BASIC SCIENCE
Neuroanatomy: an
overview
Vishy Mahadevan
Abstract
For descriptive purposes, the nervous system is divided topographi-
cally into two parts: the central nervous system and the peripheral ner-
vous system. From a functional perspective, however, this division is
somewhat artificial, given that the two ‘systems’ are intimately con-
nected, and normally function in an integrated and coordinated
manner. The central nervous system comprises the brain and spinal
cord, while the peripheral nervous system is made up of the cranial
and peripheral nerves. This article presents a broad and general over-
view of the structure and organization of the nervous system empha-
sizing several important functional features.
Keywords Autonomic nervous system; bloodebrain barrier; brain-
stem; cerebrospinal fluid (CSF); dermatome; diencephalon; functional
localization; meninges; spinal cord segment; sub-arachnoid space
It is conventional to describe the nervous system as being made
up of two principal parts: the central nervous system (CNS) and
the peripheral nervous system (PNS).
The CNS comprises the brain and spinal cord. These are
located in the cranial cavity and the vertebral (spinal) canal,
respectively, and are continuous with each other at the foramen
magnum in the skull base where the medulla oblongata of the
brainstem leaves the cranial cavity to continue as the spinal cord
(spinal medulla).
The PNS comprises 12 pairs of cranial nerves, 31 pairs of
spinal nerves, the ganglia associated with the cranial and spinal
nerves, the right and left ganglionated sympathetic chains and
the pelvic parasympathetic nerves. Within the spinal and cranial
nerves, a distinction is to be made between afferent nerve fibres
(which carry sensory information from peripheral receptors to
the central nervous system) and efferent nerve fibres (which
carry motor information from the central nervous system to the
periphery). Additionally, a distinction is made within the spinal
and cranial nerves, between visceral nerve fibres (afferent and
efferent fibres that supply the visceral organs of the cardiovas-
cular, respiratory, digestive, urinary and reproductive systems)
and somatic nerve fibres that comprise sensory and motor nerve
fibres destined for body wall (somatic) structures such as skin,
skeletal muscle, tendons, joints, etc.
The visceral nerve fibres, both afferent and efferent, form a
large part of the autonomic nervous system. The autonomic
nervous system is classified into sympathetic and para-
sympathetic divisions, each comprising afferent and efferent fi-
bres. The sympathetic and parasympathetic fibres emerge from
Vishy Mahadevan MBBS PhD FRCS (Ed & Eng) is the Barbers’ Company
Professor of Anatomy at the Royal College of Surgeons of England,
London, UK. Conflicts of interest: none.
SURGERY 36:11
the brainstem and spinal cord within specific cranial nerves
(parasympathetic fibres) or within spinal nerve roots (sympa-
thetic and parasympathetic fibres) and are distributed to visceral
smooth muscle and glandular elements. In a sense this division
into central and peripheral nervous systems is somewhat
contrived and arbitrary, as the two ‘systems’ are physically
connected, and normally function in a well-coordinated and in-
tegrated manner.
Cellular architecture of the nervous system
Notwithstanding its structural intricacy and functional
complexity, the nervous system per se consists of just two
principal cell types: neurons and neuroglia.
The neuron is the structural and functional unit of the nervous
system.
Neurons are responsible for all the special functions that are
attributed to the nervous system, such as sensory perception of
every description, neural regulation of glandular secretion,
execution and control of motor activity in muscle, cognitive
function, etc. Mature neurons have a very limited capacity for
repair and practically no capacity for replication and regeneration.
It has been estimated that the human nervous system contains
far in excess of 100 billion neurons, the majority being situated
within the brain. Furthermore each neuron may have 1000 to
10,000 synaptic connections with other neurons, accounting for
as many as 100 trillion synapses in the brain. Neurons are
specialized to transmit impulses between the nervous system and
all parts of the body, and between different regions within the
nervous system.
Although highly variable in size and shape, all neurons
possess a cell body, and extending from the cell body, an axonal
process and a variable number of dendritic processes or den-
drites. Generally speaking, dendrites carry nerve impulses to-
ward the cell body, while axons conduct impulses away from the
cell body to adjacent neurons or to muscle cells or glandular cells.
The cells of the neuroglia serve to support, protect and
nurture the neurons and maintain homoeostasis of the extracel-
lular fluid that surrounds the neurons. Neuroglial cells are
generally smaller than neurons and are up to 50 times more
numerous than the latter. Unlike neurons, neuroglial cells are
