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Neuroanatomy
Text and Atlas
 NOTICE

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Neuroanatomy
Text and Atlas
Fifth Edition

John H. Martin, PhD

Department ofMolecular, Cellular, and Biomedical Sciences
City University ofNew York School ofMedicine
City University ofNew York Graduate Center
New York, New York

Medical Photography by
Howard J. Radzyner, RBP, AIMBI, FBCA

Illustrated by
Michael E. Leonard, MA, CMI, FAMI

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Milan New Delhi Singapore Sydney Toronto
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For Carol, once more and forever
BOX FEATURES
Box 1-1 Development of the Basic Plan of the Brain and
Spinal Cord 1O
Box 1-2 C-shaped Development of the Cerebral
Hemisphere 20
Box 2-1 Anatomical Techniques for Studying the Regional
and Microscopic Anatomy of the Human Central Nervous
System 34
Box 2-2 MRI Visualizes the Structure and Function of the
Living Human Brain 36
Box3-1 Radiological Imaging ofCerebralVasculature
Box 5-1 The Patterns of Somatic Sensory Impairments
After Spinal Cord Injury 108
Box6-1 Cranial Nerve and Nuclei Historical
Nomenclature 125
Box 7-1 Lesion of Different Higher-Order Visual
Areas Produce Remarkably Specific Perceptual
Impairments 161
vi
Box9-1 Adult Neurogenesis in the Olfactory Bulb 201
Box 10-1 Lesions of the Cortical Pathways in the Brain
and Spinal Cord Produce Weakness and Paralysis and
Increased Spinal Reflexes 222
Box 11-1 Cortical Control of Swallowing and Dysphagia
After Stroke 250
Box 13-1 Inhibitory Circuitry of the Cerebellum 293
Box 14-1 Knowledge of the Intrinsic Circuitry of the Basal
Ganglia Helps to Explain Hypokinetic and Hyperkinetic
64 Motor Signs 311
Box 14-2 The Striatum Has a Compartmental
Organization 320
Box 15-1 Lesions in Diverse Locations Can Produce
Horner Syndrome 351
Box 16-1 Circuits of the Hippocampal Formation and
Entorhinal Cortex Are Important for Memory 365
CONTENTS

Preface xi Neurons ofthe Raphe Nuclei Use Serotonin as

Acknowledgments xiii Their Neurotransmitter 33
Guide to Using This Book xv Guidelines for Studying the Regional Anatomy and
Interconnections of the Central Nervous System 33
The Splnal Cord Has a Central Cellular Region
SECTION I THE CENTRAL NERVOUS Surrounded by a Region That Contains

I SYSTEM Myellnated Axons

The Direction of Information Flow Has Its Own
33

SetofTerms 38
1. Organization of the Central Nervous System 3 Surface Features of the Brain Stem Mark Key
Neurons and Glla Are the Two Prlnclpal Cellular Internal Structures 38
Constituents of the Nervous System 5 The Organization of the Medulla Varies
All Neurons Have a Common Morphological Plan 6 From Caudal to Rostral 39
Neurons Communicate With Each Other at Synapses 7 The Pontine Nuclei Surround the Axons of the
Gllal Cells Provide Structural Support for Neurons and Cortlcosplnal Tract In the Base of the Pons 40
Additionally Serve a Broad Set of Diverse Functions 7 The Dorsal Surface of the Midbrain Contains the Col lieu Ii 40
The Nervous System Consists of Separate Perlpheral The Thalamus Transmits Information From Subcortlcal
and Central Components 9 Structures to the Cerebral Cortex 41
The Spinal Cord Displays the Simplest Organization The Internal Capsule Contains Ascending
of All Seven Major Divisions 11 and Descending Axons 43
The Brain Stem and Cerebellum Regulate Body Cerebral Cortex Neurons Are Organized Into Layers 45
Functions and Movements 12 The Cerebral Cortex Has an Input-Output Organization 45
The Dlencephalon Consists of the Thalamus and Hypothalamus 14 The Cytoarchitectonic Map of the Cerebral Cortex Is the
Basis for a Map of Cortical Function 45
The Cerebral Hemispheres Have the Most Complex
Shape of All Central Nervous System Divisions 14
3. Vasculature of the Central Nervous System
The Subcortical Components of the Cerebral Hemispheres
Mediate Diverse Motor, Cognitive, and Emotional Functions 14 and the Cerebrospinal Fluid 53
The Four Lobes of the Cerebral Cortex Each Have Neural Tissue Depends on Continuous
Distinct Functions 15 Arterial Blood Supply 55
Cavities Within the Central Nervous System Contain The Vertebral and Carotid Arteries Supply
Cerebrospinal Fluid 19 Blood to the Central Nervous System 55
The Central Nervous System Is Covered by Three The Splnal and Radlcular Arteries Supply
Meningeal Layers 19 Blood to the Splnal Cord 55
An Introduction to Neuroanatomical Terms 21 The Vertebral and Basilar Arteries Supply
Blood to the Brain Stem 57
2. Structural and Functional Organization The Internal Carotid Artery Has Four Principal Portions 60
of the Central Nervous System 27 The Anterior and Posterior Clrculatlons Supply the
The Dorsal Column-Medial Lemniscal System and Dlencephalon and Cerebral Hemispheres 61
Corticospinal Tract Have a Component at Each Level Collateral Clrculatlon Can Rescue Brain Regions Deprived
of the Neuraxis 30 of Blood 61
The Modulatory Systems of the Brain Have Diffuse Deep Branches of the Anterior and Posterior Clrculatlons
Connections and Use Different Neurotransmitters 31 Supply Subcortical Structures 62
Different Functional Areas of the Cerebral Cortex Are
Neurons ln the Basal Forebraln and Dlencephalon
Su pplled by Different Cerebral Arteries 62
Contain Acetylcholine 32
The Su bstantla Nlgra and Ventral Tegmental Area Contain Cerebral Veins Drain Into the Dural Sinuses 64
Dopaminergic Neurons 33 The Blood-Brain Barrier Isolates the Chemlcal
Neurons in the Locus Ceru leus Give Rise to a Environment of the Central Nervous System From That
Noradrenerglc Projection 33 of the Rest of the Body 67

