroid plexus. Note that the blue-colored struc-
(yellow area) where the lower cranial nerve nuclei are located. At the mid- tures correspond to gray matter as the cerebellar cortex, pontine nuclei
line is the 12th (hypoglossal) nerve nuclus, which occupies the somatic (arrowheads), olives (o), and the tegmentum (the area in front of the 4th
motor area. Lateral to this is the visceral motor and visceral sensory (auto- ventricle) where the nuclei of the lower cranial nerves (9th-12th) are lo-
nomic) territory where the motor nucleus of the 10th (vagus) nerve are cated. The white tracts appear as brown structures as the pyramids (Py),
located, as well as the solitary tract (Sol) and the nucleus ambiguus (Amb). medial lemniscus (ML), medial longitudinal fasciculus (black arrow), and
Further laterally, the spinal extension of the sensory nucleus of the 5th inferior cerebellar peduncles (icp); the 12th cranial nerve nuclei are located
nerve and the spinotrigeminal pathway (thick arrow) are visualized. Hypo- at the back in the tegmentum and projects on the floor of the 4th ventricle
glossal eminence (asterisk); O, olive; Py, pyramid; ML, medial lemniscus. forming the hypoglossal eminence (open arrow). (Color version of figure is
(Color version of figure is available online.) available online.)
202 M.Á. Fernández-Gil et al

Figure 12 (A) Axial anatomic section tinged with Mulligan’s method through the mid-pons. The central pons is
located between the middle cerebellar peduncles (mcp), just in the entry zone of the 5th cranial nerve (white
arrows). White matter tracts are well depicted in dark brown color: the corticospinal tracts (cst) at the middle of
the basis pontis are well seen. The medial lemniscus shows a horizontal disposition and serves as the limit between
the tegmentum (posteriorly) and basis (anteriorly). Scp: superior cerebellar peduncle. Pontine nuclei are colored
in blue and occupy the great part of the basis pontis (arrowheads). IV, 4th ventricle; median longitudinal
fasciculus (black arrow); median eminence: asterisk. (B) Axial 3-T high-resolution PD-weighted image. The
magnetic resonance appearance of the brainstem structures correlates exquisitely with the anatomic slice. The
inferior extension of the wings of the ambient cisterns can produce 2 high-intensity areas on either side of the 4th
ventricle and superior cerebellar peduncles (open arrow). (Color version of figure is available online.)

plex may also be seen. Additionally, proton density (PD)-


weighted images demonstrate medial lemniscus, cortico-
spinal tracts, olives, and cerebellar peduncles (Figs. 5, 6,
and 8).
In the tegmentum of medulla oblongata are located the nuclei
of the last 4 cranial nerves (9th, 10th, 11th, and 12th). The
cranial nerve nuclei have a constant relationship going from
medial to lateral in the brainstem because of the embryologic
development of the brainstem (Table 1). In an axial schematic

Figure 14 View into the anterior sector of cerebellopontine angle and axial
anatomic section through the middle of the pons. Cca, cavernous carotid
artery; ps, pituitary stalk; ON, optic nerve; ba, basilar artery. Cisternal por-
tions of 5th, 6th, 7th, 8th, and 12th cranial nerves. Note the motor branch
of the 5th cranial nerve (thin arrow) accompanying the thicker sensory root
Figure 13 Anteriorviewoftheponsandmedullaoblongatagrossspecimen. (arrowhead) and reaching the brainstem just in the junction between the
There are many transverse fibers (pontocerebellar tract) crossing the sur- pons and the middle cerebral peduncle (mcp). Entrance to the Meckel’s
face of the pons (arrows) to arrive at the cerebellum through the middle cave (curved arrow). Cst, Corticospinal tracts; ML, medial lemniscus; scp,
cerebellar peduncle (mcp). The basilar groove may be appreciated in the superior cerebellar peduncles; IV, superior portion of the 4th ventricle; roof
middle (asterisk). Note the decussation of pyramidal tract at the inferior of the 4th ventricle formed by the superior medullary velum (smv); locus
end of the pyramids (curved arrow). (Color version of figure is available caeruleus (black arrow). White arrows: deep portions of the Ambiens cis-
online.) tern. (Color version of figure is available online.)
Anatomy of the brainstem 203

drawing of the medulla the most medially positioned is the


somatic motor nucleus, slightly more lateral is the visceral motor
type, and more laterally is the visceral sensory and somatic sen-
sory types (Figs. 9, 10, and 11).

Glossopharyngeal Nerve Nucleus


(9th) and Vagus Nerve Nucleus (10th)
Glossopharyngeal and vagus nerves are sensory and motor and
share nuclei and function. Therefore, they are considered as a
group. Both nerves have 3 nuclei: (1) the main motor nucleus, (2)
the parasympathetic nucleus, and (3) the sensory nucleus. The
main motor nucleus lies deep in the reticular formation of the me-
dulla oblongata and is formed by the nucleus ambiguus (the supe-
rior end for the ninth and the rest for the 10th), which is situated
dorsal to the inferior olivary nucleus. The efferent fibers supply the
striated musculature of the pharynx (stylopharyngeus muscle, 9th;
constrictor muscles, 10th) and larynx (10th).
The parasympathetic nucleus of the CN9 is also called the infe-
rior salivary nucleus. It receives afferent fibers from the hypothala-
mus through the descending autonomic pathways, from the olfac-
tory system through the reticular formation and information
concerning taste from nucleus of the solitary tract. The postgangli-
onic parasympathetic fibers innervate the parotid gland. Parasym-
pathetic fibers of the CN10 originate from the dorsal motor nucleus
Figure 15 Drawing of the anterior view of the brainstem. mb,
of the vagus nerve, which lies lateral to the hypoglossal nucleus and
mamillary body; cp, cerebral peduncle; P, pons; mcp, medial
protrudes into the floor of the 4th ventricle. The cells of the dorsal
cerebellar peduncle; 3rd, oculomotor nerve; 4th, trochlear nerve;
5th, trigeminal nerve; 6th, abducent nerve; 7th, facial nerve; 8th,
vagal nucleus receive afferent fibers from the hypothalamus
vestibulocochlear nerve; 9th, glossopharyngeal nerve; 10th, va- through the descending autonomic pathways, and its efferent fibers
gus nerve; 11th, spinal nerve; 12th, hypoglossal nerve; arrow, provide innervation to the involuntary muscle of the heart, bronchi,
posterior perforated substance; arrowhead, decussation of the esophagus, stomach, liver, pancreas, proximal intestines, and other
pyramids. (Color version of figure is available online.) viscera.

Figure 16 (A) Sagittal turbo spin echo (TSE) T2-weighted and (B) sagittal SET1-weighted images through the brainstem clearly
depict the midline structures such us hypothalamus (asterisk), infundibulum (arrow) and pineal gland (white arrow), quadrigem-
inal plate (open arrow), superior medullary velum (arrowhead). Note the CSF fluid void passing through the aqueduct and Monro
foramina (thick arrows). P, pons; dotted line; hypothalamic sulcus.
204 M.Á. Fernández-Gil et al

The sensory nucleus of both nerves is the nucleus of the The Pons
tractus solitarius. Sensations of taste travel through the pe-
ripheral axons of glossopharyngeal and the vagus nerve. They The pons is located between the midbrain cranially and the
enter the brainstem together and form the tractus solitarius, medulla oblongata caudally. It received its name because of
whose fibers terminate at the nucleus solitarius. Afferent im- the appearance it presents on the anterior surface, which is
pulses from the carotid sinus, a baroreceptor situated at the similar to a bridge connecting the right and left cerebellar
bifurcation of the common carotid artery, also travel with the
glossopharyngeal nerve and end in the nucleus of the tractus
solitarius. They are connected with the dorsal motor nucleus
of the vagus nerve. On this basis, the carotid sinus reflex
involves glossopharyngeal and vagus nerves, which assist in
the regulation of arterial blood pressure.8 The glossopharyn-
geal and vagus nerves exit the medulla oblongata as a series of
rootlets in a groove between the olive and inferior cerebellar
peduncle called the retro-olivary or posterior lateral sulcus
(Figs. 4, 6, 9, 10, and 15).

