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Figure 32 Immediately after neural tube closure, the
embryonic central nervous system (middle) consists
of the developing spinal cord and three vesicles that
will comprise the adult brain. Almost all of the
neural tube is formed by primary neurulation
(stippled area marked 1°). Secondary neurulation
(2°) forms the spinal cord from midsacral levels to
the conus medullaris, the caudal tip of the spinal
cord. The rostral end of the neural tube, the lamina
terminalis (*), ends up at a site that is buried deep
INTERPRET THIS DIAGRAM: within the adult brain (left From a posterior view
and right from a side view). The junction between
the developing spinal cord and brain is the
spinomedullary junction. Note that drawings are
schematic and not to scale.
THE THREE BULGES OR VESCICLES OF THE NEURONAL The three vesicles destined to become the brain are,
TUBE AT THE END OF THE 4TH GESTATIONAL WEEK ARE from front to back:
THE: • Prosencephalon, which will develop into the
forebrain
• Mesencephalon, which will develop into the
midbrain
• Rhombencephalon, which will develop into the
hindbrain
WHAT HAPPENS AT THE FIFTH WEEK OF GESTATION? At about the fifth week of gestation, the
WHERE DID THE LAMINAE TERMINALIS GO? prosencephalon divides into the telencephalon and
the diencephalon (Fig. 33B). Almost immediately
after forming, the single telencephalic vesicle
invaginates at the midline to become two
telencephalic hemispheres. Since the points of
attachment for the hemispheres are off the midline,
the lamina terminalis, representing the front end
of the neural tube, is now located at the front end of
the diencephalon (Fig. 3 3C).
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Figure 33 In the three vesicle stage of neural
development, the embryonic brain (A) consists of
the prosencephalon (P), mesencephalon (M), and
rhombencephalon (R). B: The prosencephalon
divides into a caudally located diencephalon (D) and
rostrally situated telencephalon (T), which together
with the mesencephalon and rhombencephalon,
comprise the fourvesicle stage. C: The single
telencephalic vesicle invaginates along the midline
to form the laterally displaced left (Tl) and right (Tr)
cerebral hemispheres. The rostral end of the neural
tube, the lamina terminalis (* in A and C), ends up
deep within the adult brain. From the diencephalon
emerges the optic vesicle (ov), which will develop
into the optic nerve and retina. The derivatives of
the telencephalon, diencephalon, mesencephalon,
and rhombencephalon are listed.
IN THE ADULT, WHAT ENCOMPRISES THE In the adult, the telencephalon includes both
TELENCEPHALONIC STRUCTURES? hemispheres of cerebral cortex, the core
components of the basal ganglia, and the amygdala.
IN THE ADULT BRAIN WHAT ARE THE DIENCEPHALIC The diencephalon
STRUCTURES? contains the thalamus and hypothalamus.
EXPLAIN THE MESENCEPHALIC AND The mesencephalon develops into the adult
RHOMBENCEPHALIC STRUCTURES IN THE ADULT midbrain, and the rhombencephalon develops into
BRAIN. the adult pons, medulla, and cerebellum (Fig. 33C).
The anterior third or so of the rhombencephalon
gives rise to the pons and cerebellum, whereas the
posterior portion develops into the medulla.
IN SUMMARY, BY THE END OF THE FIFTH WEEK, THE • Telencephalon ≈ cerebral cortex, basal ganglia,
EMBRYONIC HUMAN BRAIN CONTAINS amygdala
FOUR VESICLES. FROM ROSTRAL TO CAUDAL: • Diencephalon ≈ thalamus, hypothalamus + retina
and optic nerves
• Mesencephalon = midbrain
• Rhombencephalon = hindbrain = pons, medulla,
and cerebellum
The spinal cord and the four regions of the brain
comprise the five divisions of the adult CNS.
THE TERRITORY ALLOTTED TO THE DORSAL
TELENCEPHALON IS GREATLY EXPANDED IN THE HUMAN
WHAT DISRUPTS THE ORDERLY CELLULAR THE NEED FOR A MASSIVE CORTICAL VOLUME
ARRANGEMENT IN THE HUMAN UP TO THIS POINT IN TELENCEPHALIC EXPANSION.
TERM OF NEURONAL DEVELOPMENT
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Figure 35 In the human, the cerebral cortex is
greatly expanded. A–B: Topdown (A) and side (B)
views illustrate the expansion of embryonic dorsal
telencephalic tissue from its point of attachment to
the diencephalon at the lamina terminalis (red star).
The dorsal telencephalon extends in all directions
until it
covers the diencephalon and mesencephalon
completely and also reaches the midline.
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Figure 37 The embryonic lumen of the neural tube
becomes the adult ventricular system. A: A
midsagittal
view of the embryonic brain shows the lumen of the
neural tube lined by proliferating cells.