capable of replicating themselves. They invariably proliferate by
multiplication to fill in gaps caused by loss of neurons. Four
types of neuroglial cells are recognized in the CNS: astrocytes,
oligodendrocytes, microglial cells and ependymal cells. Astro-
cytes are identifiable by their star-shaped cell bodies which
possess many narrow, elongated processes. Astrocytes play an
important role in the metabolism of neurotransmitters and help
maintain an optimal level of ionic potassium for generation of
nerve impulses. They also help form the bloodebrain barrier.
Oligodendrocytes are smaller than astrocytes and possess fewer
processes. Their processes along with those of astrocytes form a
supporting network for neurons. A very important function of
oligodendrocytes is the production of lipoprotein wrappings
termed myelin sheaths for the peripheral processes of neurons in
the central nervous system. Microglial cells are phagocytic cells
derived from monocytes. They protect the nervous system by
engulfing and destroying microbial invaders and also remove any
debris resulting from the death of cells. Ependymal cells form the
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 BASIC SCIENCE
inner lining of the ventricles of the brain and central canal of the
spinal cord. They are an admixture of squamous cells and
columnar cells, some of which may be ciliated.
The PNS contains two types of neuroglial cells: neuro-
lemmocytes (Schwann cells) that produce myelin sheaths around
the axons of neurons in the peripheral nervous system and sat-
ellite cells that support neuronal cell bodies within the ganglia of
peripheral nerves.
A cardinal feature of the central nervous system is the great
degree of functional localization that exists within it. This term
denotes the allocation of specific functions to well-circumscribed,
discrete regions in the brain and spinal cord.
The brain and spinal cord are surrounded by three, concen-
trically arranged layers of protective membranes: dura mater (the
outermost layer), arachnoid mater (the middle layer) and pia
mater (the innermost layer). Between the arachnoid mater and
pia mater is the subarachnoid space in which is contained cere-
brospinal fluid (CSF).
Besides serving as a ‘cushion’ for the brain and spinal cord
within their respective bony enclosures, CSF plays an important
role in the exchange of nutrients and waste metabolites between
the blood and neurons of the brain and spinal cord.
The salient features of the production and circulation of ce-
rebrospinal fluid, as well as the pattern of blood supply to the
brain and spinal cord are addressed in this article.
Central nervous system
Brain (Figures 1 and 2)
On a functional and morphological basis, the brain may be
pictured as being made up of four major divisions:
 cerebral hemispheres (right and left)
Precentral gyrus Central sulcus Postcentral gyrus Supramarginal gyrus
Superior
frontal gyrus
Middle
frontal gyrus
Opercular
Parts of inferior Triangular
frontal gyrus Orbital
Lateral
sulcus
Orbital gyri
Superior middle
poral gyri
and inferior temporal
Pons
Flocculus
Pyramid
Figure 1 Lateral aspect of left half of brain showing left cerebral and cerebellar hemispheres and brainstem
SURGERY 36:11
 diencephalon (comprising, mainly, the thalamus and
hypothalamus)
 cerebellum
 brainstem (comprising, from below upwards, the medulla
oblongata, pons and midbrain).
By another convention, based on embryological development,
the brain may be divided into forebrain (comprising the cerebral
hemispheres and diencephalon), midbrain and hindbrain (made
up of pons, medulla oblongata and cerebellum).
Cerebral hemispheres: each cerebral hemisphere comprises the
cerebral cortex and the basal nuclei (basal ganglia), and various
afferent and efferent neural fibre tracts. Within each cerebral
hemisphere lies a cavity, the lateral ventricle, containing CSF.
The cortex of the cerebral hemisphere is divided geographi-
cally into four major lobes e frontal, parietal, temporal and oc-
cipital, each containing discrete areas which subserve specific
functions (functional localization) (Figure 1).
The frontal lobe harbours the primary motor area which oc-
cupies a large part of the precentral gyrus. It receives afferents from
the cerebellum and is concerned with the initiation and perfor-
mance of voluntary movements. The frontal lobe of the dominant
hemisphere (the left cerebral hemisphere in the majority of in-
dividuals) also includes Broca’s speech area (the motor speech area).
The parietal lobe features the primary somatosensory cortex
on the postcentral gyrus. This area receives afferent fibres from
the thalamus and is concerned with the recognition of all forms
of somatic sensation.
The temporal lobe features the auditory cortex whose afferent
fibres are derived from the medial geniculate body of the thal-
amus. It is concerned with the perception of auditory stimuli. The
Superior
parietal
lobule
Angular
gyrus
Parieto-
occipital
sulcus
Inferior
parietal
lobule
Occipital
lobe
Lunate