vii
viii Contents

CSF Serves Many Diverse Functions 69 Cranlal Nerve Nuder are Organized Into
Most of the CSF Is Produced by the Choroid Plexus 71 Distinctive Columns 126
CSF Circulates Throughout the Ventricles Functlonal Anatomy of the Trlgemlnal and
and Subarachnoid Space 71 Vlscerosensory Systems 127
CSF Is Drawn From the Lumbar Cistern 71 Separate Trigeminal Pathways Mediate Touch and
The Dura I Sinuses Provide the Return Path for CSF 73 Pain and Temperature Senses 127
The Vlscerosensory System Originates From the
Caudal Solitary Nucleus 130
SECTION II I SENSORY SYSTEMS Regional Anatomy of the Trlgemlnal and
Vlscerosensory Systems 131
4. Somatic Sensation: Spinal Mechanosensory Separate Sensory Roots Innervate Different Parts of
Systems 79 the Face and Mucous Membranes of the Head 131
Somatic Sensations 81 TheThreeTrlgemlnal Nuclei Are Present at
All Levels of the Brain Stem 133
Functional Anatomy of the Splnal Mechanosensory System 82 The Caudal Solitary and Parabrachial Nuclei Are
Mechanical Sensations Are Mediated by the Dorsal Key Brain Stem Vlscerosensory Integrative Centers 137
Column-Medial Lemniscal System 82 Somatic and Visceral Sensation Are Processed by
Regional Anatomy of the Spinal Mechanosensory System 82 Separate Thalamlc Nuclei 137
The Peripheral Axon Terminals of Dorsal Root Ganglion
Neurons Contain the Somatic Sensory Receptors 82 7. The Visual System 145
Dermatomes Have a Segmental Organization 83 Functional Anatomy of the Visual System 147
The Spinal Cord Gray Matter Has a Dorsoventral Anatomically Separate Visual Pathways Mediate Perception
Sensory-Motor Organization 89 and Ocular Reflex Function 147
Mechanoreceptor Axons Terminate in Deeper Portions The Pathway to the Primary Visual Cortex Is Important
of the Splnal Gray Matter and In the Medulla 89 for Perception of the Form, Colar, Location, and
The Ascending Branches of Mechanoreceptive Sensory Motion of Visual Stlmull 147
Fibers Travel in Dorsal Columns 90 The Pathway to the Midbrain Is Important in Voluntary
The Dorsal Column Nuclei Are Somatotoplcally Organized 90 and Reflexive Control of the Eyes 147
The Decussation of the Dorsal Column-Medial Lemniscal
Regional Anatomy of the Visual System 149
System Is in the Caudal Medulla 90
Mechanosensory Information Is Processed in the The Visual Field of Each Eye Partially Overlaps 149
Ventral Posterior Nucleus 90 Optical Properties of the Eye Transform Visual Stlmull 149
The Primary Somatic Sensory Cortex Has a The Retina Contains Three Major Cell Layers 149
Somatotopic Organization 93 Each Optic Nerve Contains All of the Axons of
The Primary Somatic Sensory Cortex Has a Ganglion Cells In the lpsllateral Retina 151
Columnar Organization 94 The Superior Colliculus Is Important in Ocular Motor
Higher-Order Somatic Sensory Cortical Areas Are Control and Spatial Orientation 153
Located In the Parietal Lobe, Parletal Operculum, The Lateral Geniculate Nucleus Transmits Retinotopically
and Insular Cortex 95 Organized Information to the Primary Visual Cortex 154
The Magnocellular, Parvocellular, and Konlocellular
s. Somatic Sensation: Spinal Systems for Pain, Systems Have Differential Laminar Projections in the
Primary Visual Cortex 155
Temperature, and Itch 99 The Primary Visual Cortex Has a Laminar and
Functlonal Anatomy of the SpinaI Protective Systems 101 Columnar Organization 156
Pain, Temperature, and Itch Are Mediated by the Hlgher-OrderVlsual Cortical Areas Analyze
Anterolateral System 101 Distinct Aspects of Visual Stimuli 158
Visceral Pain Is Mediated by Dorsal Horn Neurons Object Recognition ls Transmitted by the Ventral Stream
Whose Axons Ascend in the Dorsal Columns 101 and Spatial Localization and Action, by the Dorsal Stream 160
Regional Anatomy of the Splnal Protective Systems 105 The Visual Field Changes in Characteristic Ways After
Small-Diameter Sensory Fibers Mediate Pain, Damage to the Visual System 160
Temperature, and Itch 105
Small-Diameter Sensory Fibers Terminate Primarily 8. The Auditory System 167
In the Superflclal Laminae of the Dorsal Horn 105 Functlonal Anatomy of the Auditory System 169
Anterolateral System Projection Neurons Are Parallel Ascending Auditory Pathways Are Involved in
Located In the Dorsal Horn and Decussate tn Different Aspects of Hearing 169
the Ventra I Commissure 106 Regional Anatomy of the Auditory System 171
Vascular Lesions of the Medulla Dtfferentlal ly Affect The Auditory Sensory Organs Are Located Within the
Somatic Sensory Function 109 Membranous Labyrinth 171
Descending Pain Suppression Pathways Originate The Coch Iear Nuclei Are the First Central Nervous
From the Brain Stem 110 System Relays for Auditory Information 173
Several Nudei in the Thalamus Process Pain, The Su perlor Ollvary Complex Processes Stlmulr
Temperature, and Itch 111 From Both Ears for Horizontal Sound Localization 173
Limbic and Insular Areas Contain the Cortical The Ollvocochlear System Regu Iates Auditory Sensitivity
Representations of Pain, Itch, and Temperature in the Periphery 173
Sensations 113 Auditory Brain Stem Axons Ascend in the Lateral Lemniscus 175
The Inferior Colllculus Is Located ln the Mldbraln Tectum 175
6. Somatic Sensation: Trigeminal and The Medial Geniculate Nucleus Is the Thalamic Auditory
Viscerosensory Systems 119 Relay Nucleus 176
Cranlal Nerves and Nude! 121 The Primary Auditory Cortex Comprises Several Tonotopically
Organized Representations Within Heschl's Gyri 177
Important Differences Exist Between the Sensory and
Caudal Secondary and Hlg her-Order Auditory Areas
Motor Innervation of Cranial Structures and Those of the
Give Rise to Projections for Distinguishing the Location
Umbs and Trunk 124
of Sounds 178
There Are Seven Functional Categories of Cranial Nerves 125
 Contents ix

