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VETERINARY NEUROANATOMY AND CLINICAL ISBN: 978-0-7216-6706-5

NEUROLOGY

Copyright © 2009, 1983, 1977 by Saunders, an imprint of Elsevier Inc.

All rights reserved. No part of this publication may be reproduced or transmitted in any form
or by any means, electronic or mechanical, including photocopying, recording, or any ­information
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Notice

Neither the Publisher nor the Authors assume any responsibility for any loss or injury and/or
damage to persons or property arising out of or related to any use of the material c­ ontained
in this book. It is the responsibility of the treating practitioner, relying on independent
expertise and knowledge of the patient, to determine the best treatment and method of
application for the patient.

The Publisher

Library of Congress Cataloging-in-Publication Data

DeLahunta, Alexander, 1932-

Veterinary neuroanatomy and clinical neurology / Alexander de Lahunta, Eric Glass.—
3rd ed.
    p. ; cm.
Includes bibliographical references and index.
ISBN 978-0-7216-6706-5 (hardcover : alk. paper) 1. Veterinary neurology. 2. Veterinary
anatomy. 3. Neuroanatomy. I. Glass, Eric. II. Title.
[DNLM: 1. Anatomy, Veterinary. 2. Nervous System Diseases—veterinary. 3. Nervous
System—anatomy & histology. SF 895 D278v 2009]
SF895. D44 2009
636.089’18—dc22

978-0-7216-6706-5

Vice President and Publisher: Linda Duncan

Publisher: Penny Rudolph
Developmental Editor: Shelly Stringer
Publishing Services Manager: Julie Eddy
Project Manager: Laura Loveall
Designer: Teresa McBryan

Printed in China.

Last digit is the print number: 9 8 7 6 5 4 3 2

 Foreword

I
t is my profound privilege to write this foreword to the interpret MRIs of the brain and spinal cord, a technique that
third edition of Veterinary Neuroanatomy and Clinical came to the forefront of the specialty during the last decade
Neurology, a remarkable collaboration between Drs. of his career. As his resident, I had the unique opportunity
Alexander de Lahunta and Eric Glass. This anxiously antic- to observe Dr. de Lahunta develop an immediate and nat-
ipated textbook recaptures and updates the multifaceted ural mastery of MR imaging, which undoubtedly evolved
aspects of veterinary neurology, advanced neurodiagnos- from his vast experience studying gross and histopathologic
tics, neuroanatomy, and neuropathology, many aspects specimens from clinical cases. In the third edition, Dr. de
of which Dr. de Lahunta himself defined during his dis- Lahunta’s unique appreciation for the complexities of the
tinguished career at Cornell University. The third edition diseased nervous system, combined with Dr. Glass’ clinical
undoubtedly will withstand the test of time as an invaluable perspective derived from an immense caseload in specialty
contribution to the disciplines of veterinary internal medi- practice, seamlessly merge the disciplines of neuroanatomy
cine, neurology, and neurosurgery. and neuropathology together with clinical neurology and
The emphasis of the third edition remains the neuroana- state of the art neurodiagnostics.
tomic diagnosis, and this is accomplished elegantly in par- A foreword to this textbook would be remiss to ignore
allel with descriptions of key differential diagnoses and Dr. de Lahunta’s warmth, selflessness, and humanistic qual-
current diagnostic techniques. The textbook is referenced ities. Dr. de Lahunta has always found time to answer innu-
meticulously, providing the reader the opportunity to fur- merable formal and informal consultation requests from all
ther explore individual neurologic disorders. Hundreds of over the world, help students with their studies and proj-
color illustrations, case photographs, and web-based case ects, and mentor interns and residents, all the while mak-
videos are utilized to provide an invaluable resource for ing landmark contributions to the discipline of veterinary
the student, general practitioner, or specialist wishing to neurology. He has set the bar incredibly high for academic
develop or expand their understanding of neuroanatomy neurologists. We should all aspire to share of our time and
and clinical neurology. experiences with others as Dr. de Lahunta has done so grace-
Those who have been taught by or worked with Dr. de fully for over four decades.
Lahunta invariably conclude that he is one of the greatest In closing, the third edition of Veterinary Neuroanatomy
veterinary teachers of the past several generations. Dr. de and Clinical Neurology provides the critical building blocks
Lahunta’s innumerable college, national, and international necessary to gain competence in the discipline of veter-
teaching awards recognize his tremendous contributions to inary neurology. On behalf of neurologists world wide,
veterinary education. His inherent ability to teach such a I would like to thank Drs. de Lahunta and Glass for col-
complex topic is largely due to his steadfast commitment laborating on this new edition which promises to be the
to a vertically integrated learning process whereby neuro- new treatise of neuroanatomic localization and clinical
embryology, neuroanatomy, clinical neurology, and neu- neurology.
ropathology are taught as a continuum. For 42 years, Dr. Congratulations and please accept our sincere gratitude
de Lahunta preached that a basic understanding of neuro- for all of the time, energy, and wisdom you have shared
anatomy is indispensible for one to determine the anatomic with all of us through the years.
localization and, in turn, to generate the correct differen-
tial diagnosis in neurological patients. Dr. de Lahunta rec-
ognized early in his career that neuroanatomy and clinical
neurology are inseparable, and the third edition is a testa-
ment to this philosophy.
The inseparability of neuroanatomy and clinical neurol- Scott J. Schatzberg, DVM, PhD, DACVIM (Neurology)
ogy may be best epitomized by Dr. de Lahunta’s ability to University of Georgia College of Veterinary Medicine

vi
 Preface

T
his third edition is a revision based on my 42 years of The emphasis of this third edition is the anatomic ­diagnosis,
experience in teaching a vertically integrated course which is the basis for the successful practice of clinical
in veterinary neuroanatomy and clinical neurology neurology and is presented in the form of case examples.
to first year veterinary students at the Cornell University Disease descriptions are presented in the form of differen­
College of Veterinary Medicine. The clinical information tial diagnoses of individual case examples. Although many
evolved from my experience as a consultant to the Teaching neurological disorders will be described, this revision makes
Hospital that received patients with neurological disorders, no attempt to present a complete synopsis of the veterinary
consulting with veterinary practitioners, and my involve­ neurological literature. This has been adequately covered in
ment with the neuropathological studies of hospital patients other textbooks of veterinary neurology. A unique feature of
and specimens sent to me by veterinary practitioners and this text is the linking of the case descriptions to videos of
pathologists. My studies were greatly enhanced by my these patients which are available on a website that is orga­
interaction with Drs. John Cummings and Brian Summers. nized by the chapters of this text. There are 380 videotapes
I consider these two colleagues as close friends and brilliant on this website that show most of the neurological disor­
veterinary scientists. ders that are described in this text. This can be accessed at:
When I first organized a course in neuroanatomy for the www.neurovideos.vet.cornell.edu.
veterinary curriculum at Cornell University in 1963, I was My coauthor, Dr. Eric Glass, is a former student of mine
strongly influenced by two outstanding veterinary patholo­ who graduated from the College of Veterinary Medicine at
gists who in 1956 and 1965 published their experiences in Cornell University in 1995. He is the senior neurologist at
correlating neurological signs with the location of lesions Red Bank Veterinary Hospital, a very active specialty prac­
in the nervous system.1,2 These were Dr. Jack McGrath at tice in Tinton Falls, New Jersey, where he has ten years of
the University of Pennsylvania and Dr. Tony Palmer at practice experience. Eric brings a clinician’s perspective to
Cambridge University. They set a standard that I wanted to the understanding of neuroanatomy. I am honored to have
pursue. I felt strongly that teaching neuroanatomy solely his contributions to this textbook. Eric and I have agreed
for the sake of the anatomical training was not a useful on most areas of controversy in clinical neurology and have
expenditure of the student’s limited time or that of the busy presented our combined opinions in this text. We present
teacher. It is the one system that needs to be closely cor­ these controversies as well as differences of opinion as chal­
related with the study of the clinical signs to be useful to lenges for the present and future veterinary neurologists to
the student as a veterinary practitioner. The teaching of support or deny.
­clinical veterinary medicine is the primary objective of the Alexander de Lahunta
­curriculum at Cornell University. Drs. McGrath and Palmer
­set this standard that became the basis for my entire profes­
sional career and the correlated course that I taught to first
year veterinary students for 42 years. This course provided
all the useful neuroanatomy and clinical neurology simul­
taneously. This has been an exciting experience for me that 1. McGrath, JT: Neurologic examination of the dog with clini­
has been rewarded by observing the success of my students copathologic observations, 1956, Philadelphia, PA, Lea and
in this clinical specialty and by observing the satisfaction Febiger.
expressed by the students who have mastered the ability to 2. Palmer, AC: Introduction to animal neurology, 1965, Phila­
diagnose neurological disorders. delphia, PA, F.A. Davis.

vii
C h a p t e r

1 Introduction

OBJECTIVE FUNCTIONAL SYSTEMS Proprioception

General Proprioception
ACCURATE DIAGNOSIS SENSORY (AFFERENT)
Special Proprioception
Malformations Somatic Afferent
Inflammations General Somatic Afferent MOTOR (EFFERENT)
Injuries Special Somatic Afferent General Somatic Efferent
Neoplasias Visceral Afferent General Visceral Efferent
Degenerations General Visceral Afferent
Special Visceral Afferent FURTHER READING
NEURON

Objective should first attempt to determine whether all the clinical

signs can be explained by a lesion at one site in the nervous
This book was written primarily for the veterinary student system, a focal lesion, because they are more common than
and secondarily for the veterinary practitioner. It is orga- multifocal or diffuse disorders. Based on this anatomic diag-
nized to provide the veterinary student with an anatomic nosis, you will next establish a list of disorders that must
basis and sufficient information about the development, be able to affect the anatomic location of the lesion. This
organization, and function of the nervous system to be able is the differential diagnosis. You will learn various ways to
to understand and diagnose the more common disorders of remind yourself of the disorders to consider. One way is the
the nervous system of domestic animals. For the most part, MIIND system (malformation, injury, inflammation, neo-
these disorders are described in the chapter that discusses plasia, and degeneration).
the functional system primarily affected by the disorder.
Malformations
Accurate Diagnosis Malformations are the disorders that result from abnormal
development of the nervous system.
The major objective of this book is to teach enough of the
morphologic and physiologic features of the nervous sys-
tem to enable the student to make an accurate localization
Inflammations
of the lesion in the nervous system. This is the anatomic Inflammations involve a pathologic process and a reaction
­diagnosis. The differential diagnosis is totally dependent of blood vessels and tissues to physical, chemical, and bio-
on the anatomic diagnosis and that, in turn, determines logic agents—the reaction of a tissue to an irritant. In the
the ancillary procedures that will be prioritized to arrive at nervous system, this commonly refers to the tissues’ reac-
the most accurate presumptive clinical diagnosis and the tion to a microorganism or an immune system abnormality.
­subsequent selection of treatment. Suppurative inflammation is characterized by a neutrophilic
The diagnosis of clinical neurologic disorders starts with response and the products of necrosis of tissue and inflam-
recognition of the problem—the clinical signs exhibited by matory cells usually caused by a bacterium, protozoa, or
the patient and your neurologic examination. This visual fungus. Nonsuppurative inflammation is characterized by
and hands-on experience is difficult to learn by reading a lymphocytic or monocytic response and is usually caused
text descriptions. Direct contact with the affected patient by a viral agent or an immune system abnormality.
is the ideal teaching model but is impractical in a ­teaching
environment. The most effective alternative is to visualize
the clinical signs using video technology. This third edition
Injuries
includes linkage to a website consisting of 381 videos that Injuries occur when nervous tissue undergoes traumatic
provide the student with the classical appearance of the com- ­disturbance deriving from external or internal sources.
mon disorders of the nervous system of domestic animals. These cause acute or chronic displacements and disruptions
The anatomic diagnosis is determined by the nature of the or vascular impairment of the nervous tissue, which may
problem; that is, clinical signs that you have observed. You result in hemorrhage, edema, or parenchymal necrosis.


 V eterinary N euroanatomy and C linic al N eurology

Neoplasias a stimulus from the internal or external environment is

converted into an impulse in the neuron. The axon is the
Neoplasias are uncontrolled growth of cells. Primary cen- cell process composed of neurofilaments that course from
tral nervous system (CNS) neoplasias include the uncon- the dendritic zone to the telodendron. The telodendron
trolled growth of nervous tissue cells—neurons, glia, and is the termination of the neuron where the impulse leaves
ependyma. Metastatic neoplasia of the nervous system is the neuron. It is often referred to as the synapse. This syn-
the spread of primary neoplasms in other body tissues to apse may lie at an effector organ or at another neuron. The
the nervous system. cell body consists of the nucleus and the major organelles
­necessary for the neuron to function and may be located
anywhere along the axon.
Degenerations For example, a sensory neuron in the peripheral nervous
Degenerations include the deterioration of cells due to lack system for general proprioception may have its dendritic
of blood supply (ischemia), abnormal cellular metabolism zone in a neuromuscular spindle in a skeletal muscle where
caused by an inherited cellular defect, exposure to exoge- it is stimulated by a stretching of the muscle. The axon
nous toxins, and abnormalities in other body systems (renal courses toward the spinal cord through a specific periph-
disorders with uremia, diffuse liver disorders with hyper- eral nerve, then through the dorsal or ventral branch of one
ammonemia, cardiorespiratory disorders with hypoxia). of the spinal nerves and into its dorsal root. It then enters
Abiotrophy is cell degeneration due to an intrinsic defect the spinal cord and passes into the dorsal gray column of
in the essential metabolism necessary for the survival and that spinal cord segment to synapse on a second neuron
function of that cell, the neuron. in a nucleus within that gray column. The telodendron is
Do not forget to consider the breed of your patient and the nerve ending at the synapse on another neuron in that
the possible inherited disorders that must be considered. nucleus. The neuronal cell body is located in the spinal gan-
You will prioritize these disorders in your differential on glion associated with the dorsal root that the axon coursed
the basis of signalment, history, and course of the clinical through to reach the spinal cord. It is actually intercalated
signs and the characteristics of the various disorders being in the axon at this point (Fig. 1-1).
considered. The dendritic zone and cell body of a motor neuron in
Based on this ranking of the differential diagnosis, the the peripheral nervous system that is innervating a skel-
most useful ancillary procedures will be selected to further etal muscle are closely associated and are located in the
confirm or deny the diagnosis under consideration. This ventral gray column of a segment of the spinal cord. The
selection is especially critical now that neuroimaging by axon leaves the cell body in that gray column and courses
computed tomography and magnetic resonance are avail- through the white matter of that segment to enter the
able to veterinarians. These procedures require general ventral root of that segment. It continues into the spinal
anesthesia and the costs to the owner of the patient are con- nerve of that segment and its dorsal or ventral branch
siderable. Therefore, it is crucial that the correct anatomic and then travels in a specific peripheral nerve to reach
diagnosis be made prior to the selection of the ancillary pro- the skeletal muscle cells being innervated. Here the axon
cedures. Your knowledge of the characteristics of the disor- ends in a telodendron at the neuromuscular ending in a
ders of the nervous system will then permit you to offer a motor end-plate.
therapy where it is appropriate and a prognosis. Within the CNS, a neuron of the dorsal spinocerebel-
lar tract is an example of a sensory or afferent neuron to
the cerebellum. Its dendritic zone and cell body are closely
Neuron associated in a nucleus in the dorsal gray column of the
spinal cord. The impulse is initiated here by a synapse with
The nervous system is composed of primary functional the telodendron of a sensory general ­ proprioceptive neu-
cells—the neurons and supporting cells, which include the ron of the peripheral nervous system. The axon courses
glia and ependyma. In this book the neuron is defined as through the gray matter into the white matter of the lateral
consisting of a dendritic zone, axon, cell body, and telo- funiculus to join other axons in a tract on the dorsal super-
dendron. The dendritic zone is the receptor portion, where ficial surface of the lateral funiculus. This axon continues

DR
C

B
A D
Figure 1-1 Diagram of a general somatic afferent (GSA) Skin GSA
neuron and a general somatic efferent (GSE) neuron in a A
spinal nerve (SN). A, Dendritic zone. B, Cell body. C, Axon. SN B
D, Telodendron. DR, Dorsal root; SM, skeletal muscle;
VR, ventral root. C
GSE VR
D

SM
 Introduction 

c­ ranially in this dorsal spinocerebellar tract. It traverses the

Table 1-1 Functional Classification of the
portion of the spinal cord cranial to the spinal cord seg-
Nervous System
ment where it originated, then continues into the medulla
where it enters the cerebellum via the caudal cerebellar
peduncle. It courses through the cerebellar medulla and System Function and Anatomic Location
into the white matter of a cerebellar folium. It enters the 1. Afferent (A): Sensory
adjacent granular layer of the cerebellum and terminates
in a telodendron that synapses with the dendritic zone of a Somatic (S)
granule cell neuron. General (GSA) Temperature, touch, noxious stimuli
All spinal nerves, cranial nerve V
Within the CNS, the Purkinje neuron of the cerebellum
Special (SSA) Vision: Cranial nerve II
is an example of an efferent neuron in the cerebellar cortex. Hearing: Cranial nerve VIII
Its dendritic zone consists of a branched structure located in Visceral (V)
the molecular layer of the cerebellar cortex on the surface of General (GVA) Organ content, distention, chemicals
a folium. Here the telodendria of the granule cell neurons Spinal nerve splanchnic branches
synapse at sites on these branches to initiate the impulse in Cranial nerves VII, IX, X
the Purkinje neuron .The cell body is located in the Purkinje Special (SVA) Taste: Cranial nerves VII, IX, X
neuronal layer of the cerebellar cortex. The axon arises from Olfaction: Cranial nerve I
this cell body and courses through the granular layer into Proprioception
and through the white matter of that cerebellar folium and General (GP) Muscle and joint movement
enters the white matter of the cerebellar medulla. Here All spinal nerves, cranial nerve V
the axon ends in a telodendron on the dendritic zone of Special (SP) Vestibular system: Cranial nerve VIII
another efferent neuron located in a nucleus of the cerebel-
2. Efferent (E): Motor
lar medulla.
Somatic (S)
General (GSE) Striated skeletal muscle
All spinal nerves
Functional Systems Cranial nerves III, IV, V, VI, VII, IX, X, XI, XII
Visceral (V)
This book is organized primarily by functional systems General (GVE) Smooth and cardiac muscle and glands
Sympathetic: All spinal nerves, splanchnic
rather than by regions of the nervous system or by the chief
nerves
clinical neurologic complaint. It is our opinion that for
Parasympathetic: Sacral spinal nerves
teaching purposes, this is the most effective way to learn the Cranial nerves III, VII, IX, X, XI
organization of the nervous system and provide the basis
for understanding the disorders that affect the various com-
ponents of the nervous system. Most of these functional
systems are derived from a classification of the peripheral
nervous system based on its functional components. The
Special Somatic Afferent
sensory portion has extensive components in the CNS. The The special somatic afferent (SSA) system involves special-
classification is outlined in Table 1-1. ized dendritic zone receptor organs limited to one area
deep to the body surface but stimulated by changes in the
external environment. These include light to the eyeball
(cranial nerve II) and air waves indirectly to the membra-
SENSORY (AFFERENT) nous labyrinth of the inner ear (cranial nerve VIII, cochlear
division).
The sensory, or afferent, portion of the peripheral nervous
system is classified on the basis of the location of the den-
dritic zone in the body. This is the site of the origin of the
impulse.
Visceral Afferent
The visceral afferent system has its dendritic zone in the
wall of the various viscera of the body. This is tissue derived
Somatic Afferent mostly from splanchnopleura and is stimulated by changes
The somatic afferent system has its dendritic zone on or in the internal environment.
near the surface of the body derived from the ­somatopleura,
where it receives the various stimuli from the external
environment.
General Visceral Afferent
The general visceral afferent (GVA) system is composed of
General Somatic Afferent neurons distributed by the seventh, ninth, and tenth cranial
The general somatic afferent (GSA) system comprises the nerves to visceral structures in the head and by the tenth
neurons distributed primarily by the fifth cranial nerve cranial nerve and spinal nerves to the viscera of the body
to the surface of the head and all the spinal nerves to cavities and blood vessels throughout the neck, trunk, and
the ­surface of body and limbs that are sensitive to touch, limbs. This widely distributed system is stimulated primarily
­temperature, and noxious stimuli. by the distention of visceral walls and chemical changes.
 V eterinary N euroanatomy and C linic al N eurology

