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NEUROSCIENCE FOR
NEUROSURGEONS
Edited by Farhana Akter, Nigel Emptage,
Florian Engert and Mitchel S Berger
CAMtikllXiL Mcdkint
Neuroscience for Neurosurgeons
Published online by Cambridge University Press

Neuroscience
for Neurosurgeons
Edited by
Farhana Akter
Harvard University
Nigel Emptage
University of Oxford
Florian Engert
Harvard University
Mitchel S. Berger
University of California–San Francisco

Shaftesbury Road, Cambridge CB2 8EA, United Kingdom
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education, learning and research at the highest international levels of excellence.
www.cambridge.org
Information on this title: www.cambridge.org/9781108831468
DOI: 10.1017/9781108917339
© Cambridge University Press & Assessment 2024
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place without the written permission of Cambridge University Press & Assessment.
First published 2024
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Library of Congress Cataloging-in-Publication Data
Names: Akter, Farhana (Surgeon), editor. | Emptage, Nigel, editor. | Engert, Florian,
editor. | Berger, Mitchel S., editor.
Title: Neuroscience for neurosurgeons / edited by Farhana Akter, Nigel Emptage,
Florian Engert, Mitchel S. Berger.
Description: Cambridge, United Kingdom; New York, NY: Cambridge University
Press, 2023. | Includes bibliographical references and index.
Identifiers: LCCN 2023017267 | ISBN 9781108831468 (hardback) | ISBN
9781108917339 (ebook)
Subjects: MESH: Neurosurgical Procedures | Nervous System Diseases –
physiopathology | Nervous System – physiology
Classification: LCC RD592.8 | NLM WL 368 | DDC 617.4/8092–dc23/eng/20230703
LC record available at https://lccn.loc.gov/2023017267
ISBN 978-1-108-83146-8 Hardback
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remain, accurate or appropriate.
.............................................................................................................................
Every effort has been made in preparing this book to provide accurate and up-to-date
information that is in accord with accepted standards and practice at the time of
publication. Although case histories are drawn from actual cases, every effort has been
made to disguise the identities of the individuals involved. Nevertheless, the authors,
editors, and publishers can make no warranties that the information contained herein is
totally free from error, not least because clinical standards are constantly changing through
research and regulation. The authors, editors, and publishers therefore disclaim all liability
for direct or consequential damages resulting from the use of material contained in this
book. Readers are strongly advised to pay careful attention to information provided by the
manufacturer of any drugs or equipment that they plan to use.

Contents
List of Contributors vii
Section 1 Basic and Computational 13 Brain Metastases: Molecules to Medicine 193

Ethan S. Srinivasan, Vadim Tsvankin,
Neuroscience Eric W. Sankey, Matthew M. Grabowski,
1 Neuroanatomy 1 Pakawat Chongsathidkiet,
Farhana Akter, Charles Reilly, Christophe and Peter E. Fecci
Dupre, and Shifa Hossain 14 Benign Adult Brain Tumors and Pediatric Brain
2 Cerebral Autoregulation 51 Tumors 214
Parisa Nikrouz, Xingping Qin, Farhana Akter Shun Yao, Umar Raza, and Farhana Akter
3 Neuroimmune Interactions 59 15 Biomechanics of the Spine 234

Allison Chang Gaetano De Biase and Kingsley Abode-Iyamah
4 Anatomy and Physiology of the Neuron 72 16 Degenerative Cervical Myelopathy 239

Oluwatobi Ariyo and Farhana Akter T.J. Florence, Joel S. Beckett,
and Langston T. Holly
5 Synaptic Transmission 84
17 Spondylolisthesis 248
Katrina Hon, Farhana Akter, and Nigel Emptage
Yike Jin, Ann Liu, Ravi Medikonda,
6 Sensory Pathways 99 and Timothy F. Witham
Rachel Chau, Megan Chau, and Farhana Akter
18 Radiculopathy 254
7 Somatosensory and Somatic Motor Systems 121 Yingda Li and Michael Y. Wang
Maria Kaltchenko and Farhana Akter
19 Spinal Tumors 267
8 Neuron Models 134 Ziev B. Moses, Matthew Trawczynski,
Kumaresh Krishnan and John E. O’Toole
9 An Introduction to Artificial Intelligence 20 Acute Spinal Cord Injury and Spinal
and Machine Learning 146 Trauma 278
Shun Yao, Ye Wu, and Farhana Akter Dominique M. O. Higgins, Pavan
S. Upadhyayula, Michael Argenziano,
10 Artificial Intelligence in Neuroscience 158
and Paul McCormick
Will Xiao, Mengmi Zhang, and Gabriel
Kreiman 21 Traumatic Brain Injury 291
Kristin A. Keith and Jason H. Huang
11 Probability and Statistics 167
Zachary T. Miller 22 Vascular Neurosurgery 300
Karol P. Budohoski and Adib A. Abla
23 Pediatric Vascular Malformations 314
Section 2 Clinical Neurosurgical Diseases Alaa Montaser and Edward R. Smith
12 Glioma 184
24 Craniofacial Neurosurgery 326
David M. Ashley and Justin T. Low
John T. Smetona and John A. Persing

Contents
25 Hydrocephalus 335 29 Neuroradiology: Focused Ultrasound

Benjamin C. Reeves, Jason K. Karimy, Phan in Neurosurgery 382
Q. Duy, and Kristopher T. Kahle Massimiliano Del Bene, Roberto Eleopra,
Francesco Prada, and Francesco DiMeco
26 Peripheral Nerve Injury Response
Mechanisms 348 30 Magnetic Resonance Imaging
Andrew S. Jack and Line Jacques in Neurosurgery 398
David J. Segar, Jasmine A. Thum, Dhiego
27 Clinical Peripheral Nerve Injury Models 355
Bastos, and Alexandra J. Golby
Andrew S. Jack, Charlotte J. Huie, and Line
Jacques 31 Brain Mapping 410
Anthony T. Lee, Cecilia Dalle Ore, and Shawn
28 The Neuroscience of Functional
L. Hervey-Jumper
Neurosurgery 369
Joseph S. Bell, T. J. Florence, Maya Harary,
Maxwell D. Melin, Hiro Sparks, and Nader
Pouratian
Index 422
vi

Contributors
Cecilia Dalle Ore

Adib A. Abla
University of California San Francisco, San Francisco,
University of California San Francisco, California, USA
CA, USA
Kingsley Abode-Iyamah
Gaetano De Biase
Mayo Clinic, Jacksonville, FL, USA
Mayo Clinic, Jacksonville, FL, USA
Farhana Akter
Massimiliano Del Bene
Harvard University, Cambridge, MA, USA
Fondazione IRCCS Istituto Neurologico Carlo Besta, Milan,
Michael Argenziano Italy and European Institute of Oncology IRCCS, Milan,
Columbia University College of Physicians and Surgeons, Italy
New York, NY, USA
Francesco DiMeco
Oluwatobi Ariyo Fondazione IRCCS Istituto Neurologico Carlo Besta,
Harvard University, Cambridge, MA, USA Milan, Italy, University of Milan, Milan, Italy, and Johns
Hopkins Medical School, Baltimore, MD, USA
David M. Ashley
Duke University School of Medicine, Durham, NC, Christophe Dupre
USA Harvard University, Cambridge, MA, USA
Joel S. Beckett Phan Q. Duy

David Geffen UCLA School of Medicine, CA, USA Yale School of Medicine, New Haven, CT, USA
Joseph S. Bell Roberto Eleopra

UCLA Department of Neurosurgery, Los Angeles, CA, Fondazione IRCCS Istituto Neurologico Carlo Besta,
USA Milan, Italy
Dhiego Bastos Nigel Emptage

Brigham and Women’s Hospital, Boston, MA, USA University of Oxford, Oxford, UK
Karol P. Budohoski Peter E. Fecci

University of Utah, Utah, USA Duke University Medical Center, Durham, NC, USA and
Duke University Center for Brain and Spine Metastasis,
Allison Chang Durham, NC, USA
T.J. Florence
Megan Chau David Geffen UCLA School of Medicine, CA, USA
Northwestern University, Evanston, IL, USA
Alexandra J. Golby
Rachel Chau Brigham and Women’s Hospital, Boston, MA, USA
Matthew M. Grabowski
Pakawat Chongsathidkiet Cleveland Clinic Neurological Institute, Cleveland, OH,
Duke University Medical Center, Durham, NC, USA USA
vii

List of Contributors
Maya Harary Science Center, College of Medicine, Temple, TX,

UCLA Department of Neurosurgery, Los Angeles, CA, USA USA
Shawn L. Hervey-Jumper Gabriel Kreiman

University of California San Francisco, San Francisco, Harvard Medical School, Boston, MA, USA
CA, USA
Kumaresh Krishnan
Dominique M. O. Higgins Harvard University, Cambridge, MA, USA
Columbia University College of Physicians and Surgeons,
New York, NY, USA Anthony T. Lee
University of California San Francisco, San Francisco,
Langston T. Holly CA, USA
David Geffen UCLA School of Medicine, CA, USA
Yingda Li
Katrina Hon Westmead Hospital, Sydney, Australia
Ann Liu
Shifa Hossain Johns Hopkins Hospital, Baltimore, MD, USA
Justin T. Low
Jason H. Huang Duke University School of Medicine, Durham, NC, USA
Baylor Scott and White Health, Medical Center, Temple,
TX, USA and Texas A&M University Health Science Paul McCormick
Center, College of Medicine, Temple, TX, USA Columbia University College of Physicians and Surgeons,
New York, NY, USA
Charlotte J. Huie
University of California San Francisco (UCSF), San Ravi Medikonda
Francisco, CA, USA Johns Hopkins Hospital, Baltimore, MD, USA
Andrew S. Jack Maxwell D. Melin

University of Alberta, Edmonton, AB, Canada and UCLA-Caltech Medical Scientist Training Program, Los
University of California San Francisco (UCSF), San Angeles, CA, USA
Francisco, CA, USA Alaa Montaser
Line Jacques Boston Children’s Hospital, Boston, MA, USA
University of California San Francisco (UCSF), San Ziev B. Moses
Francisco, CA, USA Rush University Medical Center, Chicago, IL, USA
Yike Jin Zachary T. Miller
Johns Hopkins Hospital, Baltimore, MD, USA Harvard University, Cambridge, MA, USA
Kristopher T. Kahle Parisa Nikrouz
Yale School of Medicine, New Haven, CT, USA Maidstone and Tunbridge Wells NHS Trust,
Maria Kaltchenko Kent, UK
Harvard University, Cambridge, MA, USA John E. O’Toole
Jason K. Karimy Rush University Medical Center, Chicago, IL, USA
Yale School of Medicine, New Haven, CT, USA John A. Persing
Kristin A. Keith Yale School of Medicine, New Haven, CT, USA
Baylor Scott and White Health, Medical Center, Nader Pouratian
Temple, TX, USA and Texas A&M University Health UT Southwestern Medical Center, Dallas, TX USA
viii

List of Contributors
Francesco Prada Jasmine A. Thum

Fondazione IRCCS Istituto Neurologico Carlo Besta, Brigham and Women’s Hospital, Boston, MA,
Milan, Italy, University of Virginia Health Science USA
Center, Charlottesville, VA, USA, and Focused
Ultrasound Foundation, Charlottesville, VA, USA Matthew Trawczynski
Rush University Medical Center, Chicago, IL, USA
Xingping Qin Vadim Tsvankin
Harvard School of Public Health, Boston, MA, USA Colorado Brain and Spine Institute, Denver, CO, USA
Umar Raza Vadim Tsvankin

National University of Medical Sciences (NUMS), Colorado Brain and Spine Institute, Denver, CO,
Rawalpindi, Pakistan USA
Benjamin C. Reeves Pavan S. Upadhyayula

Yale School of Medicine, New Haven, CT, USA Columbia University College of Physicians and Surgeons,
New York, NY, USA
Charles Reilly
Harvard University, Cambridge, MA, USA Michael Y. Wang
University of Miami Miller School of Medicine, Miami, FL,
Eric W. Sankey USA
Duke University Medical Center, Durham, NC, USA
Timothy F. Witham
David J. Segar Johns Hopkins Hospital, Baltimore, MD, USA
Brigham and Women’s Hospital, Boston, MA, USA
Ye Wu
John T. Smetona Nanjing University of Science and Technology, Nanjing,
Yale School of Medicine, New Haven, CT, USA Jiangsu, China
Edward R. Smith Will Xiao
Boston Children’s Hospital, Boston, MA, USA Harvard Medical School, Boston, MA, USA
Hiro Sparks Shun Yao
UCLA David Geffen School of Medicine, Los Angeles, CA, The First Affiliated Hospital, Sun Yat-sen University,
USA Guangzhou, Guangdong, China
Ethan S. Srinivasan Mengmi Zhang
Duke University Medical Center, Durham, NC, USA Harvard Medical School, Boston, MA, USA
ix

Section 1 Basic and Computational Neuroscience
Chapter
Neuroanatomy
1 Farhana Akter, Charles Reilly, Christophe Dupre, and Shifa Hossain
1.1 Anatomical Planes and Orientation 1.2 Vascular Supply to the Brain
of the Brain The brain requires 15–20% of the resting cardiac output
and is exquisitely sensitive to oxygen deprivation. Two
The neuroaxis of humans and other bipedal orthograde
main pairs of arteries supply blood to the brain: the
animals is different from that of quadruped animals.
internal carotid arteries and the vertebral arteries.
Example axes include the anteroposterior axis, rostrocau-
Within the cranial vault, an anastomotic circle, called
dal axis and the dorsoventral axis (Figure 1.1). The brain
the Circle of Willis (Figure 1.3), forms from the terminal
is usually visualized in sections cut through three orthog-
branches of these arteries.
onal planes: sagittal (longitudinal), coronal, and trans-
verse (axial) planes.
Dorsal view Anterior Ventral view Anterior
Superior frontal gyrus Posterior Posterior
Superior frontal sulci
Lateral view
Central sulcus
Parietal lobe
Occipital
Frontal lobe lobe
Temporal lobe
Cerebellum
Anterior Posterior Anterior Posterior
Figure 1.1 Planes and orientations of the brain.