sulcus
Cerebellum
Olive
Medulla
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 BASIC SCIENCE
Corpus
callosum
Thalamus
Cerebral
aqueduct
Midbrain
Fourth
ventricle
Cerebellum
Pons
Medulla
Figure 2 Medial aspect of left half of sagittally divided brain
cortical region just above and behind this area on the dominant
hemisphere is called Wernicke’s area and is of considerable
importance in the sensory aspects of language comprehension.
The occipital lobe lies behind the parietal and temporal lobes
(Figures 1 and 2) and features on its medial surface the primary
visual cortex. This area receives afferent fibres from the lateral
geniculate body of the ipsilateral half of the thalamus.
The basal nuclei (also known as basal ganglia) are compact
masses of grey matter (neuronal cell bodies) that are situated
deep in the substance of each cerebral hemisphere and comprise
the corpus striatum (composed of the caudate nucleus, the pu-
tamen and the globus pallidus), the amygdaloid body and the
claustrum. Together with the cerebellum, the basal nuclei are
involved in the coordination and control of movement.
Diencephalon: the diencephalon comprises principally the thal-
amus and hypothalamus, which are coextensive (Figure 2). A
median, vertically orientated, cleft-like space between the right
and left halves of the diencephalon is termed the 3rd ventricle. In
addition to the thalamus and hypothalamus, the diencephalon
includes two small but functionally important regions: the epi-
thalamus and ventral thalamus. The epithalamus is the dorsal
portion of the diencephalon and contains the pineal body. The
ventral thalamus (also known as subthalamus) contains the
subthalamic nucleus, which is one of the basal ganglia.
The thalamus is an ovoid mass of grey matter which forms the
upper part of the lateral wall of the 3rd ventricle. The thalamus is
the principal sensory relay station which projects impulses from
the main sensory pathways onto the cerebral cortex. It does this
via a number of thalamic radiations in the internal capsule.
The hypothalamus forms part of the lateral wall and floor of
the 3rd ventricle. The hypothalamus performs numerous regu-
latory functions. As the so called ‘head ganglion’ of the auto-
nomic nervous system, it is concerned with the regulation of
autonomic (both sympathetic and parasympathetic) activity.
The hypothalamus plays an important role in the control of
endocrine secretions by the formation of releasing factors or
release-inhibiting factors which act directly on the subjacent pitu-
itary gland. In addition to its major influence on the autonomic
nervous system and on pituitary function, the hypothalamus also
SURGERY 36:11
Cerebral
hemisphere
Choroid plexus
of third
ventricle
Interventricular
foramen
Septum
pellucidum
Hypothalamus
plays a significant role in temperature regulation, water and elec-
trolyte balance, regulation of appetite and sleepewake patterns.
Cerebellum: the cerebellum is the largest part of the hindbrain
and occupies most of the posterior cranial fossa (see Figures 1, 2
and 6). It is made up of the right and left cerebellar hemispheres
and a median vermis.
The cerebellar hemispheres are connected to the brainstem by
way of three pairs of cerebellar peduncles: superior, middle and
inferior cerebellar peduncles. Ventrally, the cerebellum is related
to the 4th ventricle and to the medulla and pons (Figure 2).
Functions of the cerebellum e the principal function of the
cerebellum is to regulate and maintain balance, and to co-
ordinate timing and precision of body movements. The cere-
bellum has multiple connections with the cerebral cortex, retic-
ular formation in the brainstem, thalamus and vestibular nuclei.
Through these intricate connections, the cerebellum constantly
monitors proprioceptive sensory input from joints, muscles and
tendons, and accordingly refines and co-ordinates the contrac-
tions of skeletal muscles. However, unlike the cerebral cortex of
the primary motor area, the cerebellum is incapable of initiating
movement, nor is the cerebellum involved in the conscious
perception of somatic or visceral sensations.
Brainstem: extending from just above the tentorial hiatus to
just below the foramen magnum, the brainstem is a stalk-like
structure which is continuous superiorly with the dienceph-
alon, and inferiorly with the spinal cord. From above down-
wards, the brainstem comprises, successively, the midbrain
(mesencephalon), pons and medulla oblongata (Figure 2). The
brainstem and cerebellum are situated in the posterior cranial
fossa.
The brainstem serves three major functions:
 It houses the nuclei of all but 2 of the 12 pairs of cranial
nerves (the exceptions being the cranial nerve pairs I and II
which may both be regarded as peripheral extensions of
the forebrain).
 It acts as a ‘thoroughfare’ for the various ascending and
descending nerve tracts running to and from the cerebral
cortex, and for other tracts that project to the cerebellum.
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 BASIC SCIENCE