Special Visceral Afferent General Visceral Efferent

The special visceral afferent (SVA) system contains the neu- The general visceral efferent (GVE) system has its teloden-
rons in the seventh, ninth, and tenth cranial nerves, whose dria in involuntary smooth muscle of viscera derived from
dendritic zones are limited to the specialized receptors for splanchnic mesoderm as well as blood vessels, cardiac mus-
taste, and the first cranial nerve, whose dendritic zones are cle, and glands. This system is the LMN of the autonomic
localized in the caudal nasal mucosa for olfaction. nervous system, which has components in all segments
of the brain and spinal cord. In some books, the GVE sys-
tem is considered to be the entire autonomic system; we
Proprioception believe that to be an inappropriate concept that defies the
The modality of general proprioception is sometimes true functional totality of this autonomic system, which
included in the GSA system. In this book we consider it includes peripheral afferent components and a plethora of
as a separate system because of its clinical significance. nuclei and tracts at all levels of the CNS.
Disorders that affect this system express clinical signs dif- The GVE system, unlike the GSE system, is a two-neuron
ferent from those that affect the GSA system as we have system in that there are two neurons between the CNS and
classified it. Proprioception is the system responsible for the target organ. A synapse occurs in a peripheral ganglion
detecting changes in the position of the head, neck, trunk, between these two neurons. There are two divisions of this
and limbs. system: sympathetic and parasympathetic. They are further
described in Chapter 7, which is devoted to the GVE sys-
tem. These GVE neurons are distributed in the third, sev-
General Proprioception enth, ninth, tenth, and eleventh cranial nerves and all the
The general proprioception (GP) system has its dendritic spinal nerves.
zones widely distributed in receptor organs located in In previous editions of this book and in older books of
muscles, tendons, and joints deep to the body surface. developmental anatomy, a special visceral efferent system
It is distributed widely throughout all the spinal nerves was described for the innervation of striated skeletal muscle
and the fifth cranial nerve. The receptors are sensitive in the head derived from branchial arch mesoderm. This
to changes in the lengths and positions of the structures classification has now been deleted because there is no dif-
they innervate. ference between this head skeletal muscle and the muscles
in the rest of the body. All this muscle is now considered to
be innervated by the GSE system.
Special Proprioception
The special proprioception (SP) system’s dendritic zones are
limited to receptors specialized to respond to positions and Further Reading
movements of the head. They are located in a portion of the
membranous labyrinth of the inner ear. These neurons con- For further reading, the following list of textbooks is highly
cerned with the orientation of the head in space are in the recommended. For all aspects of canine neuroanatomy,
vestibular division of the vestibulocochlear nerve ­ (cranial the most extensive and thorough descriptions are found in
nerve VIII). Miller’s Anatomy of the Dog by Howard Evans. From our per-
spective, this should be considered the gold standard. The
text by Tom Jenkins, Functional Mammalian Neuroanatomy,
Motor (Efferent) is an easy read, has many simplified line drawings of vari-
ous neurologic concepts, and is based primarily on the dog.
The motor, or efferent, portion of the peripheral nervous For the study of brain sections in all three planes, The Brain
system is classified on the basis of where the motor neuron of the Dog in Section by Marcus Singer is unsurpassed. This is
terminates, which is the site of the telodendron. This periph- a superb text for ­correlation with MR images.
eral motor system is also referred to as the lower motor neu- For further reading in veterinary clinical neurology, the
ron (LMN) because it is the final neuron to ­ innervate the six texts listed are all well written and provide thorough
muscle cell. There are somatic and visceral components of coverage of their areas of concern. Cheryl Chrisman’s text
the efferent system. is limited to small animals and is usefully organized by the
patient’s problem, the chief complaint. Kyle Braund’s text
is organized like a dictionary and is useful for looking up
General Somatic Efferent short, succinct reviews of any domestic animal’s neurologic
In the general somatic efferent (GSE) system, the teloden- problem. All species are included in the handbook written
dron is located in striated skeletal muscle throughout the by Mike Lorenz and Joe Kornegay, although their personal
entire body derived from somatic mesoderm, somites, and experience is mostly with small animals. The only text in
head somitomeres. The cell body and dendritic zone of these veterinary neurology that is limited to large animals is that
GSE neurons are in the spinal cord ventral gray column and written by Joe Mayhew. This is a superb book that represents
in nuclei in the brainstem. Their axons are in the ventral his extensive personal experience. The original textbook
root and spinal nerves or in cranial nerves, and they course of veterinary neurology written by Ben Hoerlein, which
through various named peripheral nerves to terminate in a included an extensive section on surgical and medical treat-
muscle cell at the neuromuscular ending (junction). These ment, was revised by John Oliver and Joe Mayhew and was
GSE neurons are found in all the spinal nerves and in all the expanded to include some large-animal neurology. In our
cranial nerves except cranial nerves I, II, and VIII. opinion the most current and most inclusive ­ veterinary
 Introduction 

­ eurology text for small animals is the text published in

n Lorenz MD, Kornegay JN: Handbook of veterinary neurology, ed 4,
2003 and edited by Curtis Dewey. For spinal surgery, the Philadelphia, 2004, Saunders.
textbook by Nick Sharp and Simon Wheeler details the Mayhew IG: Large animal neurology, Philadelphia, 1989, Lea &
contemporary aspects of spinal surgery for small animals. Febiger.
Oliver JE, Hoerlein BF, Mayhew IG: Veterinary neurology,
This text is supported by excellent diagrams and photos.
Philadelphia, 1987, Saunders.
The only textbook devoted to veterinary neuropathology is
Platt SR, Olby NJ: BSAVA manual of canine and feline neurology,
the one written by Brian Summers with the help of John ed 3, Gloucester, UK, 2004, British Small Animal Veterinary
Cummings and myself (Alexander de Lahunta). In addi- Association.
tion to the descriptive ­ neuropathology, this text includes Sharp NJ, Wheeler SJ: Small animal spinal disorders: diagnosis and
considerable clinical correlations as well as descriptions of surgery, ed 2, Philadelphia, 2005, Elsevier.
pathogenesis.
Veterinary Neuropathology
Suggested readings Summers BA, Cummings JF, de Lahunta A: Veterinary
neuropathology, New York, 1995, Mosby.
Domestic Animal Neuroanatomy
Jenkins TW: Functional mammalian neuroanatomy, ed 2, Other References
Philadelphia, 1978, Lea & Febiger. Crosby EC, Humphrey T, Lauer EW: Correlative anatomy of the
Evans HE: Miller’s anatomy of the dog, ed 3, Philadelphia, 1993, nervous system, New York, 1962, Macmillan.
Saunders. Fankhauser R, Luginbuhl H: Pathologische Anatomie des zentralen
Singer M: The brain of the dog in section, Philadelphia, 1962, und peripheren Nervensystem der Haustiere, Berlin, Germany,
Saunders. 1968, Verlag Paul Perey.
Frauchiger E, Fankhauser R: Neuropathologie des Menschen and
Veterinary Clinical Neurology der Tiere, Berlin, Germany, 1957, Springer-Verlag.
Braund KG: Clinical syndromes in veterinary neurology, ed 2, Innes JRM, Saunders LZ: Comparative neuropathology, New York,
St Louis, 1994, Mosby. 1962, Academic Press.
Chrisman CL: Problems in small animal neurology, ed 2, Nickle R, Schummer A: Seiferle E: Lehrbuch der Anatomie der
Philadelphia, 1991, Lea & Febiger. Haustiere, Band IV, Nervensystem, Sinnesorgane, Endokrine
Chrisman CL, et al: Neurology for the small animal practitioner, Drusen, Berlin, Germany, 1975, Verlag Paul Perey.
Jackson, WY, 2003, Teton NewMedia. Papez JW: Comparative neurology, New York, 1929, TY Crowell.
Dewey CW: A practical guide to canine and feline neurology, Ames,
IO, 2003, Iowa State Press.
C h a p t e r

2 Neuroanatomy by Dissection

TRANSVERSE BRAIN SECTIONS

T
he neuroanatomic components of this textbook are with Figs. 2-2 through 2-17. The images are 2 mm thick, but to
based on and complement the dissection of the ner- select those that best demonstrate the anatomic features, the
vous system described in Guide to the Dissection of the intervals between images vary.
Dog by H. E. Evans and A. de Lahunta (ed 6, Philadelphia,
2004, Elsevier). The peripheral nerves are described and
dissected along with the regions of the body in which they
are found. The dissection of the brain and spinal cord is
found in the last section, titled “Nervous System.” The
split head of the embalmed dog used for the dissection of 2 3 45 6 7 8 9 10 11 12 13 14 15 16 17
the entire dog is also used to demonstrate the blood ves-
sels and meninges. A separate entire preserved dog brain
is provided to each group of students for the dissection.
The spinal cord can be dissected on the embalmed dogs
or presented as prosections. On completion of the brain
dissection, an additional preserved domestic animal brain
is provided to each group for the study of the transverse
sections.
In the Cornell curriculum this dissection is performed
simultaneously with lectures and discussions of nervous sys-
tem development, cerebrospinal fluid, and ­malformations,
including hydrocephalus, which are the subjects of Chapters
3 and 4 of this book.
The nomenclature used in this third edition, as in Guide
to the Dissection of the Dog, adheres to that published in the 2 3 45 6 7 8 9 10 11 12 13 14 15 16 17
fifth edition of Nomina Anatomica Veterinaria in 2005, unless
otherwise stated.

Transverse Brain Sections

The following transverse brain sections in Figs. 2-2 through
2-17 are arranged from rostral to caudal through the brain
at irregular intervals, as indicated on the drawings in Fig.
2-1. In these sections the white matter has been stained
and appears black, whereas the gray matter is relatively
unstained.
Figs. 2-18 through 2-33 are transverse plane (axial) pro- Figure 2-1 Dorsal and left lateral views of the brainstem, indicating the
ton density MR images of a normal adult dog for comparison approximate levels of the following transverse sections.


Neuroanatomy by Dissection 7

Figure 2-2
 1. Cingulate gyrus
2. Corpus callosum
3. Body of fornix
4. Column of fornix
5. Septal nuclei
6. Lateral ventricle
  7. Body of caudate nucleus
8. Internal capsule
9. Centrum semiovale
10. Globus pallidus
  11. Putamen
12. External capsule
13. Rostral commissure
14. Lateral olfactory tract
15. Optic nerve

Figure 2-3
1. Cingulate gyrus
2. Corpus callosum
3. Body of fornix
4. Crus of fornix
5. Lateral ventricle
6. Caudal (tail) caudate nucleus
7. Centrum semiovale
8. Internal capsule
9. Thalamocortical projection fibers
10. Corticopontine, corticobulbar,
and corticospinal projection fibers
11. Globus pallidus
12. Putamen
13. External capsule
14. Claustrum
15. Amygdaloid body
16. Pyriform lobe
17. Optic tract
18. Endopeduncular nucleus
19. Hypothalamic nuclei
20. Column of fornix
21. Mamilothalamic tract
22. Third ventricle
23. Interthalamic adhesion
24. Thalamic nuclei
25. Stria habenularis thalami
8 V eterinary N euroanatomy and C linic al N eurology

Figure 2-4
1. Cingulate gyrus
2. Corpus callosum
3. Hippocampus
4. Crus of fornix
5. Lateral ventricle
6. Parahippocampal gyrus
7. Lateral rhinal sulcus, caudal part
8. Internal capsule
9. Thalamocortical projection
10. Lateral geniculate nucleus
11. Thalamic nuclei
12. Habenular nucleus
13. Habenulointerpeduncular tract
14. Third ventricle
15. Interthalamic adhesion
16. Zona incerta
17. Crus cerebri
18. Optic tract
19. Subthalamic nucleus
20. Mammillary body
21. Mamillothalamic tract
22. Caudal hypothalamic region
23. Adenohypophysis
24. Neurohypophysis

Figure 2-5
1. Cingulate gyrus
2. Splenium of corpus callosum
3. Hippocampus
4. Crus of fornix
5. Lateral ventricle
6. Parahippocampal gyrus
7. Lateral rhinal sulcus, caudal part
8. Optic tract
9. Lateral geniculate nucleus
10. Medial geniculate nucleus
11. Pretectal nuclei
12. Pineal body
13. Caudal commissure
14. Mesencephalic aqueduct
15. Parasympathetic nucleus of
oculomotor nerve
16. Medial lemniscus
17. Substantia nigra
18. Crus cerebri
Neuroanatomy by Dissection 

Figure 2-6
1. Cingulate gyrus
2. Splenium of corpus callosum
3. Hippocampus
4. Crus of fornix
5. Lateral geniculate nucleus
6. Optic tract
7. Brachium of rostral colliculus
8. Rostral colliculus
9. Commissure of rostral colliculus
10. Mesencephalic aqueduct
11. Central gray substance
12. Reticular formation
13. Medial geniculate nucleus
14. Medial lemniscus
15. Substantia nigra
16. Crus cerebri
17. Oculomotor nerve fibers
18. Parasympathetic nucleus of
oculomotor nerve
19. Medial longitudinal fasciculus
20. Red nucleus

Figure 2-7
1. Mesencephalic aqueduct
2. Central gray substance
3. Rostral colliculus
4. Brachium of caudal colliculus
5. Medial geniculate nucleus
6. Reticular formation
7. Medial lemniscus
8. Substantia nigra
9. Crus cerebri
10. Interpeduncular nucleus
11. Oculomotor nucleus
12. Medial longitudinal fasciculus
13. Red nucleus
14. Ventral tegmental decussation
(rubrospinal neurons)
15. Rubrospinal tract
10 V eterinary N euroanatomy and C linic al N eurology

Figure 2-8
1. Commissure of caudal colliculus
2. Mesencephalic aqueduct
3. Central gray substance
4. Caudal colliculus
5. Brachium of caudal colliculus
6. Lateral lemniscus
7. Reticular formation
8. Nucleus of trochlear nerve
9. Medial longitudinal fasciculus
10. Decussation of rostral cerebellar
peduncle
11. Rubropsinal tract
12. Medial lemniscus
13. Interpeduncular nucleus
14. Crus cerebri
15. Pontine nuclei
16. Transverse fibers of pons

Figure 2-9
1. Medial longitudinal fasciculus
2. Fourth ventricle
3. Trochlear nerve
4. Caudal colliculus
5. Rostral cerebellar peduncle
6. Lateral lemniscus
7. Nucleus of lateral lemniscus
8. Middle cerebellar peduncle
9. Trigeminal nerve
10. Transverse fibers of pons
11. Pontine nuclei
12. Longitudinal fibers of pons
13. Medial lemniscus
14. Reticular formation
15. Locus ceruleus
Neuroanatomy by Dissection 11

Figure 2-10
1. Medial longitudinal fasciculus
2. Rostral medullary velum
3. Fourth ventricle
4. Rostral cerebellar peduncle
5. Middle cerebellar peduncle
6. Trigeminal nerve
7. Lateral lemniscus
8. Reticular formation
9. Medial lemniscus
10. Pontine nuclei
11. Longitudinal fibers of pons
12. Transverse fibers of pons

Figure 2-11
1. Medial longitudinal fasciculus
2. Rostral medullary velum
3. Rostral vestibular nucleus
4. Rostral cerebellar peduncle
5. Middle cerebellar peduncle
6. Vestibulocochlear nerve
7. Cochlear nucleus
8. Trigeminal nerve
9. Nucleus of spinal tract of trigeminal
nerve, pontine sensory portion
10. Motor nucleus of trigeminal nerve
11. Facial nerve
12. Dorsal nucleus of trapezoid body
13. Medial lemniscus
14. Trapezoid body
15. Pyramid
16. Reticular formation
12 V eterinary N euroanatomy and C linic al N eurology

Figure 2-12
1.
Medial longitudinal fasciculus
2.
Abducent nerve fibers
3.
Genu of facial nerve
4.
Ventrolateral coursing facial nerve
fibers
5. Medial vestibular nucleus
6. Vestibulocerebellar fibers
7. Lateral vestibular nucleus
8. Caudal cerebellar peduncle
9. Flocculus
10. Cochlear nuclei
11. Vestibulocochlear nerve
12. Spinal tract of trigeminal nerve
13. Nucleus of spinal tract of trigeminal
nerve
14. Dorsal nucleus of trapezoid body
15. Trapezoid body
16. Pyramid
17. Medial lemniscus