Accessory Nerve Nucleus (11th)


The accessory nerve is a motor nerve that is formed by the
union of a cranial and spinal root. It arises from the medulla
and the upper cervical spinal cord to the level of C5 by a
series of rootlets that join to form a single nerve.
The cranial root is formed by the axons of the nerve cells of
the nucleus ambiguus. The efferent fibers of the nucleus
emerge from the anterior surface of the medulla oblongata
between the olive and the inferior cerebral peduncle in com-
pany with 9th and 10th fibers (Figs. 4, 6, 10, and 15).
The spinal root is formed from axons of nerve cells in the
spinal nucleus, which is situated in the anterior gray column
of the spinal cord in the upper 5 cervical segments. The 2
roots unite and leave the skull through the jugular foramen.
Then, the roots separate and the cranial root join the vagus
nerve. The spinal root separates and runs downward and
laterally to supply the sternocleidomastoid and the trapezius
muscles. The accessory nerve thus brings about movements
of the soft palate, pharynx, and larynx and controls the move-
ment of 2 large muscles in the neck.

Hypoglossal Nerve Nucleus (12th)


Hypoglossal nerve is a motor nerve and supplies all the intrinsic
muscles of the tongue and the styloglossus, the hyoglossus and the Figure 17 Axial drawings showing the pons and corresponding 5th,
genioglossus muscles. The hypoglossal nucleus is situated close to 6th, 7th, and 8th cranial nerve nuclei. (A) The 8th cranial nerve
nuclei are located at the lower level, at the acoustic area, a zone
the midline immediately beneath the floor of the lower part of the
dorsal and lateral adjacent to the medial cerebellar peduncles. The
4th ventricle and medial to the dorsal nucleus of the 10th cranial vestibulocochlear nucleus is composed of the cochlear (arrows) part
nerve. The nucleus produces a bulge into the floor of the 4th ven- and the acoustic part (arrowheads). (B) at the medium level we can
tricle termed the hypoglossal eminence (Figs. 7 and 10).9 It receives see the 6th and the 7th nerve nuclei. The 6th is located medial and
cortico nuclear fibers of both hemispheres. However, the cells re- dorsal and the 7th is more anterior and lateral. The axons of the 7th
sponsible for supplying the genioglossus muscle only receive cor- nucleus travel backward and pass around the 6th cranial nerve; this
tico nuclear fibers from the opposite cerebral hemisphere. produces the facial colliculus (an eminence on the floor of the 4th
Axons pass ventrally through the medullary tegmentum be- ventricle). (C) Cranial segment through the trigeminal nuclei. The
sensory nucleus is larger than the motor and is located as well as the
tween the median longitudinal fascicle, medial lemniscus, acces-
7th cranial nerve nucleus in the autonomic space (visceral sensory
sory olivary nucleus, and pyramidal tract medially, and the olive and visceral motor). ML, medial lemniscus; TL, trigeminal lemnis-
laterally. It exits, as multiple rootlets, in the preolivary sulcus, cus; LL, lateral lemniscus; ST, spinothalamic tract; mlf, medial lon-
between the olive and the pyramid (Fig. 9). At the end the nerve gitudinal fasciculus; SpTr, spinptrigeminal tract; RF, reticular for-
sends branches to the muscles of the tongue.10 mation. (Color version of figure is available online.)
Anatomy of the brainstem 205

inantly the medial lemniscus. Two dark dots just anterior to


the 4th ventricle represent medial longitudinal fasciculi. The
dark structure just lateral to the 4th ventricle corresponds to
the superior cerebellar peduncle. The inferior extension of
the wings of the ambient cisterns can produce 2 high-inten-
sity areas on either side of the 4th ventricle and superior
cerebellar peduncles (Fig. 12).5-7 There are 4 cranial nerve
nuclei located at the pons: 5th, 6th, 7th, and 8th. The only
pure motor is the 6th, which are located at the midline. The
5th and 7th are mixed motor and sensitive and are located
more laterally (Table 1). The 8th nerve nucleus is the pure
sensory and is the more posterior and lateral (Fig. 9).

Trigeminal Nerve Nuclei (5th)


The trigeminal nerve is the largest cranial nerve and is made
Figure 18 Axial T2-weighted balanced gradient echo image showing up of 2 components: motor and sensory. It is the great sen-
the course of the trigeminal nerve crossing the prepontine cistern
sory nerve of the head and face and provides motor innerva-
(arrow). It is the thickest cranial nerve and, in its course, it reaches
Meckel’s cave (MC), where its ganglion is located.
tion to the muscles of mastication (Figs. 9 and 17C). The
motor component arises from the motor nucleus, which is
located in the upper part of the pons, close to its dorsal
hemispheres (Figs. 1 and 16). The proper pons is the part surface, and along the line of the lateral margin of the rhom-
between the trigeminal nerves, which emerge on each side. It boid fossa. The nucleus of the sensory component has 3
has a convex anterior surface that shows many transverse subnuclei: (1) the main sensory nucleus, (2) the mesence-
fibers that converge on each side to form the middle cerebel- phalic nucleus, and (3) the spinal nucleus.
lar peduncles (Figs. 12 and 13). In the middle of its anterior The main sensory nucleus lies in the lateral pontine teg-
surface there is a shallow groove, the basilar groove, where mentum, lateral to the motor nucleus and anterolateral to the
lodges the basilar artery (Figs. 13 and 14). 4th ventricle at the level of the root entry zone. The sensa-
Between the pons and the medulla oblongata there is other tions of touch and pressure are conveyed by nerve fibers that
groove where emerge, from medial to lateral, the roots of terminate in the main sensory nucleus. The main sensory
abducent, facial and vestibulochochear nerves (6th, 7th, and nucleus extends cranially continuing with the mesencephalic
8th cranial nerves, respectively; Figs. 14 and 15). The poste- nucleus and caudally forming the nucleus of the spinal tract
rior surface of the pons is out of sight as a result of the (Figs. 9 and 17).
cerebellum and form the superior half of the floor of the 4th The mesencephalic nucleus is composed of a column of
ventricle (Figs. 7 and 16). unipolar nerve cells situated anterolateral to the 4th ventricle
Axial PD-weighted images of the pons demonstrate dark ascending along the lateral margin of the periaqueductal gray
corticospinal and corticobulbar tracts. The horizontal dark matter. It extends to the level of the inferior colliculus and is
band posterior to the corticospinal tracts represents predom- responsible for conveying propioceptive information from