 It contains the reticular formation (a fine and diffuse
network of nerve cells and nerve fibres) and the reticular
activating system. The reticular formation spans the entire
length of the brainstem and harbours the ‘vital centres’ e
important reflex centres which regulate respiratory and
cardiovascular function. The reticular formation and
reticular activating system regulate the individual’s level of
awareness and wakefulness. Damage to the reticular for-
mation in the upper part of the brainstem may cause the
patient to be in a state of prolonged coma.
Spinal cord (Figure 3)
The spinal cord (spinal medulla) in the adult is approximately 45
cm long. It is continuous above with the medulla oblongata at the
foramen magnum and ends at the level of the lower border of the
1st lumbar vertebra.
The spinal cord bears a deep longitudinal anterior midline
fissure, and a shallow posterior midline sulcus. On either side, is a
posterolateral sulcus along which the posterior (sensory) nerve
roots of spinal nerves are arranged serially along the length of the
spinal cord. The anterior (motor) nerve roots of the spinal nerves
emerge serially along the anterolateral aspect of the length of the
spinal cord, on either side.
The anterior and posterior nerve roots unite within the
appropriate intervertebral foramen to form a spinal nerve which,
soon after, divides into anterior and posterior rami, each con-
taining both motor and sensory fibres.
Below the termination of the spinal cord, the roots of the
lumbar, sacral and coccygeal nerves continue downwards within
the vertebral canal, as the cauda equina, lying in the subarach-
noid space, as far as the lower sacral level.
A cross section of the spinal cord (Figure 3) shows the central
canal around which is the H-shaped grey matter packed with
nerve cell bodies arranged in a highly orderly manner. Sur-
rounding the grey matter is the white matter. The latter contains
the long ascending and descending tracts.
On either side of the midline the grey matter presents a
somewhat bulbous anterior (or ventral) horn packed with motor
nerve cells, and a slender posterior (or dorsal) horn that extends
almost to the periphery of the section. The posterior horn is
capped by the substantia gelatinosa in which terminate many of
the incoming sensory fibres entering the cord through the pos-
terior nerve roots. In the large anterior horn lie the motor cells
which give rise to the fibres of the anterior roots.
In that part of the spinal cord that corresponds to the thoracic
and upper lumbar segments, and sacral segments, the grey
matter features, additionally, a lateral horn on each side, con-
taining the cells of origin of the sympathetic system in the
thoracic and upper lumbar segments, and the cells of origin of
the pelvic parasympathetics in the sacral segments.
The white matter of the spinal cord has: descending (motor)
tracts of which the most important is the lateral corticospinal
tract (crossed pyramidal motor tract); and ascending sensory
tracts of which the most important are the lateral and anterior
spinothalamic tracts carrying pain and temperature sensations,
and the posterior column fibres that are as yet uncrossed and
carry sensations of fine touch, vibration and proprioception
(position sense).
All sensory fibres running up in the spinal cord eventually
relay in the thalamus before being projected onto the relevant
area of cerebral cortex.
Knowledge of the internal architecture of the spinal cord, and
the location of various fibre tracts is of much value and useful-
ness in the clinical examination and diagnosis of spinal cord
lesions.
Blood supply of the brain and spinal cord (Figure 4)
The brain receives its blood supply from the right and left in-
ternal carotid arteries (each a terminal branch of the corre-
sponding common carotid artery) and from the right and left
vertebral arteries (each a branch of the ipsilateral subclavian
artery). Each vertebral artery courses upwards along its side of