Figure 2-13
1. Fastigial cerebellar nucleus
2. Interposital cerebellar nucleus
3. Lateral cerebellar nucleus
4. Nodulus
5. Flocculus
6. Medial vestibular nucleus
7. Caudal vestibular nucleus
8. Acoustic stria
9. Caudal cerebellar peduncle
10. Spinal tract of trigeminal nerve
11. Nucleus of spinal tract of trigeminal
nerve
12. Facial nucleus
13. Dorsomedial coursing facial nerve
fibers
14. Reticular formation
15. Pyramidal tract
16. Medial lemniscus
Neuroanatomy by Dissection 13

Figure 2-14
1. Medial longitudinal fasciculus
2. Hypoglossal nucleus
3. Radix of hypoglossal nerve
4. Parasympathetic nucleus of vagus
nerve
5. Nucleus of solitary tract
6. Lateral cuneate nucleus
7. Caudal cerebellar peduncle
8. Spinal tract of trigeminal nerve
9. Nucleus of spinal tract of trigeminal
nerve
10. Deep arcuate fibers
11. Nucleus ambiguus
12. Olivary nucleus
13. Pyramidal tract
14. Medial lemniscus
15. Reticular formation

Figure 2-15
1. Central canal
2. Hypoglossal nucleus
3. Radix of hypoglossal nerve
4. Parasympathetic nucleus of
vagus nerve
5. Nucleus gracilis
6. Medial cuneate nucleus
7. Lateral cuneate nucleus
8. Spinal tract of trigeminal nerve
9. Nucleus of spinal tract of trigeminal
nerve
10. Nucleus ambiguus
11. Dorsal spinocerebellar tract
12. Olivary nucleus
13. Pyramidal tract
14. Medial lemniscus
15. Medial longitudinal fasciculus
16. Reticular formation
17. Nucleus of solitary tract
18. Solitary tract
19. Accessory nerve
14 V eterinary N euroanatomy and C linic al N eurology

Figure 2-16
1. Nucleus gracilis
2. Medial cuneate nucleus
3. Fasciculus gracilis
4. Spinal tract of trigeminal nerve
5. Nucleus of spinal tract of trigeminal
nerve
6. Medial longitudinal fasciculus
7. Pyramidal decussation
8. Spinocerebellar tracts

Figure 2-17
1. Fasciculus gracilis
2. Fasciculus cuneatus
3. Spinal tract of trigeminal nerve
4. Nucleus of spinal tract of trigeminal
nerve: dorsal gray column, first
cervical spinal cord segment
5. Rubrospinal tract
6. Lateral pyramidal (corticospinal) tract
7. Vestibulospinal tract
8. Ventral median fissure
9. Spinocerebellar tracts
 Neuroanatomy by Dissection 15

Figure 2-18
1. Olfactory bulb

3
Figure 2-19
1. Optic nerve
2. Olfactory peduncle
3. Frontal lobe
2
1
16 V eterinary N euroanatomy and C linic al N eurology

Figure 2-20
1. Optic nerve 3
2. Longitudinal cerebral fissure
3. Genu corpus callosum
4. Internal capsule
2

11
Figure 2-21
1. Third ventricle
2. Lentiform nucleus
10
3. Rostral commissure 9
4. Body of fornix at bend to columns 8
of fornix
5. Caudate nucleus 5 7
6. Internal capsule 4
7. Lateral ventricle
6
8. Corpus callosum 2 3
9. Centrum semiovale
10. Cingulum in cingulate gyrus
11. Falx cerebri 1
 Neuroanatomy by Dissection 17

Figure 2-22
13 1. Pituitary gland
2. Amygdaloid body in piriform lobe
3. Optic tract
11 4. Thalamus
6
10 5 12 5. Caudate nucleus
6. Lateral ventricle
7 7. Third ventricle
4 8. Hypothalamus
9 9. Interthalamic adhesion
3 10. Body of fornix
8 11. Corpus callosum
7 2 12. Internal capsule
13. Centrum semiovale
1

17 Figure 2-23
1. Cerebral arterial circle
2. Mamillary bodies
3. Optic tract
16 5 15 4. Third ventricle
5. Hypothalamus
12 13 6. Thalamus
8 11 14 7. Interthalamic adhesion
4 18 8. Body of fornix
10 9. Amygdaloid body in piriform lobe
6 7 10. Thalamocortical fibers
11. Caudate nucleus
4 12. Lateral ventricle
5 3 13. Corpus callosum

1 2 14. Internal capsule

9 15. Centrum semiovale
16. Cingulum in cingulate gyrus
17. Falx cerebri
18. Stria habenularis thalami
18 V eterinary N euroanatomy and C linic al N eurology

Figure 2-24
1. Parahippocampal gyrus
2. Hippocampus
3. Lateral ventricle 11
4. Third ventricle, caudal part 7 6
5. Thalamus
6. Crus of fornix 10
5 9
7. Choroid plexus in lateral ventricle
8. Internal capsule, just rostral to crus
cerebri
4
9. Thalamocortical fibers
10. Subarachnoid space 2 3
11. Corpus callosum 8
12. Trigeminal nerve
1
12

11

Figure 2-25
1. Crus cerebri
2. Mesencephalic tegmentum
3. Mesencephalic aqueduct 6
4. Subarachnoid space 8 5
5. Crus of fornix 10
6. Lateral ventricle
7. Substantia nigra 4
8. Hippocampus 12
9. Medial geniculate nucleus 3
9
10. Lateral geniculate nucleus 2
11. Falx cerebri 8
12. Pretectal nucleus
13. Trigeminal nerve
7 1
13
 Neuroanatomy by Dissection 19

Figure 2-26
1. Transverse fibers of pons, rostral part

9 6 2. Crus cerebri at longitudinal fibers of pons

3. Mesencephalic tegmentum
10 4. Mesencephalic aqueduct
5. Rostral colliculus
7 5 6. Lateral ventricle
9 7. Subarachnoid space

8 4 8. Brachium of caudal colliculus

9. Hippocampus
10. Crus of fornix
3 11. Trigeminal nerve

11 2
1

Figure 2-27
10 1. Transverse fibers of pons
2. Longitudinal fibers of pons
11 3. Mesencephalic tegmentum
9 4. Mesencephalic aqueduct
7 5. Caudal colliculus
6 6. Commissure of caudal colliculus
5 4 7. Subarachnoid space
8. Tympanic bulla
9. Hippocampus
3 10. Lateral ventricle
11. Splenium of corpus callosum
2
1

8
20 V eterinary N euroanatomy and C linic al N eurology

Figure 2-28
1. Pyramid
2. Pontine tegmentum
3. Trigeminal nerve
4. Caudal colliculus 12
5. Rostral cerebellar vermis and
rostral medullary velum
11
6. Commissure of caudal colliculus 6
7. Fourth ventricle 4 5
8. Middle cerebellar peduncle
10
9. Rostral cerebellar peduncle 9 7
10. Trochlear nerve in rostral 8
medullary velum 3
11. Subarachnoid space 2
12. Lateral ventricle
1

Figure 2-29
1. Pyramid
2. Rostral medulla
3. Fourth ventricle
4. Cerebellar vermis
5. Tympanic bulla
6. Cochlea in inner ear
7. Middle cerebellar peduncle 8 4
8. Rostral cerebellar peduncle 7 3
2 6
1

5
 Neuroanatomy by Dissection 21

6 Figure 2-30
12 1. Pyramid
2. Medulla
3. Fourth ventricle
4 4. Cerebellar vermis
5 5. Tentorium cerebelli osseum
11 6. Falx cerebri
7. Vestibulocochlear nerve–cochlear nuclei
10 8. Paraflocculus of cerebellum
9. Confluence of cerebellar penducles
8 9 3
10. Cerebellar medulla
11. Cerebellar hemisphere
7 12. Occipital lobe
2
1

10
Figure 2-31
7 1. Pyramid
2. Medulla
3. Fourth ventricle
4. Caudal cerebellar peduncle
9 8 5. Interposital cerebellar nucleus
6. Subarachnoid space
5 7. Cerebellar vermis
7 4 8. Cerebellar medulla
6 3 9. Cerebellar hemisphere
10. Occipital lobe
2
1
22 V eterinary N euroanatomy and C linic al N eurology

Figure 2-32
1. Pyramid
2. Medulla
3. Fourth ventricle near the obex
4. Caudal cerebellar peduncle
5. Cerebellar vermis 6
6. Paramedian lobule
7. Subarachnoid space 5
4
3
7
2
1

Figure 2-33
1. Pyramid at deccussation
2. Central canal
3. General proprioceptive nuclei
4. Subarachnoid space
5. Junction of medulla and first cervical
spinal cord segment
4
6. Branch of basilar or vertebral artery 3
5 2
1
6
C h a p t e r

3 Development of the Nervous

System: Malformation

NEURAL TUBE TELENCEPHALON: CEREBRUM Duplication of the Prosencephalon

Holoprosencephaly-Arrhinencephaly
Cell Differentiation Cell Bodies
Hydranencephaly
Axons Lissencephaly
MEDULLA SPINALIS: SPINAL CORD
MALFORMATIONS Cerebellar Malformations
Neural Crest Complex Malformation of Calves
Brain Malformations Occipital Bone Malformation and
MYELENCEPHALON: MEDULLA Hydrocephalus
OBLONGATA Syringohydromyelia
Hypoplasia of the Prosencephalon: Calvarial Ossification
METENCEPHALON: CEREBELLUM Cerebral (Telencephalic) Aplasia Spinal Cord Malformations
AND PONS (Anencephaly) Spina Bifida: Meningomyelocele
Meningocele: Meningoencephalocele
MESENCEPHALON: MIDBRAIN Pathogenesis of Malformations
Exencephaly case example 3-1
DIENCEPHALON: INTERBRAIN Lipomeningocele

Neural Tube the sacral, and a variable number of lumbar spinal cord
­segments. An opening may persist at the caudal end of the
The central nervous system is a tubular structure origi- neural tube, allowing communication with the subarach-
nating from a proliferation of ectod ermal epithelial cells noid space of the leptomeninges at the conus medullaris.
referred to as the neurectoderm, which is located dorsal to The rostral end of the neural tube develops rapidly and
the ­notochord along the axis of the embryo. This thickened ­produces three vesicles, from rostral to caudal: the pros-
ectoderm, known as the neural plate, invaginates along this encephalon, mesencephalon, and rhombencephalon (Fig.
axis, forming a groove until the lateral extremities of the 3-3). Early in its development the prosencephalon has lateral
original plate, the neural folds, meet centrally and fuse over enlargements, the optic vesicles, which grow laterally to con-
the neural groove to form a neural tube and canal. As the tact the overlying skin ectoderm. The further development of
neural tube forms, it separates from the nonneural ectoderm this primordial eye is described in Chapter 14, Visual System.
which grows over the dorsum of the tube to fuse along the Two additional swellings emerge from the rostral prosenceph-
midline. As this fusion and separation of ectodermal layers alon and grow out of the neural tube on each side laterally and
occurs, a longitudinal column of ectodermal epithelial cells dorsally. These telencephalic vesicles completely overgrow the
arises from the junction of nonneural and neural ectoderm original vesicular system and form the ­cerebral hemispheres.
and separates from these two structures when the neural The portion of the prosencephalon that remains at the ­rostral
tube is formed. These two bilateral columns, situated dor- end of the neural tube is the ­diencephalon. The optic ­vesicles
solateral to the neural tube throughout its length, are the remain associated with the diencephalon. The neural canal
columns of neural crest cells (Fig. 3-1). within the diencephalon is the third ventricle. It communi-
Closure of the neural tube progresses rostrally and cates rostrolaterally with the neural canal of each telencepha-
caudally from the level of the site of development of the lon (cerebral hemisphere), which is the lateral ventricle (first
rhombencephalon, the most caudal division of the brain. and second ventricles). This small communication on each
The caudal closure forms the majority of the spinal cord. side is the interventricular foramen. The nuclei of the thal-
Closure of the brain portion of the neural tube may initially amus and hypothalamus develop in the diencephalon. The
occur at multiple sites and progress rostrally and caudally. neurohypophysis is a ventral outgrowth of the dienceph-
The locations of these sites vary among species of animals. alon. The cerebral cortex and basal nuclei develop in the
Prior to complete closure, the most rostral opening is the telencephalon.
rostral neuropore (Fig. 3-2). The caudal portion of the spinal The neural canal of the mesencephalon is reduced to a
cord develops from the caudal end of the closed neural tube narrow tubular space called the mesencephalic aqueduct.
as an extension of a column of neuroepithelial cells that From the rostral rhombencephalon, the cerebellum
grows caudally on the midline between the notochord and or dorsal metencephalon develops dorsally. The remain-
skin ectoderm. A cavitation of this column of cells produces ing ventral metencephalon becomes the pons. The caudal
an extension of the neural tube and its neural canal. This rhombencephalon forms the myelencephalon or medulla
portion of the neural tube will ultimately form the ­caudal, oblongata. The fourth ventricle is the lumen of the neural

23
24 V eterinary N euroanatomy and C linic al N eurology

Neural plate

Neurectoderm

Neural
fold

Ectoderm Neural
groove

Notochord

Neural crest—Primordial ganglion

Neural canal

Neural tube

Figure 3-1 Development of the neural ­tube—transverse sections.

canal in the rhombencephalon. It communicates with the

Cell Differentiation
meningeal spaces that develop around the neural tube by In the first stage of development within the wall of the
way of openings that arise in the wall of the neural tube neural tube, the cells, which are commonly referred to as
caudal to the developing cerebellum. These openings are neuroepthelial or neurectodermal cells, are organized in a
called the lateral apertures (see Figs. 4-1 and 4-2). The neu- pseudostratified arrangement; thus the neural tube is one cell
ral canal continues caudally as the central canal of the in thickness. The cell membrane of each cell spans the full
spinal cord. width of the neural tube but the nuclei are located at various
levels within each cell. These cells are all mitotically active,
increasing the thickness of the wall of the tube. The nuclei
migrate within the cytoplasm of each cell, and their position
is dependent on the cell’s stage of ­mitosis.
During interphase, the nuclei are located on the exter-
nal surface of the neural tube. Chromosomal DNA duplica-
tion occurs with the nucleus in that position. As the nucleus
enters mitosis, it migrates through the cytoplasm to the
neural canal’s luminal surface. The peripheral portion of the
Rostral cytoplasm and the cell membrane also retract to the lumi-
neuropore nal position where cell division is completed. The two new
daughter cells extend their cytoplasm and cell membranes
back to the external surface of the neural tube. The nucleus
migrates back to the periphery again. In this position, this
cell can undergo another mitosis or it can differentiate.
Because the nucleus is at the external surface during inter-
phase, differentiation occurs at the external surface of the
neural tube. Thus, in a short time a new layer of differentiat-
ing cells appears on the external surface of the actively divid-
ing layer. The mitotic layer of cells is the germinal layer. The
cells undergoing differentiation form the mantle layer.
Caudal The cells that are differentiated are of two types: imma-
neuropore
ture neurons and spongioblasts. Immature neurons are the
primary parenchymal cells of the nervous system. They
are often referred to as neuroblasts but this is a misnomer
because once a neuron is formed it will not divide again
as the term neuroblast implies. The differentiated imma-
ture neuron grows extensively, forming long processes in
Figure 3-2 Dorsal view of neural tube closure. becoming a mature functioning cell, but it does not divide
 Development of the Nervous System: Malformation 25

Prosencephalon:
Telencephalon
Diencephalon
Optic cup

Mesencephalon

Rhombencephalon:
Metencephalon
Myelencephalon

A B
Mesencephalon Future mesencephalic aqueduct

Metencephalon

Myelencephalon III
Diencephalon

IV
Telencephalon

Otocyst Neurohypophysis
C Optic cup D

III
Telencephalon
(cerebrum)
III

IV
IV Metencephalon
(cerebellum and pons)

Myelencephalon
(medulla)
Spinal cord
E F
Figure 3-3 Development of brain vesicles. A, Three vesicle stages. B-F, Five vesicle stages, III, IV—ventricles.

again. Spongioblasts are the progenitors of the neurectoder- germinal layer of proliferating neuroepithelial cells. This
mal supporting cells of the nervous system, the neuroglia proliferative mitotic activity will ultimately be exhausted,
(glue). Two of the three forms of glial cells are derived from reducing the germinal layer to a single layer of cells ranging
these spongioblasts: astrocytes and oligodendrocytes (Fig. 3-4). from squamous to columnar and called ependymal cells.
The third glial cell is the microglial cell, which is mesoder- These ependymal cells line the entire lumen of the neural
mal in origin. It is a monocyte that enters the nervous sys- tube, which includes the ventricular system in the brain and
tem from its blood supply. the central canal of the spinal cord. Peripheral to this ger-
As the primitive neurons and spongioblasts are differ- minal layer in the embryonic neural tube is the thick layer
entiated and grow and the neurons produce processes, the of differentiated cells, the immature neurons, and spongio-
­neural tube becomes arranged in three concentric layers blasts. They form the mantle layer, which will ultimately
(Fig. 3-5). Adjacent to the lumen of the neural tube is the become the gray ­ matter of the definitive spinal cord, the
26 V eterinary N euroanatomy and C linic al N eurology

Blood vessel and astrocyte

Spongioblast

Central sensory neuron

Neurons

Motor neuron

Oligodendrocyte

Spongioblast

Figure 3-4 Mitosis and differentiation of neuroepithelial cells.

nuclei of the brainstem, the nuclei and cortex of the cer- These axons will be myelinated by the oligodendroglial cells
ebellum, and the basal nuclei and cerebral cortex of the ­forming tracts in the white matter.
telencephalon. The latter requires a migration of these neu- From the mesencephalon caudally, a longitudinal groove,
rons from the mantle layer to the external surface of the the sulcus limitans, appears in the lateral wall of the neural
neural tube. The external layer of the neural tube is the mar- canal. Thus the neural canal can be artificially divided into
ginal layer; it is composed primarily of the growing axonal dorsal and ventral portions by an imaginary dorsal plane at
processes of the neuronal cell bodies in the mantle layer. the level of this sulcus. The dorsal portion is called the alar
Embryonic Definitive