Rostral Secondary and Higher-Order Auditory Areas 11. Cranial Nerve Motor Nuclei and
Give Rise to Projections for Processing the Linguistic
Characteristics of Sounds 179
Brain Stem Motor Functions 237
Damage to Frontotemporal Regions in the Left Organization of Cranlal Motor Nuclel 239
Hemisphere Produces Aphasias 179 There Are Three Columns of Cranial Nerve Motor Nuclei 239
Neurons in the Somatic Skeletal Motor Column
9. Chemical Senses: Taste and Smell 185 Innervate Tongue and Extraocular Muscles 239
The Branchiomeric Motor Column Innervates Skeletal
The Gustatory System: Taste 187
Muscles That Develop From the Branchial Arches 239
The Ascending Gustatory Pathway Projects to
The Autonomic Motor Column Contains Parasympathetic
the lpsilateral Insular Cortex 187
Preganglionic Neurons 241
Regional Anatomy of the Gustatory System 188
The Functlonal Organization of the Cortlcobulbar Tract 242
Branches of the Facial, Glossopharyngeal, and
The Cranial Motor Nuclei Are Controlled by the
Vagus Nerves Innervate Different Parts of the
Cerebral Cortex and Diencephalon 242
Oral Cavity 188
Biiaterai CortlcobulbarTract Projections Innervate
The Solitary Nucleus Is the First Central Nervous System
the Hypoglossal Nucleus, Trigeminal Nucleus, and
Relay for Taste 189
Nucleus Amblguus 243
The Parvocellular Portion of the Ventral Posterior
Cortical Projections to the Facial Motor Nucleus
Medial Nucleus Relays Gustatory Information to
Have a Complex Pattern 243
the Insular Cortex and Operculum 190
Regional Anatomy of Cranial Motor Nuclei
The Olfactory System: Smell 191
and CorticobulbarTract 245
The Olfactory Projection to the Cerebral Cortex
Lesion of the Genu of the Internal Capsule Interrupts
Does Not Relay Through the Thalamus 191
the Cortlcobulbar Tract 246
Regional Anatomy of the Olfactory System 194 The Trigeminal Motor Nucleus Is Medial to the Main
The Primary Olfactory Neurons Are Located In the Trigeminal Sensory Nucleus 246
Nasal Mucosa 194 The Fibers of the Faclal Nerve Have a Complex Trajectory
The Olfactory Bulb Is the First Central Nervous System Through the Pons 247
Relay for Olfactory Input 197 The Glossopharyngeal Nerve Enters and Exits From the
The Olfactory Bulb Projects to Structures on the Rostral Medulla 248
Ventral Brain Surface Through the Olfactory Tract 197 A Level Through the Mid-Medulla Reveals the Locations
The Primary Olfactory Cortex Receives a Direct Input of Six Cranial Nerve Nuclel 249
From the Olfactory Bulb 197 The Spinal Accessory Nucleus Is Located at the Junction
Olfactory and Gustatory lnfonnatlon Interacts In the of the Spinal Cord and Medulla 2S2
Insular and Orbitofrontal Cortex for Sensing Flavors 200
12. The Vestibular System and Eye Movements 257
SECTION Ill I MOTOR SYSTEMS Functlonal Anatomy of the Vestlbular System
An Ascending Pathway From the Vestibular Nuclei to
259