Section 1 Basic and Computational Neuroscience
Chapter
Neuroanatomy
1 Farhana Akter, Charles Reilly, Christophe Dupre, and Shifa Hossain
1.1 Anatomical Planes and Orientation 1.2 Vascular Supply to the Brain
of the Brain The brain requires 15–20% of the resting cardiac output
and is exquisitely sensitive to oxygen deprivation. Two
The neuroaxis of humans and other bipedal orthograde
main pairs of arteries supply blood to the brain: the
animals is different from that of quadruped animals.
internal carotid arteries and the vertebral arteries.
Example axes include the anteroposterior axis, rostrocau-
Within the cranial vault, an anastomotic circle, called
dal axis and the dorsoventral axis (Figure 1.1). The brain
the Circle of Willis (Figure 1.3), forms from the terminal
is usually visualized in sections cut through three orthog-
branches of these arteries.
onal planes: sagittal (longitudinal), coronal, and trans-
verse (axial) planes.
Dorsal view Anterior Ventral view Anterior
Superior frontal gyrus Posterior Posterior
Superior frontal sulci
Lateral view
Central sulcus
Parietal lobe
Occipital
Frontal lobe lobe
Temporal lobe
Cerebellum
Anterior Posterior Anterior Posterior
Figure 1.1 Planes and orientations of the brain.
https://doi.org/10.1017/9781108917339.001 Published online by Cambridge University Press

F. Akter et al.
Figure 1.2 Planes and orientations of the

Sagittal body.
Coronal
Rostral
Dorsal
Ventral
Transverse
Caudal
The brachiocephalic trunk arises from the aorta and supplies the frontal lobes and medial aspects of the parie-
bifurcates into the right common carotid and right subcla- tal and occipital lobes (Figure 1.4).
vian artery. The left common carotid artery and subclavian The middle cerebral artery is the largest cerebral
artery branches off directly from the aortic arch. The artery and supplies the somatosensory and motor cortex,
common carotid arteries divide at the level of the thyroid basal ganglia, and the cerebral white matter. It can be
cartilage (C4) into external and internal carotid arteries. divided into four main surgical segments, denominated
The carotid sinus, a dilation at the base of the internal M1 (sphenoidal/horizontal), M2 (insular), M3 (opercu-
carotid artery, is a baroreceptor (stretch receptor) that lar), and M4 (cortical) segments. M1 gives rise to the
detects changes in systemic blood pressure. Intimately lenticulostriate perforating end arteries supplying the
related to it, is the carotid body, made up of glomus type basal ganglia and internal capsule. Occlusion of these
I chemoreceptor cells and glomus type II supporting cells. vessels can cause lacunar infarcts, the most common
The chemoreceptors are primarily sensors of partial pres- type of ischemic stroke. Hypertensive lipohyalinosis of
sure of oxygen (PaO2). The central chemoreceptors in the these vessels can lead to the formation of Charcot-
medulla oblongata are the primary sensors of the partial Bouchard aneurysms and these are a principal cause of
pressure of carbon dioxide (PaCO2) and pH, however the intracerebral hemorrhage. The cortical branches of MCA
carotid bodies also play a secondary role. arise from all of its segments and supply most of the
The external carotid artery divides into the superficial lateral surface of the brain and include the anterior tem-
temporal artery and the maxillary artery within the poral arteries from M1, lateral frontobasal artery from
parotid gland and gives rise to six branches (superior M2, and parietal branches from M4. The vertebral
thyroid artery, lingual artery, facial artery, ascending arteries arise from the subclavian artery and pass though
pharyngeal artery, occipital artery, and posterior auricu- the foramen transversarium in the cervical vertebrae. The
lar artery). The maxillary artery gives rise to the middle vertebral arteries enter the cranium via the foramen mag-
meningeal artery, which passes through the foramen spi- num and merge to form the basilar artery. Branches of
nosum to enter the cranial cavity. this artery include the anterior and posterior inferior
The internal carotid artery enters the cranial cavity via cerebellar arteries supplying the brainstem and the cere-
the carotid canal in the petrous part of the temporal bone, bellum alongside the superior cerebellar arteries. The
passes through the cavernous sinus and penetrates the basilar artery bifurcates into the posterior cerebral
dura into the subarachnoid space. Distal to the cavernous arteries and supplies the occipital lobes.
sinus, it gives rise to the following branches: ophthalmic
artery, posterior communicating artery, anterior choroi- 1.2.1 Circle of Willis
dal artery, and anterior cerebral artery. It then continues
An anastomotic circle is formed around the optic chiasm
as the middle cerebral artery. The anterior cerebral artery
between the anterior cerebral arteries (via the anterior

Neuroanatomy
Figure 1.3 Circle of Willis.

Anterior
Anterior
Middle communicating
cerebral
cerebral artery
artery
artery
Ophthalmic
artery
Internal
carotid Anterior
artery choroidal
artery
Posterior
communicating
artery
Posterior
cerebral
artery
Superior
Pontine arteries cerebellar artery
Basilar artery
Anterior
inferior
cerebellar
artery
Vertebral artery
Posterior
Anterior inferior
spinal artery cerebellar
artery
communicating artery) and the posterior cerebral arteries The dural venous sinuses are valveless structures
(via the posterior communicating arteries). found between the periosteal and meningeal layer of the
dura mater and drain into the internal jugular vein. The
1.2.2 Cerebral Venous Drainage straight, superior, and inferior sagittal sinuses are found
in the falx cerebri of the dura mater and converge at the
The superficial system of veins draining the cerebral cortex
confluence of sinuses. From here the transverse sinus,
include the superior cerebral veins, middle cerebral veins,
located bilaterally in the tentorium cerebelli, curves into
inferior cerebral veins, superior and inferior anastomotic
the sigmoid sinus and then to the internal jugular vein,
veins. The superficial veins drain into the superior and
which exits at the jugular foramen. The great cerebral
inferior sagittal sinuses (Figure 1.4).
vein and the inferior sagittal sinus continue as the straight
The deep veins include subependymal veins, the great
sinus.
cerebral vein, and medullary veins. The deep veins drain
The cavernous sinus is clinically relevant due to its
into the great cerebral vein and then to the straight or
vulnerability as a site of infection. It contains the internal
transverse sinuses.

F. Akter et al.
(a)
Pericallosal artery
Orbital branches Calcarine artery

Posterior cerebral artery
Anterior cerebral artery Superior cerebellar
artery
Basilar artery
(b)
Anterior cerebral artery
Precentral artery
Central artery
Postcentral artery
Angular artery
Temporo-occipital
artery
Middle cerebral artery
Anterior temporal artery
Superior cerebellar artery
Anterior inferior cerebellar
artery
Basilar artery Posterior inferior cerebellar
artery
Vertebral artery
(c)
Superior sagittal sinus
Superior anastomotic vein

Inferior sagittal sinus
Superficial
middle Inferior anastomotic vein
cerebral vein Great cerebral vein
Anterior Straight sinus
cerebral
vein Confluence of sinuses
Superior petrosal sinus Transverse sinus
Inferior petrosal sinus
Sigmoid sinus
Occipital sinus
Internal jugular vein
Figure 1.4 Arterial supply to the brain and venous drainage.
carotid artery, abducens nerve, oculomotor nerve, troch- reverse blood flow to the sinus, and is therefore
lear nerve, and the ophthalmic and maxillary branches of a potential route of intracranial infection. The cavernous
the trigeminal nerve. The sinus is connected to the valve- sinus also receives venous drainage from the central vein
less facial vein via the superior ophthalmic vein, allowing of the retina, the sphenoparietal sinus, the superficial

Neuroanatomy
middle cerebral vein, and the pterygoid plexus. These solutes from the parenchyma to the deep cervical lymph
empty into the superior and inferior petrosal sinuses nodes. The lymphatic system also helps maintain the
and, ultimately, into the internal jugular vein. The left water and ion balance of the ISF and allows communi-
and right cavernous sinuses are connected in the midline cation with the immune system. The lymphatic vessels
by the anterior and posterior intercavernous sinuses. are associated with the superior sagittal and transverse
sinus and the dural middle meningeal arteries. It is likely
1.2.3 Lymphatics of the Brain that the lymphatic system works in conjunction with
other efflux pathways, such as via the dural arachnoid
It has been a longstanding belief that the vertebrate
granulations.
brain does not contain a classic lymphatic drainage
system. Instead, it was thought the brain has
a “glymphatic system,” a perivascular pathway that 1.3 Bones of the Skull
allows cerebrospinal fluid (CSF) to recirculate through The skull is a supportive protection for the brain and is
the brain interstitium along perivascular spaces near the formed by intramembranous ossification of cranial and
cerebral arteries. This pathway was also thought to be facial bones. The calvarium is the roof of the cranium and
responsible for the clearance of interstitial solutes along is comprised of the frontal, occipital, and parietal bones
perivascular channels near the veins, mediated by astro- (Figure 1.5). The base of the cranium is comprised of the
glial water channels. However, recent findings have pro- ethmoid, occipital, temporal, parietal, frontal, and sphe-
vided evidence of a functional lymphatic system in the noid bones. The junction between the latter four bones is
brain, which acts to transport interstitial fluid (ISF) and known as the pterion and is of clinical relevance because
(a) Figure 1.5 The skull and facial

skeleton.

(b) Figure 1.5 (cont.)
(c)

(d) Figure 1.5 (cont.)
(e) Palatine Process (Maxilla) Maxilla
Zygomatic Process (Maxilla)

Palatine Bone
Zygomatic Bone
Pterygoid Process
(Sphenoid Bone) Frontal Bone
Zygomatic Process Sphenoid Bone

(Temporal Bone)
Petrous Portion Inferior Nasal

(Temporal Bone) Concha
Vomer
Mandibular Fossa
(Temporal Bone)
Styloid Process
(Temporal Bone)
Squamous Portion Temporal Bone

(Temporal Bone)
Tympanic Portion
(Temporal Bone)
Mastoid Process Foramen Magnum

(Temporal Bone)
Occipital Condyle Parietal Bone

(Occipital Bone)
External Occipital
Protuberance Occipital Bone
(Occipital Bone)

F. Akter et al.
Anterior
fontanelle Squamous
suture Coronal
suture
Anterior
fontanelle Posterior
Coronal
fontanelle
suture
Frontal
bones Parietal
bones
Parietal Lambdoid
Frontal
bones suture
bones
Nasal
bone Sagittal suture Occipital
Posterior Frontal sature
Zygomatic bone
fontanelle
bone
Lambdoid
Maxilla Frontal suture
Parietal
Lambdoid bones bones
suture
Occipital
Greater wing Temporal
Mandible Mastoid bone
of sphenoid bone
fontanelle Coronal
Sphenoidal
suture
fontanelle
Lateral view Superior view
Coronal Suture Sphenoparietal

Suture
Sphenofrontal
Squamosal
Suture
Suture
Sphenosquamosal
Suture
Parietomastoid
Suture
Frontoethmoidal
Suture
Frontolacrimal
Lambdoidal Suture
Suture Frontonasal
Suture
Frontomaxillary
Suture
Nasomaxillary
Suture
Ethmoidolacrimal
Suture
Maxillolacrimal
Suture
Ethmoido-
Occipitomastoid Suture maxillary
Suture
Temporozygomatic
Suture
Frontozygomatic
Suture
Zygomaticomaxillary Suture
Figure 1.6 Sutures of the brain.
it overlies the middle meningeal artery, which can rup- zygomatic arch. Other areas prone to damage are the
ture following fractures to this region, leading to the sutures, and these include the coronal, sagittal, and
formation of extradural hematomas. lambdoid sutures (Figure 1.6). In neonates, incom-
Cranial fractures may be accompanied by facial bone pletely fused sutures give rise to fontanelles – the frontal
fractures and should be sought for when assessing the fontanelle between the coronal and sagittal sutures and
trauma patient. The most common facial fractures the occipital fontanelle between the sagittal and lamb-
include those of the nasal bone, maxilla, mandible, and doid sutures.

Neuroanatomy
Cribriform plate (CNI)
Optic canal (CNII)

Superior orbital fissure (CNIII, CNIV, CNV1, CNVI)
F. Rotundum (CNV2)
F. Ovale (CNV3) F. lacerum (artery & nerve of pterygoid canal, greater &
deep petrosal nerve, meningeal branches of ascending
pharyngeal artery, emissary veins)
Internal acoustic meatus (CNVII, CNVIII)
Jugular foramen (CNIX, CNX, CNXI)

Hypoglossal canal (CNXII)
Foramen magnum (medulla oblongata,

vertebral artery, accessory nerves, spinal
arteries)
F. spinosum (middle meningeal artery

& vein, meningeal branch of Carotid canal (internal carotid
mandibular nerve) artery, carotid plexus)
Figure 1.7 Superior view of the skull base showing the foramina.
1.4 The Cranial Fossae part of the fossa is the sella turcica, a bony prominence
which supports the pituitary gland within the hypophy-
The cranial cavity is divided into three regions known as
sial fossa. The posterior wall of the sella turcica is formed
fossae. The anterior cranial fossa overlies the nasal and
by the dorsal sellae, which separate the middle cranial
orbital regions and accommodates parts of the frontal
fossa from the posterior cranial fossa. The lateral parts of
lobe. It is made up of the frontal, ethmoid, and sphenoid
the middle cranial fossa are formed by the greater wings
bones. The frontal crest on the frontal bone and the crista
of the sphenoid bone and the squamous and petrous parts
galli of the ethmoid bone are the sites of attachment for
of the temporal bones and provide structural support to
a part of the dura mater that divides the cerebral hemi-
the temporal lobes. Both the sphenoid and temporal
spheres, known as the falx cerebri. The cribriform plate
bones contain numerous foramina for transmitting ves-
supporting the olfactory bulb is lateral to the crista galli,
sels and nerves. The posterior cranial fossa is comprised
and contains a foramen transmitting the olfactory nerve
of the occipital bone and the temporal bones and contains
and two ethmoidal foramina (anterior and posterior,
the brainstem and cerebellum. The foramina of the skull
transmitting the anterior and posterior ethmoidal vessels
are most considered in the context of the cranial nerves
and nerves, respectively). The plate is very thin and can
and are shown in Figure 1.7.
fracture following facial trauma, resulting in CSF rhinor-
rhea and anosmia.
The anterior fossa is separated from the middle fossa 1.5 Layers of the Scalp
by the lesser wing of the sphenoid bone. The anterior The scalp contains several layers, and these include
clinoid processes of these bones are the site of attachment skin, dense connective tissue, epicranial aponeurosis,
for the tentorium cerebelli (dura mater dividing the cere- loose areolar connective tissue, and the periosteum
brum and cerebellum). The middle cranial fossa consists (Figure 1.8). The scalp is supplied by branches of the
of the sphenoid and temporal bones. Within the central external carotid artery (superficial temporal, posterior

F. Akter et al.
Figure 1.8 Layers of the scalp.
auricular, and occipital arteries) and the ophthalmic and in between houses the dural venous sinuses. The
artery (supraorbital and supratrochlear arteries). meningeal layer folds inward to form the dural reflections
Venous drainage includes a superficial system following and forms compartments. These compartments are the
the arterial supply (superficial temporal, occipital, poste- falx cerebri, the tentorium cerebelli, the falx cerebelli
rior auricular, supraorbital, and supratrochlear veins). (separates the cerebellar hemispheres), and the dia-
The temporal region of the skull is drained by the pter- phragma sellae (allows the passage of the pituitary
ygoid venous plexus, which drains into the maxillary stalk). Blood collecting in between the skull and the
vein. The scalp veins connect to the diploic veins of the outer periosteal layer is known as an extradural hema-
skull via valveless emissary veins, allowing a connection toma and usually occurs due to damage to the middle
between the scalp and the dural venous sinuses. The meningeal artery. A subdural hematoma occurs due to
nerves innervating the scalp include the trigeminal damage of the cerebral veins between the dura and the
nerve branches (supratrochlear, supraorbital, zygomati- arachnoid mater.
cotemporal, and auriculotemporal nerves). The cervical
nerve branches supplying the scalp include the lesser
occipital nerve, the greater occipital nerve, the great 1.6.2 Arachnoid Mater
auricular nerve, and the third occipital nerve. The arachnoid mater consists of avascular connective
tissue. Below it lies the subarachnoid space containing
1.6 Meninges CSF, which re-enters the circulation via the dural venous
sinuses through small projections called arachnoid
The brain and spinal cord are covered with membranous
granulations.
layers called the meninges. From outer to inner these are
the dura, arachnoid, and pia mater.
1.6.3 Pia Mater
1.6.1 Dura Mater This is a highly vascularized layer that adheres to the
This layer is found underneath the bones and consists of brain surface and follows the contours of the brain into
the outer periosteal layer and the inner meningeal layer the gyri of the cerebral hemispheres and the folia of the
10