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D–H: Transverse cartoons (left) through the
embryonic spinal cord (D), hindbrain (E), midbrain
(F), diencephalon (G, H), and telencephalon (H) are
arranged next to photomicrographs of brain slices
from the same regions in the adult (right). The
central lumen (red stipple; red asterisk in D) at every
level of the neuraxis is marked. Note that the brain
slices in D–G are stained for myelin so that white
matter is dark and gray matter is light; the slice in H
is unstained. Although the lumen adopts different
shapes, it is located on the midline in the spinal
cord, hindbrain, midbrain, and diencephalon.
However, the
telencephalic hemispheres are located lateral to the
diencephalon, and, consequently, the lumen divides
and
courses laterally into each telencephalic
hemisphere. The narrow foramina of Monro (fM)
connect the lumen of the diencephalon to the
lumen of each telencephalic hemisphere.
HOW IS CSF FORMED? CSF is essentially blood that has been filtered
through choroid epithelium, a specialized type of
cuboidal epithelium located at specific spots in the
ventricular system. Blood is delivered to the choroid
epithelium from a rich supply of capillaries in the
pia. Choroid epithelium, capillaries, and pia are
tightly invested tissues that comprise the choroid
plexus (Fig. 38A).
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Figure 38 A: The choroid plexus is comprised of
outpouchings of vascularized pia lined with choroid
epithelium. Choroid epithelial cells have small
protuberances called microvilli on their apical
surface. Due to hydrostatic pressure, blood from the
capillaries of the pia traverses the basal lamina and
is filtered by the choroid epithelium to produce
cerebrospinal fluid (CSF). The outside of the brain is
located on the pial side of choroid plexus, and the
ventricle is located on the choroidal side
B: Choroid plexus develops in specific
areas of the developing neural tube (red lines)
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WHERE IN THE BRAIN WILL YOU NEVER FIND CHOROID Since choroid plexus must border on the periphery,
PLEXUS? it is not present in either the central canal or the
cerebral aqueduct, both of which are tubes
surrounded on all sides by parenchyma.
WHAT DOES THE VELUM INTERPOSITUM HAVE TO DO For example, the third ventricle is surrounded by
WITH THE LOCATION OF CHOROID PLEXUS? parenchyma everywhere except along the roof of
the diencephalon. Recall that the velum
interpositum sits in the space above the
diencephalon and below the telencephalic
hemispheres. Capillaries arising from blood vessels
in the velum interpositum dive down into the roof of
the third ventricle, are invested in choroidal
epithelium, and form the third ventricle’s
complement of choroid plexus. Thus, choroid plexus
is present in the roof of the
third ventricle but not along its sides or floor.
Figure 310 A: The cerebellum develops from the
rhombic lip, located in the anterior portion of the
rhombencephalic vesicle. B: The cerebellum grows
caudally back from the rhombic lip, eventually coveri
most of the medulla as well as all of the pons in the
adult (D). C: The cerebellum is attached to the
hindbrain only through the cerebellar peduncles.
Photographs in C–D kindly provided by Peter Pytel,
MD, University of Chicago.
Figure 311 The corpus callosum and internal capsule are two major tracts that alter the layout of the brain.
The corpus callosum is the major commissural tract that carries axons linking the two cerebral hemispheres and
the internal capsule forms a physical join between each cerebral hemisphere and the thalamus. A: A
coronal section through the brain shows both major tracts. B: A topdown cartoon shows the corpus
callosum joining the two hemispheres. The internal capsule runs between the lateral edge of the
diencephalon and the medial edge of each cerebral hemisphere. From the topdown perspective, the axons
of the internal capsule run in and out of the page and are therefore depicted as transversely cut. C: A coronal
cartoon through the brain (at the level indicated by the dotted arrow) shows that the velum interpositum (vi)
is situated between the corpus callosum and the roof of the diencephalon. The internal capsule runs through
the subcortical portion of the telencephalon (stippled area) and outlines the lateral edge of the
diencephalon. D–E: Horizontal (D) and sagittal (E) sections through the brain show the internal capsule
running between the diencephalon (D) and the subcortical structures (white asterisks) of the telencephalon
(T). Abbreviations: corpus callosum, cc; internal capsule, ic; lateral ventricle, lv. Photographs in A, D, and
E kindly provided by Peter Pytel, MD, University of Chicago.
WHAT CONNECTS THE TELENCEPHALON TO THE In fact, beyond the small attachments surrounding
DIENCEPHALON? the foramina of Monro, the internal capsules are
theonly connection between the telencephalon and
diencephalon. Thus, cutting the internal capsule on
either side allows the telencephalic cap to be
removed from the underlying diencephalon and
brainstem.