Marginal White

Mantle Grey

Germinal Ependymal

Sulcus limitans

GSA Roof plate

GP
SP Alar plate
SSA (sensory)
SVA
GVA

GVE
Basal plate
(motor)
GSE

Floor plate

Figure 3-5 Functional organization of the neural tube. GP, General proprioception; GSA, general somatic afferent;
GSE, general somatic efferent; GVA, general visceral afferent; GVE, general visceral efferent; SP, special proprioception;
SSA, special somatic afferent; SVA, special visceral afferent.
 Development of the Nervous System: Malformation 27

plate and the ventral portion the basal plate. Functionally layer, and the general somatic efferent column is located ventrally in
the alar plate mantle layer is concerned predominantly the basal plate of the mantle layer. Because the relative size of
with sensory systems, and the basal plate mantle layer with the components of each spinal cord segment depends on the
motor systems (see Fig. 3-5). volume of tissue being innervated, at the levels of the limbs
the spinal cord segments responsible for their innervation are
enlarged forming the cervical and lumbosacral intumescences.
Medulla Spinalis: Spinal The ultimate growth to maturity of a neuron in the peripheral
Cord (See Fig. 2-17) nervous system depends on its appropriate innervation of a
muscle cell (general somatic efferent [GSE]) or formation of a
In this text, the term spinal cord is used rather than medulla peripheral receptor (general somatic afferent [GSA], general
spinalis, the nomenclature preferred by Nomina Anatomica proprioception [GP]). The lack of such innervation results in
Veterinaria. The spinal cord provides the best example of the degeneration of that neuron. In the cervical and thora-
the symmetric development of the neural tube by layers. columbar regions where appendages are not innervated, the
Ventral growth of the two basal layers and associated mar- immature primitive neurons in the basal plate mantle layer
ginal layers beyond the level of the floor plate (Fig. 3-6) and the adjacent spinal ganglion that fail to innervate struc-
leaves a separation between the two sides, which is the ven- tures will degenerate by a process of cell death referred to as
tral median fissure. The mantle and marginal layers of the apoptosis. The shape of the ­ventral gray column depicts this
alar plates grow dorsally. The dorsal marginal ­layers fuse on process.
the median plane to form a dorsal median septum that may In the basal plate mantle layer, the GSE neurons located
be poorly defined. The external margin of this septum forms medially innervate the axial skeletal muscles. Those located
the dorsal median sulcus. This midline growth displaces the laterally innervate the appendicular skeletal muscles.
roof plate region ventrally, resulting in a reduction of the Within these areas of the ventral gray column, the GSE neu-
neural canal to form the small central canal of the spinal ronal cell bodies can be further grouped according to the
cord lined by ependymal cells. The mantle layer of the alar specific peripheral nerve that contains their axon and by
plate becomes the dorsal gray column (also referred to as the ­specific muscles innervated.
horn), and that of the basal plate becomes the ventral gray The growth of axons of the basal plate neurons through
column. The mantle zone at the plane of the sulcus limitans the marginal layer and outside the neural tube forms the ven-
becomes the intermediate gray column (see Fig. 3-6). tral root and part of the spinal nerve and further branching
Not only is there a gross topographic differentiation of of the peripheral nerves. This includes the general somatic
function of primitive neurons between the alar and basal efferent neurons located in the ventral gray column and the
plates, but within the mantle layer of each plate, neurons are general visceral efferent neurons located in the intermedi-
further arranged in functional columns. The general visceral ate gray column adjacent to the sulcus limitans. These latter
afferent and general visceral efferent neurons are located adja- general visceral efferent (GVE) neurons are the preganglionic
cent to each other in their respective gray columns on either lower motor neurons of the autonomic nervous system. This
side of the dorsal plane through the sulcus limitans. The intermediate gray column is only present in the thoracic,
­general somatic afferent and general proprioceptive neuronal cranial lumbar and sacral spinal cord segments. In the other
columns are located dorsally in the alar plate of the mantle segments, it was present in the embryo but subsequently

Layers Roof plate

Germinal Alar
mantle lamina

Mantle
Basal
mantle
Marginal
lamina

Floor plate

Dorsal median septum

Gray columns
GSA
GP Dorsal
GVA
GVE Intermediate

GSE Ventral

Ventral median fissure

Figure 3-6 Development and functional organization of the spinal cord.

28 V eterinary N euroanatomy and C linic al N eurology

degenerated due to the absence of a target organ or biochem- tube. Along the developing spinal cord segments, these cells
ical attractant. These GVE neurons terminate in ganglia in ­provide the neurons that form the spinal ganglia at each
the peripheral nervous system that contain cell bodies of segment. Adjacent to each somite a proliferation of neural
the postganglionic neurons in this two-neuron lower motor crest cells forms the segmental spinal ganglion (see Figs. 3-1
neuron system (Figs. 3-7 and 3-8; see also Fig. 3-6). and 3-7). One portion of the axon that emerges from each
of these cell bodies grows centrally into the spinal cord seg-
ment to enter the alar plate dorsal gray column forming the
Neural Crest dorsal root. The other portion of the axon grows distally to
The neural crest cells are the cell bodies in the longitudi- form a ­sensory component of the spinal nerve and further
nal column of cells that formed dorsolateral to the neural branches of the peripheral nerves. The point of penetration

Dorsal gray column Axon of dorsal root

Neural crest cells

of spinal ganglion

Intermediate
gray column

Ventral gray column

Axons of
ventral root

Neural crest cells

Neural groove

Primordial ganglion
Ectoderm

Neural canal

Neural tube

Spinal ganglion

Figure 3-7 Spinal ganglia development from the neural crest.

 Development of the Nervous System: Malformation 29

Melanoblasts Spinal ganglion

Sympathetic trunk
ganglion

Abdominal autonomic
plexus ganglion Adrenal medulla

Autonomic neuron

Dorsal funiculus

Dorsal root

Lateral funiculus
Spinal ganglion

Spinal nerve
Ventral funiculus Ventral root

Sympathetic trunk ganglion

Abdominal autonomic
plexus ganglion

Adrenal

Viscera

Figure 3-8 Neural crest contribution to the development of GVE neurons.

of the marginal white matter layer of the spinal cord seg- the same endocrine substance, norepinephrine, that is the
ment by the axons in the dorsal and ventral roots divides neurotransmitter released at the telodendron of the sym-
the spinal cord white matter processes into three regions pathetic postganglionic axon derived from the neural
called funiculi. These are the dorsal, lateral, and ventral crest cells. Although the melanoblasts and GVE neurons
funiculi on each side of the spinal cord. seem unrelated, their common denominator is the unique
The formation of spinal ganglia is only one of many out- ­metabolism of tyrosine, which provides melanin for the
comes of neural crest differentiation. Prior to its segrega- melanocytes and the norepinephrine for the neuron. In
tion into spinal gangli, an early migration of cells from this addition, there is extensive migration of the neural crest
­neural crest column provides melanoblasts to the somitic cells into the wall of the developing gastrointestinal tract.
dermatome and adjacent epidermis as well as to the cell These will form the postganglionic neurons of the parasym-
bodies of postganglionic axons in the two-neuron GVE pathetic portion of the GVE lower motor neuron as well
system. as interneurons; and they form the glial cells that develop
These latter cell bodies form the ganglia of the sympa- in the wall of the bowel, creating what is referred to as the
thetic trunk and the abdominal plexus sympathetic ganglia enteric nervous system. The latter is extremely extensive
as well as the medullary cells in the adrenal gland (see Fig. and complex and presumably is entirely derived from the
3-8). These adrenal medullary cells do not grow any pro- neural crest cells. A similar migration forms the postgan-
cesses but synthesize and elaborate into the blood stream glionic ­parasympathetic ­neurons for the urogenital system.
30 V eterinary N euroanatomy and C linic al N eurology

In addition to these nervous system structures, the neural as neurobiotaxis. As a result of this migration, the axons
crest contributes to the formation of bone and cartilage in leaving this facial nucleus initially course dorsomedially to
the skull and derivatives of the branchial arches; to the wall the floor of the fourth ventricle before turning to course
of the great vessels at the base of the heart; and to thyroid ventrolaterally to leave the medulla and form the facial
parafollicular (C) cells, odontoblasts, a portion of the lep- nerve. The GSE cell bodies of cranial nerves IX, X, and XI
tomeninges and the lemmocytes, Schwann cells, that form undergo a similar migration ventrolaterally and accumulate
the myelin of the peripheral nervous system. This is an in nucleus ambiguus, which is well named because it is a
amazing display of developmental capabilities in an initial poorly defined nucleus. The preganglionic neurons of the
column of cells. When a student is asked about the origin of parasympathetic portion of the GVE system are located in
a structure, if there is any doubt, relying on neural crest is a an interrupted column just ventromedial to the sulcus limi-
worthwhile consideration! tans, similar to their location in the spinal cord. Their axons
leave the medulla in cranial nerves VII, IX, X, and XI.
The sensory components of cranial nerves associated
Myelencephalon: Medulla with the medulla arise primarily from primitive neurons
Oblongata (See Figs. 2-11 through 2-16) that develop from the column of neural crest cells, with a
few arising from ectodermal cells that proliferate from bran-
In this text, the term medulla oblongata is shortened to medulla. chial arch ectoderm. The latter areas are referred to as cranial
The medulla is the most caudal portion of the brainstem and placodes. These two sources form the sensory ganglia of cra-
is continuous caudally with the spinal cord. The basic forma- nial nerves VII, IX, and X, which are concerned with general
tion of the medulla involves only a slight modification of the visceral afferent and special visceral afferent (taste) function.
development described for the spinal cord. The narrow mid- Ectodermal cells derived from the otic placode form the sen-
dorsal roof plate of the initial neural tube (see Figs. 3-5 and sory ganglia of cranial nerve VIII, which is concerned with
3-6) is stretched extensively instead of being obliterated by special proprioception for vestibular system function and
the proliferating alar plate and marginal tissue as it is in the with the special somatic afferent system for auditory func-
spinal cord. Imagine grasping the midline roof plate of the tion. These cranial nerve VIII ganglia are located in the inner
neural tube with both hands and then pulling your hands ear within the petrosal portion of the temporal bone. Their
apart sideways. This would stretch out a thin layer of neural axons course into the alar plate region of the medulla to syn-
tube (roof plate) and displace the entire alar and basal plates apse on cell bodies comparable to the dorsal gray column
to a lateral and ventral position (Fig. 3-9). This would enlarge cell bodies in the spinal cord (see Fig. 3-9).
the lumen of the neural tube to form the fourth ventricle of The leptomeninges that surround the entire developing
the medulla, which is covered dorsally by only a single cell central nervous system (CNS; neural tube) arise from neu-
layer of neuroepithelial cells. At this site these cells will not ral crest cells and adjacent mesodermal cells. These menin-
enter mitosis but will remain as a single layer of ependymal ges contain the bulk of the blood vessels that supply the
cells. The sulcus limitans that is present on the ventrolateral CNS and the roots of the peripheral nerves. Dorsal to the
wall of the fourth ventricle provides the plane of division of stretched-out roof plate of the fourth ventricle, the capil-
the medulla into a ventromedial basal plate and a dorsolat- lary blood vessels proliferate to form the two longitudinal
eral alar plate, which have the same functional significance rows of a dense capillary bed. The adjacent ependymal cells
as in the ­spinal cord development. enlarge into cuboidal cells, and the entire structure (cuboi-
Throughout the brainstem, the mantle layer of the neural dal ependymal cells, pia mater, and capillary bed) hangs
tube is broken up into nuclei that are collections of ­neuronal down into the lumen of the fourth ventricle (Fig. 3-10). This
cell bodies with a common purpose, and they are interspersed is called the choroid plexus of the fourth ventricle. By strict
with neuronal processes. Some nuclei are more distinct than definition, only the proliferated capillary bed is the plexus.
others. The functional columns described in the spinal cord Thus the choroid plexus of the fourth ventricle comprises
have a similar location in the brainstem. In addition, there two sagittal lines parallel to and on either side of the median
are neurons in the medulla that are organized into functional plane. These extend from the caudal part of the fourth ven-
groups that are present only in cranial nerves (see Fig. 3-9). tricle rostrally to the level of the cerebellar peduncles where
In domestic animals, cranial nerves VI through XII and each plexus turns laterally. At this point there is an opening
the trapezoid body are part of the medulla. The rostral that develops in the medullary roof plate, called the lateral
­border of the medulla is the caudal border of the pontine aperture. This aperture allows communication between the
transverse fibers. In the primate and some nondomesticated lumen of the fourth ventricle and the subarachnoid space
species, the transverse fibers of the pons expand caudally that develops in the leptomeninges. At the level of this lat-
to cover the trapezoid body, so cranial nerves VI through eral aperture, the choroid plexus protrudes from the lumen
VIII are included with V in the pons. Cranial nerves VI, VII, of the fourth ventricle out through the aperture, where it
IX, X, XI, and XII contain general somatic efferent neu- is visible on each side at the cerebellomedullary angle (see
rons. The medullary nuclei of cranial nerves VI and XII are Fig. 3-10). The aperture itself is invisible grossly because it is
located in an interrupted column along the median plane filled with this choroid plexus. The choroid plexus is a major
adjacent to the fourth ventricle. The hypoglossal nucleus is site of formation of cerebrospinal fluid (see Chapter 4).
very long. The GSE neuronal cell bodies of the facial nerve In domestic animals, the lateral aperture is the only com-
have migrated from their initial formation in the mantle munication between the ventricular system of the brain
layer to form the facial nucleus in a ventrolateral position, and the subarachnoid space, which makes it critical for the
which is closer to their common source of sensory stimuli maintenance of normal intracranial pressure. Primates have
coursing into the medulla in the spinal tract of the trigem- an additional aperture located caudally in the caudal med-
inal nerve. This phenomenon of migration is referred to ullary velum of the fourth ventricle.
 Development of the Nervous System: Malformation 31

Roof plate (ependyma)

Sulcus limitans

GSA
GP
SP
SSA

SVA

GVA

GVE Alar plate

Basal plate
GSE
Floor plate

SP:
Vestibular
CN VIII
GSA
GP Fourth ventricle
SSA:
SP
Cochlear
SSA SVA:
GVA:
SVA CN VII, IX, X
GVA Visceral receptors
Taste
GVE
GVE:
GSE
CN VII, IX
Lacrimal and salivary
glands via autonomic
ganglia
CN X
Smooth muscle and glands
of throacic and abdominal
organs via autonomic
intramural ganglia
GSE:
CN VII
Facial mm
CN IX, X
Palatal pharyngeal mm
CN X, XI
Laryngeal mm
GSE:
CN VI
Extraocular mm
CN XII
Tongue mm

Figure 3-9  Functional organization of cranial nerves VI to XII in the myelencephalon. CN VI, Abducent; CN VII,
facial; CN VIII, vestibulocochlear; CN IX, glossopharyngeal; CN X, vagus; CN XI, accessory; CN XII, hypoglossal; GP,
general proprioception; GSA, general somatic afferent; GSE, general somatic efferent; GVA, general visceral afferent;
GVE, general visceral efferent; SP, special proprioception; SSA, special somatic afferent; SVA, special visceral afferent.

Metencephalon: Cerebellum of the pons, and form a small, well-defined motor nucleus.
AND PONS (See Figs. 2-9 and 2-10) These neurons function in the general somatic efferent sys-
tem and innervate the muscles of mastication derived from
The initial development of the metencephalon is comparable the somitomeres in branchial arch 1. The sensory neurons
to that of the myelencephalon. Cranial nerve V, the trigemi- in cranial nerve V are derived primarily from neural crest
nal nerve, is associated with this segment of the brainstem cells and form the trigeminal ganglion. Most of these neu-
(Fig. 3-11). Its motor neurons arise in the basal plate mantle rons are GSA and their dendritic zones are widely spread over
layer, migrate a short way ­ventrolaterally into the ­parenchyma the entire surface of the head and to the inner ­surface of the
32 V eterinary N euroanatomy and C linic al N eurology

Capillary network form the sensory pontine nucleus of the trigeminal nerve in
Ependyma
the pons and the nucleus of the spinal tract of the trigeminal
nerve in the medulla. This spinal tract and nucleus extends
Pia mater the full length of the medulla, where caudally they meet the
comparable functional neurons developing in the first cervi-
cal spinal nerves and spinal cord segment (see Fig. 3-11).
The cerebellum, or dorsal metencephalon, is formed pri-
Fourth ventricle
marily from the proliferation of the germinal epithelial cells
of the alar plate, forming the rhombic lip (see Figs. 3-11 and
Medulla 3-12). This growth dorsolaterally from each side overgrows
the roof plate of the fourth ventricle so that the cerebellum
forms part of the dorsal boundary of the fourth ventricle
in the metencephalon. The development of the cerebellar
Cerebellar peduncles
cortex and nuclei are described in Chapter 13. The ventral
Choroid plexus metencephalon is the pons. A ventral migration of alar plate
mantle layer neurons forms the pontine nucleus (see Fig.
3-12). The axons of these neurons cross the midline and
course dorsally into the cerebellum. This forms the trans-
verse fibers of the pons, which demarcate the ventral sur-
face of the pons and the middle cerebellar peduncle.

Mesencephalon: Midbrain
(See Figs. 2-5 through 2-9)
Symmetric proliferation of the walls of the neural tube in
Medullary velum Lateral aperture the mesencephalon reduces the size of the neural canal to
of fourth ventricle a narrow tube, the mesencephalic aqueduct. This is smaller
Cerebellomedullary rostrally, where it joins the third ventricle of the dienceph-
angle
alon, and larger caudally, where it is continuous with the
Figure 3-10 Development of the roof plate and choroid plexus of the fourth ventricle beneath the rostral medullary velum.
fourth ventricle. Cranial nerves III (oculomotor) and IV (trochlear) are
associated with the midbrain. These contain primarily GSE
upper respiratory and digestive systems. A smaller compo- neurons that innervate extraocular muscles. The cell bodies
nent is composed of general proprioceptive neurons for the are in nuclei derived from the basal plate mantle layer. They
muscles and joints in the head region. These sensory neurons do not migrate but remain adjacent to the median plane
greatly ­outnumber the motor neurons. Therefore, when these ventral to the aqueduct, which is in the same ­topographic
sensory axons enter the alar plate region of the metencepha- nuclear column as the abducent and hypoglossal GSE nuclei
lon, they spread out for a short distance rostrally and for a in the medulla (Fig. 3-13). The oculomotor nucleus also con-
long distance caudally, forming the spinal tract of trigemi- tains the neuronal cell bodies of the preganglionic parasym-
nal nerve in the pons and medulla. These axons terminate pathetic innervation to the ­ constrictor muscle of the iris.
in telodendria at synapses in the alar plate neurons, which These are derived from the same basal plate mantle layer.