Neuroanatomy
cerebellum. The pia mater and arachnoid mater are A major anatomical component of the SNS are a pair
joined by connective tissue in the subarachnoid space of nerve fibers that span the skull to coccyx, known as the
and together are known as leptomeninges. There are sympathetic chains. There are two types of neurons
compartments where the two are not in close approxima- involved in sympathetic signal transmission and these
tion, resulting in naturally enlarged CSF filled pools are the short preganglionic neurons that originate from
called the subarachnoid cisterns. T1 to L2–L3, which synapse with postganglionic neurons
that extends to the rest of the body. At the synapses,
1.7 Organization of the Sympathetic the preganglionic neurons release acetylcholine, which
activates the nicotinic acetylcholine receptors in the post-
and Parasympathetic Nervous Systems ganglionic neurons to release norepinephrine, and these
The autonomic nervous system is composed of the sym- subsequently bind to adrenergic receptors in the target
pathetic nervous system (SNS) and the parasympathetic tissue leading to sympathetic effects (Figure 1.9). The
nervous system (PNS) and acts to regulate the body’s postganglionic neurons of sweat glands release acetylcho-
unconscious actions. The SNS stimulates the so-called line to activate muscarinic receptors. The chromaffin cells
“fight or flight” response and the PNS is involved in of the adrenal medulla act as a postganglionic neuron and
“rest and digest” responses (Box 1.1). release norepinephrine and epinephrine.
Box 1.1
Organ Parasympathetic Response Sympathetic Response

Iris Miosis and accommodation due to constriction of the Pupil dilation (adrenergic innervation to the dilator
sphincter muscles via the short ciliary nerves originating pupillae muscle via the long ciliary nerves, arising from the
from the Edinger–Westphal nucleus of cranial nerve III superior cervical ganglion)
Salivary Increased watery secretion via cranial nerves IX (parotid Reduced saliva secretion (innervation via fibers arising from
glands gland) and chorda tympani of VII (submandibular and the superior cervical ganglion resulting in norepinephrine
sublingual glands) leading to acetylcholine release onto release acting on alpha- and beta-adrenergic receptors)
M3 muscarinic receptors
Lacrimal Increased secretion (preganglionic fibers reach the Reduced secretion (innervation via fibers originating in the
glands pterygopalatine ganglion via the greater petrosal nerve superior cervical ganglion, which reach the
and the nerve of the pterygoid canal to synapse with pterygopalatine ganglion via the internal carotid plexus
postganglionic fibers) and the deep petrosal nerve)
Heart Negative chronotopy, inotropy and reduced conduction Positive chronotropy, inotropy and increased conduction
velocity and coronary artery vasoconstriction via the vagus velocity via the cardiac nerves from the lower cervical and
nerve upper thoracic ganglia
Lung Bronchial muscle contraction (vagus nerve) Bronchial muscle relaxation (thoracic sympathetic ganglia)
Stomach Increased peristalsis and motility and pyloric sphincter Reduced gastric motility and peristalsis and pyloric
relaxation allowing gastric emptying via the vagus nerve sphincter constriction preventing gastric emptying via the
celiac plexus (T5–T12)
Gallbladder Contraction (vagus nerve) Relaxation (T7–T9 through the celiac plexus)
Internal Relaxation Constriction
urethral
sphincter
Detrusor Contraction (via pelvic splanchnic nerves to allow bladder Relaxation (via sympathetic branches from the inferior
muscle of emptying) hypogastric plexus to allow bladder filling)
bladder
Penis Erection (pelvic nerve) Ejaculation (peristaltic contraction of vas deferens, seminal
vesicles, and prostatic smooth muscles via the hypogastric
nerve and ejaculation via the pudendal nerve)
Adrenal Norepinephrine and epinephrine secretion
medulla
11

F. Akter et al.
Sympathetic nervous system
Cholinergic fiber
Neuron
Effector
organ
Adrenergic fiber
Preganglionic fiber Postganglionic fiber
Ganglion Nicotinic
CNS
cholinergic receptor
Muscarinic
cholinergic receptor
Adrenal Effector Adrenergic
medulla organ cholinergic receptor
CNS
Acetylcholine
Norepinephrine/
epinephrine
Parasympathetic nervous system
Effector
Preganglionic fiber organ
Postganglionic fiber
CNS Ganglion
Figure 1.9 Sympathetic and parasympathetic fibers. CNS; central nervous system.
Sympathetic nerves arise in the spinal cord in the nerve to the pharyngeal plexus, the superior cardiac
intermediolateral nucleus of the lateral gray column. branch, and the gray rami communicantes. The middle
Axons leave the spinal cord through the anterior root ganglion may be absent, but when present is found ante-
and pass near the sensory ganglion to enter the anterior rior to the inferior thyroid artery. Its postganglionic fibers
rami of spinal nerves. The axons terminate at the para- are the gray rami communicantes, the thyroid branches,
vertebral or prevertebral ganglia. The main prevertebral and the middle cardiac branch. The inferior cervical
ganglia (celiac, mesenteric and aorticorenal ganglia) are ganglion is situated anteriorly to the C7 vertebra. Its
located anterior to the aorta and vertebral column and branches are the gray rami communicantes, branches to
receive preganglionic axons via the splanchnic nerves. the subclavian and vertebral arteries, and the inferior
The paravertebral ganglia are located bilaterally ventro- cardiac nerve. Damage to sympathetic fibers en route to
lateral to the vertebral column. There are three paraver- the head and neck can lead to Horner’s syndrome, a
tebral ganglia and these are the superior cervical condition presenting with partial ptosis of the upper
ganglion, middle cervical ganglion and inferior cervical eyelid, miosis (constricted pupil) and hemi-facial anhi-
ganglion. drosis (absence of sweating).
1.7.1 Sympathetic Ganglia Supplying 1.7.2 Parasympathetic Ganglia Supplying

the Head and Neck: Cervical Ganglia the Head and Neck
There are three cervical ganglia that supply the head and The parasympathetic fibers supplying the head and neck
neck (the superior, middle, and inferior cervical ganglia). are found in four brainstem nuclei associated with a cranial
The superior ganglion is found posterior to the carotid nerve. They synapse in a peripheral ganglion near the
artery and gives rise to a number of postganglionic target viscera. There are four parasympathetic ganglia
nerves: the internal and external carotid nerves, the located within the head –ciliary, otic, pterygopalatine, and
12

Neuroanatomy
submandibular. They receive fibers from the oculomotor, three expansions (vesicles) develop: the forebrain (pros-
facial, and glossopharyngeal nerves (the vagus nerve only encephalon), the midbrain (mesencephalon), and the
innervates structures in the thorax and abdomen). hindbrain (rhombencephalon). Further division sepa-
The ciliary ganglion is located within the bony orbit. rates the prosencephalon into the diencephalon (thala-
Its preganglionic fibers are from the Edinger–Westphal mus and hypothalamus) and the telencephalon
nucleus, associated with the oculomotor nerve. Its post- (cerebrum). The mesencephalon consists of the tectum,
ganglionic fibers leave the ganglion via the short ciliary cerebral aqueduct, tegmentum, and the cerebral pedun-
nerves to innervate the sphincter pupillae and the ciliary cles. The rhomboencephalon consists of the pons, the
muscles. Sympathetic nerves from the internal carotid medulla, and the cerebellum. The cavities within the
plexus and sensory fibers from the nasocilary nerve pass primary brain vesicles are precursors of the ventricular
through the ganglion without synapsing. system. The caudal parts of the neural tube form the
The pterygopalatine ganglion is located within the spinal cord.
pterygopalatine fossa and is supplied by fibers from the
superior salivatory nucleus (associated with the facial 1.9 Forebrain
nerve). Its postganglionic fibers join branches of the max-
The structures in the forebrain include the cerebral cortex
illary nerve to supply the lacrimal gland, the nasophar-
and subcortical structures of the limbic system including the
ynx, and the palate.
amygdala, hypothalamus, thalamus, hippocampus, basal
Sympathetic fibers from the internal carotid plexus
ganglia, and cingulate gyrus.
and sensory branches from the maxillary nerve pass
through the pterygopalatine ganglion without synapsing.
The submandibular ganglion is located inferiorly to 1.10 Cerebrum
the lingual nerve and is supplied by fibers from the The largest part of the brain is the cerebrum, containing
superior salivatory nucleus. These fibers are carried two hemispheres separated by the falx cerebri of the dura
within a branch of the facial nerve, the chorda tympani. mater. The visible surface of the cerebral hemisphere is
This nerve travels along the lingual branch of the man- a folded sheet of neural tissue called the cerebral cortex,
dibular nerve to reach the ganglion and leaves the gan- characterized by sulci (depressions) and gyri (elevations).
glion to the submandibular and sublingual glands. Some of the larger folds include the lateral sulcus (also
Sympathetic fibers from the facial artery plexus pass known as the Sylvian fissure), which separates the tempo-
through the submandibular ganglion. They are thought ral lobe from the frontal and parietal lobes, the central
to innervate glands in the base of the oral cavity. sulcus (dividing the frontal and parietal lobes), and the
The otic ganglion is located inferiorly to the foramen parieto-occipital sulcus, which separates the occipital and
ovale within the infratemporal fossa. It is medial to the parietal lobes (Figure 1.10). The two cerebral hemispheres
mandibular branch of the trigeminal nerve. The gan- are connected by a white-matter tract called the corpus
glion is supplied by fibers from the inferior salivatory callosum. The tissue separating the two hemispheres is
nucleus (associated with the glossopharyngeal nerve). called the longitudinal fissure. The septum pellucidum
Parasympathetic fibers travel within the lesser petrosal continues from the corpus callosum to the fornix and
nerve, a branch of the glossopharyngeal nerve, to reach separates the anterior horns of the lateral ventricles. The
the otic ganglion. The parasympathetic fibers travel calcarine fissure separates the occipital lobe into the infe-
along the auriculotemporal nerve (a branch of the man- rior lingual gyrus and the superior cuneus.
dibular division of the trigeminal nerve) to provide The cerebrum is made up of gray matter (containing
secretomotor innervation to the parotid gland. cell bodies and dendrites) and white matter (consisting of
Sympathetic fibers from the superior cervical chain glial cells and myelinated axons). The cerebrum can be
pass through the otic ganglion, where they travel with divided into four lobes. The frontal lobe subserve deci-
the middle meningeal artery to innervate the parotid sion-making and executive control. The parietal lobe is
gland. vital for sensory perception and integration. The occipital
lobe is the visuospatial processing area of the brain for
1.8 Structures of the Brain color, form and motion. The temporal lobe contains
The nervous system forms during the third week of cortical areas that process auditory stimuli, encoding of
development. At the cranial end of the neural tube, memory and language comprehension.
13

F. Akter et al.
Pre central sulcus Central sulcus Post central sulcus
Cingulate gyrus
Corpus callosum
Thalamus
Septum
pellucidum Parieto-occipital
sulcus
Calcarine
fissure
Fornix
Olfactory bulb
Lateral sulcus
Optic chiasm
Temporal lobe
(medial surface)
Cut edge of brainstem
Hippocampus
Figure 1.10 Structures of the brain (medial surface).
1.10.1 Brodmann’s Map into two streams. The ventral stream is from the primary
visual cortex to the temporal lobe and is important for
The cerebral cortex can also be subdivided into 52 func-
pattern and object recognition. The dorsal stream from
tional regions, numbered by the neuroanatomist
the striate cortex into the parietal lobe is responsible for
Korbinian Brodmann based on cytological structure.
spatial recognition of motion and location. The primary
The primary motor cortex (Brodmann area 4) is anterior
auditory cortex is responsible for recognition of auditory
to the central sulcus (Figure 1.10). It contains large neu-
stimuli and is in the lateral temporal lobe. Wernicke’s
rons called Betz cells, which send axons to the spinal cord
area (Brodmann area 22) is located in the superior tem-
and is important for planning and execution of move-
poral gyrus of the dominant cerebral hemisphere. It is
ments. The topographic map of the motor cortex is
important for comprehension of written and spoken lan-
arranged with an “overrepresentation” of neurons
guage, and therefore any damage to this area leads to
responsible for complex motor behaviors (Figure 1.11).
fluent but nonsensical speech. Broca’s area (Brodmann
The premotor cortex (Brodmann area 6) also plays a role
areas 44 and 45) is located in the dominant prefrontal
in planning movement; however, its function is less well
cortex and is involved in language processing and speech
understood. The supplementary motor area (Brodmann
production. Lesions in this region lead to expressive
area 6) contributes to the control of movement. The
aphasia, where the patient retains comprehension but
primary somatosensory cortex (Brodmann areas 3, 2,
cannot create fluent speech.
and 1) is found in the parietal lobe, posterior to the
central sulcus. It processes afferent somatosensory input
and helps integrate sensory and motor information 1.10.2 Layers of The Cerebral Cortex
required for skilled movement. The primary visual cortex The cerebral neocortex is arranged in six layers. The
(Brodmann area 17) is at the occipital pole. The extra outermost layer is the molecular layer (layer I), con-
striate cortex is adjacent to the visual cortex and processes taining fibers that run parallel to the cortical surface
specific features of visual information. It can be separated with very few neurons. Layer II is the outer granular
14

Neuroanatomy
Primary motor
Supplementary
cortex
motor area
Primary somatosensory cortex
Premotor Posterior parietal cortex
cortex
Prefrontal
cortex Extrastriate
cortex
Primary visual
Wernicke's striate cortex
area
Primary
Broca's area Mid-sagittal view
auditory cortex
Lateral view
Trunk Lower
extremity
Upper extremity
Face
Coronal view
Figure 1.11 Topographic map of the primary motor cortex.
layer, containing small, rounded neurons. Below this is the lower motor neurons in the ventral horn of the
the outer pyramidal layer (layer III), containing pyram- spinal cord. Finally, layer VI is the innermost, multi-
idal neurons. Next is the inner granular layer (layer IV), form layer containing morphologically heterogenous
containing spiny stellate and pyramidal cells. It is a population of neurons and sends efferent fibers to the
major input layer and receives specific sensory inputs thalamus.
from thalamocortical afferent fibers. The primary sen- Certain parts of the cortex are arranged differently:
sory cortex has a well-developed layer IV, but the out- the hippocampus has three layers and the cingulate gyrus
put layers are less well developed. Layer IV is well has four to five layers.
developed in the primary visual cortex and contains
the stria of Gennari, a band of myelinated axons that 1.10.3 Vascular Supply
runs parallel to the surface of the cerebral cortex. The
The branches of the anterior, middle, and posterior cere-
inner pyramidal later (layer V) contains many output
bral arteries are responsible for the blood supply to the
neurons. The primary motor cortex has a very enlarged
cerebrum. The venous drainage of the cerebrum is via
layer V containing large Betz cells, which send axons to
cerebral veins that empty into the dural venous sinuses
the contralateral motor nuclei of cranial nerves and to
(Figure 1.4).
15

F. Akter et al.
Figure 1.12 The ventricular

system
Third ventricle
Cerebral aqueduct
Fourth ventricle
Spinal canal
Lateral ventricle
(beneath overlying cortex)
1.11 Ventricles
The ventricles (Figure 1.12) are lined by ependymal cells,
ciliated glial cells, which form the choroid plexus, a struc-
ture where CSF is produced (Figure 1.13). This provides
hydromechanical protection by acting as a shock
absorber and providing buoyancy.
The ventricular system is composed of four connect-
ing cavities derived from the neural tube. The right and
left ventricles and the third ventricle are part of the
forebrain, while the fourth ventricle is part of the
hindbrain.
During development, the fluid-filled cavity of the pri-
mary vesicles become the lateral ventricles. They commu-
nicate with the third ventricle via the foramen of Monro Capillaries
in the diencephalon. The third ventricle communicates Ependymal cells
with the fourth ventricle in the hindbrain via the cerebral Figure 1.13 Histology of the choroid plexus
16