Posterior median sulcus and septum

Gracile tract
S
L Posterior white column
Cuneate tract T
C Posterior nerve root
Dorsolateral tract
I
Posterior Posterior horn
spinocerebellar
tract V Lateral corticospinal tract

Lateral reticulospinal tract

Anterior VII
spinocerebellar X Denticulate ligament
tract S IX Lateral white column
L IX Lateral horn
Anterolateral VIII
tract T Anterior horn
C Lateral vestibulospinal tract

Anterior nerve root

Anterior median fissure Medial reticulospinal tract

Anterior white column
Anterior corticospinal tract

Figure 3 Cross section of spinal cord showing major descending and ascending tracts

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 BASIC SCIENCE
the neck, through the cervical vertebral transverse processes
before entering the posterior cranial fossa through the foramen
magnum. The two vertebral arteries then unite in front of the
brainstem to form the basilar artery. The latter runs upward and
ends by dividing into the right and left posterior cerebral arteries.
Each internal carotid artery enters the cranial cavity through the
temporal bone of its side and eventually divides into the anterior
and middle cerebral arteries. Shortly before its termination the
internal carotid artery gives off the all-important ophthalmic ar-
tery which enters the orbit through the optic foramen (accom-
panying the optic nerve) to supply all the orbital contents. By
means of three narrow communicating arteries, the two internal
carotid arteries and the two posterior cerebral arteries form an
‘arterial ring’ in the middle cranial, termed the circle of Willis
(Figure 4).
The basilar artery and the two vertebral arteries together
supply all of the cerebellum and the brainstem (i.e. the contents
of the posterior cranial fossa) and through the two posterior ce-
rebral arteries, a good deal of the medial surface of the cerebral
hemispheres including the visual cortex on the occipital lobe
(Figure 4). The two internal carotids supply the remainder of the
brain. Knowledge of the vascular territories of individual vessels
aids in the interpretation of clinical signs caused by occlusion of
cerebral vessels.
Anterior cerebral artery
Anterior communicating artery
Infundibulum
Oculomotor nerve
Basilar artery
Abducens nerve
Facial nerve
Anterior inferior cerebellar artery
Vertebral artery
Posterior inferior cerebellar artery
Figure 4 Arterial supply of brain and spinal cord
SURGERY 36:11
Capillaries in the brain and spinal cord (CNS) differ signifi-
cantly from those elsewhere in the body. Thus the endothelial
cells that line the capillaries in the CNS are non-fenestrated and
form tight junctions with their neighbours. The basement
membrane underlying the endothelial cells in brain capillaries
tends to be significantly thicker. The relative impermeability of
brain capillaries is further enhanced by the enveloping foot
processes of astrocytes that surround the brain capillaries. These
unique features of brain capillaries account for the so called
bloodebrain barrier that functions to protect the internal milieu
of the brain by allowing only certain substances in blood plasma
to be transported across the endothelial cells into the extracel-
lular fluid. There are however specific regions within the where a
bloodebrain barrier would prove a hindrance to the function of
that part of the brain. In these regions (exemplified by the pos-
terior pituitary, median eminence of the hypothalamus, pineal
body and area postrema in the floor of the 4th ventricle) the
bloodebrain barrier is absent.
Venous blood from the brain drains into an interconnected
system of intracranial dural venous sinuses (Figure 5) which are
endothelium-lined channels located, in the main, between the
dura and the endocranium (the name given to the periosteum
that lines the inner surface of the cranial cavity). The intracranial
venous sinuses are valveless and eventually drain into the right
Optic nerve
Internal carotid artery
Middle cerebral artery
Choroidal artery
Posterior cerebral artery
Superior cerebellar artery
Labyrinthine artery
Anterior spinal artery
601 Ó 2018 Elsevier Ltd. All rights reserved.