Trigeminal tract (CN V)

Metencephalon (pons)

Figure 3-11 Development of the metencephalon: surface view and Trigeminal ganglion
sagittal view of the afferent portion of cranial nerve V.
Developing
cerebellum
Rhombic
lip
Velum
of
fourth Pons
ventricle
 Development of the Nervous System: Malformation 33

Roof plate of
fourth ventricle
Purkinje cell
Rhombic lip
Granular cells (source of cerebellum)
Cerebellar
nuclei Trigeminal ganglion
Germinal cells
GSA: Figure 3-12 Development of the metencephalon:
Basal CN V transverse section, pontine nucleus.
Sensory from head
Alar

GSE:
Pontine nuclei
CN V
Motor to muscles
of mastication

Tectum:
Corpora quadrigemina
Tectum Rostral colliculus (visual reflex)
Alar Caudal colliculus (auditory reflex)

Basal GSE:
Tegmentum CN IV
To dorsal oblique muscle
Substantia nigra Figure 3-13 Development of the mesencephalon: transverse
Crus cerebri section.
Tegmentum:
Red nucleus
Reticular formation
GSE:
CN III
To extraocular muscles
GVE:
CN III
To smooth muscle of eyeball
via ciliary ganglion

The alar plate proliferates dorsally to form the tectum of the mesencephalic aqueduct and rostrally at the level of the
the midbrain, which is divided into paired rostral and caudal interventricular foramina, which connect to each telence­
colliculi, collectively known as the corpora quadrigemina. phalic lateral ventricle (Fig. 3-14). On the dorsal median
These are associated with visual and auditory reflex func- plane of the diencephalon, there is no proliferation of the
tion, respectively. The crus cerebri on the ventral surface of neural tube epithelial cells, leaving a single-cell-thick roof
the midbrain results from the caudal growth of descend- plate, where a small choroid plexus develops in two parallel
ing axons from telencephalic projection neurons. These are lines similar to those in the medulla. At the interventricu-
continuous from the internal capsule in the diencephalon. lar foramina each of these is continuous with the choroid
plexus that develops in each lateral ventricle (Fig. 3-15).
In the diencephalon, a plethora of nuclei are formed from
Diencephalon: Interbrain the mantle layer, which are dispersed diffusely through this
(See Figs. 2-3 through 2-6) brain segment forming a complex of nuclei and neuronal
processes.
Rostral to the mesencephalon, the sulcus limitans is no lon- These nuclei form the thalamencephalon, hypothala-
ger evident in the neural tube and the diencephalon and mus, and subthalamus. The thalamencephalon consists
telencephalon are considered to be developments of the of the thalamus, metathalamus, and epithalamus, which
alar plate. The symmetric development of the lateral walls comprise those nuclei located dorsal to the ventral por-
of the neural tube in the diencephalon reduces the neural tion of the third ventricle. The hypothalamus includes the
canal to a vertical slit on the median plane, the third ven- nuclei located on the sides and floor of the ventral portion
tricle. Adhesion of the developing thalamus in the center of the third ­ventricle. The subthalamic nuclei are located
forms the interthalamic adhesion and separates the third ­ventrolaterally in the diencephalon. A ventral outgrowth of
ventricle into a small dorsal component and a larger ven- the ­hypothalamus, including an extension of the third ven-
tral component. These two portions converge caudally at tricle, forms the neurohypophysis. The neurohypophysis
34 V eterinary N euroanatomy and C linic al N eurology

Lateral ventricle
Choroid plexus Choroidal
fissure

Roof
plate
Third
ventricle
Third
ventricle

A B
Cut edge of roof plate
Interthalamic adhesion
Choroidal fissure
Interventricular foramen

Epithalamus

Lamina
Thalamus
terminalis

B C

Hypothalamus D

Optic chiasm A Mesencephalic aqueduct

Third ventricle Hypophysis

Mesencephalic aqueduct

Sulcus
limitans

D
Figure 3-14 Relationship of the diencephalon and mesencephalon. A, Transverse section of mid-diencephalon.
B, Transverse section of caudal diencephalon. C, Transverse section of rostral mesencephalon. D, Transverse section
of caudal mesencephalon.

becomes associated with a dorsal extension of the adjacent II, the special somatic afferent neurons of the visual system.
oral ectoderm, the hypophyseal (Rathke’s) pouch, to form the By definition, a nerve is a collection of axons outside the
hypophysis (pituitary gland). The optic vesicles that initially CNS that are myelinated by Schwann cells, which arise from
grew out of the prosencephalon will form the neural layer of the neural crest. Optic nerves are misnamed because they
the eye and optic nerves, which become ­associated with the develop as extensions of the prosencephalon. They form in
­diencephalon (see Fig. 3-3). The axons that grow caudally the optic stalk that extends from the diencephalon to the
from the retina in the optic nerve will form the optic tracts optic cup. Their axons are myelinated by CNS oligodendrog-
of the diencephalon, and many will terminate in a nuclear lial cells. This is important to remember because the optic
area of the thalamus. These optic nerves form cranial nerve nerves are affected by diseases that are specific to the CNS.
 Development of the Nervous System: Malformation 35

Choroid plexus

Lateral ventricle

Interventricular
Third ventricle foramen
A
A
Lateral ventricle B

Choroid
plexus

Third ventricle

Fornix
Choroidal fissure

Cut edge of roof plate

Arrow in
lateral ventricle

Lamina terminalis
Third ventricle

Figure 3-15 Development of choroid plexus and ventricular system of diencephalon and telencephalon.

Therefore, optic neuritis is a form of encephalitis. A polyneu- to form the two cerebral hemispheres that comprise the
ritis does not affect the optic nerves. cerebrum. The lamina terminalis is located on the median
plane between these two outgrowths. The telencephalic
­vesicles grow out of the prosencephalon a short distance ros-
Telencephalon: Cerebrum trally and then in a large curve caudally and ventrally. The
(See Figs. 2-2 through 2-7) neural canal in each telencephalon is the lateral ­ventricle,
which communicates with the diencephalic third ­ventricle
The rostral boundary of the brainstem is the lamina termi- via the interventricular foramen on each side of the lamina
nalis of the diencephalon. It is the rostral boundary of the ­terminalis (see Figs. 3-15 and 3-18).
third ventricle. The optic chiasm is located at the ventral At one aspect of the medial wall of the telencephalic
portion of this lamina, and the rostral commissure develops ­vesicle, the neuroepithelial layer of the neural tube does
in and remains in this lamina. It is at this level that the tel- not proliferate and remains a single layer of cells that
encephalic vesicles grow out of the original ­prosencephalon become ependymal cells comparable to the roof plate of
36 V eterinary N euroanatomy and C linic al N eurology

the ­myelencephalon over the fourth ventricle and the roof portions of the cerebrum. The neopallium is a more recent
plate of the diencephalon over the third ventricle. As the evolutionary brain development and makes up the surface
rest of the telencephalon proliferates and differentiates, this of all the gyri of the cerebrum (Fig. 3-16).
telencephalic roof plate will be attached to the crus of the Comparative evolutionary studies show the continual
hippocampal fornix on one side and the stria terminalis on development of the neopallium in higher animals, relegat-
the other side. The choroid plexus of each lateral ventricle ing the archipallium and paleopallium to a lesser portion
develops in this roof plate as was described for the choroid anatomically.
plexus in the medulla. This is a curved structure similar to The surface of the amphibian cerebrum is smooth, lack-
the structures that it is attached to, and it protrudes into the ing any gyri. It is composed of the archipallium dorsally, the
lumen of the lateral ventricle. At each interventricular fora- paleopallium laterally, and the basal nuclei ventrally. In the
men, each lateral ventricular choroid plexus is continuous advanced reptile, the basal nuclei have receded from the ven-
with the choroid plexus of the third ventricle. tral surface and have been replaced by the paleopallium on
An extensive development of projection axons occurs the ventral surface. A small lateral area is neopallium and the
from diencephalic thalamic neurons to the cerebrum and dorsal area is archipallium. In mammals, the neopallium has
telencephalic neurons to the brainstem. This gives rise to a overgrown the other divisions of the cerebral cortex so that
thick layer of myelinated processes, white matter, between the paleopallium is entirely on the ventral surface of the cere-
the diencephalon and telencephalon that is known as the brum, ventral to the rhinal sulcus, and the archipallium is
internal capsule. rolled medially into the lateral ventricle as an internal gyrus,
Telencephalic neuronal cell bodies and white matter can the hippocampus. Continual development of the neopallium
be organized as follows. in higher mammals has resulted in the characteristic gyri and
sulci observed over most of the exposed surface of the cere-
brum. The rhinal sulcus separates the neopallium from the
Cell Bodies paleopallium. This is characteristic of all of the domestic ani-
The telencephalic neuronal cell bodies are located in one of mal species, but most laboratory rodents and all birds have
two general locations. One is on the external surface of the no gyri because in these animals the neopallium is unfolded,
entire telencephalon, forming the various layers of the cere- so the cerebrum has a smooth surface. Although at birth a few
bral cortex. The other is deep to the surface in subcortical gyri and sulci are present in the puppy brain, they increase
basal nuclei. These are often incorrectly called basal ganglia. remarkably during the first 3 to 6 weeks of life.
Remember that ganglia are collections of neuronal cell bodies
outside the CNS. Such collections inside the CNS are nuclei
or cortices. Cortices are located superficially, and their neu-
Axons
ronal cell bodies are in a continuous arrangement. Examples The axons of telencephalic neurons form three groups of
of basal nuclei are the caudate nucleus, globus pallidus, puta- processes on the basis of their destinations. The association
men, claustrum, and amygdaloid body. The cerebral cortex axons course between cortical areas within one cerebral
can be divided into three regions based on evolutionary and hemisphere. They can be short and course between adjacent
anatomic features. The archipallium (pallium is a synonym gyri or long and traverse the entire cerebral hemisphere, but
for cortex) is the hippocampus, which is an internal gyrus, they never leave that hemisphere. Projection axons leave
an area of cerebral cortex that has been rolled into the lateral the cerebral hemisphere where their cell bodies are located
ventricle and is not visible on the surface of the cerebrum. and enter the brainstem via the internal capsule. They ter-
The paleopallium is the olfactory system and is composed minate in nuclei in various parts of the brainstem, with a
of the olfactory bulbs, the peduncles, and the piriform lobe few reaching the spinal cord. Commissural axons cross from
cortex. In animal evolution, these are the most primitive one cerebrum to the other (Fig. 3-17). All of these axons are

Cerebral cortex

Subcortical-basal
nuclei Neopallium

Lateral ventricle

Caudate nucleus
Figure 3-16 Development of the
telencephalon.
Internal capsule

Lentiform nucleus
Lateral rhinal sulcus
Amygdala Paleopallium (olfactory bulb,
peduncle, and pyriform lobe cortex)

Archipallium (hippocampus and fornix)

 Development of the Nervous System: Malformation 37

their axons grow centrally, forming the white matter on the

inside of the gray matter. Radial astrocytes participate in
this migration by guiding the neurons to the surface of the
neural tube. Ultimately, six layers of neurons will populate
Commissural the cortex and are numbered from superficial (layer I) to
fibers deep (layer VI). The first neurons to migrate to the devel-
(corpus callosum) oping cerebral cortex will form layer VI. As more neurons
arrive, they pass by those already there to form the rest of
the ­layers in a reverse sequence. Thus the last to arrive form
Projection fibers layer I. The basal nuclei are formed by neurons that migrate
(internal capsule) only a short distance from the mantle layer into the devel-
oping white matter so that they remain in a subcortical
position. Remnants of the telencephalic germinal layer per-
sist throughout the life of the animal, forming the subepen-
Association fibers
dymal layer, which consists of a variably sized population of
(long and short)
small cells that are thought to be a continuous source of glia
and neurons throughout the life of the animal. Postnatal
neurogenesis is now a well-recognized event, especially in
the olfactory system and the hippocampus. This subepen-
dymal-subventricular layer is thought to be the source of
some of the glial neoplasms that arise in the brain.
Figure 3-17 Development of the neuronal processes in the telencephalon, The development of the choroid plexuses of the lateral,
white matter. third, and fourth ventricles is similar, but the adult mor-
phology varies. The choroid plexuses of the lateral and
third ventricle are small and form a thin, undulating veil
intermixed in the white matter of each gyrus, which is the that projects into the ventricle. The fourth ventricle cho-
corona radiata, and in the centrum semiovale, which is roid plexus is more robust and lobulated, especially where it
the mass of white matter in the center of the cerebrum dor- projects through the lateral apertures.
sal to the lateral ventricle. The semioval appearance of this This is especially evident in horses and cattle. In addition, in
structure can be appreciated only in a dorsal plane section older horses there often is an accumulation of cholesterol crys-
of the cerebral hemisphere. tals resulting from chronic bleeding that enlarges the choroid
There are three groups of commissural axons, all of which plexus. The accumulation of these crystals may present a spar-
initially develop in the lamina terminalis (Fig. 3-18). The ros- kling appearance in the choroid plexus. Occasionally, in the
tral commissure is located ventrally in the lamina terminalis choroid plexus of the lateral ventricle, this chronic bleeding
and courses primarily between paleopallial structures and results in such a large accumulation of cholesterol crystals and
basal nuclei (the amygdaloid body) on each side. This com- associated chronic inflammation that the mass causes neuro-
missure remains at this site dorsal to the optic chiasm in the logic signs associated with increased intracranial pressure. This
fully developed brain. Another small group of ­commissural cholesterinic granuloma can be unilateral or bilateral (Figs.
axons courses between the archipallium (hippocampus) of 3-19 and 3-20) and can be diagnosed by computed tomogra-
each side. This commissure migrated caudodorsally as the phy (CT) or magnetic resonance (MR) imaging. They can be
telencephalon developed to reach a position dorsal to the very large when neurologic signs occur. The choroid plexus
caudal aspect of the diencephalon. can give rise to choroid plexus papillomas and carcinomas.
The largest group of commissural axons forms the cor- The latter can spread to other sites by way of the cerebrospi-
pus callosum, which also expands dorsally as the cerebrum nal fluid (CSF) in the subarachnoid space. Meningiomas arise
develops so that it is positioned between the other two com- from arachnoid cells and expand in the subarachnoid space.
missures. The corpus callosum serves primarily to connect Meningiomas that bulge into the ventricular system arise from
the neopallial areas of each cerebral hemisphere. It begins the arachnoid cells associated with the choroid plexus.
in the lamina terminalis and as the telencephalic vesicle
expands, the corpus callosum enlarges and extends cau-
dally dorsal to the diencephalon. Near the median plane
it is located between the neopallial cingulate gyrus dorsally
Malformations
and the archipallial hippocampus and body of the fornix
ventrally. Laterally in each cerebral hemisphere, the ­corpus
Brain Malformations
callosum forms the roof of the lateral ventricle. The ­septum
pellucidum develops dorsally in the lamina terminalis
Hydrocephalus
between the genu of the corpus callosum and the rostral Many circumstances can cause the ventricular system to
body of the fornix (see Fig. 3-18). enlarge. It is usually most evident in the lateral ventricles, and
In the telencephalon, the neural tube germinal layer the most common cause is an interference with the flow of
­ultimately is replaced by the ependyma of the lateral ven- CSF from the ventricles into the subarachnoid space, which
tricle. Except for the area of the basal nuclei, the mantle and results in increased CSF pressure. This hypertensive hydro-
­marginal layers reverse their positions. This is the result of cephalus causes extensive degeneration of the ­telencephalon,
the migration of the newly formed primitive neurons to the especially the neopallium. The subject of hydrocephalus is
surface of the neural tube to form the cerebral cortex, where considered in the discussion of CSF in Chapter 4.
 Corpus callosum Hippocampal commissure

Corpus callosum

Septum
pellucidum

Diencephalon

Optic chiasm
Rostral commissure

Lamina terminalis

Fornix
Corpus callosum

Hippocampal commissure

Septum pellucidum

Rostral commissure
Third ventricle Corpus callosum
Splenium

Optic chiasm

Genu Body of fornix

Septum pellucidum Hippocampus

Columna fornicis
Crus fornicis

Figure 3-18 Development of telencephalic commissural pathways.

Figure 3-19 Transverse section of the brain of a 14-year-old Morgan horse with Figure 3-20 Transverse section of the brain of a 10-year-old Percheron horse
a unilateral cholesterinic granuloma of the choroid plexus in the left lateral ventricle. with bilateral cholesterinic granulomas of the choroid plexuses of the lateral ventricles.
 Development of the Nervous System: Malformation 39

Hypoplasia of the Prosencephalon: Cerebral

(Telencephalic) Aplasia (Anencephaly)
Calves have been observed at birth with a failure of the
telencephalic vesicles to develop; this is associated with a
small opening in the calvaria on the midline at the level of
the orbits where fluid, presumably CSF, emerges (Fig. 3-21).
The calvarial defect here is called a cranioschisis or cranium
bifidum. The skin around this opening is continuous with a
malformed diencephalon inside the small cranial cavity. There
are no cerebral hemispheres. The brainstem and cerebel-
lum are present but reduced in size and abnormally shaped.
There are no recognizable thalamic geniculate nuclei or
­mesencephalic colliculi (Figs. 3-22 and 3-23). The eyes are
well developed and all cranial nerves are ­ present except
for the first, the olfactory neurons. In human babies this Figure 3-22 Dorsal view of the brain of a calf with prosencephalic hypoplasia.
is called anencephaly, an incorrect term because it implies Note the complete absence of any cerebral hemispheres.
that there is no brain development at all. No ­ single term
best describes this malformation.
Telencephalic (cerebral) aplasia accounts for the most
extensive part of the malformation. One hypothesis about
the mechanism involved in this brain malformation is as
follows. If some factor caused the neurectoderm of the pros-
encephalon to fail to separate from the skin ectoderm at the
level of the rostral neuropore, that could prevent the out-
growth of the telencephalic vesicles and create a defect in
the closure of the calvaria, resulting in cerebral aplasia and
cranioschisis, respectively. CSF could leak from the third
ventricle of the diencephalon, which remains attached to
the surface skin. The cause of this malformation in animals

Figure 3-23 Lateral view of the brain of a calf with prosencephalic hypoplasia.
Note the abnormally shaped brainstem and cerebellum.

is unknown but is one of the neural tube defects thought to

result from a deficiency in folic acid in the diet of the human
mother very early in development, before pregnancy may
be diagnosed. Hyperthermia has also been implicated as a
cause of this neural tube defect.
These calves are usually unable to get up at birth. Video
3-1 shows a newborn Guernsey calf with this malformation.
When supported, these calves struggle to stand and exhibit
considerable voluntary movement. They are blind because
of the lack of any cerebral visual cortex for perception, but
the cranial nerves function normally. Spinal reflexes are
intact and surprisingly, there is some evidence of nocicep-
tion (the perception of noxious stimuli), which supports
the idea that some “conscious” perception can occur at the
level of the diencephalon. The evidence for this is the occa-
sional voluntary struggling that occurs when forceps pres-
sure is applied to the digits as a noxious stimulus or when
forceps are pressed against the nasal septum, and occasion-
ally a loud noise elicits a voluntary response. We have not
observed this malformation in other species of domestic
animals.