Figure 19 TSE T2-weighted axial high-resolution image at the level of the pons. The cisternal portions of the 6th (open
arrow), 7th (arrow), and 8th (arrowhead). At the IAC the 2 portions of the vestibulocochlear nerve may be seen: the
posterior nerve runs toward the semicircular channels and the anterior 1 toward the cochlea. Both portions fuse at the
CPA cistern to form the bundle of the vestibulocochlear nerve. Note the bulging of the genu of the facial nerve on
the floor of the 4th ventricle forming the facial colliculi (asterisk).
206 M.Á. Fernández-Gil et al

The proper trigeminal nerve arises as separate small motor


and large sensory roots from the ventrolateral surface of the
pons, an area called the “root entry zone” (Figs. 12 and 14).
The roots travel anteriorly through the prepontine cistern
along the medial aspect of the petrous apex and enter a dural
compartment (trigeminal or Meckel’s cave), where it expands
to form the trigeminal ganglion (Figs. 17C and 18).
The trigeminal ganglion divides in its anterior border into
its major divisions: ophthalmic (V1), mandibular (V2), and
maxillary (V3). The motor branch joins the V3 division and
exits the skull base through the foramen ovale to innervate
the muscles of mastication, mylohyoid, tensor tympani, ten-
sor veli palatini, and anterior belly of the digastric muscles.

Abducent Nerve Nucleus (6th)


The abducent nucleus is situated in the pontine tegmentum,
Figure 20 Macroscopic anterior view of the midbrain tinged with
just ventral to the floor of the 4th ventricle (Figs. 9 and 17B).
Mulligan’s method. Asterisk, interpeduncular fossa; PC, pes cerebri;
P, pons; mt, mammilary tubercle. (Color version of figure is avail-
The landmark to identify its location on cross-sectional im-
able online.) ages is the facial colliculus, which is a small protuberance in
the floor of the 4th ventricle produced by the axons of the
facial nerve when loop around the 6th nerve nucleus to turn,
the masticator muscles, the hard palate, teeth, periodontium, afterward, anteriorly (Fig. 7). Motor fibers of the abducent
and temporomandibular joint (Fig. 9). nerve run ventrally through the pontine tegmentum to
The spinal nucleus extends inferiorly through the whole emerge from its ventral surface in the groove between the
length of the medulla oblongata and into the upper part of the lower border of the pons and the medullary pyramids (Figs.
spinal cord as far as the second cervical segment. It continues 9, 17B, and 19). The abducent nerve is entirely motor and
inferiorly with the dorsal gray matter of the dorsal horn of the supplies the lateral rectus muscle and therefore, is responsi-
cord. The sensations of pain and temperature are conveyed ble for turning the eyeball laterally.
by nerve-fibers that form the spinotrigeminal pathway and
end up in the spinal trigeminal nucleus (Figs. 9 and 10).11 Facial Nerve Nucleus (7th)
The motor nucleus of the trigeminal nerve receives cortico The facial nerve nucleus comprises both motor and sensory
nuclear fibers from both cerebral hemispheres. It also re- components as well as a parasympathetic motor component.
ceives fibers from the reticular formation, the red nucleus, The main motor nucleus lies deep in the reticular formation
the tectum and the medial longitudinal fasciculus. In addi- at the ventrolateral aspect of the tegmentum of the caudal
tion, it receives fibers from the mesencephalic nucleus, pons. It is located in proximity to the spinal nucleus of the
thereby forming a monosynaptic reflex arc. The cells of the trigeminal nerve, and near the median eminence of the floor
motor nucleus give rise to the axons that form the motor root. of the 4th ventricle.6 The fibers course upward to the 6th

Figure 21 (A, B) 3T Axial high-resolution PD-weighted images through the midbrain. It is recognized by the “mouse
ears.” White matter tracts appear as dark structures: Corticospinal tracts (arrowheads), medial lemniscus (open arrow),
and medial longitudinal fasciculus (arrow). The white fibers of the superior cerebellar peduncles (scp) have decussated
and run superiorly to reach the red nucleus (asterisk).
Anatomy of the brainstem 207

central gyri. The output from these neurons is carried via the
corticobulbar tracts, which roam along the genu of the inter-
nal capsule. This component is the supranuclear contribu-
tion to the facial nerve function. Keeping this in mind, we
must consider that there is a difference in the transmission of
neural orders to the upper face and lower face: the cortico
nuclear fibers that supply the muscles of the upper part of the
face partially decussate and consequently, originate in both
cerebral hemispheres, whereas the fibers that supply the
muscles of the lower part of the face decussate completely,

Figure 22 Axial IR T1-weighted images through the midbrain.


(A) superior colliculi level; (B) inferior colliculi level. This sequence
clearly separates the gray and white substance. It depicts the basis
(brighter) located anteriorly and the tegmentum (darker) located
posteriorly. Grey substance appears as dark gray colored structures
as the periaqueductal grey matter (arrow) and the substantia nigra
(arrowhead). The basis composed by the pyramidal pathway is in-
deed white (open arrow). The medial lemniscus, spinothalamic
tract, and trigeminal lemniscus join and run upward together. At
axial images they show a boomerang shape (curved arrow) and serve
as a limit between the basis and the tegmentum. Interpeduncular
fossa (asterisk).

nerve nucleus, curve laterally around it, forming the facial


colliculus, and then, they move ventral and slightly lateral,
between the motor nucleus of the facial nerve and spinal
nucleus and tract of the trigeminal nucleus (5th) to exit the
brainstem at the cerebellopontine angle (CPA). This turn Figure 23 Schematic drawings though the midbrain. (A) superior colliculi
around the 6th nerve nucleus is occasionally referred to as the level and (B) inferior colliculi level. BA, basilar artery; Py, pyramidal tracts;
internal genu of the facial nerve (Figs. 9 and 17B).9 RN, red nucleus; mlf, medial longitudinal fasciculus; ML, medial lemnis-
The motor nucleus of the CN7 is in charge of the voluntary cus; LL, lateral lemniscus; 3rd, oculomotor nerve nucleus; E-W, Edinger–
control of facial muscles. Voluntary facial movement begins Westphal nucleus; 4th, trochlear nucleus; RF, reticular formation. (Color
at the level of the motor cortex, in the precentral and post- version of figure is available online.)
208 M.Á. Fernández-Gil et al

Figure 24 (A) Coronal fluid attenuation inversion recovery and (B) coronal SE T2-weighted sequences show the
rhomboid shape of the 4th ventricle. In that plane we can see the superior (arrows), medial (asterisk), and inferior
(arrowhead) cerebellar peduncles, which constitute the lateral boundaries.