Thalamostriate
and choroidal veins
Anterior
cerebral vein
Superficial middle
cerebral vein
Deep middle
cerebral vein
Cavernous sinus
Superior petrosal sinus
Inferior petrosal sinus
Figure 5 The cerebral venous system showing superficial and deep cerebral veins and their relationship to the network of intracranial dural venous
sinuses
and left internal jugular veins which leave the cranial cavity
through the skull base to run down the neck.
The spinal cord derives its blood supply from the anterior and
posterior spinal arteries which are branches of the vertebral ar-
teries. Blood supply to the lower half of the spinal cord is supple-
mented by the spinal branches of the posterior intercostal arteries
which are, in turn, branches of the descending thoracic aorta.
Floor of the cranial cavity (Figure 6)
The floor of the cranial cavity is the upper surface of the base of
skull. In a dry skeleton it is revealed by removing the cranial
vault (skull cap).
The floor of the cranial cavity presents a terraced arrangement
of three regions (areas). In anterior to posterior sequence, these
are the anterior cranial fossa, middle cranial fossa and posterior
cranial fossa. The anterior fossa is the shallowest and the
smallest, whereas the posterior fossa is the deepest and largest of
the three areas (Figure 6).
Anterior cranial fossa: the central part of the floor of the anterior
cranial fossa is a depressed, perforated plate of bone, the cribri-
form plate of the ethmoid (Figure 6). This forms the highest part
of the roof of the nasal cavity, and is traversed on either side of
the midline by the corresponding olfactory nerve filaments, on
their way from the roof of the nasal cavity to the olfactory bulb in
the anterior cranial fossa. Lateral to the cribriform plate, on
either side, the floor of the anterior cranial fossa is somewhat
raised and forms the roof of the orbit.
The anterior cranial fossa houses, in addition to the right and left
olfactory pathways, the frontal lobes of the cerebral hemispheres
and the anterior cerebral arteries as the latter course beneath the
frontal lobes. The frontal lobes lie just above the orbital roof.
Middle cranial fossa: the middle cranial fossa floor shows a
platform-like elevation in the centre (Figure 6) This is the body of
the sphenoid bone. Its upper surface is slightly concave and
SURGERY 36:11
BASIC SCIENCE
Superior anastomotic vein Superior sagittal sinus
Inferior sagittal
sinus
Internal
cerebral
vein
Great
cerebral
vein
Basal vein
Straight
sinus
Inferior
anastomotic
vein
Transverse sinus
Occipital sinus
Sigmoid sinus
constitutes the pituitary fossa. Situated immediately above the
pituitary fossa is the pituitary gland which is suspended from the
hypothalamus by the pituitary stalk. On either side of the central
elevation, the floor of the middle cranial fossa accommodates the
corresponding temporal lobe of the cerebral hemisphere.
Posterolateral to the pituitary fossa on either side lies the corre-
sponding trigeminal ganglion, and directly lateral to the pituitary
fossa on either side is the cavernous sinus.
Anteriorly, on either side of the midline, the middle cranial
fossa communicates with the ipsilateral orbit by means of two
openings: the larger, wider and laterally located one being the
superior orbital fissure and the smaller opening being the optic
foramen. The latter transmits the optic nerve (ensheathed in the
three meningeal layers) and ophthalmic artery, while the supe-
rior orbital fissure transmits the oculomotor (III), trochlear (IV)
and abducens (VI) nerves, the ophthalmic nerve and the
ophthalmic veins. On each side, the floor of the middle cranial
fossa shows three significant foramina. From front to back these
are the foramen rotundum, foramen ovale and foramen spino-
sum, which transmit respectively the maxillary nerve, mandib-
ular nerve and the middle meningeal artery.
Posterior cranial fossa: the posterior cranial fossa is the deepest
and largest of the three fossae (Figure 6). It houses the entire
cerebellum. Lying anterior to the cerebellum in the median part
of the posterior cranial fossa is the brainstem comprising, from
above downwards, the midbrain, pons and medulla oblongata.
The medulla oblongata leaves the posterior cranial fossa through
a large opening in the centre of the fossa’s floor, the foramen
magnum, to become the spinal cord. The ‘slope’ of bone which
forms the anterior wall of the posterior cranial fossa, and lies in
front of the brainstem, is called the clivus. The cerebellum is
roofed by a large, thick, double-layered sheet of infolded dura
mater termed the tentorium cerebelli. Above the tentorium cer-
ebelli (and therefore outside the posterior cranial fossa) lie the
occipital lobes of the cerebral hemispheres.
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 BASIC SCIENCE
Anterior
cranial
fossa
Middle
cranial
fossa
Posterior
cranial
fossa
Figure 6 Floor of cranial cavity showing anterior, middle and posterior cranial fossae
Three significant openings, on each side, lead away from the
posterior cranial fossa. These are the internal acoustic (auditory)
meatus, the jugular foramen and the hypoglossal canal. The in-
ternal acoustic meatus transmits the facial (VII) and vestibulo-
cochlear (VIII) cranial nerves. The jugular foramen transmits the
glossopharyngeal (IX), vagus (X) and accessory (XI) cranial
nerves, and also the sigmoid and inferior petrosal venous si-
nuses. The hypoglossal canal transmits the hypoglossal (XII)
cranial nerve.
Peripheral nervous system
Cranial nerves
There are 12 pairs of cranial nerves, all of which leave or enter
the cranial cavity through foramina or fissures in its floor or
walls. By convention, the cranial nerves are designated in roman
numerals. The first and second cranial nerve pairs are unlike the
other ten, in that they do not have their ‘origins’ in the brain-
stem. For this reason, the first and second cranial nerves are
generally regarded by neurophysiologists as fibre tracts of the
forebrain, rather than as true nerves.
The numbered cranial nerves (with their corresponding
names) are as follows: I (olfactory), II (optic), III (oculomotor),
IV (trochlear), V (trigeminal), VI (abducens), VII (facial), VIII
(vestibulo-cochlear), IX (glossopharyngeal), X (vagus), XI
(accessory) and XII (hypoglossal).