Meningocele: Meningoencephalocele
Figure 3-21 Calf with prosencephalic hypoplasia. Note the area in the center A newborn animal with a large skin-covered, soft swelling
of the calvaria devoid of hair. The malformed diencephalon is fused to the skin here, outside the calvaria, usually on the midline, has a meningo-
and CSF leaked out of the third ventricle at this site. cele or meningoencephalocele. The latter has brain tissue in
40 V eterinary N euroanatomy and C linic al N eurology

the swelling along with the meninges and is by far the most
common of these two malformations, but microscopic exam-
ination of the wall of the swelling is commonly required to
appreciate the brain component (Figs. 3-24 and 3-25). The cal-
varial defect is called cranioschisis or cranium bifidum. This
is usually on the midline and at any level of the length of the
calvaria. This cranioschisis may be very large with both cere-
bral hemispheres bulging into the meningoencephalocele, or
it can be small and involve an extension from the brainstem
into the meningoencephalocele. In the large meningoenceph-
alocele, the protruding cerebral hemispheres are usually very
thin and fluid-filled due to dilation of their lateral ventri-
cles, which results in the meningoencephalocele’s being soft
and fluctuant. Usually the cause is unknown. It is seen in all
­species of domestic animal but may be more common in pigs.
In one kitten, a meningoencephalocele was associated with
the queen’s being treated orally with griseofulvin throughout
the ­gestation period for a ringworm infection (Fig. 3-26).26
A very large meningoencephalocele occurs as part of
an inherited craniofacial malformation in Burmese cats
(Fig. 3-27).31 Prior to its recognition as an inherited disor-
der, it was quite common in this breed in many different
l­ocations in the United States. It became apparent when
a number of breeders attempted to breed for Burmese cats
with ­shortened faces. In addition to the large meningoen- Figure 3-25 Head of a stillborn Belgian foal with a small
cephaloceles that often hung over their shortened faces, meningoencephalocele. The hairless mass protruded through a small
these kittens had significant facial malformations: the upper midline cranioschisis. On microscopic examination of the mass, brain tissue
jaws and nasal areas were shortened and had no recogniz- including choroid plexus was found adjacent to the dermis.
able nares, planum nasale, or nasal cavities. There were no
olfactory bulbs or peduncles in the malformed cerebrums.
There was a philtrum on either side of the rostral midline of
the face and vibrissae on both sides of the cleft (Fig. 3-28).
Duplication was also evident in the two pairs of maxillary
bones and two sets of maxillary canine teeth. There was
no gross evidence of any eyes but one pair of orbits was
present. No optic nerves or chiasm were associated with
the diencephalon. The mandibles were prominent and the
tongue was enlarged and protruded from the mouth. These
facial deformities represent the result of abnormal cranial
neural crest development. This is a prime example of what
can happen when breeders select for a characteristic that
in reality is a malformation, the shortened face; it is clear

Figure 3-26 Newborn kitten with a meningoencephalocele. The queen had

been treated with griseofulvin during gestation.

that more ­ serious and life-threatening malformations can

result. Selection for a longer cat skull shape will eliminate
the problem. The malformation that occurs in Manx cats
that are bred for the absence of a tail is a similar failure of
judgment by misinformed breeders. This is described with
the spinal cord malformations in this chapter.

Exencephaly
Exencephaly is now defined as brain tissue that protrudes out
Figure 3-24 Newborn Holstein calf with a large meningoencephalocele. of the cranial cavity and is not covered by skin (Fig. 3-29).
 Development of the Nervous System: Malformation 41

found that exencephaly is much less common than meningo-

celes or meningoencephaloceles. Both malformations require
a cranioschisis, which is the opening in the calvaria where the
bones have not fused.

Lipomeningocele
A lipomeningocele, as the term suggests, is a meningo-
cele that has a large amount of fat associated with it. It can
occur along the midline of the calvaria or the vertebral col-
umn through a cranioschisis or spina bifida, respectively.
It consists of fat-filled meningeal tissue covered by skin
that is continuous with the falx cerebri in the head and the
dura of the spinal cord in the vertebral column (Fig. 3-30).
With no associated neural tube malformation, there are no
­neurologic signs in these animals. These are rare malforma-
tions and the cause is unknown.

Figure 3-27 Newborn Burmese kitten with a large meningoencephalocele

associated with facial duplication. Duplication of the Prosencephalon
Prosencephalic duplication occurs most commonly in calves
and is associated with varying degrees of duplication in the
face that are called diprosopus or dicephalus (Fig. 3-31). The

Figure 3-28 Another Burmese kitten with a smaller meningoencephalocele.

Note the duplication of the philtrum and whisker pads.

Figure 3-30 A 2-year-old Holstein cow with a calvarial lipomeningocele. This

malformation was associated with the meninges of the falx cerebri with no cerebral
involvement. This cow was clinically normal.

Figure 3-29 Two Pomeranian puppies with exencephaly.

There is no stratified squamous epithelium over the brain,

only the connective tissue of the meninges. This can be deter-
mined only by microscopic study of the protruding tissue. Figure 3-31 Dicephalic Holstein calf. Besides the enlarged midline orbit containing
When these malformations are studied microscopically, it is two fused eyes, there is an orbit and a complete eye on each side of the head.
Exploring the Variety of Random
Documents with Different Content
This is the daie, wherein the Lord God came to the crosse. But as the
crosse shall never more come to him: so let never the hot or cold fit of
this ague come anie more unto this man, In nomine patris ✠ & fi✠lii, &
spiritus ✠ sancti ✠. Then saie seven and twentie Pater nosters, and as
manie Aves, and use this three daies togither. ❈ Otherwise:/
 Fécana, cagéti, daphnes, gebáre, gedáco, 272.
Gébali stant, sed non stant phebas, hecas,* & hedas. This is t
mystical
Everie one of these words must be written upon a peece of bread, englishe
Nota.
and/ be given in order one daie after another to the sicke bodie, and so [* 1584
must he be cured. This saith Nicholas Hemingius he chanced to read in hec as]

the schooles in jest; so as one noting the words, practised the medicine 196.
in earnest; and was not onelie cured himselfe, but also cured manie
others thereby. And therefore he concludeth, that this is a kind of a
miraculous cure, wrought by the illusion of the divell: whereas in truth, it
will fall out most commonlie, that a tertian ague will not hold anie man
longer than so, though no medicine be given, nor anie words spoken. ❈ Fernel
Otherwise: This word, Abra cadabra written on a paper, with a certeine
figure joined therewith, and hanged about ones necke, helpeth the ague.
❈ Otherwise: Let the urine of the sicke bodie made earlie in the
morning, be softlie heated nine daies togither continuallie, untill all be
consumed into vapor. ❈ Otherwise: A crosse made of two litle twigs
joined togither, wherewith when the partie is touched, he will be whole;
speciallie if he weare it about his necke. ❈ Otherwise: Take a like
quantitie of water out of three ponds of equall bignesse, and tast thereof
in a new earthen vessell, and drinke of it when the fit commeth.
In the yeare of our lord 1568, the Spaniards and Italians received from Notab
the pope, this incantation following; whereby they were promised both the Sp
Italian
remission of sinnes, and good successe in their warres in the lowe
countries. Which whether it be not as prophane and impious, as anie
witches charme, I report me to the indifferent reader. ✠ Crucem pro
nobis subiit ✠ & stans in illa sitiit ✠ Jesus sacratis manibus, clavis
ferreis, pedibus perfossis, Jesus, Jesus, Jesus: Domine libera nos ab hoc
malo, & ab hac peste: then three Pater nosters, and three ave Maries.
Also the same yeere their ensignes were by the authoritie aforesaid
conjured with certeine ceremonies, & consecrated against their enimies.
And if you read the histories of these warres, you maie see what victorie
they gained hereby. Item, they baptised their cheefe standard, and gave
it to name S. Margaret, who overthrew the divell. And bicause you shall
under/stand the mysterie hereof, I have the rather set it downe 273.
elsewhere, being indeed worth the reading.

¶ For a bloudie flux, or rather an issue of bloud.

TAke a cup of cold water, and let fall thereinto three drops of the same
bloud, and betweene each drop saie a Pater noster, and an Ave, then
drinke to the patient, and saie; Who shall helpe you? The patient must
answer S. Marie. Then saie you; S. Marie stop the issue of bloud. ❈ He mu
by non
Otherwise: Write upon the patients forhead with the same bloud; she pe
Consummatum est. ❈ Otherwise: Saie to the patient; Sanguis mane in the cu
te, sicut fecit Christus in se; Sanguis mane in tua vena, sicut Christus in by pat
sua pœna; Sanguis mane fixus, sicut Christus quando fuit crucifixus: ter.
❈ Otherwise, as followeth.
In the bloud of Adam death was taken ✠
In the bloud of Christ it was all to shaken ✠
And by the same bloud I doo thee charge,
That thou doo runne no longer at large. ❈ Otherwise.

Christ was borne at Bethelem, and suffered at Jerusalem, where his

bloud was troubled. I command thee by the vertue of God, and through/ 197.
the helpe of all saincts, to staie even as Jordan did, when John baptised
Christ Jesus; In nomine patris ✠ & filii ✠ & spiritus |sancti ✠ ❈
Otherwise: Put thy nameles finger in the wound, and make therwith
three crosses upon the wound, and saie five Pater nosters, five Aves,
and one Credo, in the honour of the five wounds. ❈ Otherwise: Touch See J.
11. co
that part and saie, De latere ejus exivit sanguis & aqua. ❈ Otherwise: In
nomine patris ✠ & filii ✠ & spiritus sancti ✠ &c. Chimrat, chara, sarite,
confirma, consona, Imohalite. ❈ Otherwise: Sepa ✠ sepaga ✠ sepagoga
✠ sta sanguis in nomine patris ✠ podendi ✠ & filii ✠ podera ✠ &
spiritus sancti ✠ pandorica ✠ pax tecum, Amen.

¶ Cures commensed and finished by witchcraft.

THere was a jollie fellowe that tooke upon him to be a notable surgion,
in the dutchie of Mentz, 1567. to whom there resorted a Gentleman
that had beene vexed with sicknesse, named/ Elibert, having a kerchiefe 274.
on his head, according to the guise of sicke folke. But the surgion made
him pull off his kerchiefe, and willed him to drinke with him freelie. The
sickeman said he durst not; for he was forbidden by physicke so to doo.
Tush (said this cunning man) they know not your disease: be ruled by
me, and take in your drinke lustilie. For he thought that when he was
well tippled, he might the more easilie beguile him in his bargaine, and
make his reward the greater, which he was to receive in part aforehand.
When they had well droonke, he called the sicke man aside, and told him
the greatnes and danger of his disease, and how that it grew by meanes
of witchcraft, and that it would be universallie spread in his house, and
among all his cattell, if it were not prevented: and impudentlie
persuaded the sicke man to receive cure of him. And after bargaine The su
made, he demanded of the sicke man, whether he had not anie at most i
setteth
home, whom he might assuredlie trust. The sicke man answered, that he knaver
had a daughter and a servant. The cousener asked how old his daughter
was? The patient said, twentie. Well (said the cousener) that is fit for our
turne. Then he made the mother and father to kneele on their knees to
their daughter, and to desire hir in all things to obey the physician, and
that she would doo in everie thing as he commanded hir; otherwise hir
father could not be restored to his health. In which respect hir parents
humblie besought hir on their knees so to doo. Then he assigned hir to
bring him into his lodging hir fathers haire, and hir mothers, and of all
those which he kept in his house, as well of men and women, as also of
his cattell. When she came therewith unto him, according to the match
made, and hir parents commandement, he lead hir downe into a lowe
parlor, where having made a long speech, he opened a booke that laie
on the boord, and laieth thereon two knives acrosse, with much
circumstance of words. Then conjureth he, and maketh strange A pret
characters, and at length he maketh a circle on the ground, wherein he conjur
causeth hir to sticke one of those conjured knives; and after manie more
strange words, he maketh hir sticke the other knife beside it. Then fell
downe the maid in a swoone for feare; so as he was faine to frote hir
and put a sop into hir mouth, after the receipt whereof she was sore
troubled and amazed. Then he made hir brests to be uncovered, so as
when/ they were bare, he dal/lied with them, diverslie and long together. 275.
Then he made hir lie right upward, all uncovered and bare belowe hir 198.
pappes. Wherein the maid being loth to obeie him, resisted, and in
shame forbad that villanie. Then said the knave; Your fathers destruction
is at hand: for except you will be ruled, he and all his familie shall
susteine greater greefe and inconvenience, than is yet happened unto
him. And no remedie, except you will seeke his utter overthrowe, I must
have carnall copulation with you, and therewithall fell into hir bosome,
and overthrew hir and hir virginitie. So did he the second daie, and
attempted the like on the third daie. But he failed then of his purpose, as Ad vad
the wench confessed afterwards. In the meane time he ministred so urna q
cadit.
cruell medicines to the sicke man, that through the torments therof he
feared present death, and was faine to keepe his bed, whereas he
walked about before verie well and lustilie. The patient in his torments
calleth unto him for remedie, who being slacke and negligent in that
behalfe, made roome for the daughter to accompanie hir father, who
asked hir what she thought of the cure, and what hope she had of his
recoverie. Who with teares remained silent, as being oppressed with
greefe; till at the last in abundance of sorrowe she uttered the whole
matter to hir father. This dooth Johannes Wierus report, saieng, that it
came unto him by the lamentable relation of the father himselfe. And
this is here at this time for none other purpose rehearsed, but that men
may hereby learne to take heed of such cousening merchants, and
knowe what they be that take upon them to be so cunning in witchcraft;
least they be bewitched: as maister Elibert and his daughter were.

¶ Another witchcraft or knaverie, practised by the same surgion.

THis surgion ministred to a noble man, that laie sicke of an ague,

offering unto him three peeces of a roote to be eaten at three
Three
the fir
with c
morsels; saieng to the first: I would Christ had not beene borne; unto the se
the second: I would he had not suffered; unto the third: I would he had his pas
not risen againe. And then putting them about the sicke mans necke, third w
resurre
said; Be of good cheere. And if he lost them, whosoever tooke them up,
should therewithall take awaie/ his ague. ❈ Otherwise: Jesus Christ, 276.
which was borne, deliver thee from this infirmitie ✠ Jesus Christ which
died ✠ deliver thee from this infirmitie ✠ Jesus Christ which rose againe
✠ deliver thee from this infirmitie. Then dailie must be said five Pater
nosters, and five Aves.

¶ Another experiment for one bewitched.

ANother such cousening physician persuaded one which had a

timpanie, that it was one old viper, and twoo yoong mainteined in his
A cous
physic
foolish
bellie by witchcraft. But being watched, so as he could not conveie vipers
into his ordure or excrements, after his purgations: at length he told the
partie, that he should suffer the paines of childbirth, if it were not
prevented; and therefore he must put his hand into his breech, and rake
out those wormes there. But the mother of the sicke partie having
warning hereof, said she could doo that hir selfe. So the cousener was
prevented, and the partie died onelie of a timpanie, and the knave ran
awaie out of the countrie.

¶ Otherwise.
MOnsieur Bodin telleth of a witch, who undertaking to cure a woman
bewitched, caused a masse to be soong at midnight in our ladies
John.

chap/pell. And when she had overlien the sicke partie, and breathed 199.
certeine words upon hir, she was healed. Wherein Bodin saith, she
followed the example of Elias the prophet, who raised the Sunamitie. Kakoze
And this storie must needs be true: for goodman Hardivin Blesensis his
host at the signe of the lion told him the storie.

¶ A knacke to knowe whether you be bewitched, or no, &c.

ITbewitched
is also expedient to learne how to know whether a sicke man be
or no: this is the practise thereof. You must hold molten
Mal. m
quæ.
Barth.
lead over the sicke bodie, and powre it into a porrenger full of water; novo
and then, if there appeare upon the lead, anie image, you may then Mal. m
knowe the partie is bewitched./

The xix. Chapter. 277.

That one witchcraft maie lawfullie meete with another.

COTUS, Hostiensis, Gofridus, and all the old canonists agree, that
it is lawfull to take awaie witchcraft by witchcraft, Et vana vanis
contundere. And Scotus saith, It were follie to forbeare to Scotus
encounter witchcraft by witchcraft; for (saith he) there can be none distinc
imperi
inconvenience therein; bicause the overthrower of witchcraft assenteth
not to the works of the divell. And therefore he saith further, that it is
meritorious so to extinguish and overthrow the divels workes. As though
he should saie; It maketh no matter, though S. Paule saie; Non facies
malum, ut indè veniat bonum, Thou shalt not doo evill, that good maie
come thereof. Humbertus saith, that witchcraft maie be taken awaie by Dist. 4
that meanes whereby it was brought. But Gofredus inveieth sore against
the oppugners thereof. Pope Nicholas the fift gave indulgence and leave Gofred
to bishop Miraties (who was so bewitched in his privities, that he could sua.
not use the gift of venerie) to seeke remedie at witches hands. And this
was the clause of his dispensation, Ut ex duobus malis fugiatur majus,
that of two evils, the greater should be avoided. And so a witch, by
taking his doublet, cured him, and killed the other witch: as the storie
saith, which is to be seene in M. Mal. and diverse other writers.

The xx. Chapter.