giving innervation from the opposite cerebral hemisphere. The vestibular nerve is the nerve of equilibrium. It arises
The knowledge of this pathway help in the clinical examina- from bipolar cells in the vestibular ganglion, ganglion of
tion and location of a facial lesion because a supranuclear Scarpa, which is situated in the upper part of the outer end of
facial palsy would have a corresponding contralateral central the internal auditory meatus. The cochlear nerve is the nerve
lesion, and would likely spare the upper face.12 However, of hearing and conducts nerve impulses concerned with
another involuntary pathway exists; it is separated and con- sound from the organ of Corti in the cochlea. Both vestibular
trols mimetic or emotional changes in facial expression. This and cochlear nerve leave the internal auditory canal at the
other pathway forms part of the reticular formation. porus acusticus and link to form the vestibulocochlear nerve
The sensory nucleus is the upper part of the nucleus of the bundle (8th), which course the CPA cistern with the facial
tractus solitarius (solitarius nucleus or gustatory nucleus) nerve (7th; Figs. 15 and 19).13,14
and lies poserolateral and close to motor nucleus of the 7th. It The vestibulocochlear nerve enters the anterolateral brain-
is the end point for axons originating in the geniculate gan- stem at the pontomedullary junction. Vestibular nerve-fibers
glion of the petrous bone. This axons form the sensory root, terminate in the vestibular nuclear complex (Fig. 17A). The
also called nervus intermedius (pars intermedii of Wrisberg). vestibular complex consists of 4 nuclei (lateral, superior, me-
The parasympathetic nuclei are the superior salivatory and dial, and inferior) and is located along the lateral floor of the
lacrimal nuclei, which lie posterolateral to the main motor 4th ventricle in the lower pons (Figs. 7 and 9). Axons from
nucleus, just slightly superior to the gustatory nucleus, at the these nuclei go along 3 roads: (1) to cerebellum via the ves-
inferior pons. They give rise to the motor parasympathetic tibulocerebellar tract to aid in coordination, (2) descend un-
fibers supplying the submandibular, sublingual and lacrimal
glands (Fig. 9).
The sensory root is composed of both a sensory compo-
nent and a parasympathetic motor component. The end
function of the sensory root is (1) to provide taste function to
the anterior two-thirds of the tongue, (2) to allow sensory
control of lacrimation, and (3) to control the stapedial reflex.
Associated with this functionality is the control of the output
of the sublingual and submandibular glands.
The 2 roots of the facial nerve (motor and sensory) emerge
at the lower border of the pons in the recess between the olive
and the inferior peduncle. The motor part is more medial and
the sensory part is lateral to the former. Immediately poste-
rior to the lateral side of the sensory part is the 8th nerve
(Figs. 15 and 19).
Vestibulocochlear Nerve Nuclei
The vestibulocochlear nerve consists of 2 distinct parts, the
vestibular nerve and the cochlear nerve. They are concerned Figure 25 Coronal cut of a brain specimen tinged with Mulligan’s
with the transmission of afferent information from the inter- method. Some brainstem structures may be identified: cn, caudate
nal ear to the CNS related to balance and hearing. Both are nucleus; lv, lateral ventricle; ic, internal capsule; th, thalamus; bg,
sensory in function and course together until they reach their basal ganglia; sn, substantia nigra; ec, external capsule. (Color ver-
respective nuclei in the brainstem. sion of figure is available online.)
Anatomy of the brainstem 209

crossed to the spinal cord forming the vestibulospinal tract, menturn. From this both stations they run cranially form-
and (3) to join the nucleus of the cranial nerves III, IV, VI via ing the lateral lemniscus. Most neurons from the dorsal
the medial longitudinal fasciculus providing conjugate gaze cochlear nucleus tend to extend directly to the contralat-
and conjugate eye reflexes.15 eral lateral lemniscus and only a few auditory fibers ascend
The fibers of the cochlear nerve enters the brainstem at in the ipsilateral lateral lemniscus (Fig. 32).
the lower border of the pons on the lateral side of the
emerging facial nerve and are separated from it by the
vestibular nerve. When enters the pons the nerve bifur-
The Midbrain
cates to synapse with the secondary neurons which are The midbrain connects the pons to the diencephalon. It com-
situated in the dorsal cochlear and ventral cochlear nu- municates with the cerebellum via the superior cerebellar
cleus, located on the surface of the inferior cerebellar pe- peduncles (Figs. 1 and 20). On the anterior surface of the
duncle, lateral to the restiform body. The dorsal cochlear midbrain there is a deep depression in the midline, the inter-
nucleus process high-frequency sound input (outer fibers peduncular fossa, which is bounded on either side by the
from the cochlear base). The ventral nucleus senses low- crus cerebri (Fig. 20). Many small blood vessels perforate the
frequency sound (Fig. 17A).16 floor of the interpeduncular fossa, and this region is termed
Some axons of the ventral cochlear nucleus synapse the posterior perforated substance (Fig. 15). The part situ-
with the ipsilateral superior olivary nucleus and others ated posterior to the cerebral aqueduct is the tectum and on
cross the midline to form the trapezoid body, the major its surface there are 4 colliculi (corpora quadrigemina; Figs. 9
acoustic pathway decussation, located in the pontine teg- and 16). These are rounded eminences that are divided into

Figure 26 (A) Axial IR T1-weighted image through the midbrain-diencephalon. The posterior margin of the thalami (PV,
pulvinar) hang over the superior colliculi (Sc). (B) Axial IR T1-weighted image through the diencephalon. The 3rd
ventricle and the structures, which surround it, form the diencephalon. The thalami are at both sides of the 3rd
ventricle (arrowheads) and the internal capsule separates (arrows) them from the basal ganglia (Bg). (C) Axial DP-
weighted and (D) axial T2-weighted images show the corticospinal tracts that run downward through the posterior
limb of the internal capsule (arrow). On T2-weighted images the corticospinal pathway is seen as 2 high intensity dots
positioned at the posterior limb of the internal capsule (thick arrow).
210 M.Á. Fernández-Gil et al

the superior colliculus, ventral to the cerebral aqueduct and


dorsal to the medial longitudinal fasciculus (MLF) and red
nucleus. The accessory parasympathetic nucleus (Edinger–
Westphal nucleus) is situated immediately dorsal to the main
motor nucleus.17

Figure 27 Reticular formation distribution along the brainstem. It is con-


fined to the tegmentum and the cranial nerve nuclei are immersed into it.

superior and inferior pairs by a vertical and transverse


groove. The superior colliculi are centers for visual reflexes
and are connected, via the superior brachium, to the lateral
geniculate body and the optic tract. The inferior colliculi are
lower auditory centers and are connected to the medial genic-
ulate body by the inferior brachium.
In the midline below the inferior colliculi emerge the
trochlear or 4th cranial nerve.5-7 Axial sections of the mid-
brain are recognized by the “mouse ears” of the cerebral pe-
duncles anterolaterally and by the cerebral aqueduct poste-
riorly (Fig. 22). The thin boomerang shaped structures
correlate predominantly with the medial lemniscus but also
with the spinothalamic tract and a part of the trigeminal
leminiscus. The lateral lemniscus has extended to the inferior
colliculus. Dark dots in front of the cerebral aqueduct repre-
sent medial longitudinal fasciculus (Figs. 21, 23, 34, and 35).
Through the upper part of the mesencephalon, the cere-
Figure 28 Scheme of the motor pathway. The motor pathway of the pyra-
bral peduncles again appear dark. Red nuclei, which lay at
midal system are the axons of the upper motor neurons which extend
the midbrain, show low intensity in both DP and T2- downward from these upper motor neurons and coalesce to form the
weighted images as well as in fluid attenuation inversion corona radiata (thick arrow). They descend passing through the posterior
recovery images as the result of its iron content (Fig. 22). Like limb of the internal capsule, where the fibers are organized so that those
in lower level of the mesencephalon, the comma-shaped dark closest to the genu are concerned with upper body, whereas those situated
region lateral to red nucleus is composed by the medial lem- more posteriorly are concerned with the lower extremity (arrow). The tract
niscus, spinothalamic tract and trigeminal lemniscus (Fig. continues through the middle three-fifths of the basis pedunculi of the
23). The dark line to the red nucleus represents the medial midbrain, where the fibers concerned with the upper body are situated
longitudinal fasciculus. medially while those concerned to the lower body are placed laterally (open
arrow). On entering the pons, the tract is broken into many bundles by the
Oculomotor Nerve Nuclei (3rd) transverse pontocerebellar fibers (short arrows). While passing through the
midbrain, pons and medulla oblongata some of the axons (corticomesen-
The oculomotor nerve (3rd) is entirely motor in function. It cephalic, corticopontine and corticobulbar tracts) cross the midline to ter-
has 2 motor nuclei: (1) the main motor nucleus and (2) the minate at the motor cranial nuclei of the contralateral side (dotted arrows).
accessory parasympathetic nucleus, which form together the In the medulla oblongata, the bundles become grouped together along the
oculomotor complex. The main oculomotor nucleus lies in anterior border to form a swelling known as the pyramid (curved arrow).
the periaqueductal gray matter in the midbrain at the level of (Color version of figure is available online.)
Anatomy of the brainstem 211