Neuroanatomy
aqueduct (of Sylvius), which is continuous with the cen- globus pallidus and then to the thalamus and cerebral
tral canal of the spinal cord. The fourth ventricle is also cortex, forming a subcortical loop. The internal segment
connected to the subarachnoid space by a median aper- projects to the motor areas of the thalamus and the
ture (the foramen of Magendie) and two lateral apertures medial nucleus of the thalamus. The external segment
(the foramina of Luschka). projects to the subthalamic nucleus (STN), which also
projects to the internal globus pallidus.
1.12 Higher Association Areas The neostriatum also projects to the substantia nigra
par compacta (SNPC), which contains dopaminergic
of the Cortex neurons and to the pars reticulata, which contains
Higher association areas of the cortex are involved in mainly GABAergic neurons and is one of the output
complex processing of various sensory modalities, cog- nuclei of the basal ganglia to the thalamus (alongside
nition and emotion. The prefrontal and limbic associa- the internal globus pallidus) and plays a vital role in
tion areas are important for regulation of cognition, movement execution. The neostriatum is supplied by
abstract reasoning, complex emotions and self-aware- small branches of the middle and anterior cerebral
ness. The cingulate and parahippocampal gyri are arteries. The neostriatum and internal capsule are com-
involved in expression of emotions and formation of monly affected in stroke. Damage to the internal capsule
memories. The hippocampal formation is important leads to contralateral weakness. The most affected
for declarative memory. It contains the Cornu region of the internal capsule is the genu, where corti-
Ammonis (CA) regions, the subiculum, and the dentate cospinal fibers to the head, neck, and part of the upper
gyrus, and is found in the medial temporal lobe in the limb are located.
inferior horn of the lateral ventricle. The amygdala is
a subcortical nucleus in the medial temporal lobe and is 1.13.1.1 Connections of the Basal Ganglia
connected to the orbitofrontal cortex, the hypothala- The striatum receives input from the cerebral cortex
mus, and the nucleus accumbens. and the SNPC (Figure 1.15). The striatum sends inhibi-
tory connections to the internal and external globus pal-
1.13 Limbic System lidus. The external region therefore disinhibits the STN,
The limbic system is a part of the brain involved in which then sends excitatory input to the internal globus
behavioral and emotional responses. There are several pallidus. This sends inhibitory input to the thalamus,
important structures within the limbic system and these leading to inhibition of information flow to the cerebral
include the basal ganglia, thalamus, hypothalamus, hip- cortex.
pocampus, amygdala, and the cingulate gyrus. The basal ganglia also project to the medial dorsal
nucleus of the thalamus, which then projects to the pre-
frontal association cortex. This region is involved in
1.13.1 Basal Ganglia higher cortical and executive function.
The basal ganglia are situated at the base of the forebrain Parkinson’s disease results from degeneration of
and top of the midbrain. They are a group of subcortical dopaminergic neurons of the SNPC and excessive inhibi-
nuclei connected to the cerebral cortex, thalamus, and the tion of the thalamus. The SNPC projects to both direct
brainstem. The basal ganglia are responsible for control and indirect pathways in the striatum. Due to the pres-
of voluntary motor movements, procedural learning, cog- ence of two different types of dopamine receptors, the net
nition, and emotion. They consist of several distinct effect is to excite the direct pathway and inhibit the indi-
structures (Figure 1.14) and these include the caudate rect pathway. However, the loss of the neurons in
nucleus and putamen (together known as the neostria- Parkinson’s disease upsets the fine balance of these path-
tum) separated by the internal capsule). The internal ways and reduces excitation of the motor cortex, resulting
capsule is the site of the passage of many fibers including in poverty of movement.
the efferent corticobulbar fibers, corticospinal fibers,
efferent corticopontine fibers, and afferent thalamocorti-
cal fibers.
1.13.2 Thalamus
The caudate receives inputs from the prefrontal cortex The thalamus is located in the forebrain and contains
and the putamen receives inputs from the sensorimotor nuclei with connections to the cerebral cortex, the hippo-
cortex. They send outputs to the external and internal campus, the mammillary bodies, and the fornix.
17

F. Akter et al.
Figure 1.14 Cross-

section of the basal
ganglia.
Fiber groups: Cell groups:
Cerebral cortex
Corpus callosum
Septal area
Fornix Caudate nucleus
Cortical
white matter Putamen
Internal capsule
Globus pallidus
The thalamus is divided into three major nuclear memory, regulation of sleep, and wakefulness. Lesions
groups (anterior, medial, and ventral) (Figure 1.16). in the anterior thalamus can lead to obstruction of the
The ventral group contains ascending somatosensory interventricular foramen of Monro and lesions in the
relays (ventroposterior nuclei) and relays from the posteriomedial thalamus can obstruct the third ventricle
cerebellum and basal ganglia (ventrolateral). It also and cerebral aqueduct, leading to the development of
contains the motor association areas (ventro-anterior hydrocephalus.
nuclei). The lateral and medial geniculate nuclei are
found posteriorly. The anterior group projects to the 1.13.3 Cingulate Gyrus
cingulate gyrus and receives input from the mammil-
The cingulate gyrus is situated above the corpus callosum
lary bodies of the hypothalamus. The medial nuclei
(Figure 1.10). The anterior part relays signals between the
and the pulvinar receive input from the cerebral cor-
right and left hemispheres and is involved in autonomic
tex and form the cortico-thalamo-cortical relays,
functions and cognitive processes such as reward behav-
which project to areas of the association cortex.
ior, empathy, and emotion. The posterior part becomes
These are the prefrontal cortex and the temporal–
continuous with the most medial part of the temporal
parietal–occipital association cortex, which mainly
lobe, the parahippocampal gyrus. The cingulate gyrus is
receive input from the medial nuclei and the pulvinar,
thought to be involved in retrieving episodic memory
respectively.
information. Dysfunction of this gyrus is found in schiz-
The thalamus is supplied by the posterior cerebral
ophrenia and depression. Deep brain stimulation of the
artery and branches of the posterior communicating
subgenual cortex of the gyrus is used to treat intractable
artery. The thalamus is involved in learning, episodic
18

Neuroanatomy
Figure 1.15 Basal ganglia connections.
Figure 1.16 Nuclei of the thalamus.
Ventral
posterior
Centromedial medial
nucleus (VPM)
(CM)
Ventral
Lateral Lateral posterior
dorsal posterior lateral
(LD) (LP) (VPL)
Medial
nuclei Internal
medullary
Anterior
nuclei Pulvinar
Medial
geniculate
body
Ventral Lateral
anterior geniculate
Ventral Ventral body
(VA)
lateral posterior
(VL) (VP)
19

F. Akter et al.
Figure 1.17 Cornu Ammonis

axons of the hippocampus.
depression. Output fibers are mainly to the parahippo- hippocampus has been described as similar to a seahorse
campal gyrus and are linked by the cingulum, a white- or a ram’s horn (Cornu Ammonis). The abbreviation
matter tract. CA is used to name the different regions: CA1, CA2,
CA3, and CA4 (Figure 1.17).
1.13.4 Hippocampus 1.13.4.1 Dentate Gyrus
The hippocampus is a seahorse-shaped structure and the
The dentate gyrus contains granule cells and axons called
hippocampal formation refers to the hippocampus
mossy fibers, which synapse with the pyramidal cells in
proper (Cornu Ammonis), the dentate gyrus, and the
the CA3 field of the hippocampus. They also contain
subiculum. It is found in the temporal lobe, medial to
some pyramidal cells in the polymorphic cell layer.
the inferior horn of the lateral ventricle.
The hippocampus has been studied extensively and is 1.13.4.2 Hippocampal Inputs
used as a model system for electrophysiological studies,
The hippocampus receives information from the lateral
particularly for investigating neural plasticity. Damage to
perforate and medial perforate pathways in the entorh-
the hippocampus is commonly seen in patients with
inal cortex (Figure 1.18), the prefrontal cortex, the ante-
dementia, particularly Alzheimer’s disease.
rior cingulate gyrus, the pre- mammillary region, and
Hippocampal tissue is made up of layers and these
the reticular formation of the brainstem. It also receives
include, from outer to inner: an external plexiform layer;
input from the thalamus to field CA1 and from the
a stratum oriens layer containing basal dendrites and
serotonin, norepinephrine, and dopamine systems.
basket cells; a pyramidal cell layer containing the pri-
The medial septal nucleus sends cholinergic and γ-ami-
mary cells of the hippocampus; a stratum radiatum
nobutyric acid (GABA)-ergic inputs to the
layer; and the stratum lacunosum-moleculare layers
hippocampus.
containing the perforate pathway made up of pyramidal
The largest input and output pathway of the hippo-
cell apical dendrites and afferent fibers from the entorh-
campus is via the fornix, which connects it to other
inal cortex. The external plexiform layer contains the
structures including the mammillary bodies of the hypo-
alvear pathway, and this contains pyramidal cell axons
thalamus, prefrontal cortex and the lateral septal area.
through which information from the hippocampus is
The entorhinal cortex (part of the parahippocampal
passed to the inferior horn of the lateral ventricle before
gyrus) receives output from the deeper layers of the
reaching the entorhinal cortex. In addition, the hippo-
hippocampus and gives input to the superficial layers.
campus also contains distinct regions. The shape of the
20

Neuroanatomy
Hippocampal gyrus
Septum pellucidum Cingulate gyrus (Ammon's horn)
Indusium griseum
Temporal horn Parahippocampal gyrus
Corpus callosum lateral ventricle
PC
Fornix
Tail caudate
Subcallosal
area EC
CA2 CA1
CA3
Paraterminal
gyrus
Mammillary CA4 Subiculum
body
Anterior Fimbria Hippocampal sulcus
commissure Hippocampus
Amygdala Parahippocampal Choroid plexus
gyrus Dentate gyrus
Limbic gyrus Intralimbic gyrus Fornix and inner arc Fimbria of hippocampus
Figure 1.18 Hippocampal connections.
The fornix has two branches: the precommissural and in the entorhinal cortex, which generate a map of firing
the postcommissural pathways. The former connects to the patterns covering entire regions. The spatial firing
septal nuclei, preoptic nuclei, ventral striatum, orbital cor- sequences appear to be stored in the hippocampus during
tex, and anterior cingulate gyrus. The postcommissural exploration and can be retrieved at a later time. The
branch connects to the anterior nucleus of the thalamus dorsal hippocampus is responsible for spatial memory,
and mammillary bodies of the hypothalamus. In Korsakoff’s verbal memory, and learning conceptual information,
syndrome, these bodies are damaged and hence patients and there appear to be more place cells in the dorsal
have trouble learning new memories. The anterior thalamic region. The ventral hippocampus functions in fear con-
nuclei connect to the cingulate cortex, which connects back ditioning and affective processes.
to the entorhinal cortex and creates the Papez circuit, which
is involved in learning, memory, and emotion. 1.13.4.3.2 Explicit Versus Implicit Memory
Declarative or explicit memories are available in the con-
1.13.4.3 Hippocampus and Memory sciousness as semantic facts or episodic memories
Memory is the ability to acquire, store, and retrieve infor- (Figure 1.19). The areas of the brain involved are the
mation and are formed through learning. The process of hippocampus, the neocortex, and the amygdala. Non-
learning activates engram cells, a population of cells that declarative memories, also known as implicit memories,
have undergone cellular changes as a result of learning are memories of skills, and rely on the basal ganglia and
and when reactivated by the original stimulus leads to cerebellum. Short-term memories (seconds to minutes)
memory recall. are brief memories including working memory and rely
heavily on the prefrontal cortex. These memories can be
1.13.4.3.1 Spatial Memory consolidated into long-term memory.
The hippocampus is well known to be involved in spatial Formation of explicit memory begins with consolida-
memory and navigation. Within the hippocampus are tion and storage of encoded information in the
found place cells, which contain information regarding hippocampus. Over time, certain memories can be trans-
the spatial context in which a memory took place in one ferred to the neocortex as general knowledge. The amyg-
specific location. These cells cluster in place fields and fire dala interacts with the hippocampus and neocortex to
action potentials when an animal passes a certain loca- stabilize a memory and form new memories related to fear.
tion. Place-cell responses have been seen in the pyramidal One of the earlier findings of how the hippocampus
cells of the hippocampus and the granule cells of the impacts memory consolidation came from the study of
dentate gyrus. This is in contrast to the grid cells found Henry Molaison, a patient who had a bilateral temporal
21

F. Akter et al.
Long-Term Memory
Declarative Non-declarative
(explicit, conscious) (implicit, unconscious)
Priming (exposure
Episodic (e.g., to one stimulus
Semantic (e.g., Procedural (e.g., influences a Classical
first day at response to a
capital of Japan) riding a bike) conditioning
school) subsequent
stimulus)
Figure 1.19 Declarative versus non-declarative memory.
lobectomy to treat his epileptic seizures. The surgery led stimulates a cascade of short- and long-term changes. In
to deficits including an inability to complete tasks that the short term, there is insertion of AMPA receptors
required long-term episodic memory and an inability to (Figure 1.20), and in the long term there are changes in
create new long-term memories. His spatial orientation transcription factors and increased protein translation.
was also severely affected. His long-term memory of This forms the basis of LTP.
events many years before the accident was largely intact On the contrary, LTD is induced by prolonged low-
and his procedural memory and intelligence were unaf- frequency stimulation, which leads to prolonged calcium
fected. This showed that the medial temporal lobe con- activation of phosphatase enzymes and an eventual
taining the hippocampus is important for declarative but removal of AMPA receptors and pruning of spines.
not procedural memory consolidation.
Neurophysiological theories of synaptic plasticity 1.13.4.3.3 Implicit Memory
based on the cellular models that underpin synaptic plas- Implicit memory involves several different brain regions.
ticity including long-term potentiation (LTP) and long- Classical conditioning involves various sensory and motor
term depression (LTD) (see Chapter 5 for further details) systems. Operant conditioning involves the striatum and
can be used to explain the neural basis of learning and cerebellum and fear conditioning involves the amygdala.
memory. Donald Hebb was the first to postulate that Classic (Pavlov) conditioning refers to learning that
activation or inactivation of extant synaptic contacts occurs when a neutral stimulus becomes associated with
depends on the synchronous impulse activity of pre- a stimulus that naturally produces a behavior. It was first
and postsynaptic nerve cells. observed by the Russian physiologist Ivan Pavlov, who
One of the most fascinating features of the hippocam- exposed dogs to sounds (neutral stimulus) immediately
pus is its capacity for plasticity. Action potentials travel before receiving food (natural stimulus producing
down the Schaffer collaterals and stimulate the release behavior). Initially, the dogs began to salivate only
of glutamate into the synaptic cleft, which stimulates when they saw the food; however, later, the dogs learned
the α-amino-3-hydroxy-5-methyl-4-isoxazo-lepropionic to associate the sound with the food and began salivating
acid (AMPA) and N-methyl-D-aspartate (NMDA) recep- as they heard the sound. The food is the unconditioned
tors. Once the AMPA receptors open, sodium travels into stimulus as it naturally leads to a behavior. The condi-
the postsynaptic membrane and elicits a membrane tioned stimulus is the neutral tone, which upon repeated
depolarization. With a large enough stimulation, a large presentation leads to the same behavior as the uncondi-
amount of glutamate will be released, allowing a large tioned stimulus. Classic conditioning involves enhanced
membrane depolarization that removes the magnesium synaptic strength due to presynaptic facilitation, where
block typically seen at the NMDA receptor. This allows there is increased release of neurotransmitters from the
both sodium and calcium to enter the postsynaptic mem- presynaptic cells under the action of an additional
brane. The calcium activates protein kinases and neuron.
22

Neuroanatomy
Normal synaptic transmission Induction of long-term potentiation
Presynaptic
terminal
Glutamate
Na+ Mg2+ blocks Na+ Ca2+