the Thalamus Is Important for Perception, Orientation,

10. Descending Motor Pathways and the and Posture 259
Motor Function of the Spinal Cord 209 The Vestibular System Regulates Blood Pressure in
Functional Anatomy of the Motor Systems Response to Changes in Body Posture and Gravity 260
for Limb Control and Posture 211 The Vestibular Nuclei Have Functionally Distinct
Diverse Central Nervous System Structures Descending Spinal Projections for Axial Muscle Control 262
Comprise the Motor Systems 211 Functlonal Anatomy of Eye Movement Control 262
Many Cortical Regions Are Recruited Into Action The Extraocu lar Motor Neurons Are Located in Three
During Visually Guided Movements 213 Cranial Nerve Motor Nuclei 262
Functional Anatomy of the Descending The Vestlbuloocular Reflex Maintains Direction of
Motor Pathways 213 Gaze During Head Movement 262
Multiple Parallel Motor Control Pathways Originate Voluntary Eye Movements Are Controlled. by Neurons
From the Cortex and Brain Stem 213 in the Frontal Lobe and the Parietal-Temporal-Occipital
Three Rules Govern the Logic of the Organization of Association Cortex 262
the Descending Motor Pathways 215 Regional Organization of the Vestibular and Eye
Two Laterally Descending Pathways Movement Control Systems 264
Control Limb Muscles 215 Vestibular Sensory Organs Are Contained Within the
Four Medially Descending Pathways Control Axial Membranous Labyrinth 264
and Girdle Muscles to Regulate Posture 218 The Vestibular Nuclei Have Functionally Diverse
Regional Anatomy of the Motor Systems and Projections 266
the Descending Motor Pathways 220 The Extraocu lar Motor Nuclei Are Located Adjacent to
the MLF in the Pons and Midbrain 267
The Cortical Motor Areas Are Located in the
Parasympathetic Neurons In the Mldbraln Regulate Pupil Size 270
Frontal Lobe 220
Eye Movement Control Involves the Integrated
The Projection From Cortical Motor Regions Passes
Functions of Many Brain Stem Structures 271
Through the Internal Capsule En Route to the
The Ventral Posterior Nucleus of the Thalamus
Brain Stem and Spinal Cord 224
The Corticospinal Tract Courses in the Base of Transmits Vestibular Information to the Parietal
the Mldbrain 225 and Insular Cortica I Areas 271
The Pontine and Medullary Reticular Formation Multiple Areas of the Cerebral Cortex Function
in Eye Movement Control 273
Gives Rise to the Reticulospinal Tracts 225
The Lateral Cortlcospinal Tract Decussates
in the Caudal Medulla 227
13. The Cerebellum 277
The Intermediate Zone and Ventral Horn of the Gross Anatomy of the Cerebellum 279
Spinal Cord Receive Input From the Descending Functional Anatomy of the Cerebellum 279
Pathways 228 The Cerebellum Has a Basic Circuit 279
x Contents

All Three Functlonal Divisions of the Cerebellum
Display a Similar Input-Output Organization 279
Damage to the Cerebellum Produces Limb Motor Signs
on the Same Side as the Lesion 287
Regional Anatomy of the Cerebellum 289
Spinal Cord and Medullary Sections Reveal Nuclel
and Paths Transmitting Somatic Sensory Information to
the Cerebellum 290
The Inferior Olivary Nucleus Is the Only Source
of Climbing Fibers 291
TheVestlbulocerebellum Receives Input From Primary
and Secondary Vestibular Neurons 291
The Pontlne Nuclei Provide the Major Input to the
Cerebrocerebelium 291
The Intrinsic Circuitry of the Cerebellar Cortex Is
the Same for the Different Functlonal Divisions 291
The Deep Cerebellar Nuclei Are a Major Source of
Input to the Brain Stem and Cortical Motor Pathways 295
A Major Part of the Dentate Nucleus Is Important
for Nonmotor Functions 295
The Cortlcopontlne Projection Brings Information
From Diverse Cortical Areas to the Cerebellum for
Motor Control and Nonmotor Functions 295
14. The Basal Ganglia 303
Organization and Development of the Basal Ganglla 305
Separate Components of the Basal Ganglia Process Incoming
Information and Mediate the Output 305
The Complex Shapes and Fractionation of Basal Ganglia
Components Are Understood by How the
Basal Ganglia Develop 305
Functional Anatomy of the Basal Ganglia 308
Direct and Indirect Pathways Form Common Circuits
Throughout All Functlonal Divisions of the Basal Ganglia 308
Knowledge of Basal Ganglia Connections and
Neurotransmitters Provides Insight Into Their
Function in Health and Disease 310
Functionally Distinct Parallel Circuits Course
Through the Basal Ganglia 314
Integration of Information Between the Basal Ganglia
Loops Is Needed for Adaptive Behaviors 314
Regional Anatomy of the Basal Ganglia 314
The Anterior Limb of the Internal Capsule Separates
the Head of the Caudate Nucleus From the Putamen 316
The Three Components of the Striatum Are Located at
the Level of the Anterior Horn of the Lateral Ventricle 316
The External Segment of the Globus Pal lldus and
the Ventral Pallidum Are Separated bythe
Anterior Commlssure 319
The Ansa Lenticularis and the Lenticular Fascicu lus Are
OutputTracts ofthe Internal Segment of the Globus Pallidus 319
Lesion of the Subthalamlc Nucleus Produces Hemlballlsm 321
The Substantia Nigra Contains Two Anatomical Divisions 321
The Pedunculopontlne Nucleus Is Part ofa Parallel
Path From the Basal Ganglia to Brain Stem
Locomotor Control Centers 322
Stlmu latlon-Based Treatments for Movement and
Nonmovement Disorders Demand a Precise Knowledge
of the Region al Anatomy of the Basal Gang Ila 322
The Vascular Supply of the Basal Ganglia Is
Provided by the Middle Cerebral Artery 323
The Parasympathetic and Sympathetic Divisions
of the Autonomic Nervous System Originate From
Different Central Nervous System Locations 337
Hypothalamic Nuclei Coordinate Integrated Visceral
Responses to Body and Envlronmental Stlmull 340
The Hypothalamus Coordinates Circadian Responses,
Sleep, and Wakefulness 342
Regional Anatomy of the Hypothalamus 344
The Preoptic Area Influences Release of Reproductive
Hormones From the Anterior Pituitary 344
Section Through the Median Eminence Reveals
Parvocellular and Magnocellular Nuclei 344
The Posterior Hypothalamus Contains the
Mammillary Bodies 346
Descending Autonomic Fibers Course in the Periaqueductal
Gray Matter and In the Lateral Tegmentum 346
Nuclei in the Pons Are Important for Bladder Control 347
Dorsolateral Brain Stem Lesions Interrupt
Descending Sympathetic Fibers 347
Preganglionic Neurons Are Located in the Lateral
Intermediate Zone of the Spinal Cord 350
16. The Limbic System and Cerebral Circuits
for Reward, Emotions, and Memory 357
Anatomical and Functional Overview of Neural
Systems for Reward, Emotions, and Memory 359
The Limbic Association Cortex Is Located on the Medial
Surface ofthe Frontal, Parletal, and Temporal Lobes 359
The Hippocampal Formation Plays a Role in
Consolldatlng Explicit Memories 362
The Amygdala Contains Three Major Functional
Divisions for Emotions and Their Behavioral Expression 366
The Mesollmblc Dopamine System and Ventral
Striatum Are Important in Reward 368
Connections Exist Between Components ofthe
Limbic System and the Three Effector Systems 368
All Major Neurotransmitter Regulatory Systems
Have Projections to the Limbic System 370
Reglonal Anatomy of Neural Systems for Emotions,
Learning, and Memory, and Reward 371
The Nucleus Accumbens and Olfactory Tubercle
Comprise Part of the Basal Forebrain 371
Basal Forebrain Cholinergic Systems Have Diffuse
Limbic and Neocortlcal Projections 371
The Cingulum Courses Beneath the Cingulate and
Parahlppocampal Gyrl 374
The Three Nuclear Divisions of the Amygdala Are
Revealed in Coronal Section 374
The Hlppocampal Formation Is Located In the
Floor of the Inferior Hom of the Lateral Ventricle 376
A Sagittal Cut Through the Mammillary Bodies
Reveals the Fornix and Mammillothalamic Tract 378
Nuclei in the Brain Stem Link Telencephalic and
Dlencephal le Limbic Structures With the Autonomic
Nervous System and the Spinal Cord 380
SECTION v I ATLAS
Atlas I: Surface Topography of the Central
Nervous System 387