Who are privileged from witches, what bodies are aptest to be
bewitched, or to be witches, why women are rather witches than
men, and what they are.
OW if you will know who and what persons are privileged from
witches, you must understand, that they be even such as Mal. m
cannot be bewitched. In the number of whome first be the quæ.
in/quisitors, and such as exercise publike justice upon them. Howbeit,* a 278.
justice in Essex, whome for diverse respects I have left unnamed, not * Whe
long since thought he was bewitched, in the verie instant whiles he more i
booke
examined the witch; so as his leg was broken therby, &c: which either in prin
was false, or else this rule untrue, or both rather injurious unto Gods
providence. Secondlie, such as observe dulie the rites and ceremonies of
holie church, and worship them with reverence, through the sprinkling of
holie water, and receiving consecrated salt, by the lawfull use of candles
hallowed on Candelmas daie, and greene leaves consecrated on Palme
sundaie (which things they saie the/ church useth for the qualifieng of 200.
the divels power) are preserved from witchcraft. Thirdlie, some are
preserved by their good angels, which attend and wait upon them.
But I maie not omit here the reasons, which they bring, to prove what
bodies are the more apt and effectuall to execute the art of fascination.
And that is, first they saie the force of celestiall bodies, which
indifferentlie communicate their vertues unto men, beasts, trees, stones,
&c. But this gift and naturall influence of fascination maie be increased in
man, according to his affections and perturbations; as thorough anger,
feare, love, hate, &c. For by hate (saith Vairus) entereth a fierie L. Vair
inflammation into the eie of man, which being violentlie sent out by fascin.
beams and streames, &c: infect and bewitch those bodies against
whome they are opposed. And therefore he saith (in the favour of
women) that that is the cause why women are oftener found to be
witches than men. For (saith he) they have such an unbrideled force of
furie and concupiscence naturallie, that by no meanes it is possible for
them to temper or moderate the same. So as upon everie trifling Much
occasion, they (like brute beasts) fix their furious eies upon the partie eiebiti
of who
whom they bewitch. Hereby it commeth to passe, that whereas women elswhe
having a mervellous fickle nature, what greefe so ever happeneth unto
them, immediatlie all peaceablenes of mind departeth; and they are so
troubled with evill humors, that out go their venomous exhalations,
ingendred thorough their ilfavoured diet, and increased by meanes of
their pernicious excrements, which they expell. Women are also (saith
he) monethlie filled full of superfluous humors, and with them/ the
melancholike bloud boileth; whereof spring vapors, and are carried up, 279.
and conveied through the nosethrels and mouth, &c: to the bewitching
of whatsoever it meeteth. For they belch up a certeine breath, wherewith
they bewitch whomsoever they list. And of all other women, leane, Who a
hollow eied, old, beetlebrowed women (saith he) are the most infectious. likelie
and to
Marie he saith, that hot, subtill, and thin bodies are most subject to be bewitc
bewitched, if they be moist, and all they generallie, whose veines, pipes,
and passages of their bodies are open. And finallie he saith, that all
beautifull things whatsoever, are soone subject to be bewitched; as
namelie goodlie yoongmen, faire women, such as are naturallie borne to
be rich, goodlie beasts, faire horsses, ranke corne, beutifull trees, &c.
Yea a freend of his told him, that he saw one with his eie breake a
pretious stone in peeces. And all this he telleth as soberlie, as though it
were true. And if it were true, honest women maie be witches, in
despight of all inquisitors: neither can anie avoid being a witch, except
shee locke hir selfe up in a chamber.

The xxi. Chapter.

What miracles witchmongers report to have beene done by witches
words, &c: contradictions of witchmongers among themselves, how
beasts are cured herby, of bewitched butter, a charme against
witches, and a counter charme, the effect of charmes and words
proved by L. Vairus to be woonderfull.
F I should go about to recite all charmes, I should take an infinite
worke in hand. For the witching writers hold opinion, that anie
thing al/most maie be therby brought to passe; & that whether 197.[2
the words of the charme be understandable or not, it skilleth not: so the
charmer have a steddie intention to bring his desire about. And then
what is it that cannot be done by words? For L. Vairus saith, that old L. Vair
women have infeebled and killed children with words, and have made fascin.
women with child miscarrie;/ they have made men pine awaie to death, 280.
they have killed horsses, deprived sheepe of their milke, *transformed
men into beasts, flowne in the aire, tamed and staied wild beasts, driven * Acco
all noisome cattell and vermine from corne, vines and hearbs, staied Ovids
Proteu
serpents, &c: and all with words. In so much as he saith, that with which
certeine words spoken in a bulles eare by a witch, the bull hath fallen alledge
downe to the ground as dead. Yea some by vertue of words have gone therefo
aqua,
upon a sharpe sword, and walked upon hot glowing coles, without hurt; modò
with words (saith he) verie heavie weights and burthens have beene cervus
lifted up; and with words wild horsses and wild bulles have beene
tamed, and also mad dogs; with words they have killed wormes, and
other vermine, and staied all maner of bleedings and fluxes: with words
all the diseases in mans bodie are healed, and wounds cured; arowes
are with wonderfull strangenesse and cunning plucked out of mens
bones. Yea (saith he) there be manie that can heale all bitings of dogs,
or stingings of serpents, or anie other poison: and all with nothing but
words spoken. And that which is most strange, he saith, that they can
remedie anie stranger, and him that is absent, with that verie sword
wherewith they are wounded. Yea and that which is beyond all
admiration, if they stroke the sword upwards with their fingers, the
partie shall feele no paine: whereas if they drawe their finger
downewards thereupon, the partie wounded shall feele intollerable
paine. With a number of other cures, done altogither by the vertue and
force of words uttered and spoken.
Where, by the waie, I maie not omit this speciall note, given by M.
Mal. to wit, that holie water maie not be sprinkled upon bewitched Mal. M
beasts, but must be powred into their mouthes. And yet he, and also quæ. 2
Nider, saie, that It is lawfull to blesse and sanctifie beasts, as well as Nider
men; both by charmes written, and also by holie words spoken. For præce
præce
(saith Nider) if your cow be bewitched, three crosses, three Pater 11.
nosters, and three Aves will certeinlie cure hir: and likewise all other Nider
ceremonies ecclesiasticall. And this is a sure Maxime,* that they which fornica
are delivered from witchcraft by shrift, are ever after in the night much Mal. M
molested (I beleeve by their ghostlie fathers.) Also they loose their 2. cap
monie out of their pursses and caskets: as M. Mal. saith he knoweth by [* Ital
experience./ Also one generall rule is given by M. Mal. to all butter wives, 281.
and dairie maides, that they neither give nor lend anie butter, milke, or A good
cheese, to anie witches, which alwaies use to beg therof, when they starve
wome
meane to worke mischeefe to their kine or whitmeats. Whereas indeed
there are in milke three substances commixted; to wit, butter, cheese, Mal. M
and whaie: if the same be kept too long, or in an evill place, or be 2. quæ
sluttishlie used, so as it be stale and sower, which happeneth sometimes
in winter, but oftener in summer, when it is set over the fier, the cheese
and butter runneth togither, and congealeth, so as it will rope like
birdlime, that you maie wind it about a sticke, and/ in short space it will 198.[2
be so drie, as you maie beate it to powder. Which alteration being
strange, is woondered at, and imputed to witches. And herehence
sometimes proceedeth the cause, why butter commeth not, which when
the countrie people see that it commeth not, then get they out of the
suspected witches house, a little butter, whereof must be made three
balles, in the name of the holie trinitie; and so if they be put into the
cherne, the butter will presentlie come, and the witchcraft will cease; Sic
ars deluditur arte. But if you put a little sugar or sope into the cherne,
among the creame, the butter will never come: which is plaine
witchcraft, if it be closelie, cleanlie, and privilie handled. There be
twentie severall waies to make your butter come, which for brevitie I
omit; as to bind your cherne with a rope, to thrust thereinto a red hot
spit, &c: but your best remedie and surest waie is, to looke well to your
dairie maid or wife, that she neither eat up the creame, nor sell awaie
your butter.

¶ A charme to find hir that bewitched your kine.

PUtpasture
a paire of breeches upon the cowes head, and beate hir out of the
with a good cudgell upon a fridaie, and she will runne right to
A ridic
charm
the witches doore, and strike thereat with hir hornes.

¶ Another, for all that have bewitched anie kind of cattell.

WHen anie of your cattell are killed with witchcraft, hast you to the
place where the carcase lieth, and traile the bowels of the beast
unto your house, and drawe them not in at/ the doore, but under the 282.
threshhold of the house into the kitchen; and there make a fier, and set
over the same a grediron, and thereupon laie the inwards or bowels; and
as they wax hot, so shall the witches entrailes be molested with
extreame heate and paine. But then must you make fast your doores,
least the witch come and fetch awaie a cole of your fier: for then ceaseth
hir torments. And we have knowne saith M. Mal. when the witch could
not come in, that the whole house hath beene so darkened, and the aire
round about the same so troubled, with such horrible noise and
earthquakes; that except the doore had beene opened, we had thought
the house would have fallen on our heads. Thomas Aquinas, a principall
treator herein, alloweth conjurations against the changelings, and in
diverse other cases: whereof I will saie more in the word Iidoni.

¶ A speciall charme to preserve all cattell from witchcraft.

ATpaschall
Easter you must take certeine drops, that lie uppermost of the holie
candle, and make a little waxe candle thereof: and upon
In anie
observ
festiva
some sundaie morning rath, light it, and hold it, so as it maie drop upon else yo
and betweene the hornes and eares of the beast, saieng: In nomine all.
patris, & filii, et duplex s s: and burne the beast a little betweene the
hornes on* the eares with the same wax: and that which is left thereof, [* or]
sticke it in crossewise about the stable or stall, or upon the threshold, or
over the doore, where the cattell use to go in and out, and for all that
yeare your cattell shall never be be/witched. ❈ Otherwise: Jacobus de 199.[2
Chusa Carthusianus sheweth, how bread, water, and salt is conjured,
and saith, that if either man or beast receive holie bread and holie water
nine daies together, with three Pater nosters, and three Aves, in the
honour of the trinitie, and of S. Hubert, it preserveth that man or beast
from all diseases, and defendeth them against all assaults of witchcraft,
of satan, or of a mad dog, &c.
Lo this is their stuffe, mainteined to be at the least effectuall, if not
wholsome, by all papists and witchmongers, and speciallie of the last
and proudest writers. But to proove these things to be effectuall, God
knoweth their reasons are base and absurd. For they write so, as they
take the matter in question as granted,/ and by that meanes go awaie 283.
therewith. For L. Vairus saith in the beginning of his booke, that there is
no doubt of this supernaturall matter, bicause a number of writers agree L. Vair
herein, and a number of stories confirme it, and manie poets handle the fascin.
same argument, and in the twelve tables there is a lawe against it, and
bicause the consent of the common people is fullie with it, and bicause
immoderate praise is to be approoved a kind of witchcraft, and bicause
old women have such charmes and superstitious meanes as preserve
themselves from it, and bicause they are mocked that take awaie the
credit of such miracles, and bicause Salomon saith; Fascinatio Sapi. 4
malignitatis obscurat bona, and bicause the apostle saith; O insensati Gali. 3
Psal. 1
Galatæ, quis vos fascinavit? And bicause it is written, Qui timent te,
videbunt me. And finallie he saith, least you should seeme to distrust
and detract anie thing from the credit of so manie grave men, from
histories, and common opinion of all men: he meaneth in no wise to
proove that there is miraculous working by witchcraft and fascination;
and proceedeth so, according to his promise.

The xxii. Chapter.

Lawfull charmes, or rather medicinable cures for diseased cattell. The
charme of charmes, and the power thereof.
UT if you desire to learne true and lawfull charmes, to cure Direct
diseased cattell, even such as seeme to have extraordinarie meane
cattell
sicknesse, or to be bewitched, or (as they saie) strangelie
taken: looke in B. Googe his third booke, treating of cattell, and happilie
you shall find some good medicine or cure for them: or if you list to see
more ancient stuffe, read Vegetius his foure bookes thereupon: or, if you
be unlearned, seeke some cunning bullocke leech. If all this will not
serve, then set Jobs patience before your eies. And never thinke that a
poore old woman can alter supernaturallie the notable course, which God
hath appointed among his/ creatures. If it had beene Gods pleasure to 284.
have permitted such a course, he would no doubt have both given notice
in his word, that he had given such power unto them, and also would
have taught remedies to have prevented them.
Furthermore, if you will knowe assured meanes, and infallible
charmes, yeelding indeed undoubted remedies, and preventing all maner
of witchcrafts, and also the assaults of wicked spirits; then despise first
all cou/sening knaverie of priests, witches, and couseners: and with true 200.[2
faith read the sixt chapter of S. Paule to the Ephesians, and followe his
counsell, which is ministred unto you in the words following, deserving
worthilie to be called by the name insuing:

The charme of charmes.

Finallie my brethren, be strong in the Lord, and in the power of his A char
might. Put on the whole armour of God, that you may stand against the charm
out of
assaults of the divell. For we wrestle not against flesh and bloud, but chapte
against principalities and powers, & against worldlie governors the Paule
princes of the darkenes of this world, against spirituall wickednes, which Ephes
are in the high places. For this cause take unto you the armour of God,
that you may be able to resist in the evill daie; and having finished all
things, stand fast. Stand therefore, and your loines girded about with
veritie, and having on the brestplate of righteousnes, &c: as followeth in
that chapter, verses 15. 16. 17. 18. 1 Thes. 5. 1 Pet. 5, verse. 8. Ephes.
1. and elsewhere in the holie scripture.

¶ Otherwise.

IFlearned,
you be unlearned, and want the comfort of freends, repaire to some
godlie, and discreet preacher. If otherwise need require, go to
a learned physician, who by learning and experience knoweth and can
discerne the difference, signes, and causes of such diseases, as
faithlesse men and unskilfull physicians impute to witchcraft.//

The xxiii. Chapter. 285. 2

A confutation of the force and vertue falselie ascribed to charmes and
amulets, by the authorities of ancient writers, both divines and
physicians.
Y meaning is not, that these words, in the bare letter, can doo
anie thing towards your ease or comfort in this behalfe; or that
it were wholesome for your bodie or soule to weare them about
your necke: for then would I wish you to weare the whole Bible, which
must needs be more effectuall than anie one parcell thereof. But I find
not that the apostles, or anie of them in the primitive church, either
carried S. Johns gospell, or anie Agnus Dei* about them, to the end they [* Ital
might be preserved from bugges: neither that they looked into the foure
corners of the house, or else in the roofe, or under the threshhold, to
find matter of witchcraft, and so to burne it, to be freed from the same;
according to the popish rules. Neither did they by such and such verses Mal. M
or praiers made unto saints, at such or such houres, seeke to obteine 2. qu.
grace: neither spake they of anie old women that used such trades.
Neither did Christ at anie time use or command holie water, or crosses,
&c: to be used as terrors against the divell, who was not affraid to
assault himselfe, when he was on earth. And therefore a verie vaine
thing it is to thinke that he feareth these trifles, or anie externall matter. 1. Tim
Let us then cast awaie these prophane and old wives fables. For(as Origin
Job.
Origen saith) Incantationes sunt dæmonū irrisiones idololatriæ fæx,
animarum infatuatio, &c.
Chrysostome saith; There be some that carrie about their necks a J. Chry
peece of a gospell. But* is it not dailie read (saith he) and heard of all Matth.
men? But if they be never the better for it, being put into their eares, * Mark
how shall they be saved, by carrieng it about their necks? And further he was no
service
saith; Where is the vertue of the gospell? In the figure of the letter, or in
Idem.
the understanding of the sense? If in the figure, thou dooest well to
weare it about thy/ necke; but if in the understanding, then thou 286.
shouldest laie it up in thine heart. Augustine saith; Let the faithfull
ministers admonish and tell their people, that these magicall arts and Augus
incantations doo bring no remedie to the infirmities either of men or ultim.
cattell, &c.
The heathen philosophers shall at the last daie confound the infidelitie
and barbarous foolishnes of our christian or rather anti-christian and
prophane witchmongers. For as Aristotle saith, that Incantamenta sunt
muliercularum figmenta: so dooth Socrates (who was said to be cunning
herein) affirme, that Incantationes sunt verba animas decipientia
humanas. Others saie; Inscitæ pallium sunt carmina, maleficium, &
incantatio. Galen also saith, that such as impute the falling e/vill, and Galen.
such like diseases to divine matter, and not rather to naturall causes, are comitia
witches, conjurers, &c. Hippocrates calleth them arrogant; and in 285. [
another place affirming that in his time there were manie deceivers and Hippoc
couseners, that would undertake to cure the falling evill, &c: by the morbo
power and helpe of divels, by burieng some lots or inchantments in the
ground, or casting them into the sea, concludeth thus in their credit, that
they are all knaves and couseners: for God is our onlie defender and
deliverer. O notable sentence of a heathen philosopher!/
 ¶ The xiii. Booke. 287.

The first Chapter.

The signification of the Hebrue word Hartumim, where it is found written in the
scriptures, and how it is diverslie translated: whereby the objection of Pharaos
magicians is afterward answered in this booke; also of naturall magicke not evill
in it selfe.
ARTUMIM is no naturall Hebrue word, but is borrowed of some other nation:
howbeit, it is used of the Hebrues in these places; to wit, Gen. 4. 1. 8. 24.
Exod. 7. 13. 24. & 8. 7. 18. & 9. 11. Dan. 1, 20. & 2. 2. Hierome sometimes Hierony
translateth it Conjectores, sometimes Malefici, sometimes Arioli: which we for the Gen. 41
In Exod
most part translate by this word witches. But the right signification heereof may be Dan. 1,
conceived, in that the inchanters of Pharao, being magicians of Aegypt, were called
Hartumim. And yet in Exodus they are named in some Latine translations Venefici.
Rabbi Levi saith, it betokeneth such as doo strange and woonderfull things,
naturallie, artificiallie, and deceitfullie. Rabbi Isaac Natar affirmeth, that such were so
termed, as amongst the Gentiles professed singular wisedome. Aben Ezra
expoundeth it, to signifie such as knowe the secrets of nature, and the qualitie of
stones and hearbs, &c: which is atteined unto by art, and/ speciallie by naturall 203.
magicke. But we, either for want of speach, or knowlege, call them all by the name/
and terme of witches. 288.

Certeinlie, God indueth bodies with woonderfull graces, the perfect knowledge
whereof man hath not reached unto: and on the one side, there is amongst them
such mutuall love, societie, and consent; and on the other side, such naturall
discord, and secret enimitie, that therein manie things are wrought to the
astonishment of mans capacitie. But when deceit and diabolicall words are coupled
therewith, then extendeth it to witchcraft and conjuration; as whereunto those
naturall effects are falselie imputed. So as heere I shall have some occasion to say The aut
somewhat of naturall magicke; bicause under it lieth hidden the venome of this word intentio
the mat
Hartumim. This art is said by some to be the profoundnesse, and the verie absolute hereaft
perfection of naturall philosophie, and shewing foorth the active part thereof, & discour
through the aid of naturall vertues, by the convenient applieng of them, works are
published, exceeding all capacitie and admiration; and yet not so much by art, as by
nature. This art of it selfe is not evill; for it consisteth in searching foorth the nature,
causes, and effects of things. As farre as I can conceive, it hath beene more
corrupted and prophaned by us Christians, than either by Jewes or Gentiles.