Additional
receptor AMPA
receptor
AMPA NMDA
receptor receptor
Signaling
cascade
Postsynaptic
terminal
During low-frequency synaptic transmission, Mg2+ blocks High-frequency transmission expels Mg2+ from the
the NMDA receptor. NMDA receptor, allowing Na+ and Ca2+ influx. Ca2+ then
triggers a signaling cascade, increasing the number of
AMPA receptors at the synapse.
Figure 1.20 Long-term potentiation.
Habituation is a type of non-associative learning where Operant conditioning, is learning that occurs based
a repetitive stimulus leads to a decrement in the response on the consequences of behavior (e.g. reinforcement or
intensity. Sensitization is the increment in response inten- punishment). This is a voluntary behavioral response and
sity in response to a stimulus. Experiments performed in the according to the law of effect, the behavior is strength-
sea slug, Aplysia californica, by Eric Kandel were crucial in ened or weakened by the learner based on their desired
establishing the importance of synaptic changes to learning result. The early studies on operant conditioning were
and memory. It was demonstrated that upon stimulation of performed by Edward Thorndike and B. F. Skinner. The
the siphon receptors, the motor neuron is activated directly “Skinner box” was designed to study the principles of
or indirectly through the excitatory interneuron as the gill is animal behavior in a controlled environment.
withdrawn. However, with repeated stimulation, there is
reduced release of synaptic transmitters from the sensory 1.13.5 Amygdala
neurons and therefore there is less withdrawal of the
The amygdala is one of two almond-shaped clusters of
gill (habituation). This would typically occur when an ani-
nuclei found medially within the temporal lobe and is
mal repeatedly encounters a harmless stimulus. However,
responsible for emotions such as fear and aggression.
when the animal encounters a harmful stimulus, a vigorous
Emotion is a subjective state and consists of a physical
response occurs to not only the harmful stimulus but also
response involving the autonomic motor and endocrine
the harmless stimulus (sensitization). When the head or tail
systems. It also requires conscious registration, and this
of the Aplysia is stimulated by an electric shock, there is
involves both the cerebral cortex and the amygdala. The
activation of the facilitatory interneurons. These synapse
amygdala is also involved in memory formation, reward
with the sensory neurons of the head, and there is increased
processing, and decision making.
transmitter release leading to excitation of the excitatory
interneurons and the motor neurons to the gill 1.13.5.1 Inputs
(Figure 1.21). Dishabituation refers to the restoration of
The amygdala receives inputs from the hypothalamus,
a full-strength response that was weakened by habituation.
septal area, and the orbital cortex.
23

F. Akter et al.
Interneuron
Sensory neuron
(facilitatory)
Head
Motor
neuron
Interneuron
(excitatory)
Gill
Siphon Sensory neuron
Figure 1.21 Aplysia californica experiments performed to demonstrate habituation and sensitization.
1.13.5.2 Outputs a tone, with an aversive unconditional stimulus, such as an

The major outputs of the amygdala include the ventral electric foot shock, and responds by freezing even when it
amygdalofugal pathway, the stria terminalis, the hippocam- is not receiving the aversive stimulus. Fear extinction refers
pus, the entorhinal cortex, and the dorsomedial nucleus of to the reduction in the conditioned fear responses after
the thalamus. The amygdala connects to the hypothalamus repeated presentation of the conditioned stimulus without
via the stria terminalis and the ventral amygdalofugal path- the unconditioned stimulus that had elicited the fear.
way, the latter of which also connects to the anterior olfac-
tory nucleus, the anterior perforated substance in the 1.13.6 Nucleus Accumbens
forebrain, the piriform cortex, the orbitofrontal cortex, the The nucleus accumbens is part of the striatum. It receives
anterior cingulate cortex, and the ventral striatum. This input from the orbitofrontal cortex, the cingulate cortex,
pathway is particularly important in associative learning and the amygdala and is involved in emotion and
and the linking of behavior with reward and punishment. motivation.
The stria terminalis continues as precommissural and post-
commissural branches. The precommissural branch travels
to the septal area and the postcommissural branch to the
1.13.7 Prefrontal Cortex
hypothalamus. The stria terminalis also projects to the The prefrontal cortex is located in the frontal lobe and is
habenula (involved in reward and aversion processing), implicated in personality development, decision making,
which is part of the epithalamus. See section 1.15. and reasoning. Its role was discovered from the symp-
toms and signs that Phineas Gage (an American railroad
1.13.5.3 Amygdala and Fear Conditioning construction worker) developed following an accident
that involved a rod being driven into his brain resulting
The amygdala is involved in Pavlovian fear conditioning,
in irreversible damage to his frontal lobe.
a behavioral paradigm that involves learning to associate
The medial part of the prefrontal cortex connects with
stimuli with certain adverse effects. In fear conditioning,
the amygdala, the hippocampus, and the temporal lobe.
an animal associates a neutral conditional stimulus, such as
24

Neuroanatomy
Figure 1.22 Hypothalamic

nuclei.
Paraventricular nucleus Lateral hypothalamic area

Dorsomedial nucleus
Preoptic nucleus Posterior hypothalamic area
Anterior hypothalamic area
Suprachiasmatic nucleus
Supraoptic nucleus
Mammillary body
Arcuate nucleus
Ventromedial nucleus
The lateral part connects to the basal ganglia, the premo- regional groups of nuclei (Figure 1.22): the preoptic region
tor cortex, the supplemental motor area, the thalamus, (which contains the preoptic nucleus), the supraoptic
and the cingulate cortex. The orbitofrontal region forms region (which contains suprachiasmatic, supraoptic, para-
connections with the amygdala, the medial part of the ventricular, and anterior nuclei), the tuberal region (which
thalamus, the hypothalamus, and the basal ganglia. contains dorsomedial, ventromedial, arcuate, premammil-
lary and lateral tuberal nuclei), and the mammillary region
1.13.8 Olfactory Bulb (which contains mammillary and posterior nuclei). The
medial preoptic nucleus secretes gonadotropin-releasing
The olfactory nerve projects to the olfactory bulb in the
hormone (GnRH), which is responsible for the secretion
forebrain. The bulb is separated from the olfactory epi-
of luteinizing hormone (LH) and follicle-stimulating hor-
thelium by the cribriform plate. It contains neurons
mone (FSH) by the pituitary. The paraventricular and
involved in olfaction and receives sensory input from
supraoptic nuclei both produce the peptide hormones
axons of the olfactory receptor neurons in the olfactory
and antidiuretic hormone (ADH). The paraventricular
epithelium and outputs to the mitral cell axons. It then
nucleus also releases thyrotropin-releasing hormone
sends olfactory information to the amygdala, the orbito-
(TRH), corticotropin-releasing hormone (CRH), and
frontal cortex, and the hippocampus. It also receives
somatostatin. TRH is responsible for the formation and
information from the amygdala, the neocortex, the hip-
secretion of the thyroid-stimulating hormone (TSH) in the
pocampus, the locus coeruleus, and the substantia nigra.
pituitary gland, which in turn regulates the production of
The bulb is divided into the main and accessory parts.
thyroid hormones in the thyroid gland. TRH also stimu-
The main bulb connects to the amygdala via the piriform
lates the release of prolactin from the pituitary gland.
cortex of the primary olfactory cortex. Associative learn-
Corticotropin-releasing hormone activates the release of
ing between certain odors and behaviors takes place in the
adrenocorticotropic hormone (ACTH) from the pituitary
amygdala. The accessory bulb forms a parallel pathway.
gland. Somatostatin regulates the endocrine and digestive
systems. The anterior nucleus is involved in thermoregu-
1.13.9 Hypothalamus lation by stimulating the PNS, and causes hyperthermia if
The hypothalamus is bordered by the optic chiasm anteri- damaged. The suprachiasmatic nucleus regulates the cir-
orly and the mammillary bodies posteriorly. The function cadian rhythm and pineal gland function. The ventrome-
of the hypothalamus is to maintain homeostasis. It links dial nuclei mediate satiety. The lateral nuclei mediate
the nervous system to the endocrine system via the pitui- hunger via orexin neurons. The arcuate nuclei secrete
tary gland, which is located below it. It contains four prolactin, growth hormone releasing hormone (GHRH),
25

F. Akter et al.
and GnRH. Dorsomedial nuclei regulate blood pressure and clock maintained in a 24-hour pattern by clock genes
heart rate. The posterior nuclei release vasopressin and are such as Per, tim, and Cry. The retina sends inputs to the
involved in thermoregulation via the SNS. The mammillary SCN, which can regulate the circadian rhythm and sleep
nuclei are involved in memory. by releasing a number of hormones such as norepineph-
rine, and this can stimulate the pineal gland to release
1.13.9.1 Hypothalamus and Sleep melatonin. Activation of the ventrolateral preoptic
Sleep is a reversible state of reduced consciousness that is (VLPO) area is also important in initiating sleep.
important for regulation of inflammatory processes, Wakefulness is regulated by ascending arousal pathways
removal of toxins via the glymphatic system, maintenance activating the cortical system and uses chemicals such as
of a reduced metabolic rate and memory consolidation. norepinephrine, serotonin, dopamine, acetylcholine, his-
Sleep deprivation has negative consequences for the cardi- tamine, and orexin.
ovascular system, endocrine regulation, glucose tolerance
and mental health. Sleep can be measured using polysom- 1.14 Pituitary Gland
nography (PSG), which has been used to reveal distinct
The pituitary gland is functionally and anatomically
sleep states. Normally, people transition between non-
linked to the hypothalamus (via the infundibulum;
rapid eye movement (NREM) stages and rapid eye move-
Figure 1.23) and is responsible for releasing several hor-
ment (REM) stages and this is regulated by reciprocal
mones. It is located in the sella turcica of the sphenoid
inhibition of monoaminergic and cholinergic neurons.
bone and is covered by the diaphragma sellae. The gland
There is increased activity of cholinergic neurons and
is divided into the anterior (adenohypophysis) lobe –
decreased activity of adrenergic and serotonergic neurons
derived from an outpouching of the roof of the pharynx
during REM sleep and this is reversed in NREM sleep.
called Rathke’s pouch – and the posterior lobe. The ante-
NREM sleep occurs during the transition from being
rior lobe is divided into three parts: Pars anterior (hor-
awake to sleeping and is characterized by light sleeping
mone secretion), Pars intermedia, and Pars tuberalis. The
with slowing of brain waves. Rapid eye movement sleep is
posterior (neurohypophysis) lobe releases two hormones
characterized by rapid eye movements with faster breath-
(oxytocin and vasopressin) that are initially produced in
ing and increased heart rate and blood pressure.
the hypothalamus.
The sleep cycle is regulated by the circadian rhythm
The anterior pituitary is supplied by the superior
and is controlled by the suprachiasmatic nucleus (SCN)
hypophyseal artery (a branch of the internal carotid
of the hypothalamus. The circadian rhythm is a biological
Figure 1.23 The pituitary gland.

Hypothalamic
Third Ventricle
Supraoptic Nucleus of Brain
Hypothalamic
Neurons
Optic Chiasm
Superior
Hypophyseal
Artery Hypothalamic
Hypophyseal Tract
Hypophyseal
Portal Veins
Inferior
Hypophyseal
Anterior Artery
Pituitary
Secretory Cells of Posterior

Adenohypophysis Pituitary
Venule
26

Neuroanatomy
Figure 1.24 Cross-section of the midbrain at the level of the superior colliculus.
artery), which forms a capillary network and a plexus 1.16 Midbrain

called the hypophyseal portal system. The infundibulum
The midbrain (mesencephalon) is part of the brainstem
and posterior pituitary gland are supplied by the superior
and lies above the pons and below the forebrain
hypophyseal artery, the infundibular artery, and the infe-
(Figure 1.24). It is comprised of two parts: the tectum
rior hypophyseal artery. Both lobes are drained by the
and the cerebral peduncles (crus cerebra and tegmen-
anterior and posterior hypophyseal veins.
tum). The tectum houses four colliculi inferior to the
pineal gland and superior to the trochlear nerve. The
1.15 Epithalamus oculomotor nerve exits between the peduncles and the
The epithalamus is a dorsal segment of the diencephalon optic tract is found on the superior border.
and contains the pineal gland, habenula and stria medul- The cross-section of the midbrain reveals several fiber
laris. The pineal gland is responsible for secreting mela- tracts, and these include the frontopontine fibers, the
tonin, a hormone involved in the regulation of the corticospinal fibers, the corticobulbar tracts, and the tem-
circadian rhythm of the body. This gland is supplied by poropontine fibers. The substantia nigra, the tegmentum,
the posterior choroidal arteries (a branch of the posterior and the tectum can be visualized posteriorly. The cerebral
cerebral artery) and drains to the internal cerebral veins. aqueduct and the periaqueductal gray matter can be seen
The stria medullaris, contains afferent fibers from the in the midline, and the medial longitudinal fasciculus can
septal nuclei and the anterior thalamic nuclei to the be seen anteriorly. The red nuclei (which receive input
habenula, bilateral structures connected by a commis- from the cerebral cortex and cerebellum and are involved
sure. The lateral habenula is primarily involved in learn- in coordination of sensorimotor information) and decus-
ing from reward omission and aversive experiences, sation of the rubrospinal tracts (motor control, modula-
affect, cognition, and social behavior. The medial habe- tion of flexor muscle tone, reflex activity, and inhibition
nula may also be involved in fear responses, mood and of antigravity muscles) can be seen at the level of the
memory. superior colliculus. The oculomotor nucleus can be seen
27

F. Akter et al.
Thalamus
Pineal body
Anterior lobe
Hypothalamus
Tegmentum
Primary fissure
Midbrain Tectum Cerebellum
Pons
Medulla Posterior lobe
Figure 1.25 Sagittal section of the brain demonstrating the medulla and pons.
with the oculomotor nerve projecting anteriorly nuclei (responsible for coordination of movement), cor-
(Figure 1.24). The trochlear nerve can be seen at the ticospinal and corticobulbar tracts, and the dorsal pons
level of the inferior colliculi. (tegmentum), which forms part of the reticular formation
The blood supply to the midbrain is the basilar artery responsible for arousal and attentiveness and which con-
and its branches (the posterior cerebral artery, the supe- tains cranial nerves and the fourth ventricle.
rior cerebellar artery, the posterior choroidal artery, and
the interpeduncular branches). 1.17.2 Vascular Supply
The pons is supplied by the pontine artery (basilar
1.17 Hindbrain artery branch), the superior cerebellar artery, and the
anterior inferior cerebellar artery. The pons drains to
1.17.1 Pons the anterior pontomesencephalic vein, and then to the
The pons is found below the midbrain and above the basal and cerebral veins. The inferior aspects drain
medulla (Figure 1.25). It develops from the embryonic into the inferior petrosal sinus, which drains into the
metencephalon. The pons is the site of origin for several internal jugular veins.
cranial nerves (Figure 1.26).
The pons is connected to the cerebellum via the mid-
dle cerebellar peduncles. Underlying the cerebellum is the
1.18 Medulla Oblongata
fourth ventricle. The angle formed at the junction of the The medulla oblongata contains the ascending and des-
pons, the medulla, and the cerebellum is the cerebello- cending tracts and the brainstem nuclei. The inferior
pontine angle. The floor of the fourth ventricle reveals margin is marked by the origin of the first pair of cervical
some anatomical landmarks including the medial emi- spinal nerves as the medulla exits the skull through the
nence at the midline, the facial colliculus (containing the foramen magnum.
abducens nucleus and the facial motor fibers), and the Several structures can be visualized on the anterior sur-
stria medullaris (a part of the epithalamus). The pons is face. These include the pyramids, the olives, and five cranial
comprised of the ventral pons containing the pontine nerves (abducens nerve, accessory nerve, hypoglossal nerve,
28