On a functional basis cranial nerves may be classified into one
or other of three groups:
SURGERY 36:11
Cribriform plate of ethmoid
Optic canal
Superior orbital fissure
Foramen rotundum
Foramen ovale
Foramen lacerum;
sphenopetrosal fissure
Foramen spinosum
Carotid canal,
internal aperture
Internal acoustic meatus
Jugular foramen
Hypoglossal canal
Foramen magnum
 Wholly sensory cranial nerves: I (olfactory nerve), II (optic
nerve), VIII (vestibulo-cochlear nerve)
 Wholly motor cranial nerves: III (oculomotor), IV (troch-
lear), VI (abducens), XI (accessory), XII (hypoglossal)
 Mixed (motor and sensory) cranial nerves: V (trigeminal),
VII (facial), IX (glossopharyngeal), X (vagus).
The design of clinical tests undertaken to test the integrity of
individual cranial nerves is obviously determined by this
classification.
The motor (or efferent) cranial nerves arise from discrete ag-
gregations of neurons in the brainstem, termed nuclei of origin.
The sensory (or afferent) cranial nerves arise from neurons
situated outside the brain. The central processes of these neurons
enter the brain, eventually to join collections of neurons termed
nuclei of termination. Despite this fundamental distinction,
nuclei of origin and nuclei of termination are both commonly
referred to as ‘origins of cranial nerves’.
Of the 12 cranial nerve pairs, all but one are confined to the
head and neck. The exception is the vagus (Xth cranial nerve)
whose distribution extends beyond the head and neck to the
thorax and abdomen.
Spinal nerves, spinal cord segments and dermatomes
The spinal nerves are complementarily paired and symmetrically
arranged structures which originate in the central grey matter of
the spinal cord. There are 31 pairs of spinal nerves in all (8
cervical, 12 thoracic, 5 lumbar, 5 sacral and 1 coccygeal). As has
603 Ó 2018 Elsevier Ltd. All rights reserved.
 BASIC SCIENCE
already been noted, each spinal nerve is formed by the conflu-
ence of a motor (anterior) root with a sensory (posterior) root.
A horizontal section of spinal cord which gives rise to a
complementary pair of spinal nerves is termed a spinal cord
segment (Figure 3). It follows, therefore, that the spinal cord
possesses thirty one segments.
Each spinal nerve (with the notable exception of the first
cervical nerve, which has no cutaneous supply) supplies a spe-
cific area of skin. This cutaneous territory of the spinal nerve is
termed a dermatome.
Autonomic nervous system is so named because, unlike the
somatic nervous system, it is not under voluntary control. The
autonomic nervous system consists of three parts: (i) sympa-
thetic nervous system; (ii) parasympathetic system; and (iii) the
enteric nervous system. The autonomic nervous system contains
both afferent and efferent fibres. These are distributed to the
thoracic, abdominal, pelvic and perineal viscera and to vascular
smooth muscle.
Meningeal coverings of the brain and spinal cord
The brain and spinal cord are surrounded by three layers of
protective membranes, arranged concentrically (Figure 7). The
outermost layer is the dura mater, the middle layer is the
arachnoid mater and the innermost layer is the pia mater.
The pia mater is close applied to the surface of the brain and
spinal cord, and even lines all the major and minor fissures of the
brain.
The arachnoid mater is closely applied to the inner surface of
the dura mater everywhere, and never normally leaves the dura.
In places the dura in the cranial cavity forms fairly strong, large
(double-layered) infoldings. These serve to create compartments
within the cranial cavity, and perform the very important func-
tion of supporting the delicate brain, and protecting the brain
very effectively from the torsional stresses to which it would
otherwise be subjected. The most important of these dural re-
duplications are the falx cerebri and the tentorium cerebelli.
Superior
sagittal sinus
Emissary vein
Arachnoid
granulation
Venous lacuna
Cerebral vein
Falx cerebri
Figure 7 Coronal section through the vertex of the skull showing the meningeal layers and their relationship to the
dural venous sinuses
SURGERY 36:11
Between the arachnoid mater and the underlying pia mater is
the subarachnoid space in which is contained CSF. Between the
arachnoid and the overlying dura is a potential space called the
subdural space. Normally, only a thin film of fluid lies in this
space which enables the arachnoid to glide on the inner surface
of the dura. Abnormally, however, blood may collect in this
potential space resulting in a subdural haematoma.
Within the cranial cavity the dura mater is firmly fused to the
periosteum that lines the inner surface of the bony walls and
floor of the cranial cavity. Within the vertebral canal, however,
the dura is free of the wall of the canal. The space between the
two is called the epidural (or extradural) space. It contains a thin
layer of fat surrounding the dural sheath. Within the fat and
arranged circumferentially around the dural sheath is a delicate
plexus of (valveless) veins called the internal vertebral venous
plexus.
The ventricular system and the cerebrospinal fluid
circulation (Figure 8)
Developmentally, the entire central nervous system is derived
from a simple ectodermal closed but hollow tube. Notwith-
standing the eventual transformation of this simple tube into the
highly complex-looking brain and spinal cord, the original lumen
is retained in every part of the fully developed central nervous
system, and constitutes the ventricular system. It is lined with a
specialized epithelium termed ependyma.
The ventricular system comprises the right and left lateral
ventricles, the 3rd ventricle and the 4th ventricle. The right and
left lateral ventricles (each situated within the corresponding ce-
rebral hemisphere) are by far the largest components of the sys-
tem. Each lateral ventricle opens into the 3rd ventricle through an
opening called the interventricular foramen (of Monro) located in
the corresponding lateral wall of the third ventricle.
From the 3rd ventricle the CSF passes through a narrow
channel in the midbrain, the aqueduct (of Sylvius), into the 4th
ventricle.
Diploic vein
Skull
Dura mater
Subarachnoid space
Pia mater
Arachnoid mater
Cerebral cortex
604 Ó 2018 Elsevier Ltd. All rights reserved.
 BASIC SCIENCE