The second Chapter.

How the philosophers in times past travelled for the knowledge of naturall magicke,
of Salomons knowledge therein, who is to be called a naturall magician, a
distinction thereof, and why it is condemned for witchcraft.
 ANIE philosophers; as namely Plato, Pythagoras, Empedocles, Democritus,
&c: travelled over all the world, to find out & learne the knowlege of this
art; & at their returne they preached and taught, professed and published
it. Yea, it should appeere by the magicians that came to adore Christ, that the
knowledge and re/putation thereof was greater, than we conceive or make account 289.
of. But of all other, Salomon was the greatest traveller in this art, as may appeere
throughout the booke of Ecclesiastes: and speciallie in the booke of Wisedome,
where hee saith* God hath given me the true science of things, so as I knowe how * Sap. 7
the world was made, and the power of the elements, the beginning and the end, 18.
19.
and the middest of times, how the times alter, and the change of seasons, the 20.
course of the yeare, and the situation of the starres, the nature of living things, and 21.
the furiousnesse of beasts, the power of the wind, and the imaginations of men, the
diversities of plants, and the vertues of roots, and all things both secret and knowne,
&c. Finallie, he was so cunning in this art, that he is said to have bene a conjurer or
witch, and is so reputed in the Romish church at this daie. Whereby you may see, See Iidi
how fooles and papists are inclined to credit false accusations in matters of/ [Iidoni.
witchcraft and conjuration. The lesse knowledge we have in this art, the more we 204.
have it in contempt: in which respect Plato saith trulie to Dionysius; They make
philosophie a mockerie, that deliver it to prophane and rude people. Certeinlie, the
witchcraft, conjuration, and inchantment that is imputed to Salomon, is gathered out Eccle. 1
of these his words following: I applied my mind to knowledge, and to search and
seeke out science, wisedome and understanding, to knowe the foolishnesse of the
ungodlie, and the error of doting fooles. In this art of naturall magike (without great
heed be taken) a student shall soone be abused. For manie (writing by report,
without experience) mistake their authors, and set downe one thing for another.
Then the conclusions being found false, the experiment groweth into contempt, and
in the end seemeth ridiculous, though never so true. Plinie and Albert being curious
writers heerein, are often deceived; insomuch as Plinie is called a noble lier, and
Albert a rusticall lier; the one lieng by heeresaie, the other by authoritie.
A magician is indeed that which the Latines call a wise man, as Numa Pompilius A magic
was among the Romans; The Greeks, a philosopher, as Socrates was among them; describe
art disti
the Aegyptians a preest, as Hermes was; the Cabalists called them prophets. But
although these distinguished this art, accounting the one part/ thereof infamous, as 290.
being too much given unto wicked, vaine, and impious curiositie, as unto moovings,
numbers, figures, sounds, voices, tunes, lights, affections of the mind, and words;
and the other part commendable, as teaching manie good and necessarie things, as
times and seasons to sowe, plant, till, cut, &c: and diverse other things, which I will
make manifest unto you heereafter: yet we generallie condemne the whole art,
without distinction, as a part of witchcraft; having learned to hate it, before we
knowe it; affirming all to be witchcraft, which our grosse heads are not able to
conceive, and yet can thinke that an old doting woman seeth through it, &c. Wherein
we consider not how God bestoweth his gifts, and hath established an order in his
works, graffing in them sundrie vertues to the comfort of his severall creatures; and
speciallie to the use and behoofe of man: neither doo we therein weigh that art is
servant unto nature, and waiteth upon hir as hir handmaiden.
 The third Chapter.
What secrets do lie hidden, and what is taught in naturall magicke, how Gods glorie
is magnified therein, and that it is nothing but the worke of nature.
N this art of naturall magicke, God almightie hath hidden manie secret Read Pl
mysteries; as wherein a man may learne the properties, qualities, and natural.
Cardan
knowledge of all nature. For it teacheth to accomplish maters in such sort variet. A
and oportunitie, as the common people thinketh the same to be miraculous; and to de occu
be compassed none other waie, but onelie by witchcraft. And yet in truth, naturall propriet
Barthol
magicke is nothing else, but the worke of na/ture. For in tillage, as nature produceth natural.
corne and hearbs; so art, being natures minister, prepareth it. Wherein times and many o
seasons are greatlie to be respected: for Annus non arvus producit aristas. 205.
But as manie necessarie and sober things are heerein taught: so dooth it partlie (I
saie) consist in such experiments and conclu/sions as are but toies, but 291.
neverthelesse lie hid in nature, and being unknowne, doo seeme miraculous,
speciallie when they are intermedled and corrupted with cunning illusion, or
legierdemaine, from whence is derived the estimation of witchcraft. But being
learned and knowne, they are contemned, and appeere ridiculous: for that onelie is
woonderfull to the beholder, whereof he can conceive no cause nor reason,
according to the saieng of Ephesius, Miraculum solvitur unde videtur esse miraculum.
And therefore a man shall take great paines heerein, and bestow great cost to learne
that which is of no value, and a meere jugling knacke. Whereupon it is said, that a
man may not learne philosophie to be rich; but must get riches to learne
philosophie: for to sluggards, niggards, & dizzards, the secrets of nature are never
opened. And doubtlesse a man may gather out of this art, that which being Natural
published, shall set foorth the glorie of God, and be many waies beneficiall to the hath a d
end, wh
commonwealth: the first is doone by the manifestation of his works; the second, by
proveth
skilfullie applieng them to our use and service. excellen
same.

The fourth Chapter.

What strange things are brought to passe by naturall magicke.
HE dailie use and practise of medicine taketh awaie all admiration of the
woonderfull effects of the same. Manie other things of lesse weight, being
more secret and rare, seeme more miraculous. As for example (if it be true
that J. Bap. Neap. and many other writers doo constantlie affirme.) Tie a wild bull to
a figtree, and he will be presentlie tame; or hang an old cocke thereupon, and he
will immediatlie be tender; as also the feathers of an eagle consume all other
feathers, if they be intermedled together. Wherein it may not be denied, but nature
sheweth hir selfe a proper workwoman. But it seemeth unpossible, that a little fish
being but halfe a foot long, called Remora or Remiligo, or/ of some Echeneis, staieth 292.
a mightie ship with all hir loade and tackling, and being also under saile. And yet it is
affirmed by so manie and so grave authors, that I dare not denie it; speciallie, Pompan
bicause I see as strange effects of nature otherwise: as the propertie of the de inca
J. Wieru
loadstone, which is so beneficiall to the mariner; and of Rheubarb, which onelie lamiis. J
medleth with choler, and purgeth neither flegme nor melancholie, & is as beneficiall Peucer
&c.
to the physician, as the other to the mariner./

The fift Chapter. 206.

The incredible operation of waters, both standing and running; of wels, lakes, rivers,
and of their woonderfull effects.
HE operation of waters, and their sundrie vertues are also incredible, I
meane not of waters compounded and distilled: for it were endlesse to
treate of their forces, speciallie concerning medicines. But we have heere
even in England naturall springs, wels, and waters, both standing and running, of Of late
excellent vertues, even such as except we had seene, and had experiment of, we experie
Coventr
would not beleeve to be In rerum natura. And to let the physicall nature of them
passe (for the which we cannot be so thankefull to God, as they are wholsome for
our bodies) is it not miraculous, that wood is by the qualitie of divers waters heere in
England transubstantiated into a stone? The which vertue is also found to be in a
lake besides the citie Masaca in Cappadocia, there is a river called Scarmandrus, that
maketh yellow sheepe. Yea, there be manie waters, as in Pontus & Thessalia, and in
the land of Assyrides, in a river of Thracia (as Aristotle saith) that if a white sheepe Aristot.
being with lambe drinke thereof, the lambe will be blacke. Strabo writeth of the river hist. an
called Crantes, in the borders of Italie, running towards Tarentum, where mens haire
is made white and yellow being washed therein. Plinie dooth write that of what Plin. de
colour the veines are under the rammes toong, of/ the same colour or colours will colore.
the lambs be. There is a lake in a field called Cornetus, in the bottome whereof 293.
manifestlie appeareth to the eie, the carcases of snakes, ewts, and other serpents:
whereas if you put in your hand, to pull them out, you shall find nothing there.
There droppeth water out of a rocke in Arcadia, the which neither a silverne nor a
brasen boll can conteine, but it leapeth out, and sprinkleth awaie; and yet will it
remaine without motion in the hoofe of a mule. Such conclusions (I warrant you)
were not unknowne to Jannes and Jambres.

The sixt Chapter.

The vertues and qualities of sundrie pretious stones, of cousening Lapidaries, &c.
HE excellent vertues and qualities of stones, found, conceived and tried by
this art, is woonderfull. Howbeit many things most false and fabulous are
added unto their true effects, wherewith I thought good in part to trie the
readers patience and cunning withall. An Aggat (they saie) hath vertue against the
biting of scorpions or serpents. It is written (but I will not stand to it) that it maketh
a man eloquent, and procureth the favour of princes; yea that the fume thereof
dooth turne awaie tempests. Alectorius is a stone about the bignesse of a beane, as
cleere as/ the christall, taken out of a cocks bellie which hath beene gelt or made a 207.
capon foure yeares. If it be held in ones mouth, it asswageth thirst, it maketh the
husband to love the wife, and the bearer invincible: for heereby Milo was said to Ludovic
overcome his enimies. A crawpocke delivereth from prison. Chelidonius is a stone Rhodo.
lect. 11
taken out of a swallowe, which cureth melancholie: howbeit, some authors saie, it is Barthol
lib. 16.
the hearbe wherby the swallowes recover the sight of their yoong, even if their eies
be picked out with an instrument. Geranites is taken out of a crane, and Draconites
out of a dragon. But it is to be noted, that such stones must be taken out of the
bellies of the serpents, beasts, or birds,/ (wherein they are) whiles they live: 294.
otherwise, they vanish awaie with the life, and so they reteine the vertues of those
starres under which they are. Amethysus maketh a droonken man sober, and
refresheth the wit. The* corrall preserveth such as beare it from fascination or * Avice
bewitching, and in this respect they are hanged about childrens necks. But from 2. tract
124.
whence that superstition is derived, and who invented the lie, I knowe not: but I see Serapio
how readie the people are to give credit thereunto, by the multitude of corrals that 100.
waie emploied. I find in good authors, that while it remaineth in the sea, it is an Dioscor
cap. 93
hearbe; and when it is brought thence, into the aire, it hardeneth, and becommeth a
stone.
Heliotropius stancheth bloud, driveth awaie poisons, preserveth health: yea, and
some write that it provoketh raine, and darkeneth the sunne, suffering not him that
beareth it to be abused. Hyacinthus dooth all that the other dooth, and also
preserveth from lightening. Dinothera hanged about the necke, collar, or yoke of any
creature, tameth it presentlie. A Topase healeth the lunatike person of his passion of
lunacie. Aitites, if it be shaken, soundeth as if there were a little stone in the bellie
thereof: it is good for the falling sicknesse, and to prevent untimelie birth.
Amethysus aforesaid resisteth droonkenesse, so as the bearers shall be able to
drinke freelie, and recover themselves soone being droonke as apes: the same
maketh a man wise. Chalcedonius maketh the bearer luckie in lawe, quickeneth the
power of the bodie, and is of force also against the illusions of the divell, and
phantasticall cogitations arising of melancholie. Corneolus mitigateth the heate of
the mind, and qualifieth malice, it stancheth bloudie fluxes, speciallie of women that
are troubled with their flowers. Heliotropius aforesaid darkeneth the sunne, raiseth Plin. lib
shewers, stancheth bloud, procureth good fame, keepeth the bearer in health, and 10.
Albert.
suffereth him not to be deceived. If this were true, one of them would be deerer 7.
than a thousand diamonds. Solin. c

Hyacinthus delivereth one from the danger of lightening, driveth awaie poison and
pestilent infection, and hath manie other vertues. Iris helpeth a woman to speedie
deliverance, and maketh rainebowes to appeere. A Saphire preserveth the members,
and maketh them livelie, and helpeth agues and gowts, and suffereth not the bearer
to be afraid: it hath vertue against/ venome, and staieth bleeding at the nose being 295.
often put thereto. *A Smarag is good for the eiesight, and suffereth not carnall
copulation, it maketh one rich and eloquent. A Topase increaseth riches, healeth the * Rabbi
lunatike passion, and stancheth bloud. Mephis (as Aaron and Hermes/ report out of aphoris
22.
Albertus Magnus) being broken into powder, and droonke with water, maketh Isidor. l
insensibilitie of torture. Heereby you may understand, that as God hath bestowed cap. 3.
upon these stones, and such other like bodies, most excellent and woonderfull Savano
virtues; so according to the abundance of humane superstitions and follies, manie 208.
ascribe unto them either more vertues, or others than they have: other boast that
they are able to adde new qualities unto them. And heerin consisteth a part of
witchcraft and common cousenage used sometimes of the Lapidaries for gaines;
sometimes of others for cousening purposes. Some part of the vanitie heereof I will
heere describe, bicause the place serveth well therefore. And it is not to be forgotten
or omitted, that Pharos magicians were like enough to be cunning therein.
Neverthelesse, I will first give you the opinion of one, who professed himselfe a
verie skilfull and well experimented Lapidarie, as appeereth by a booke of his owne
penning, published under this title of Dactylotheca, and (as I thinke) to be had
among the bookesellers. And thus followeth his assertion:
Evax rex Arabum fertur scripsisse Neroni, Marbodeu
(Qui post Augustum regnavit in orbe secundus) in sua
dactyloth
Quot species lapidis, quæ nomina, quíve colores, pag. 5, 6
Quæq́; sit his regio, vel quanta potentia cuiq́;,
Ocult as* etenim lapidum cognoscere vires, [* Oculta
Quorum causa latens effectus dat manifestos,
Egregium quiddam volumus rarúmque videri.
Scilicet hinc solers medicorum cura juvatur.† [* ,]
Auxilio lapidum morbos expellere docta.
Nec minùs inde dari cunctarum commoda rerum
Autores perhibent, quibus hæc perspecta feruntur.
Nec dubium cuiquam debet falsúmque videri,
Quin sua sit gemmis divinitùs insita virtus:/
Evax an old Arabian king 296.
is named to have writ Englished
A treatise, and on Neros Grace Abraham
Fleming.
to have bestowed it,
(Who in the world did second reigne
after Augustus time)
Of pretious stones the sundrie sorts,
their names, and in what clime
And countrie they were to be found,
their colours and their hue,
Their privie power and secret force,
the which with knowledge true
To understand, their hidden cause
most plaine effects declare:
And this will we a noble thing
have counted be and rare./
The skilfull care of leeches learnd 209.
is aided in this case,
And hereby holpen, and are taught
with aid of stones to chase
}
Vis gemm
Awaie from men such sicknesses lapillorum
as have in them a place. pretiosoru
negatur, q
No less precise commodities occulta es
of althings else therebie rarissiméq
Are ministred and given to men, sensum c
if authors doo not lie,
To whome these things are said to bee
most manifestlie knowne.
It shall no false or doubtfull case
appeare to anie one,
But that by heavenlie influence
each pretious pearle and stone,
}
Hath in his substance fixed force
 and vertue largelie sowne.

Whereby it is to be concluded, that stones have in them cer/teine proper vertues, 297.
which are given them of a speciall influence of the planets, and a due proportion of
the elements, their substance being a verie fine and pure compound, consisting of
well tempered matter wherein is no grosse mixture: as appeareth by plaine proofe of
India and Aethopia, where the sunne being orient and meridionall, dooth more
effectuallie shew his operation, procuring more pretious stones there to be
ingendred, than in the countries that are occident and septentrionall. Unto this Manie m
opinion doo diverse ancients accord; namelie, Alexander Peripateticus, Hermes, authors
named
Evax, Bocchus Zoroastes, Isaac Judæus, Zacharias Babylonicus, and manie more lesse an
beside. and lea

The seventh Chapter.

Whence the pretious stones receive their operations, how curious Magicians use
them, and of their seales.
URIOUS Magicians affirme, that these stones receive their vertues altogether
of the planets and heavenlie bodies, and have not onelie the verie operation
of the planets, but sometimes the verie images and impressions of the
starres naturallie ingraffed in them, and otherwise ought alwaies to have graven
upon them, the similitudes of such monsters, beasts, and other devises, as they
imagine to be both internallie in operation, and externallie in view, expressed in the
planets. As for example, upon the Achate are graven serpents or venomous beasts; Plin. lib
and sometimes a man riding on a serpent: which they know to be Aesculapius, 10.
Albert.
which is the celestiall serpent, whereby are cured (they saie) poisons and stingings 2. ca. 1
of serpents and scorpions. These grow in the river of Achates, where the/ greatest Solin. c
scorpions are ingendred, and their noisomnes is thereby qualified, and by the force Diurius
cap. de
of the scorpions the stones vertue is quickened and increased. Also, if they would complex
induce love for the accomplishment of venerie, they inscribe and expresse in the complex
stones, amiable embracings and love/lie countenances and gestures, words and 210.
kissings in apt figures. For the desires of the mind are consonant with the nature of 298.
the stones, which must also be set in rings, and upon foiles of such metals as have
affinitie with those stones, thorough the operation of the planets whereunto they are
addicted, whereby they may gather the greater force of their working.
As for example, They make the images of Saturne in lead, of Sol in gold, of Luna Geor. P
in silver. Marrie there is no small regard to be had for the certeine and due times to Villang.
medici
be observed in the graving of them: for so are they made with more life, and the super M
influences and configurations of the planets are made thereby the more to abound in dactyl.
them. As if you will procure love, you must worke in apt, proper, and freendlie
aspects, as in the houre of Venus, &c: to make debate, the direct contrarie order is
to be taken. If you determine to make the image of Venus, you must expect to be
under Aquarius or Capricornus: for Saturne, Taurus, and Libra must be taken heed
of. Manie other observations there be, as to avoid the infortunate seate and place of
the planets, when you would bring a happie thing to passe, and speciallie that it be
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