Neuroanatomy
Figure 1.26 Cranial nerves emerging from the pons and medulla.
trochlear nerve, and trigeminal nerve). Posteriorly, the fas- be found laterally, with the trigeminal nucleus found
ciculus gracilis, the fasciculus cuneatus, and the posterior posterior to these tracts.
median sulcus can be seen. The decussation of the sensory pathways can be found
The internal structure of the medulla is usually best at the level of the decussation of the medial lemniscus.
appreciated in cross-section and is typically discussed at The medial lemniscus is a bundle of large myelinated
the level of the pyramidal decussation, the medial lem- axons that function as second-order neurons of the dorsal
nisci decussation (Figure 1.27), and the level of the olives. column–medial lemniscus pathway to transport sensory
At the level of the pyramidal decussation, we can see information. The medial lemniscus is formed by the
the descending motor fibers. The central portion contains crossings of the internal arcuate fibers, which are com-
the gray matter and the outer portion is made up of white posed of axons of nucleus gracilis and nucleus cuneatus.
matter, which contains the fasciculus gracilis and the Centrally the hypoglossal nucleus and laterally the medial
fasciculus cuneatus. Corresponding portions of gray mat- longitudinal fasciculus can be seen with the nucleus
ter extend to these regions and are the nucleus gracilis ambiguus. The medial longitudinal fasciculus controls
and nucleus cuneatus, respectively. The spinocerebellar horizontal eye movements by interconnecting oculomo-
tracts with the lateral spinothalamic tracts in between can tor and abducens nuclei. The nucleus ambiguus contains
29

F. Akter et al.
Hypoglossal nucleus
Dorsal motor nucleus of CN X
Medial vestibular nucleus Cochlear nucleus
Solitary nucleus
Inferior cerebellar
Medial longitudinal fasciculus peduncle
Tectospinal tract Spinal nucleus and

tract of CN V
Lateral spinothalamic tract
Inferior olivary nucleus
Medial lemniscus Pyramid
Figure 1.27 Cross-section the medulla at the level of the medial meniscus decussation.
cells bodies of motor nerves involved in swallowing and innervate muscles to produce movement, discussed in
speaking. The inferior olivary nucleus (ION) is found Chapter 7.
between the nucleus ambiguus and the pyramids and
coordinates signals from the spinal cord to the cerebel- 1.19 Cranial Nerves
lum. The climbing fiber axons leave the ION and travel to
There are 12 paired cranial nerves that arise from the
the cerebellar Purkinje cell.
brain. The first two, the olfactory and optic nerves, origi-
At the level of the olives, the central canal opens into
nate in the cerebrum and the rest arise in the brainstem.
the fourth ventricle, and at this level the inferior olivary
nucleus and inferior cerebellar peduncles can be seen.
The vestibular nuclei can be seen in the midline. 1.19.1 Olfactory Nerve
Laterally, the nucleus of the tractus solitarius (which This is the first and the shortest nerve. It is derived from
generates peristaltic activity of the gastrointestinal system the olfactory placode, a thickening of the neural ecto-
during swallowing) can be seen. derm, which give rise to the olfactory epithelium of the
nose containing the olfactory receptor neurons. These are
1.18.1 Vascular Supply bipolar cells that gives rise to unmyelinated axons, which
are found in bundles and penetrate the cribriform plate of
The vessels that supply the medulla include: the anterior
the ethmoid bone. They then enter the cranium and then
spinal artery, the posterior spinal artery, the posterior
the olfactory bulb to synapse with neurons called mitral
inferior cerebellar artery, the anterior inferior cerebellar
cells, forming the synaptic glomeruli. From here, second-
artery, and the vertebral artery.
order neurons pass to the olfactory tract, which travels to
the optic chiasm and divides into the two stria. The lateral
1.18.2 Pathways stria carries axons to the primary olfactory cortex in the
The ascending tracts refer to the neural pathways by uncus of the temporal lobe and the medial stria to the
which sensory information from the peripheral nerves is anterior commissure, where they meet the olfactory bulb
transmitted to the cerebral cortex. In some texts, ascend- of the opposite side. The primary olfactory cortex sends
ing tracts are also known as somatosensory pathways. fibers to the piriform cortex, the amygdala, the olfactory
The descending tracts are the pathways by which tubercle, and the secondary olfactory cortex.
motor signals are sent from the brain to the lower The olfactory mucosa is found on the roof of the nasal
motor neurons. The lower motor neurons then directly cavity and is made up of pseudostratified columnar
30

Neuroanatomy
epithelium containing basal cells (which form new stem rotation). The Edinger–Westphal nucleus is dorsal to
cells), sustentacular cells (for structural support, analo- the oculomotor nuclei and contains preganglionic para-
gous to glial cells), olfactory receptor cells (bipolar cells sympathetic neurons to the ciliary ganglion. From here,
consisting of dendrite processes with cilia that react to postganglionic parasympathetic fibers supply the ciliary
odors and stimulate olfactory cells), and a central process muscles for pupil constriction and the sphincter pupillae
projecting in the opposite direction through the base- for accommodation.
ment membrane. The mucosa also contains Bowman’s Damage to the oculomotor nerves leads to ptosis,
glands, which secrete mucus. a down-and-out position of the eye and a dilated pupil.
This can occur due to raised intracranial pressure, aneu-
1.19.2 Optic Nerve rysm of the posterior communicating artery, or damage
to the cavernous sinus. It can also be found in other
The optic nerve develops from the optic vesicle. It is
diseases such as multiple sclerosis and myasthenia
a part of the central nervous system (CNS) and is cov-
gravis.
ered by meninges. It is formed by the convergence of
axons from the retinal ganglion cells, which in turn
receive information from bipolar cells and the photo- 1.19.4 Trochlear Nerve
receptors of the eye. Each optic nerve leaves its respec- The trochlear nerve originates from the trochlear nuclei
tive orbit via the optic canal, enters the middle cranial in the midbrain at the level of the inferior colliculus. It has
fossa, and unite to form the optic chiasm. Here, fibers the longest intracranial route because it is the only nerve
from the nasal medial half of each retina cross to the to emerge from the dorsal brainstem, making it particu-
contralateral optic tract; however, lateral fibers remain larly vulnerable to damage. It travels within the cavernous
ipsilateral. Therefore, the right optic tract, for example, sinus and passes through the superior orbital fissure to
would contain fibers from the right temporal (lateral) innervate the superior oblique muscle, which functions to
retina but the left nasal retina. The optic tracts then depress, abduct, and intort the eye. Damage to this nerve
reach the lateral geniculate nucleus (LGN) in the leads to double vision and a characteristic head tilt toward
thalamus, which carries visual information in the optic the unaffected side.
radiation. The upper radiation carries fibers from the
superior retinal quadrants (corresponding to the infe- 1.19.5 Trigeminal Nerve
rior visual field quadrants) through the parietal lobe to
The trigeminal nerve provides sensory and motor inner-
reach the visual cortex. The lower radiation carries fibers
vation to the face. It originates from three sensory nuclei
from the inferior retinal quadrants (corresponding to
(the mesencephalic, principal sensory, and spinal nuclei)
the superior visual field quadrants), through the tempo-
and one motor nucleus extending from the midbrain to
ral lobe, via Meyer’s loop, to reach the visual cortex for
the medulla. It has three branches, the ophthalmic, the
processing of visual information.
maxillary, and the mandibular nerves, which arise from
the trigeminal ganglion to provide sensory innervation to
1.19.3 Oculomotor Nerve the face. The mandibular branch also supplies the mus-
The oculomotor nerve originates from the oculomotor cles of mastication. Clinically, the corneal reflex can be
nucleus at the level of the superior colliculus in the mid- performed to test damage to the ophthalmic nerve (which
brain of the brainstem. It travels through the dura mater acts as the afferent limb to detect the stimulus) or the
and enters the cavernous sinus. It leaves the cranium via facial nerve (which is the efferent limb causing contract of
the superior orbital fissure and divides into the superior the orbicularis oculi muscle).
branch to supply the superior rectus (which elevates the
eyeball) and the levator palpabrae superioris (which 1.19.6 Abducens Nerve
raises the upper eyelid). It travels with sympathetic fibers The abducens nerve is the sixth paired cranial nerve, with
that innervate the superior tarsal muscle (which helps to a somatic motor function to the lateral rectus muscle. It
raise the eyelid). The inferior branch supplies the remain- arises from the abducens nucleus in the pons, exiting the
ing extra ocular muscles (the inferior rectus depressing brainstem at the junction of the pons and the medulla. It
the eyeball, the medial rectus for adduction, and the then enters the subarachnoid space and pierces the dura
inferior oblique for elevation, abduction, and lateral mater, and enters the cavernous sinus through the
31

F. Akter et al.
superior orbital fissure. It can be damaged by any space- reflex, which allows the stabilization of images on the
occupying lesion, which leads to diplopia and unopposed retina while the head is moving.
adduction. Inflammation of the vestibular branch of the nerve
can lead to vertigo, nystagmus, loss of equilibrium, and
1.19.7 The Facial Nerve nausea/vomiting. Inflammation of the membranous lab-
yrinth (labyrinthitis) can cause similar symptoms and in
The facial nerve is derived from the second branchial
addition affect the cochlear nerve, leading to tinnitus.
arch. The upper motor neuron of the facial nerve is in
the primary motor cortex, with its axons descending to
the ventral and dorsal facial nucleus in the pons. The 1.19.9 Glossopharyngeal Nerve
dorsal regions supply the muscles of the upper face and The glossopharyngeal nerve begins in the medulla and
receive input from both hemispheres, and the ventral leaves the cranium via the jugular foramen. It has mixed
region supplies the muscles of the lower face and receives sensory and parasympathetic components. The sensory
mainly contralateral inputs. The facial nerve passes branch innervates the oropharynx via the pharyngeal
through the internal auditory meatus and then on to the branch, which merges with the vagus nerve to form the
facial canal to synapse on the geniculate ganglion. It gives pharyngeal plexus. It forms the afferent limb of the gag
rise to the greater petrosal nerve, supplying the lacrimal reflex (the efferent limb is via the vagus nerve). It supplies
gland, and joins the deep petrosal nerve to form the nerve the posterior one third of the tongue via the lingual
of the pterygoid canal, innervating the pterygopalatine branch and the palatine tonsils via the tonsillar plexus.
ganglion. It gives rise to the nerve to the stapedius muscle It also supplies the carotid body and sinus. The tympanic
of the ear and the chorda tympani nerve, which supplies branch of the nerve enters the middle ear and forms the
the submandibular ganglion and the anterior two thirds tympanic plexus to supply the middle ear, tympanic
of the tongue. membrane, and the eustachian tube. It supplies the sty-
The facial nerve leaves the cranium through the sty- lopharyngeus muscle of the pharynx and provides para-
lomastoid foramen and supplies the external ear, the sympathetic innervation to the parotid gland.
external auditory meatus, the posterior belly of the digas-
tric, the stylohyoid, the superior and inferior auricular, 1.19.10 Vagus Nerve
and the occipitalis muscles. It then travels to the parotid
The vagus nerve extends from the medulla and exits the
gland, and without supplying it splits into five branches
cranium, where it gives rise to an auricular branch and
(temporal, zygomatic, buccal, marginal mandibular, and
courses downward into the carotid sheath. At the base of
cervical) to supply the muscles of facial expression.
the neck, the right vagus nerve enters the thorax anterior to
the subclavian artery and the left vagus nerve passes
1.19.8 Vestibulocochlear Nerve between the left common carotid and subclavian arteries.
The vestibulocochlear nerve has two main sensory divisions: In the neck it gives rise to pharyngeal branches (which
the vestibular and cochlear nerves, which arise from the supply the pharynx and soft palate), the external superior
vestibular nuclei in the pons and medulla, and the cochlear laryngeal nerve (which supplies the cricothyroid muscle),
nuclei in the inferior cerebellar peduncle, respectively. They and the internal laryngeal branch (which supplies the
combine at the pons and emerge at the cerebellopontine laryngopharynx and part of the larynx). The right vagus
angle, exiting the cranium via the internal acoustic meatus. nerve also gives rise to the recurrent laryngeal nerve (which
The vestibular nerve is responsible for equilibrium and the supplies the intrinsic muscles of the larynx) and then forms
cochlear nerve is responsible for hearing. the posterior vagal trunk in the thorax. The left vagus nerve
The cochlea contains inner hair cells that respond to forms the anterior vagal trunk. These contribute to the
vibrations of sound and trigger action potentials from the formation of the esophageal plexus innervating the smooth
spiral ganglia. Sound frequency is coded by the position muscle of the esophagus. Cardiac branches to the heart
of the activated inner hair cells. arise in the thorax. The left vagus nerve gives rise to the left
The vestibular hair cells are found in the otoliths (the recurrent laryngeal nerve under the arch of aorta. The
saccule and the utricule), where they detect linear motion vagal trunks enter the abdomen via the esophageal hiatus
of the head, and the semicircular canals (which detect in the diaphragm and then terminate into branches to
rotational movement of the head), and coordinate bal- supply the esophagus, the stomach, and the small and
ance. They are also important for the vestibulo-ocular large intestines up to the splenic flexure.
32

Neuroanatomy
1.19.11 Accessory Nerve found on the cerebellopontine angle and is important for
vestibular function. On each side, close to the medulla, are
The accessory nerve has a somatic motor function and
tonsils, and clinically these are at risk of coning into the
supplies the sternocleidomastoid and trapezius muscles.
foramen magnum when CSF is withdrawn in patients with
It contains spinal and cranial components. The spinal
increased intracranial pressure, resulting in pressure on the
components arise from C1 to C6 spinal nerve roots,
vital respiratory and autonomic centers of the medulla.
which merge and enter the cranium via the foramen mag-
There are three main lobes of the cerebellum. The
num. After exiting the cranium, it descends to the sterno-
anterior lobe extends from the cerebellar peduncle and
cleidomastoid muscle and then to the posterior neck to
terminates at the primary fissure and continues as the
supply the trapezius. The cranial component arises in the
posterior lobe. The smallest lobe is the flocculonodular
medulla and exits the cranium via the jugular foramen. It
lobe and this lies between the posterolateral fissure (infe-
combines with the vagus nerve at the inferior ganglion.
riorly) and the cerebellar peduncles (superiorly).
1.19.12 Hypoglossal Nerve

The hypoglossal nerve supplies the extrinsic (genioglos- 1.20.1 Functional Divisions
sus, hyoglossus, styloglossus) and intrinsic muscles of the The cerebellum can be divided into three functional
tongue. The palatoglossus muscle is innervated by the areas: the cerebrocerebellum, the spinocerebellum, and
vagus nerve. The hypoglossal nerve arises from the hypo- the vestibulocerebellum (Figure 1.28).
glossal nucleus in the medulla and exits the cranium via The large division is the cerebrocerebellum, which is
the hypoglossal canal. formed by the lateral hemispheres and is responsible for
planning movements and motor learning. It also regu-
1.20 Cerebellum lates muscle activation and visually guided movements. It
receives inputs from the cerebral cortex and pontine
The cerebellum is derived from the rhombencephalon
nuclei. The fibers travel through the internal capsule
and is found in the posterior cranial fossa. It is a large
and cerebral peduncles and terminate ipsilaterally in the
structure of the hindbrain with a protruding central ver-
pons and send their axons to the contralateral middle
mis, which sits between the cerebellar hemispheres.
cerebellar peduncle. The cerebellum also receives infor-
On the ventral aspect of the brain, the hemispheres are
mation via the ascending spinocerebellar mossy fibers.
connected to the pons by the middle cerebellar peduncle. On
Fibers from the inferior olive in the medulla form the
the ventral aspect of the cerebellum is the flocculus, which is
climbing fibre input via the inferior cerebellar peduncle.
Spinocerebellum Figure 1.28 Functional divisions

Lateral Intermediate (Motor execution) of the cerebellum.
hemisphere hemisphere Vermis
Cerebrocerebellum
(Motor planning)
Dentate nuclei
Fastigial nuclei
Interposed nuclei
Vestibulocerebellum
(Equilibrium & eyeball movements)
Functional divisions of human cerebellum
33