a Olfactory organs b

Lateral ventricle
Telencephalon
Forebrain
(Prosencephalon) Rostral choroid plexus

Diencephalon

Eye

Midbrain
(Mesencephalon) III ventricle
c Lateral ventricle III ventricle
Cerebral aqueduct Choroid plexus of
Pons, cerebellum lateral ventricle
IV ventricle
(Metencephalon)
Hindbrain
(Rhombencephalon)

Medulla oblongata
Caudal choroid plexus
(Myelencephalon)

Spinal cord

Cerebral aqueduct
Choroid plexus
IV ventricle of IV ventricle

Figure 8 Organization of the ventricular system in the brain

The 4th ventricle lies between the pons/medulla ventrally and
the cerebellum dorsally, separated from the latter by a
membrane-like roof. The CSF escapes from the 4th ventricle into
the subarachnoid space by way of three natural apertures in the
membranous roof of the 4th ventricle, and then flows over the
surface of the brain and spinal cord. The three apertures are the
right and left foramina of Luschka and the midline foramen of
Magendie.
Production and circulation of CSF
CSF is produced mainly in the ventricular system of the brain,
whence it enters the subarachnoid space. The production and
circulation of CSF are constant, dynamic and energy-dependent
processes.
Besides serving as a ‘cushion’ for the brain and spinal cord
within their respective bony enclosures, CSF plays an important
role in the exchange of nutrients and waste metabolites between
the blood and neurons of the brain and spinal cord.
CSF is formed by the secretory activity of the choroid plexuses
(vascular tufts that are invaginated into parts of the ventricular
system) in the lateral, 3rd and 4th ventricles. CSF is produced
constantly at a rate of 0.5 ml/minute. It circulates through the
ventricular system of the brain and enters the subarachnoid
space through three natural openings in the membranous roof of
the 4th ventricle. From the subarachnoid space, CSF is extruded
into the system of intracranial dural venous sinuses (principally
the superior sagittal sinus) by specialized evaginations of
arachnoid, termed arachnoid villi/arachnoid granulations
(Figure 7).
Any obstruction to the flow and circulation of CSF results in a
condition termed hydrocephalus. A
FURTHER READING
Ellis H, Mahadevan V, eds. Clinical anatomy. 13th edn. WILEY
Blackwell, 2013.
Moore KL, Dalley AF, Agur AMR, eds. Clinically oriented anatomy. 7th
edn. Wolters Kluwer/Lippincott Williams & Wilkins, 2014.
Sinnatamby CS, ed. Last’s anatomy. 12th edn. Churchill Livingstone
Elsevier, 2011.

SURGERY 36:11 605 Ó 2018 Elsevier Ltd. All rights reserved.