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THE CORPUS CALLOSUM CONNECTS THE LEFT AND RIGHT
CEREBRAL HEMISPHERES
WHICH HEMISPHERE IS OFTEN RESPONSIBLE FOR The left hemisphere interpreter is not
CONFABULATION? bound by truth and, in fact, is often responsible for
confabulation
NAME EXAMPLES OF INTERHEMISPHERIC AND 1. CORPUS CALLOSUM
INTRAHEMISPHERIC COMMUNICATION 2. HIPPOCAMPAL COMMISURE
3. ANTERIOR COMMISURES
4. OPTIC CHIASM
5. POSTERIRO COMMISURE OF THE MIDBRAIN
6. ANTERIOR WHITE COMMISURE OF THE
SPINAL CANAL
DEVELOPMENTAL TERRITORIES CONFER A BASIC
FUNCTIONAL ORGANIZATION TO THE BRAIN AND SPINAL
CORD
Figure 312 In the spinal cord (A), a distinct inflection
point, the sulcus limitans, separates the alar plate,
destined to give rise to neurons that receive input
from primary sensory afferents, from the basal plate
that gives rise to motoneurons, autonomic motor
neurons, and motor interneurons. In the hindbrain
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In each telencephalic hemisphere (D), the thin rind
of the dorsal telencephalon becomes the cerebral
cortex, whereas the lateral and medial ganglionic
eminences (lge, mge) develop into the striatum,
pallidum, and portions of the amygdala.
Figure 313 A developmentally inspired cartoon of the human brain and spinal cord (sc). For illustrative
purposes, the left (Tl) and right (Tr) telencephalic hemispheres are shown unconnected to the diencephalon
(D) and the rhombencephalic cerebellum (Rc) is filleted down the middle and splayed out. The brainstem
consists of the hindbrain and the midbrain (M). The adult derivatives of the rhombencephalon include the
pons (Rp), medulla (Rm), and cerebellum (Rc). B: The sources and modalities of sensory inputs to the spinal
cord, hindbrain, diencephalon, and telencephalon are illustrated on the left. The skeletal muscle targets of
motoneurons in the spinal cord, hindbrain, and midbrain are illustrated on the right. C: The autonomic
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targets of autonomic preganglionic neurons in the spinal cord, hindbrain, and midbrain are illustrated. D: A
nonexhaustive catalog of abstract functions served by the telencephalon is listed on the left. As illustrated
on the right, damage to different levels of the neuraxis impacts an individual’s sense of self to varying
degrees. The chance for improvement following injury also varies with the level of damage.
MAPPING FUNCTIONS ONTO THE SPINAL CORD,
BRAINSTEM, AND FOREBRAIN
SENSORY INPUT TO THE SPINAL CORD: The spinal cord receives all the somatosensory input
from the body and a great portion of the
viscerosensory input from deeper structures. This
input arrives via spinal nerves. Spinal cord or nerve
damage can impair the perception of touch,
vibration, temperature, pain, and the position of the
body from the legs, trunk, arms, neck, and back of
the head.
SENSORY INPUT OF THE BRAINSTEM The brainstem receives a wide variety of sensory
input via cranial nerves:
• Somatosensory input from the face, oral cavity,
and anterior fossa
• Viscerosensory input from viscera above the
hindgut
• Special sensory inputs from the inner ear, which
supports hearing and the vestibular sense
• Special sensory input from the oral and pharyngeal
cavities that support taste
All of the sensory input received by the brainstem
enters into the hindbrain
SENSORY INPUT OF THE MIDBRAIN The midbrain does not receive any sensory input
SENSORY INPUTS THAT ENTER THE FOREBRAIN VISION AND OLFACTION Optical information from
the retina arrives in the diencephalon, and
information about odorants enters the telencephalic
olfactory bulbs
WHERE MUST SENSORY INPUTS GO, OR REACH TO Thus, sensory inputs must reach the neocortex for
ACHIEVE PERCEPTION? perception to occur.
SPINE’S RESPONSIBILITY FOR MOVEMENT? Movements of the body, from the shoulders down,
depend on spinal motoneurons and spinal nerves.
BRAINSTEM MOVEMENT CONTROL? Movements controlled by brainstem, motoneurons
and cranial nerves include:
• Shrugging and shaking the head in a “No” gesture
• Tongue movements important in chewing and
speaking
• Movements of the upper airway musculature
important to swallowing, speaking, coughing, and
the like
• Facial expression
• Movements of the eyes to enable gaze control
• Jaw movements used in chewing
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HOW DOES CNS DAMAGE AFFECT MOTOR Thus, damage to the CNS impacts movements that
MOVEMENT? depend on motoneurons leaving the CNS from
points caudal to the site of damage. Yet, all volitional
actions are initiated by neocortex. Thus, voluntary
motor control of any part of the head or body may
be affected by forebrain damage.
HOW DOES THE FOREBRAIN AFFECT HOMEOSTASIS? The forebrain, specifically the diencephalic
hypothalamus, controls the pituitary gland from
which a wide variety of hormones is secreted.
Pituitary hormones play critical roles in growth, fluid
and electrolyte balance, thermoregulation,
metabolism, mating and reproduction, and arousal
evoked by stress. The hypothalamus is also
important in setting the circadian rhythm through its
influence on both the pituitary and pineal glands
HOW DO THE SPINAL CORD AND BRAINSTEM Brainstem and spinal cord contributions to
INFLUENCE HOMESOTASIS? autonomic control are many and varied but fall into
two categories: parasympathetic and sympathetic.
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