SECTION IV I INTEGRATIVE SYSTEMS Atlas II: Myelin-Stained Sections Through

the Central Nervous System 403
15. The Hypothalamus and Answers to Clinical Cases 467
Regulation of Bodily Functions 329 Answers to Study Questions 473
Gross Anatomy of the Hypothalamus 331 Glossary 479
Functional Anatomy of the Hypothalamus 333 Index 505
Separate Parvoceli ular and Magnoceliular Neurosecretory
Systems Regulate Hormone Release From the Anterior
and Posterior Lobes of the Pituitary 333
PREFACE
Neuroanatomy plays a crucial role in the health science curric-
ulum. by preparing students to widerstand the anatomical basis
of neurology and psychiatry. Imaging the human brain, in both
the clinical and research setting, helps us to identify its basic
structure and connections. And, when the brain becomes dam-
aged by disease or trauma, imaging locali7.es the extent of the
injury. Functional imaging helps to identify the parts ofthe brain
that become active during our thoughts and actions, and reveals
brain regions where drugs act to produce their neurological and
psychiatric effects. Complementary experimental approaches in
animals-such as mapping neural connections, localizing par-
ticular neuroactive chemicals within different brain regions,
and detennining the effects of lesioning or inactivating a brain
region-provide the neuroscientist with the tools to study the
biological substrates of normal and disordered behavior. To
interpret this wealth of clinical and basic science information
requires a high level of neuroanatomical competence.
Knowledge of human neuroanatomy is becoming increas-
ingly more important for procedures to treat central nervous
system diseases. Therapeutic electrophysiological interventions
target specific brain regions, such as deep brain stimulation
(DBS) of the basal ganglia for Parkinson disease. lnterventional
neuroradiology is a chosen approach for treating many vascular
abnormalities, such as repair of arterial aneurysms. Surgery to
resect a portion ofthe temporal lobe is the treatment ofchoice to
reduce the incidence ofseizures for many patients with epilepsy.
Neurosurgeons routinely use high-resolution imaging tools to
characterize the functions and even the connections of regions
surrounding tumors, to resect the tumor safely and minimize
risk of loss of speech or motor function. Mathematical model-
ing ofbrain tissue characteristics based on high-resolution MRI
is used to guide placement of surface electrodes for transcranial
magnetic and direct current electric stimulation. Each of these
innovative approaches clearly requires that the clinical team
have a sufficient knowledge of functional neuroanatomy-that
is, to have .knowledge ofbrain functions and in whic:b. structures
these functions are localized-to design and carry out these
tasks. And this demand for knowledge of brain structure, func-
tion, and connectivity will only be more important in the future
as higher-resolution imaging and more effective interventional
approaches are developed to repair the damaged brain.
Neuroanatomy helps to provide key insights into disease
by providing a bridge between molecular and clinical neu-
ral science. We are learning the genetic and molecular bases
for many neurological and psychiatric diseases, such as amy-
otrophic lateral sclerosis, Huntington disease, and schizophre-
nia. Localizing defective genes to particular brain regions,
neural circuits, and even neuron and glial cell classes helps to
further our understanding of how pathological changes in brain
structure alter brain function. And this knowledge, in turn, will
hopefully lead to breakthroughs in treatments and even cures.
An important goal of Neuroanatomy: Thxt and Atlas is to
prepare the reader for interpreting the new wealth of human
brain images-structural, functional, and connectivity-by
developing an understanding of the anatomical localization
of brain functions. To provide a workable focus, this book is
largely restricted to the central nervous system. It takes a tra-
ditional approach to gaining neuroanatomical competence:
Because the basic imaging picture is a two-dimensional slice
through the brain (e.g., CT or MRI scan), the locations of struc-
tures and consideration of their functions are examined on
two-dimensional myelin-stained sections through the human
central nervous system.
All chapters have been revised for the fifth edition of
Neuroanatomy: Text and Atlas to reflect advances in neural
science since the last edition, with many new full color illus-
trations. Designed as a self-study guide and resource for infor-
mation on the structure and function of the human central
nervous system, this book can serve as both text and atlas for an
introductory laboratory course in human neuroanatomy.
For over 30 years, both at Columbia University's College
of Physicians and Surgeons and now at the City University of
New York's Medical School. we use this book in conjunction
with a series of neuroanatomy laboratory exercises during the
neuroscience/neurology-psychiatry teaching block in the cur-
riculum. Rather than presenting the material in a traditional
lecture format, we have successfully taught neuroanatomy in
a dynamic small group learning environment. Supplemented
with use of brain models and specimens, neuroanatomy small
group sessions complement neural science, neurology, and psy-
chiatry lecture material and round-out medical, graduate, and
allied health science students' learning experience.
xl
xii Preface
The organization of Neuroanatomy: Text and Atlas contin-
ues to parallel that of Principles of Neural Science, edited by
Eric R. Kandel, Steven A. Siegelbaum, Sarah Maclc, and John