From these nuclei, the fibers pass forward through the sinus between the oculomotor nerve cranially and the oph-
tegmentum, the MLF, the red nucleus, and the medial part of thalmic division of the trigeminal nerve (V1) caudally.
the substantia nigra, and emerge from the oculomotor sulcus
on the medial side of the cerebral peduncle to enter the in-
terpeduncular cistern (Figs. 9 and 23A). The 3rd nerve (with The 4th Ventricle
accompanying parasympathetic fibers) passes ventrally be- The 4th ventricle, or cavity of the hindbrain or rhomben-
tween the posterior cerebral and superior cerebellar arteries, cephalon, is situated in front of the cerebellum and behind
inferior to the posterior communicating artery and medial to the pons and upper half of the medulla oblongata It is lined
the free edge of the tentorium to enter the roof of the cavern- by ciliated epithelium, and is continuous below with the
ous sinus.9 The oculomotor nerve innervates all the extraoc- central canal of the medulla oblongata. Above, it communi-
ular muscles except the lateral rectus and the superior cates, through the cerebral aqueduct, with the cavity of the
oblique muscle and provides parasympathetic innervation to 3rd ventricle (Figs. 1 and 16). It possesses a roof or dorsal
the constrictor pupillae and ciliary muscles via the ciliary wall, a floor or ventral wall, and lateral boundaries.
ganglion.
Roof or Dorsal Wall
Trochlear Nerve Nucleus (4th) The upper portion of the roof is formed by the superior
The trochlear nerve is the slimmest of the cranial nerves and cerebellar peduncle and the superior medullary velum. The
the only one to leave the posterior surface of the brainstem. It superior peduncle emerge from the central white substance
is entirely motor and supplies the superior oblique muscle of of the cerebellum, pass upward and forward, forming, at first,
the eyeball allowing turning the eye downward and laterally. the lateral boundaries of the upper part of the cavity. On
The trochlear nucleus is situated in the dorsal midbrain, cau- approaching the inferior colliculi, they converge, and their
dal to the oculomotor complex and dorsal to the MLF, im- medial portions overlap the cavity and form part of its roof.
mersed in the anterior part of the gray matter that surrounds The superior medullary velum, which is a thin white matter
the cerebral aqueduct of the midbrain. The landmark to iden- lamina, fills the angular interval between both superior pe-
tify its location is the level of the inferior colliculus (Figs. 9 duncles (Fig. 9).
and 23B). The lower portion of the roof is formed by the inferior
From its origin it runs downward and backward through medullary velum. This is continued downward and for-
the tegmentum, around the aqueduct, and decussate, with ward from the central white substance of the cerebellum
the nerve of the opposite side, in the superior medullary and ends inferiorly in a thin, concave, somewhat ragged
velum emerging from the surface of the velum, immediately margin. Immediately inferior to the inferior medullary ve-
behind the inferior colliculus. lum, in the midline, there is a thin triangular shaped area
In the subarachnoid space, the nerve turns anteriorly with no nervous matter called Obex, which forms, at the
through the Ambiens cistern and runs between the tentorium same time, the close-up of the posterior median sulcus of
and the midbrain. It passes around the cerebral peduncle the medulla (Fig. 10).
parallel to the 3rd cranial nerve and, just like it, courses In the roof of the 4th ventricle there are 3 openings, ie, 1
between the superior cerebellar artery and posterior cerebral medial and 2 laterals: the medial aperture (foramen Ma-
artery. The nerve enters the lateral dural wall of the cavernous gendi), is situated immediately above the inferior angle of the

Figure 29 (A) Coronal fast spin echo T2-weighted and (B) coronal TSE T1-weighted after gadolinium injection. Both
images clearly demonstrate the corticospinal tracts passing through the internal capsule (arrowheads), pes cerebri in the
midbrain (arrows) and anterior portion of the pons (open arrow).
212 M.Á. Fernández-Gil et al

ventricle (above the Obex); and the lateral apertures (foram- boidal shape and its boundaries (superior medullary velum
ina of Luschka) are found at the extremities of the lateral and cerebellar peduncles). We can also identify in axial
recesses. By means of these 3 openings the ventricle commu- planes the facial colliculi what indicates the level of the 6th
nicates with the subarachnoid cavity owing the cerebrospinal and 7th cranial nerve nuclei (Figs. 19 and 24).
fluid to circulate.
The non-nervous part of the roof is formed by the epithe-
lial lining of the ventricle, which covers the deep surface of
The Diencephalon
the inferior medullary velum and extends on to the floor of The diencephalon, consists of the 3rd ventricle and the structures
the ventricular cavity; it is covered and strengthened by a which bound it. It extends posteriorly to the point where the 3rd
portion of the pia mater, which is named the Tela choroidea
of the 4th ventricle. The Tela choroidea has 2 highly vascular
infections, the choroid plexuses.

Floor or Ventral Wall


The floor of the 4th ventricle is hidden from view by the
cerebellum and is named, from its shape, the rhomboid fossa
(Figs. 7 and 24). The back surface of the pons and the supe-
rior half of the dorsal surface of the medulla oblongata form it
(Fig. 7). A thin layer of gray substance, continuous with that
of the medulla spinalis, covers it; superficial to this is a thin
lamina of neuroglia, which constitutes the ependyma of the
ventricle and supports a layer of ciliated epithelium.
The rhomboid fossa is divided longitudinally into 2 sym-
metric halves by a median sulcus. On either side of this sulcus
is an elevation, the medial eminence, bounded laterally by
other sulcus, the sulcus limitans or sulcus of Monro (Figs. 7
and 9). In the superior part of the rhomboid fossa corre-
sponding with its lateral limit there is a bluish-gray area, the
locus ceruleus, which owes its color to an underlying patch of
deeply pigmented nerve cells, termed the substantia ferrug-
inea (Fig. 14).
The inferior end of the medial eminence is slightly ex-
panded to form the colliculus facialis, which is produce by
the roots of the facial nerve winding around the nucleus of
the abducent nerve (Fig. 7).
At the level of the colliculus facialis the sulcus limitans
widens into a flattened depression, the superior fovea. Lateral
to the fovea is a triangular elevation named the area acustica,
where underlie the vestibular nuclei. These nuclei are astride
the pons and medulla (Figs. 7 and 9).
Winding around the inferior peduncle and crossing the
area acoustical and the medial eminence are several delicate
transverse white strands, the striae medullares, which form a
portion of the cochlear division of the acoustic nerve, which
disappear into the median sulcus. The striae medullare may
serve as a transverse line that divides the floor of the 4th
ventricle in 2 areas: the superior (corresponding to the pons)
and the inferior (corresponding to the medulla oblongata)
(Fig. 7). In the inferior part of the fossa, the medial eminence
assumes the form of a triangular area, the trigonum hypo-
glossi (Fig. 7).
Between the trigonum hypoglossi and the lower part of the
Figure 30 (A) Axial PD-weighted image and (B) axial T2-weighted
area acoustical is a triangular dark field, the ala cinerea, image show the low intensity of the compact white matter as the
which corresponds to the sensory nucleus of the vagus and corpus callosum (arrowheads) and internal capsule (arrows). Note
glossopharyngeal nerves.5-7 the normal high intensity rounded-shaped of the pyramidal path-
The 4th ventricle is hardly visualized on magnetic reso- way running through the posterior limb of internal capsule, which
nance imaging and, hence, we have to resign to see its rhom- disappears at DP-weighted image (open arrows).
Anatomy of the brainstem 213