F. Akter et al.
Figure 1.29 Deep cerebellar nuclei.
They are thought to mediate plasticity in the mossy fiber– 1.20.3 The Cerebellar Peduncles
granule cell–Purkinje cell pathway to coordinate move-
There are three peduncles that connect the cerebellum
ment. Outputs from the dentate nucleus and the other
with the brainstem on each side. The middle peduncle is
deep cerebellar nuclei leave the superior cerebellar
the largest and the superior peduncle is the major output
peduncle and then cross the midline at the lower mid-
pathway from the deep nuclei.
brain, pass through the red nucleus and ascend to termi-
nate in the thalamus. The cerebellum represents
ipsilateral movement and sensation. 1.20.4 Cerebellar Cortex
The spinocerebellum comprises the vermis and inter- The cerebellar cortex contains many tightly packed folds
mediate zone of the cerebellar hemispheres. It receives (folia). It has three well-defined layers: (i) an outer molec-
proprioceptive information and regulates body move- ular layer consisting mainly of parallel fibers that run
ments by allowing for error correction. parallel to the folia and dendrites; (ii) the Purkinje cell
The vestibulocerebellum is the functional equivalent layer containing large Purkinje cells; and (iii) a granule
to the flocculonodular lobe. It controls balance and ocular layer consisting of many small granule cells (Figure 1.30).
reflexes, and receives inputs from the vestibular system The Purkinje cells are the output cells of the cerebellar
and sends outputs back to the vestibular nuclei. cortex. They have extensive dendrites and are intersected
at right angles by the parallel fibers, allowing each
1.20.2 Deep Cerebellar Nuclei Purkinje cell to receive and make synaptic contact with
many parallel fibers along a single folium. The Purkinje
The cortical output of the cerebellum, which is typically
cells send axons out of the cortex and inhibit cells in the
inhibitory, involves the deep cerebellar nuclei (Figures 1.28
deep cerebellar nuclei.
and 1.29). There are three nuclei on each side: the dentate
nucleus, the nucleus interpositus, and the fastigial nucleus.
Some areas, such as the flocculus and the flocculonodular 1.20.5 Vascular Supply
lobe, send outputs to the vestibular nuclei in the medulla. The cerebellum is supplied by the superior cerebellar,
Together, the entire cerebellar output is transmitted to the anterior inferior cerebellar, and posterior inferior cere-
brainstem descending motor pathways and the motor bellar arteries. The venous supply to the cerebellum
areas of the cerebral cortex via the thalamus. includes the superior cerebellar veins draining to the
34

Neuroanatomy
Figure 1.30 Layers of the cerebellum.
Molecular layer
Purkinje cell layer
Granule layer
White matter
straight sinus and internal cerebral veins and inferior CSF. The pia mater surrounds the spinal cord, nerve roots,
cerebellar veins draining to the transverse sinus, superior and blood vessels and inferiorly fuses with the filum ter-
petrosal sinus, and occipital sinus. minale. The pia mater thickens to form denticulate liga-
ments which attach to the dura mater, and these help
1.21 Spinal Cord suspend the spinal cord in the vertebral canal.
The spinal cord is an organized bundle of nervous tissue 1.21.1 Neurovascular Supply
extending from the medulla oblongata through the verte-
The arterial supply to the spinal cord is via the anterior
bral canal to the L2 vertebral level and terminates as the
spinal artery (a branch of the vertebral artery) and the two
conus medullaris. Spinal nerves arise from the end of the posterior spinal arteries (branches of the vertebral artery
spinal cord and together are known as the cauda equina.
or the posteroinferior cerebellar artery), which anasto-
Compression of these leads to the neurosurgical emer-
mose in the pia mater. The segmental medullary arteries,
gency known as cauda equina syndrome and can lead to
of which the largest is the anterior segmental medullary
paralysis. The spinal cord enlarges at the level of C4–T1
artery (the artery of Adamkiewicz), also supply the spinal
(cervical enlargement) and is the site of origin of the
cord. Disruption of the blood supply to the spinal cord
brachial plexus. A second enlargement is at the level of
leads to nerve cell death and signs of weakness, paralysis,
T11–L1 (lumbar enlargement), and is where the lumbar
and loss of reflexes.
and sacral plexi originate. The spinal cord venous drainage includes an extrinsic
The spinal cord is surrounded by the spinal meninges
and an intrinsic network. The extrinsic system includes
(dura, arachnoid, and pia mater) and contains CSF. It is
radicular veins, the pial venous network, dorsal and ventral
anchored distally to the coccyx by a fibrous band of tissue
spinal veins. They are connected to the valveless vertebral
called the filum terminale. The dura mater extends from
venous plexuses, that communicate with the dural venous
the foramen magnum to the filum terminale and is sepa-
sinuses and are composed of an external plexus surround-
rated from the vertebral canal by the epidural space con-
ing the vertebral column, internal plexus within the spinal
taining the internal vertebral venous plexus. The dura
canal (linked together via segmental veins) and basiverteb-
mater also surrounds the epineurium of spinal nerves ral veins in the vertebral bodies. The intrinsic venous
that pierce the dura. The arachnoid mater is separated
system includes the sulcal (drains to the anterior spinal
from the pia mater by the subarachnoid space and contains
veins) and radial veins (drains to the posterior spinal
35

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The Project Gutenberg eBook of Radio mates
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and most other parts of the world at no cost and with almost no
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under the terms of the Project Gutenberg License included with this
ebook or online at www.gutenberg.org. If you are not located in the
United States, you will have to check the laws of the country where
you are located before using this eBook.
Title: Radio mates
Author: Benjamin Witwer
Illustrator: Frank R. Paul
Release date: December 11, 2023 [eBook #72380]
Language: English
Original publication: New York: E. P. Co., Inc, 1927
Credits: Roger Frank and Sue Clark
*** START OF THE PROJECT GUTENBERG EBOOK RADIO

MATES ***
Radio Mates
“Think of it! A twist of a switch and the living, breathing piglet slowly
dissolved before my eyes and vanished along a pair of wires to my
aerial, whence it was transferred as a set of waves in the ether to the
receiving apparatus—there to reincarnate into the living organism
once more, alive and breathing, unharmed by its extraordinary
journey!”
RADIO MATES
by Benjamin Witwer
From the telegraph to the telephone was but a step. From the telephone to
radio constituted but another such step, and we are now enjoying radio
broadcast from stations thousands of miles away. Every time you have an
X-ray photograph taken you are bombarded, not by rays, but by actual
particles that go right through the walls of the tube, which particles are just
as real as if they were bullets or bricks, the only difference being that they
are smaller. Thus our scientists lead up to the way of sending solids
through space. While impossible of achievement, as yet, it may be
possible, years hence, to send living beings through space, to be received
at distant points. At any rate, the author of this story weaves a fascinating
romance around this idea. It makes excellent reading, and the plot is as
unusual as is its entire treatment.
It was a large brown envelope, of the size commonly used for

mailing pamphlets or catalogues. Yet it was registered, and had
come by special messenger that afternoon, my landlady informed
me. Probably it was a strain of that detective instinct which is present
in most of us that delayed my opening the missive until I had
carefully scrutinized the handwriting of the superscription. There was
something vaguely familiar in its slanting exactitude, yet when I
deciphered the postmark,—“Eastport, N. Y.,”—I was still in the dark,
for I could not remember ever having heard of the place before. As I
turned the packet over, however, my pleasant tingle of anticipation
was rudely chilled. Along the flap was a sinister row of black sealing
wax blobs, which seemed to stare at me with a malignant fore-
knowledge. On closer examination, I noticed that each seal retained
the impression of a coat-of-arms, also elusively familiar.
With a strange sense of foreboding I dropped the missive on the
table. Queer what ominous significance a few drops of wax can
impart to an ordinary envelope. Deliberately I changed into smoking
jacket and slippers, poked the well laid fire and lit a pipe before
finally tearing open the seals.
There were many typewritten sheets, commencing in letter form:
54 Westervelt Ave.,
Eastport, New York,
February 15th.
Dear Cousin George:
Now that you have identified me by referring to my signature on
the last page (which I had just done) you will no doubt wonder at the
occasion for this rather effusive letter from one so long silent as I
have been. The fact of the matter is that you are the only male
relative with whom I can communicate at this time. My nephew,
Ralph, is first officer of a freighter somewhere in the Caribbean, and
Alfred Hutton, your mother’s first cousin, has not been heard from
since he embarked on that colonizing scheme in New Guinea, nearly
a year ago.
I must do all in my power to prevent the bungling metropolitan
police from implicating Howard Marsden in my disappearance. It
would take no great stretch of the imagination to do just that, and
were the State to require Marsden’s life as forfeit for my own, then
my carefully planned revenge would be utterly frustrated. I have
been cultivating the village postmaster for some weeks, ever since
this plan began to shape definitely in my mind. I am mailing this letter
at three o’clock this afternoon, for I have noticed that at that hour the
postal section of the store is generally deserted. I shall ask him if his
clock is correct, thus fixing the time in his mind. Please remember
these points. Then I shall register this letter, taking care to exhibit the
unusual collection of seals on the back. I shall manage to inform him
also that I stamped the seals with my ring and will show him the
coat-of-arms, explaining its meaning in detail. These villagers are a
curiosity-ridden lot. Upon returning home, I shall drop this same ring
into the inkwell which stands upon my desk. Finally I shall proffer my
friend the postmaster a fifty-dollar bill in paying for my registry. The
registry slip itself will be found within the hatband of my brown hat,
which I shall place in the wall safe of my study.
You are becoming more amazed as you proceed, no doubt asking
yourself if this letter is the product of a madman or a faker. Before
you have finished you will probably be assured that both
assumptions are correct. It matters little, for I will at least have firmly
established the fact that this letter was mailed by no one else but
me. As for the rest, Howard Marsden will corroborate what follows.
To begin at the beginning. As you know, or perhaps you do not
know, for I forget that our correspondence has been negligible of
late, five years ago I accompanied the Rodgers expedition into
Afghanistan. We were officially booked as a geological mission, but
were actually in search of radium, among other things. When I left, I
was practically engaged to Venice Potter, a distant relation of the
Long Island Potters, of whom you have perhaps heard. I say
“practically” engaged because the outcome of this expedition was to
furnish me with the standing and position necessary for a formal
demand for her hand. As I said, that was nearly five years ago.
Four months after my departure her letters ceased coming and
mine were returned to me unopened. Two months later I received an
announcement of her betrothal to Howard Marsden. Received it out
there in Afghanistan, when I had returned to the coast for supplies.
We’ll skip that next year, during which I stuck with the expedition. We
were successful. I returned.
Then I found out where the Marsdens were living, here in
Eastport. I’d met Marsden once or twice in the old days, but paid him
little attention at the time. He seemed but another of the moneyed
idlers; had a comfortable income from his father’s estate and was
interested in “gentleman farming,”—blooded stock and the rest. I
decided that it was useless to dig into dead ashes for the time being,
at least until I could determine the lay of the land, so to speak.
Meanwhile I had my researches to make, a theory I had evolved as a
sort of backfire to fill that awful void of Venice’s loss,—out there on
the edge of the world. Countless sleepless nights I had spent in a
feverish attempt to lose myself in scientific speculation. At last I
believed I had struck a clue to conclusions until now entirely
overlooked by eager searchers. I decided to establish my laboratory
here in Eastport, perhaps devoting any leisure hours to an
unravelling of that mystery of my sudden jilting. With a two-year-old
beard and sunbaked complexion there were few who would have
recognized me under my real name, and none in my assumed role of
“Professor Walters.”
Thus it was that I leased an old house not half a mile from Marsden’s
pretentious “farm.” I converted the entire ground floor into a
laboratory, living in solitary state upon the upper floor. I was used to
caring for myself, and the nature of my experiment being of such
potentialities, I felt that I wanted no prying servants about me.
Indeed, it has turned out to be of such international importance that I
feel no compunction whatever in utilizing it for my own selfish ends.
It could be a boon to humanity, yet its possibilities for evil in the
hands of any individual or group is so great as to render it most
dangerous to the happiness of the human kind on this small globe.
One day, some three months after I had taken up my residence in
Eastport, I had a visitor. It was Marsden. He had been attracted by
the sight of my novel aerial, just completed. By his own admission he
was an ardent “radio fan,” as they are popularly termed, I believe,
and he spent the better part of an afternoon bragging of stations he
had “logged” with his latest model radio set. Aside from my vague
suspicions of his complicity in the alienation of my beloved Venice, I
must admit that even then I felt an indefinable repulsion towards him.
There was something intangibly unwholesome about him, a
narrowness between the eyes which repelled me. Yet, although at
that time I had no plan in mind, nevertheless I encouraged him in my
most hospitable manner, for even thus early I felt, that at some time
not far distant, I might be called upon to utilize this acquaintanceship
to my own advantage.
This first visit was followed by others, and we discussed radio in
all its phases, for the man had more than a smattering of technical
knowledge on the subject and was eager to learn more. At last, one
day, I yielded to his insistence that I inspect his set and agreed to
dine at his house the following evening. By now I felt secure in my
disguise, and although I dreaded the moment when I should actually
confront my lost love once more, yet I longed for the sweet pain of it
with an intensity which a hard-shelled bachelor like you will never
understand. Enough. I arrived at the Marsden’s the next evening and
was duly presented to my hostess as “Thomas Walters.” In spite of
my private rehearsals I felt a wave of giddiness sweep over me as I
clasped that small white hand in my own after the lapse of almost
five years, for she was, if possible, lovelier than ever. I noted when
my vision cleared that her eyes had widened as they met mine. I
realized that my perturbation had been more apparent than I
imagined and managed to mutter something about my alleged “weak
heart,” a grimmer jest by far than I intended. Frantically I fortified
myself with remembrances of those barren days in Afghanistan,
where I stayed on and on, impotent to raise a hand in the salvage of
my heart’s wreckage.
We chatted politely all through that interminable meal, no morsel
of which aroused the faintest appreciation on my dry tongue. Finally
the chairs were pushed back and my host excused himself to bring
down some pieces of apparatus he had recently purchased,
concerning which he professed to desire my invaluable opinion.
No sooner had he left the room than the polite smile dropped from
Venice’s face like a discarded mask.
“Dick,” she cried, “what are you doing here?”
It was my first inkling that she suspected my true identity. I rallied
quickly, however, and allowed my self-encouraged bitterness its
outlet.
“Had I believed you would recognize me, Mrs. Marsden, I should
not have inflicted my unwelcome presence upon you, I can assure
you.”
She bit her lips and her head raised with a jerk. Then her mouth
softened again as her great eyes searched mine.
“Yes, but why—” she broke off at the sound of approaching
footsteps. Suddenly she leaned forward. “Meet me in the pine grove
to-morrow afternoon—four o’clock,” she breathed. Then her husband
entered.
The remainder of the evening I was forced to listen to Marsden’s
eager dissertation on the alleged “static eliminator” which had been
foisted upon him on his last trip to the city. Mechanically I answered
or grunted in simulated appreciation when a pause in his endless
monologue warned me that some reply was expected of me; but my
pulses were leaping in exultation because of the fleeting hope which
those few words from my lost Venice had kindled. I could not
imagine why the offer to bridge the breach of years should come
from her so voluntarily, yet it was enough for me that she
remembered and wished to see me. I cared not why.
I arrived nearly an hour early that next afternoon, for I had been
unable either to sleep or work during the interim. I shall not bore you
with the particulars of that meeting, even were I free to reveal such
sacred details. Suffice to say that after the preliminaries of doubt and
misunderstanding had been brushed away—and it was not the
simple process this synopsis would seem to infer, I can assure you—
I stood revealed as the victim of a most ingenious and thoroughly
knavish plot. Boiled down, it resembles one of those early movie
scenarios.
You remember I spoke of Venice as related to the Long Island
Potters, a branch of the family highly rated in the Social Register?
You will also remember that before I undertook that expedition I was
never particularly certain whence my next year’s expenses were to
be derived, nor to what extent, if you understand what I mean. At
about the time I was preparing for this expedition which I hoped
would make me financially and scientifically independent, this
wealthy branch of the family seriously “took up” my darling Venice,
inviting her to live with them that summer. I remember now all too
late, that even during that confusion of mind caused by the agony of
leaving my loved one, coupled with the feverish preparations for
departure, chill clouds of censure came from the aloof Potters. They
made no effort to mask their disapproval of my humble self and
prospects, yet in my blindness I had never connected them
intimately with what followed.
It was, in short, the old story of the ingenious man-on-the-ground,
the “good match,” aided and abetted by the patronesses of the “poor
relation.” The discriminating Marsden naturally fell in love with
Venice, and to his great surprise and chagrin, was decisively
repulsed by her. Never before having been refused anything he
really wanted in his comfortably arranged life, he became
passionately desirous of possessing her. Accordingly, my darling
was shown a letter, forged with such diabolical cleverness as to be
almost indistinguishable from my own hand. It purported to intrigue
me with a very ordinary female at a period coincident with the time I
had been so fervently courting my dear one.
She refused to credit the document and dispatched me a
voluminous explanation of the whole occurrence. Attributing my
silence to the exigencies of distance, she continued to write me for
over a month. When no answer arrived after nearly three long
months, she at length delivered a hastily planned ultimatum, to which
she was later persuaded to adhere through the combined pressure
of Marsden and her family, beating against the razed defences of her
broken heart. Then it was that I received the betrothal
announcement, the only communication her watchful family had
permitted to escape their net of espionage.
As the story unfolded, my heart pounded with alternate waves of