Koester (McGraw Hill). Like Principles of Neural Science, Neu-
roanatomy: Text and Atlas is aimed at medical students, and
graduate students in neuroscience, biology, and psychology
programs. The content of many of the chapters is geared to
dental students, such as a chapter focus on the trigeminal system,
as well as physical therapy and occupational therapy students
by considering the motor systems in detail. Neuroanatomy: Text
and Atlas is also being used by neurology and neurosurgery res-
idency programs, as part of their didactic learning experiences
and board certification review courses.
John H. Martin
ACKNOWLEDGMENTS
I take this opportunity to recogniu the help I received in the
preparation of the fifth edition of Neuroanatomy: Tuxt and
Atlas. I am grateful to the following friends and colleagues who
have read portions of the manuscript or have provided radio-
logical or histological mate.rials for this or previous editions:
Dimitris Agamanolis, David Amaral, Richard Axel, Berti.I. Blok.
Eric Bushong, Bud Craig. Mike Crutcher, Maurice Curtis,
Adrian Danek,AniruddhaDas, Sam David, MonydeLeon, John
Dowling, Mark Ellisman, Susan Polstein, Blair Ford, Peter Pox,
Stephen Prey, Eitan Friedman, Guido Gainotti, Lice Ghilardi,
MickeyGoldberg, James Goldman, PatGoldman-Rakic,. Suzanne
Haber, Shaheen Hamdy, Andrei Holodny. Jonathan Horton,
David Hubel, Matilde Inglese, Sharon Juliano, Joe LeDoux,
Kevin Leung, Marge Livingstone, Camillo Marra, Randy
Marshall, Etienne Olivier, Elizabeth Pimentel, Jems Pujol, Josef
Rauschecker, David Ruggiero, Neal Rutledge, Thomas Schultz,
Brian Somerville, Bob Vassar, Bob Waters, Torsten Wiesel,
Rachel Wong, and Semir Zeki. I also would like to thank Alice
Ko for help with the three-dimensional reconstructions that
provided the basis for various illustrations. I am grateful to
Dr. Frank Galliard, who created the Radiopaedia.com web-
site, for selection of many fine MR.Is illustrating neurological
dam.age. I would especially like to thank Dr. Joy Hirsch, Steve
Dashnaw, and Glenn Castilo for many of the high-resolution
MRls fust used in the fourth edition. Many thanks also to
my colleagues at the City College of New York, Liz Pimentel,
Dr. Jay Edelman, and Gary Temple, for pointing out errors and
offering countless helpful suggestions.
I would like to extend a special note of thanks to members
of the neuroanatomy teaching faculty at the College of Phf5i-
cians and Surgeons and the City University of New York Med-
ical School for many helpful discussions. For new and revised
illustrations, I thank the Dragonfly Media Group, and especially
Rob PedirkD for bringing to fruition the many facets of the
complex art program. For artwork carried over from previous
editions, I thank Michael Leonard, the original illustrator and
Dragonfly Media Group. I especially thank Howard Radzyner
for the superb photographs of myelin-stained brain sections
that have helped to define Neuroanatomy: Text and Atlas from
its first edition. At McGraw Hill, I greatly appreciate the hard
work and patience of Peter Boyle, project development editor,
and Catherine Saggese, senior production supervisor. I also
thank Garima Poddar at Cenveo Publisher Services. Finally, I
would like to thank my editor Michael Weitz fo.r his support,
patience, and guidance-not to mention timely pressure-in
the preparation of the fifth edition.
xiii
This page intentionally left blank
GUIDE TO USING THIS BOOK
Neuroanatomy: Text and Atlas takes a combined regional and
functional approach to teaching neuroanatomy: Knowledge
of the spatial interrelations and connections between brain
regions is developed in relation to the functions of the brain's
various components. The book first introduces the major
concepts of central nervous system organization. Subse-
quent chapters consider neural systems subserving particular
sensory, motor, and integrative functions. At the end of the
book is an atlas of surface anatomy of the brain and myelin-
stained histological sections, and a glossary of key terms and
structures.
Each chapter begins with a clinical case to illustrate the con-
nections and function of the key material There are key ques-
tions in the case that the reader can answer based primarily
on the chapter readings, but also on prior chapter material.
Chapters also end with a series of multiple choice review ques-
tions. Answers both to the case and review questions are at the
back of the book. Material on central nervous system develop-
ment is included in the relevant individual chapters.
Overview of Chapters
The general structural organization of the mature central
nervous system is surveyed in Chapter 1. This chapter also
introduces neuroanatomical nomenclature and fundamental
histological and imaging techniques for studying brain struc-
ture and function. The three-dimensional shapes of key deep
structures are also considered in this chapter. The functional
organization of the central nervous system is introduced in
Chapter 2. This chapter considers how different neural cir-
cuits, spanning the entire central nervous system, serve partic-
ular functions. The circuits for touch perception and voluntary
movement control are used as examples. The major neuro-
transmitter systems are also discussed.
Central nervous system vasculature and cerebrospinal fluid
are the topics of Chapter 3. By considering vasculature early in
the book, the reader can better understand why particular func-
tions can become profoundly disturbed when brain regions are
deprived of nourishment. These three chapters are intended to
provide a synthesis of the basic concepts of the structure of the