ventricle becomes continuous with the cerebral aqueduct and an- the thalami is narrow and rounded and forms de posterior
teriorly as far as the interventricular foramina (Figs. 1 and 16). The boundary of the interventricular foramen (Fig. 16).
diencephalon can be divided into 4 major parts: the epithala- The posterior extremity is expanded, directed backward
mus, the hypothalamus, the subthalamus, and the thalamus. and lateral ward, and overlaps the superior colliculus. Medi-
The epithalamus comprises the trigonum habenulæ, the ally, it presents an angular prominence, the pulvinar, which
pineal body, and the posterior commissure. The hypothala- is continued laterally into an oval swelling, the lateral genic-
mus includes the subthalamic tegmental region and the ulate body. Beneath the pulvinar, but separated from it by the
structures forming the greater part of the floor of the 3rd superior brachium, is a second oval swelling, the medial
ventricle. It is the only area exposed to the surface and con- geniculate body (Figs. 9 and 28), which forms part of the
sists of the corpora mammillaria, tuber cinereum, infundib- auditory pathway. Lateral is the lateral geniculate body,
ulum, hypophysis, and optic chiasma (Figs. 15, 16, and 26). which forms part of the visual pathway.
The subthalamic tegmental region is the upward continua- The thalamus is divided into several parts, which contain
tion of the tegmentum; it lies on the ventrolateral aspect of the groups of different thalamic nuclei. The anterior part of the
thalamus. The red nucleus and the substantia nigra are pro- thalamus contains the anterior thalamic nuclei, which receive
longed into its lower part; in front it is continuous with the the mammilothalamic tract from the mammillary nuclei. The
substantia innominata of Meynert. Medially, it is the gray sub- function of the anterior thalamic nuclei is closely associated
stance of the floor of the 3rd ventricle, the corpus subthalami- with of that of the limbic system and is concerned with emo-
cum (nucleus of luys), which is a brownish mass, lenticular in tional tone and the mechanisms of recent memory.
shape on transverse section, and situated on the dorsal aspect of The medial part of the thalamus contains the large dorso-
the fibers of the base of the cerebral peduncle (Fig. 25). medial nucleus and several smaller nuclei. The dorsomedial
The thalamus is the largest part of the diencephalon and nucleus has 2 connections with the whole prefrontal cortex of
serves as a relay station to all the main sensory tracts. It may the frontal lobe of the cerebral hemisphere. It also has similar
be regarded as a large ganglionic mass in which the ascending connections with the hypothalamic nuclei. The medial part
tracts of the tegmentum and a considerable proportion of the of the thalamus is responsible for the integration of a large
fibers of the optic tract end, and from the cells of which variety of sensory information, including somatic visceral
numerous fibers (thalamocortical) take origin, and radiate to and olfactory information and the relation of this information
almost every part of the cerebral cortex. to ones emotional feelings and subjective states.
It consists of 2 large ovoid masses, situated one on either The lateral part is subdivided in dorsal and ventral com-
side of the 3d ventricle (Figs. 25 and 26). The anterior end of ponents. The dorsal stage includes the lateral dorsal nucleus,

Figure 31 Axial PD-weighted high-resolution images performed with a 3-T equipment. The main running tracts, which
cross the brainstem, may be identified. The pyramidal pathway is marked with arrowheads; The lemniscus (ie, medial
lemniscus, spinotrigeminal lemniscus, and spinothalamic pathway) is shown by the arrows. The medial longitudinal
fasciculus is only seen in 5 images as 2 dark dots just in front of the 4th ventricle and located at both side of the midline
(open arrows). (Color version of figure is available online.)
214 M.Á. Fernández-Gil et al

the lateral posterior nucleus, and the pulvinar. The ventral


level contains the ventral anterior and the ventral lateral
nuclei, (which influences the activities of the motor cortex
because of its connection to the substantia nigra, reticular
formation, and corpus striatum), as well as the ventral pos-
teromedial (which receive the ascending trigeminal and gus-
tatory pathways) and ventral posterolateral nuclei, which re-
ceive the medial and spinal lemnisci.
For the purposes of this article, it is important to remember
several thalamic nuclei: the medial geniculate bodies, the
lateral geniculate bodies, the ventroposterolateral nuclei
(sensory relay stations from the body), and the ventropos-
teromedial nuclei (sensory relay stations from the face). The
thalamus also receives numerous fibers (corticothalamic)
from the cells of the cerebral cortex.
The superior surface of the thalamus is free, slightly con-
vex, and covered by a layer of white substance, termed the
stratum zonale. It is separated laterally from the caudate nu-
cleus by a white band, the stria terminalis and on its lateral
surface by another layer, the external medullary lamina,
which is in contact with a thick band of white substance, the
internal capsule that separates the thalamus from the lenti-
form nucleus of the corpus striatum.
The inferior surface rests upon and is continuous with the
upward prolongation of the tegmentum (subthalamic teg-
mental region; Fig. 25). The medial surface constitutes the
upper part of the lateral wall of the 3rd ventricle and is con-
nected to the corresponding surface of the opposite thalamus
by a flattened gray band, the Massa intermedia (middle or
gray commissure), which may be occasionally absent. The
hypothalamic sulcus separates the medial part of the thala-
mus from the medial part of hypothalamus (Fig. 16).5-7,18 On
axial magnetic resonance images, the diencephalon is easily
recognized by the presence of the 3rd ventricle, which is
flanked from medial to lateral by the thalami, the internal Figure 32 Spinotrigeminal tract (in green) and trigeminal lemnis-
capsule, and the lenticular nuclei (Figs. 25 and 26).19 cus (in blue). Most of the neuronal bodies containing somes-
The internal capsule can be divided into an anterior limb thetic information of the head are settled in the trigeminal gan-
and a posterior limb, which are connected by a knee or genu. glia (Gasser). From here, axons that form the central projection
Lesions of the anterior limb are clinically silent. Lesions of the of these neuronal bodies (first neuron of the chain) synapse on
genu (corticobulbar tract) produce contralateral facial paral- the neurons of the sensory nucleus, the spinal nucleus and the
mesencephalic nucleus of the trigeminal nerve. Pain and temper-
ysis. Lesions of the anterior portion of the posterior limb
ature enter the brainstem via the trigeminal nerve (CN5), facial
(corticospinal tract) produce a contralateral hemiparesis of (CN7), and vagus (CN10) nerves to synapse on the second neu-
the body. The posterior part of the posterior limb conducts ron located at the spinal nucleus. When these fibers penetrate
sensory information (its anterior part conducts general sen- they descend parallel to the nucleus forming the spinal tract of
sory information and its posterior part conducts special sen- the trigeminal nerve (open arrows) While they move down, they
sory information, such as hearing and vision.20 enter the nucleus to synapse. From here, the axons of the second
neuron project to the contralateral side crossing the midline and
ascend to constitute the trigeminal lemniscus. Touch and pro-
The Reticular Formation prioception enter the brainstem at the same time and by the same
The reticular formation (RF) is found in the tegmentum and way. The fibers that convey these sensations do not descend but
consists of an intricate mixture of neuronal cell bodies and instead synapse on the neurons of the sensory nucleus of the
trigeminal nerve (arrow). From here they traverse to the con-
fascicles of axons running in small bundles, which are ori-
tralateral side to join the trigeminal lemniscus ascending through
ented in many different directions. Thus, as its name suggest, the rest of the pons and midbrain to synapse with the 3rd neuron
it is like a network extending from the spinal cord through placed in the ventral posterolateral nucleus of the thalamus.
the medulla, the pons, the midbrain, the subthalamus, hypo- Similarly to the somatosensory pathway from the body, the axon
thalamus and the thalamus. of the 3rd neuron project to the somesthetic area in the postcen-
The RF receives impulses from most parts of the central tral gyrus of the cerebral cortex. (Color version of figure is avail-
nervous system: spinoreticular tracts, spinothalamic tracts, able online.)
Anatomy of the brainstem 215