exaltation and red rage at the treacherous Marsden. Because of
selfish duplicity, he had robbed us both of five years’ happiness, for I
had forced my darling’s admission that she had never loved him, and
now despised him as a common thief. My brief moment of delirious
joy was sharply curtailed, however, when I came to press her to
separate from this selfish swine. After some demur she confided that
he was a drug addict. She said that he had been fighting desperately
to break this habit ever since their marriage, for his jealous love of
her was the only remaining weapon with which to combat his deep
rooted vice. Deprived of his one motive, my darling earnestly
assured me that it would be a matter of but a few short years before
the white powders wrote Finis to yet another life. I could see but a
balancing of an already overdrawn account in such an event, and
said so in no uncertain terms. She did not chide me, merely patiently
explained with sweet, sad resignation that she held herself
responsible for his very life for the present. That although she could
not love and honor him as she had promised, yet she was bound to
cleave to him during this, his “worse” hour. And so we left it for the
time, our future clouded, yet with no locked door to bar the present
from us.
We met almost daily, unless Marsden’s activities interfered. At
those times I was like a raging beast, unable to work, consumed with
a livid hatred for the cunning thief who had stolen my love while my
back was turned. I could not shake her resolution to terminate this
loveless match, even though she now loathed the mate she had
once tolerated. But in spite of the formlessness of our future, my
work progressed as never before. Now my days were more than a
mere procession of dates, for each was crowned with the glow of
those few stolen moments with my darling Venice.
Came the day of my first complete success. Some weeks
previously I had finally succeeded in transmitting a small wooden ball
by radio. Perhaps I should say that I had “dissolved” it into its
vibrations, for it was not until this later day that I had been able to
materialize or “receive” it after it had been “sent.” I see you start and
re-read this last sentence. I mean just what I say, and Marsden will
bear me out, for as you shall see, he has witnessed this and other
such experiments here in my laboratory. I have explained to him as
much as I wanted him to know of the process, in fact, just enough so
that he believes that a little intensive research and experimentation
on his part will make him master of my secret. But he is entirely
ignorant of the most important element, as well as of the manner of
its employment.
Yes, after years of study and interrupted experimental research I
was enabled finally to disintegrate, without the aid of heat, a solid
object into its fundamental vibrations, transmit these vibrations into
the ether in the form of so-called “radio waves” which I then attracted
and condensed in my “receiving” apparatus, slowly damping their
short kinked vibration-rate until finally there was deposited the
homogeneous whole, identical in outline and displacement,—entirely
unharmed from its etheric transmigration!
My success in this, my life’s dream, was directly the result of our
discoveries on that bitter expedition into Afghanistan. All my life I had
been interested in the study of vibrations, but had achieved no
startling successes or keen expectations thereof until we stumbled
upon that strange mineral deposit on what was an otherwise ill-fated
trip for me. It was then that I realized that radioactive niton might
solve my hitherto insurmountable difficulty in the transmission of
material vibrations into electronic waves. My experiments thereafter,
while successful to the degree that I discovered several entirely new
principles of resonic harmonics, as well as an absolute refutation of
the quantum theory of radiation, fell far short of my hoped-for goal.
At that time I was including both helium and uranium in my improved
cathode projectors, and it was not until I had effected a more
sympathetic combination with thorium that I began to receive
encouraging results. My final success came with the substitution of
actinium for the uranium and the addition of polonium, plus a finer
adjustment which I was able to make in the vortices of my three
modified Tesla coils, whose limitations I had at first underrated. I was
then enabled to filter my resonance waves into pitch with my
“electronic radiate rays,” as I called them, with the success I shall
soon describe.
Of course, all this is no clearer than a page of Sanskrit to you, nor
do I intend that it shall be otherwise. As I have said, such a secret is
far too potent to be unloosed upon a world of such delicately poised
nations, whose jaws are still reddened from their recent ravening. It
needs no explanation of mine to envision the terrible possibilities for
evil in the application of this great discovery. It shall go with me—to
return at some future, more enlightened time after another equally
single-minded investigator shall have stumbled upon it. It is this latter
thought which has caused me to drop the hints that I have. My
earnest hope is that you will permit the misguided Marsden to read
the preceding paragraph. In it he will note a reference to an element
which I have not mentioned to him before, and will enable him to
obtain certain encouraging results,—encouraging but to further
efforts, to more frantic attempts. But I digress.
With my success on inanimate objects, I plunged the more
enthusiastically into my work. I should have lost all track of time but
for my daily tryst with Venice. Her belief in me was the tonic which
spurred me on to further efforts after each series of meticulously
conducted experiments had crumbled into failure. It was the
knowledge that she awaited me which alone upheld me in those dark
moments of depression, which every searcher into the realms of the
unknown must encounter.
Then came the night of November 28th, the Great Night. After
countless failures, I finally succeeded in transmitting a live guinea pig
through the atmosphere and “received” it, alive and well, in the
corner of my laboratory. Think of it! A twist of a switch and the living,
breathing piglet slowly dissolved before my eyes and vanished along
a pair of wires to my aerial, whence it was transferred as a set of
waves in the ether to the receiving apparatus,—there to reincarnate
into the living organism once more, alive and breathing, unharmed
by its extraordinary journey! That night I strode out into the open and
walked until dawn suddenly impressed the gray world upon my
oblivious exaltation, for I was King of the Universe, a Weaver of
Miracles.
Then it was that my great plan began to take shape. With
renewed energy I began the construction of a mammoth transmitter.
At intervals I “transmitted” stray cats and dogs of every description,
filling several books with notes wherein I recorded minutely the
varying conditions of my subjects before transmission. Invariably
their condition upon being “cohered” in the receiving tube, was
excellent. In some cases, indeed, minor ailments had entirely
disappeared during their short passage through the ether. What a
study for the medical profession!
I had, of course, told Venice the object of my researches long ago,
but had never brought her to my laboratory for reasons of discretion.
One afternoon, however, I slipped her in under cover of the heavy
downpour. After I had warmed her with a cup of tea, before her
astonished eyes I transmitted an old she-cat which was afflicted with
some sort of rheumatism or paralysis of its hind legs. When its form
began to reappear in the transparent receiving tube, my darling
gasped in awed wonder. She was rendered utterly speechless,
however, when I switched off the current and released the animal
from its crystal prison. And no wonder, for it gambolled about like a
young kitten, all trace of its former malady having entirely
disappeared! The impression upon Venice was all that I had hoped
for, and when I at length escorted her out into the dusk, I felt her
quick, awed glances flickering over me like the reverence of a shy
neophyte for the high priest.
All was set for the final act. I literally hurled myself into the
completion of my improved set. The large quantities of certain
minerals required caused me an unexpected delay. This I filled with
demonstrations in the presence of Marsden, whom I was
encouraging as a fellow radio enthusiast,—with considerable
unexpected histrionic ability on my part. It was so hard to keep my
fingers off his throat! I pretended to explain to him the important
factors of my great secret, and drilled him in the mechanical
operation of the sets. I had divulged to him also that my greatest
desire was to demonstrate my principle on a human being, and like
all great scientific explorers, proposed to offer myself as the subject.
Venice had strenuously opposed the proposal until the
demonstration on the diseased cat, and even now viewed the entire
proposition with alarm. Yet I insisted that unless applied to human
beings my entire work went for naught, and I finally succeeded in
quieting her fears to a great extent.
At last I am ready. I have told my darling how it is impossible to
transmit anything metallic by the very nature of the conflicting rays
encountered. I have bemoaned the fact that, due to the softness of
my teeth since boyhood, my mouth is one mass of metallic fillings
and crowns, rendering it impossible for me to test the efficiency of
my life’s work. As I had hoped, she has volunteered herself as the
subject for the great experiment, for her white teeth are as yet
innocent of fillings. I have demurred and refused to listen to the idea,
permitting myself to be won over only after days of earnest argument
on her part. We are not to tell Marsden, for there is no doubt that his
fanatical love for her would refuse to tolerate the mere suggestion.
Tonight it shall be accomplished. There is no other way, for that
accursed husband of hers seems to progress in neither direction. He
will be nothing but a mud-buried anchor until the end of her days,
while I—I love her. What other excuse need be offered?
But to the facts. At eight o’clock that drug-soaked love pirate
comes to officiate at my transmission through space. I shall meet
him with a chloroformed soaked rag. Later he will awake to find
himself effectively gagged, with his hands and feet firmly shackled to
the wall of a dark corner of my laboratory. These shackles consist of
armatures across the poles of large electro-magnets which I have
embedded in the walls. At 10:30, a time switch will cut off the
current, releasing the wretch, for, above all things, he must live. I
debated sending a message for his chauffeur to call for him here at
the designated hour. I have decided rather to trust to mechanical
certitude than lay my plan open to frustration because of some
human vagary.
At nine o’clock Venice comes for the great experiment. Marsden
has told her that he will remain in the city over night, at my
suggestion, so that in case I fail to materialize after being “sent” he
cannot be held in connection with my disappearance. She does not
know that I have had my teeth extracted and have been using India
rubber plates for nearly a month. By the time she has arrived, the
effects of the chloroform will have entirely worn off from my would-be
assistant, and I shall have had plenty of time to introduce myself
properly to him and explain the evening’s program which has been
so carefully arranged for his benefit.
Then he will have the excruciating pleasure of watching his
beloved wife dissolve into—nothingness! Soon thereafter he will
witness the same process repeated upon myself, for I have so
adapted the apparatus that I need no outside assistance other than a
time-clock to actuate the mechanism! Then, at the appointed hour,
the current will be shut off and the frenzied wretch will rush to the
distant switch controlling the receiving apparatus. As he throws the
metal bars into their split receptacles there will come a blinding flash,
and behold—the apparatus will have disappeared in a puff of
crystalline particles! The secret has returned whence it came!
Then will come that personally prepared hell for my mean spirited
forger. As I told you, he believes that he is in possession of enough
of the details of my secret to reconstruct the apparatus and duplicate
my success. The added details of this letter will assure him into an
idiotic confidence which will lead him on and on through partially
successful attempts. I know that no matter whether you sympathize
with my actions or not (and I am sure that you do not, for you never
have), your sense of justice will force you to show this letter to the
proper authorities in order to prevent a fatal bungling.
Meanwhile that miserable sneak will be frenzied with the
knowledge that at last, the lover he so long cheated of his loved one
is now with her, alone,—where he, her lawful husband, can never
follow. And we shall be together, unchanged, awaiting the day when
some other enlightened mortal solves Nature’s riddle, when we shall
once more assume our earthly forms, unhindered by other selfish
manbeasts.
Farewell,
Bromley Cranston.
Needless to say, I hurried to Eastport. But my trip was unnecessary. I

found Harold Marsden in a “private sanitarium” for the hopelessly
insane. There all day, and as far into the night as the opiates would
permit him, he is to be found seated before a radio set, the
earphones clamped to his head—listening. His statements,
methodically filed away by the head of the place, corresponded
wildly with the prophecies of my strange letter. Now he was listening
to fragmentary messages from those two he had seen precipitated
into space, he maintained. Listening.
And they had disappeared, utterly. I found the large seal ring in
the inkwell on the desk. Also the slip in the hatband of the hat which
had been placed in the wall safe, unlocked. The postmaster
remembered the seals on the letter my cousin had mailed, and the
approximate time he had received it. I felt my own reason wavering.
That is why, fantastic as is the whole affair, I cannot yet bear the
sound of one of those radio loud speakers. It is when that inarticulate
sound they call “static” occurs, when fragments of words and
sentences seem to be painfully attempting to pierce a hostile
medium,—that I picture that hunched up figure with its spidery
earphones,—listening. Listening. For what?
The End
Transcriber’s Note: This story appeared in the July 1927 issue of

Amazing Stories Magazine.
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Neuroscience For Neurosurgeons Feb 29 2024 - 110883146X - Cambridge University Press 1St Edition Farhana Akter Full Chapter

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Neuroscience for Neurosurgeons (Feb

Published online by Cambridge University Press

Published online by Cambridge University Press

Cambridge University Press is part of Cambridge University Press & Assessment,

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Section 1 Basic and Computational 13 Brain Metastases: Molecules to Medicine 193

3 Neuroimmune Interactions 59 15 Biomechanics of the Spine 234

4 Anatomy and Physiology of the Neuron 72 16 Degenerative Cervical Myelopathy 239

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25 Hydrocephalus 335 29 Neuroradiology: Focused Ultrasound

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Cecilia Dalle Ore

Joel S. Beckett Phan Q. Duy

Joseph S. Bell Roberto Eleopra

Dhiego Bastos Nigel Emptage

Karol P. Budohoski Peter E. Fecci

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Maya Harary Science Center, College of Medicine, Temple, TX,

Shawn L. Hervey-Jumper Gabriel Kreiman

Andrew S. Jack Maxwell D. Melin

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Francesco Prada Jasmine A. Thum

Umar Raza Vadim Tsvankin

Benjamin C. Reeves Pavan S. Upadhyayula

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1 Farhana Akter, Charles Reilly, Christophe Dupre, and Shifa Hossain

Dorsal view Anterior Ventral view Anterior

Superior frontal gyrus Posterior Posterior

Superior frontal sulci

Anterior Posterior Anterior Posterior

Figure 1.1 Planes and orientations of the brain.

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1 Farhana Akter, Charles Reilly, Christophe Dupre, and Shifa Hossain

Dorsal view Anterior Ventral view Anterior

Superior frontal gyrus Posterior Posterior

Superior frontal sulci

Anterior Posterior Anterior Posterior

Figure 1.1 Planes and orientations of the brain.

https://doi.org/10.1017/9781108917339.001 Published online by Cambridge University Press

Figure 1.2 Planes and orientations of the

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Figure 1.3 Circle of Willis.

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Orbital branches Calcarine artery

Superior anastomotic vein

Internal jugular vein

Figure 1.4 Arterial supply to the brain and venous drainage.

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(a) Figure 1.5 The skull and facial

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https://doi.org/10.1017/9781108917339.001 Published online by Cambridge University Press

(e) Palatine Process (Maxilla) Maxilla

Zygomatic Process (Maxilla)

Zygomatic Process Sphenoid Bone

Petrous Portion Inferior Nasal

Squamous Portion Temporal Bone

Mastoid Process Foramen Magnum

Occipital Condyle Parietal Bone

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Lateral view Superior view

Coronal Suture Sphenoparietal

Figure 1.6 Sutures of the brain.

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