central nervous system and its functional architecture. A funda-
mental neuroanatomical vocabulary is also established in these
chapters.
The remaining 13 chapters examine the major functional
neural systems: sensory, motor, and integrative. These chapters
reexamine the views of the surface and internal structures of the
central nervous system presented in the introductory chapters,
but now from the perspective of the difrerent functional neu-
ral systems. As these latter chapters on functional brain archi-
tecture unfold, the reader gradually builds a neuroanatomical
knowledge of the regional and functional organiution of the
spinal cord and brain, one system at a time.
These chapters on neural systems have a different organi-
zation from that of the introductory chapters: Eacll is divided
into two parts, functional and regional neuroanatomy. The
initial part. on functional neuroanatomy. considers how the
brain regions that comprise the particular neural system work
together to produce their intended functions. This part of the
chapter presents an overall view offunction in relation to struc-
ture before considering the detailed anatomical orgaruzation of
the neural system. Together with descriptions of the functions
of the various components, diagrams illustrate each system's
anatomical organization. including key connections that help
to show how the particular system accomplishes its tasks.
Neural circuits that nm through various divisions ofthe brain are
depicted in a standardized format: Representations of myelin-
stained sections through selected levels of the spinal cord and
brain stem are presented with the neural circuit superimposed.
Regional neuroanatomy is emphasized in the latter part of
the chapter. Here, structures are depicted on myelin-stained
histological sections through the brain, as well as magnetic res-
onance images (MRis). These sections reveal the locations of
major pathways and neuronal integrative regions. Typically, this
part examines a sequence of myelin-stained sections ordered
according to the flow of information processing in the system.
For example, coverage of regional anatomy of the auditory sys-
tem begins with the ear, where sounds are received and initially
processed, and ends with the cerebral cortex. where our per-
ceptions are formulated. In keeping with the overall theme of
the book. the relation between the structure and the function of
discrete brain regions is emphasized.
Emphasis is placed on the close relationship between neu-
roanatomy and neuroradiology especially through use of MRI
scans. These scans are intended to facilitate the transition
from learning the actual structure of the brain, as revealed by
xv
xvi Guide To Using This Book
histological sections, to that which is depicted on radiological
images. This is important in learning to "read" the scans, an
important clinical skill. MRI scans are presented either using
the radiological convention of showing the ventral surface of
the brain up or, when the focus is learning detailed regional
anatomy, showing the ventral surface down together with corre-
sponding myelin-stained sections. It should be recognized that
there is no substitute for actual stained brain sections for devel-
oping an understanding of regional anatomy and localization
of function. This is because current MRI resolution is not suf-
ficient to reveal the breadth of brain and spinal cord structures
whose functions need to be considered.
Atlas of the Central Nervous System
This book's atlas, in two parts, offers a complete reference of
anatomical structure. The first part presents key views of the
surface anatomy of the central nervous system. This collec-
tion of drawings is based on actual specimens but emphasizes
common features. Thus, no single brain has precisely the form
illustrated in the atlas. The second part of the atlas presents
a complete set of photographs of myelin-stained sections
through the central nervous system in three anatomical planes.
With few exceptions, the same surface views and histolog-
ical sections used in the atlas are also present in the chapters.
In this way, the reader does not have to cope with anatomical
variability and is thus better able to develop a thorough under-
standing of a limited, and sufficiently complete, set of materials.
Moreover, brain views and histological sections shown in the
chapters have identified only the key structures and those
important for the topics discussed. In the atlas, all illustrations
are comprehensively labeled as a reference. The atlas also serves
as a useful guide during a neuroanatomy laboratory.
Didactic Boxes
Selected topics that complement material covered in the
chapters are presented in boxes. In many of the boxes, a
new perspective on neuroanatomy is presented, one that
has emerged only recently from research. The neuroscience
community is enthusiastic that many of these new perspec-
tives may help explain changes in brain function that occur
following brain trauma or may be used to repair the damaged
nervous system.
Clinical Cases
Each chapter begins with a clinical case, chosen to highlight a
fascinating clinical feature of the neural system discussed in the
chapter. Whereas some of these cases are rare and not apt to be
seen in routine medical practice, they show how perception,
motor behavior, or personality and emotions can change after
a stroke or tumor damages the brain, or how brain structure
and function change after a particular gene mutation. The case
description is followed by an explanation of what structures
and neural systems are damaged that produce the neurological