medial lemniscus, vestibular, acoustic and visual pathways, terminate primarily within the medial reticular zone. A 4th
cerebelloreticular pathways. Efferent projections extend region must be added: the intermediate zone, found only
down to the brainstem and spinal cord through reticulobul- in the medulla, is situated between the medial and lateral
bar and reticulospinal tracts. Additional pathways extend to columns. It is involved in autonomic regulation of respi-
the corpus striatum, the cerebellum, the red nucleus, the ration, heart rate, and blood pressure.22,23
substantia nigra, the tectum and the nuclei of thalamus, sub- On the basis of this organization we may say that the RF is “a
thalamus and hypothalamus.21 mess well arranged,” and its functions are accurately distributed.
The RF has a complex organization, but it is not random Thus, caudal to the midpontine level, RF is involved in the
and may be divided, for teaching purposes, into 3 longi- coordination of fine movements, autonomic regulation of respi-
tudinal columns with ill-defined boundaries. This areas ration, heart rate, and blood pressure. Rostral to the midpontine
are distinguished on the basis of the their cytoarchitecture level the RF is involved in arousal, consciousness, and the wak-
but they are also functionally distinct. In addition a 4th ing state. The RF also regulates the preservation of mental estate,
zone is defined in the medulla: (1) the raphe with cells homeostasis-neurovegetative reflexes, postural reflexes, and
lying along the midline. They mediate posture, movement, taste.2 Because of its redundancy, a bilateral lesion or complete
pain, autonomic function and arousal; (2) the paramedi- section of the tegmentum is required to cause dysfunction of the
an-medial zone proyects to the cerebellum and to hypo- RF (Figs. 17, 23, and 27).
thalamus and thalamus (via the central tegmental tract)
and to the spinal cord via the reticulospinal tract. All of
these projections are associated with motor functions; (3)
The Running Tracts
the lateral zone receives multiple sensory inputs from the Several ascending and descending tracts travel the entire brain-
spinal cord, cranial nerves and cerebellum. It is thought to stem. The most important descending one is the motor path-
be an afferent association area and has short axons that way, which is composed by the corticospinal and corticobulbar
tracts. It controls the voluntary skeletal muscle activity of the
body and face. There are several important ascending tracts that

Figure 33 Spinothalamic tract (in green) and medial lemniscus (in


blue). Pain and temperature sensations enter the spinal cord by the
central axons of the neurons located at the root ganglion. These fibers of
the first neuron terminate by synapsing with cells of the posterior gray
column. The axons of the second neuron cross obliquely to the oppo-
site side and ascend in the contralateral white column as the lateral and
anterior spinothalamic tract. As the spinothalamic tracts ascend
through the medulla oblongata, they lie near the lateral surface and join
to form the spinothalamic tract or spinal lemniscus. It continues to
ascend through the lateral part of the tegmentum of the pons and
midbrain, always lateral to the medial lemniscus. Many of the fibers end
by synapsing with the 3rd neuron of the chain located in the ventral
posterolateral nucleus of the thalamus. From here, the axons of the 3rd
neuron pass through the posterior limb of the internal capsule and the
corona Radiata to reach the somesthetic area in the postcentral gyrus of
the cerebral cortex. The role of the cerebral cortex is to interpret the
quality of the sensory information at the level of consciousness of the
contralateral half of the body. Equally to the spinothalamic tract,
the sensations of touch and proprioception enter the spinal cord by the
central axons of the spinal root ganglion. The fibers pass directly to the
posterior white column and travel upward as the fasciculus gracilis and
fasciculus cuneatus, which ascend and terminate by synapsing on the
second neuron located in the ipsilateral nucleus gracilis and nucleus
cuneatus on the medulla oblongata. The axons of the second neuron,
called the internal arcuate fibers, sweep anteromedially and cross the
midline and decussate ascending as a single compact bundle, the me-
dial lemniscus, which as its name suggest, it runs through the brainstem
in a medial position, until reaching the ventral posterolateral nucleus of
the thalamus where synapse on the 3rd neuron. The axon of the 3rd
neuron pass through the posterior limb of the internal capsule and
corona Radiata to reach the somesthetic area in the postcentral gyrus of
the cerebral cortex where vibratory sense and the positions of the dif-
ferent parts of the body can be consciously recognized. (Color version
of figure is available online.)
216 M.Á. Fernández-Gil et al

The Motor Pathway (Efferent)


The motor pathway, also called the pyramidal system, arise
from the precentral gyrus of the cerebral cortex and contain
the upper motor neurons. These upper motor neurons are
arranged in a structured manner: neurons that control move-
ments of the face and mouth are located near the Sylvian or
lateral fissure, and neurons that control the muscles of the
thighs and legs are located near the medial longitudinal fis-
sure and within the central sulcus. This somatotropic ar-
rangement allows us to represent the parts of the body in this
area of the cortex, the so-called homunculus, which is a dis-
torted picture of the body with the various parts having a size
proportional to the area of the cerebral cortex devoted to their

Figure 34 Lateral lemniscus. Auditory impulses enter the brain-


stem through the 8th cranial nerve or cochlear nerve (8th). The
neuronal bodies of the axons, which capture the impulses from
the cochlea, are located at the ventral and dorsal cochlear nuclei.
Some of the fibers run to synapse on neurons located at the
ipsilateral and contralateral nuclei of the superior olivary com-
plex that also take part in the auditory pathway. From here, fibers
ascend (some are decussated). Several synapse on the neurons
placed at the nuclei of the lateral lemnisci and others continue to
ascend. Cranially, to the nuclei of the lateral lemnisci, all join to
constitute the lateral lemniscus, which further ascend to synapse
on the neurons of the inferior colliculi (the inferior colliculi take
part in the auditory pathway, whereas the superior colliculi make Figure 35 Medial longitudinal fasciculus. The movements of the eye-
it in the visual pathway). Easy to remember: the “eyes are higher balls are carried out by the extraocular muscles, which are con-
than the ears.” From the inferior colliculi, axons arrive at the trolled by neurons placed in different nuclei (3rd, 4th, 6th). The
medial geniculate body where they synapse to project to the medial longitudinal fasciculus consists of paired group of ascending
primary auditory area of the cerebral cortex at the temporal and descending axons, which connects the cranial nerve nuclei 3rd,
lobes. (Color version of figure is available online.) 4th, and 6th together, as well as the gaze centers and information
about head movement (from the 8th cranial nerve). Many of the
fibers arise in the lateral vestibular nucleus and divide into ascend-
ing and descending branches, which send terminal branches and
collaterals to the motor nuclei of the cranial and spinal nerves. In the
transport all the sensations from the periphery to the cerebral medulla oblongata and pons, the medial longitudinal fasciculus,
cortex. They comprise the somatosensory pathways. The main runs close to the middle line, near the floor of the 4th ventricle; in
sensitive tracts are the spinothalamic tract, medial lemniscus, the midbrain it is situated on the ventral aspect of the cerebral
spinotrigeminal tract, trigeminal lemniscus, lateral lemniscus, aqueduct, below the nuclei of the oculomotor and trochlear nerves.
and medial longitudinal fasciculus. (Color version of figure is available online.)
Anatomy of the brainstem 217

control: the homunculus has a very large face and mouth pyramids. The crossed fibers enter the lateral white column of
because there are many upper motor that innervate these the spinal cord to form the lateral corticospinal tract. The few
parts of the body. This initial somatotropic organization in fibers, which do not cross the midline, descend in the anterior
the cortex continues along the whole route of the pathway white column of the spinal cord as the anterior corticospinal
(Fig. 28). tract. The fibers of these both tracts synapse with the second
When one passes through the midbrain, pons and me- neuron located at the anterior gray column of the medulla.
dulla oblongata some of the axons (corticomesencephalic, The clinical appearance of a lesion affecting the corticospi-
corticopontine, and corticobulbar tracts) cross the midline nal tract depends on the level of injury. Usually it manifests as
to terminate at the motor cranial nuclei of the contralateral an upper motor neuron defect, which results in contralateral
side. Lesions involving the corticobulbar tracts before they hemiparesis accompanied by spastic paralysis, hyper-re-
have decussated within the brainstem will result in a con- flexia, and Babinski sign. If all cranial nerves are affected,
tralateral loss of motor function in the face; lesions involv- then, the lesion is above the midbrain. If the lesion is located
ing the motor nuclei themselves or exiting motor compo- in the brainstem, it will cause lower motor neuron distur-
nents of the appropriate cranial nerves will result in an bance on the level where the lesion is and long tract signs
ipsilateral hemiparesis. The greater part of these descend- below the lesion (ipsilateral cranial nerve dysfunction and
ing fibers, however, extends downward to the spinal cord contralateral hemiparesis).19,24-27
to form the corticospinal tract. The corticospinal tract may be identified on magnetic res-
At the junction of the medulla oblongata and the spinal cord, onance imaging as 2 well-defined bilaterally symmetric
most of the fibers cross the midline at the decussation of the round or oval foci increased intensity in the posterior internal

Figure 36 Axial 3-T high-resolution images through the pontomedullary junction, pons, upper pons, and midbrain
showing the medial longitudinal fasciculus (arrows). It may be seen as 2 paired dark spots just near the floor of the 4th
ventricle. It extends from the upper medulla to the midbrain connecting the 8th, 6th, 4th, and 3rd cranial nerve nuclei
with the reticular formation.
218 M.Á. Fernández-Gil et al

Table 2 The Main Somatosensory Pathways


First
Sensation Neuron Second Neuron Third Neuron Pathway
Pain and temperature Posterior root Substantia gelatinosa Ventral posterolateral nucleus Lateral spinothalamic and
ganglion of thalamus spinal lemniscus
Light touch and Posterior root Substantia gelatinosa Ventral posterolateral nucleus Anterior spinothalic and
pressure ganglion of thalamus spinal lemniscus
Discriminative touch, Posterior root Nuclei gracilis and Ventral posterolateral nucleus Medial lemniscus and
vibratory sense, ganglion cuneatus of thalamus trigeminal lemniscus
conscious muscle
joint sense

capsule. These normal focal hyperintense areas are 3 to 4 mm (of the spines) and temperature of the face go through the
in diameter, have homogeneous internal architecture and are “spinotrigeminal tract.”
without any mass effect. The high intensity indicates that the The other main sensations, proprioception and touch, are
corticospinal fibers, at this level, have lower axonal and my- transported by the lemniscus. The medial lemniscus carries
elin densities compared with the surrounding posterior cap- the information from the body and, of course, trigeminal
sular fibers, which are tightly packed myelinated with small- lemniscus transfers the sensations from the face.19,25,26,29
caliber axons (Figs. 26, 29, and 30). Because of the complexity of the brainstem organization
Normal hyperintense foci are observed usually in the ros- and its many connections, we are going to simplify the anat-
tral midbrain on T2-weighted images, but they could not be omy into those which the radiologist must know: the spino-
followed below the caudal midbrain because the corticospi- thalamic, spinotrigeminal, medial lemniscus, trigeminal lem-
nal tracts separate into several bundles in the pons and the niscus, longitudinal medial fasciculus, and lateral lemniscus.
density of the large axons decreases.28 At the pons these fibers These 6 tracts are represented and explained in detail in Figs.
may be identified in axial PD-weighted images as transversal 32-36. The main somatosensory pathways are summarized
foci of dark gray color (Figs. 30 and 31). in the Table 2.

The Somatosensory Where Is the Lesion?


Pathway (Afferent) To follow are several tips about the location of lesions:

For teaching purposes, we are going to consider that all the 1. Cerebellar and basal ganglia lesions result in motor
different ascending pathways, which are in charge of conduit problems.
the sensations, consist of 3 neurons. The first neuron has its 2. Basal ganglia disorders are specifically characterized by
cell body in the posterior root ganglion of the spinal nerve. A meaningless, unintentional, and unexpected move-
peripheral process connects with a sensory receptor ending, ments.
whereas a central process enters the spinal cord through the 3. The presence of cranial nerve involvement signifies that
posterior root to synapses on the second neuron. The second the lesion lies above the level of the foramen magnum.
neuron gives rise to an axon that cross the midline (de- 4. The presence of a radicular pain along an extremity
cussates) and ascends to a higher level of the CNS, where it suggests that the lesion lies below the level of the fora-
synapse with the 3rd neuron. The 3rd neuron is usually men magnum.
located in the thalamus and give rise to projection fibers that 5. The presence of a cranial nerve defect on 1 side and
pass to a sensory region of the cerebral cortex. On this basis, defects of motor or sensory modalities in the contralat-
the tracts passing through the brainstem are the decussated eral extremities confirms that the lesion lies at the level
axons of the second neuron. of the brainstem and not in the cerebral cortex or inter-
Many of the neurons in the ascending pathways branch nal capsule.
into the RF, which, in turn, activates the cerebral cortex, 6. Lesions of the cerebral cortex and internal capsule both
maintaining wakefulness. The main sensations that are con- result in contralateral sensory and motor deficits.29
veyed by the coming tracts are pain and temperature on the
one hand and proprioception and touch on the other. References
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