signs. Questions are posed that can be answered on the basis
of reading the case explanations and the chapter text. Detailed
answers are provided at the end of the book.
Study Questions
Each chapter ends with a set of study questions. Answers are
provided at the end of the book. A brief explanation of the
more integrative and difficult questions also is provided.
Glossary
The glossary contains a listing of key terms and structures.
Typically, these terms are printed in boldface within the
chapters. Key terms are defined briefly in the context of their
usage in the chapters. Key structures are identified by location
and function.
Additional Study Aids
This book offers three features that can be used as aids in learn-
ing neuroanatomy initially, as well as in reviewing for examina-
tions, including professional competency exams:
• Summaries at the end of each chapter, which present
concise descriptions of key structures in relation to their
functions.
• A glossary of key terms.
• The atlas of key brain views and myelin-stained histologi-
cal sections, which juxtapose unlabeled and labeled views.
The unlabeled image can also be used for self-testing, such
as for structure identification.
These study aids are designed to help the reader assimilate effi-
ciently and quickly the extraordinary amount of detail required
to develop a thorough knowledge of human neuroanatomy.
This page intentionally left blank
 Organization of t~e
Central Nervous System
CLINICAL CASE I
79-Year-Old Man With
Memory Impairment
A 79-year-old man has become forgetful, often mlsplaclng
Ttems at home, and sometimes Is confused when paying for
his groceries. His famlly reports that his forgetfulness seems
to be getting worse. On neurological examlnat1on, he reports
the correct date and knows where he Is and why he Is there;
he has normal speech. However, he ls unable to recall three
unrelated words 5 minutes after correctly repeating them.
When asked to perform simple addition and subtraction, he
ls slow and has dlfflculty. His mental status was further eval-
uated by neuropsychologlcal test1ng, which revealed addi-
tional cognitive Impairments.
Magnetic resonance Images (MRls) of the patient and a
healthy control are presented (Figure 1-1A 1-4, 81-4}. In
these Images, which were obtained using a particular MRI
protocol termed T1 weighting, white and gray matter of the
brain appear as different shades of gray and cerebrosptnal
fluid, black. Cranial fatty substances (eg, In skin and the
bony orbits) are white. Note how the ventricles, which
are fluld-fllled cavttles, are thin In the healthy brain (right
column), but dilated tn the brain of the patient Oeft column).
Note also how the gray and white matter are both thick
tn the healthy brain and th1nner In the patient's bratn. The
hlppocampal formatton (Figures 1-1A4, 84, 1-10A; see
Chapter 16) also Is atrophic In the pattent's brain. The gener-
alized cortical atrophy and ventricular enlargement are also
apparent on the other MRls.
The patient died several years later after developing
severe dementia. At autopsy, his brain was found to show
clear evidence of degeneratton compared with the brain
from a healthy person (AS, BS). The gyri of the cerebral cortex
are narrow, and the sulci are widened. By contrast, the exter-
nal characteristics of the brain stem and cerebellum were
unremarkable.
You should try to answer the following questions based
on your reading of the chapter and inspection of the images.
Note that the description of key neurological signs that fol-
low the questions also will provide the answers.
1. Why is the ventricular system affected, even though it
is a non-neuronal structure?
2. Are some brain areas more severely affected than
others In the patient?
CHAPTER CONTENTS
Neu runs and Cilia Are dte Two Principal Cellular Constitue11ts of the
Nemm S)'Stem
All Neurons Have a Common Morphological Plan
Neurons Communicate Mth Each Other at Synapses
Gllal Cells P~lde Sttuctural Support for Neurons and Addltionally Serve a
Broad Set of Diverse Fundlons
The Nenom s,stem ComistsofSep.nt.e l\qlheral and Central Components
The Spinal Cord Displays dte Simplest Organization of All Sevell Major
Dtvlslons
The Bra1n Stem and Cerellellum Regulm Body Funcdllns and Movements
The Dle11c.ephalon Consists ofdie Thalamus and Hypothalamus
The Cerebral Hemispheres Have the Most Complex Shape of All Central
Nemm S)'Stem Divisions
The Subcortical Components of the Cerebral Hemispheres Mediate Diverse
Motor, Cognitive, and Emotional Functions
The Four l.cbesoftheCerebral Cortex Each Have Distinct Functions
Cavl11es W1dltn the Central Nervom System Contain Cerebru.splnal Fluld
The Central Nervous Sptem Is Coven?d byThree Meningeal Layers
An lntruduction to Neuroana:tomic:al Terms
Box 1-1. Development of the Bask Plan of the Bra.In and Spinal Cord
Box 1-2. C;haped Development of1heCerellr.ll Hemisphere
Summary
Sele<ted Readings
Additional Referet1w
3. Among the various bra1n regions affetted by the neu-
ropathologlcal process, which Is most closely associ-
ated with the patient's memory Impairment?
4. Autopsy revealed that the density of acetylcholine-
containing neurons in a part of the forebrain was
severely reduced in the patient. What impact might
this have on the function of cortical neurons?
s. At autopsy it was discovered that the patient had large
accumulations of amyloid plaques, which contain beta
amyloid protein, as well as neurofibrillary tangles,
which consist of an abnormal form of the microtu-
bule-associated protein tau. What is the significance of
these neuropathological findings?
-Continued next page
3
4 Section I • The Central Nervous System

A1

A1,B1
A2,B2
A3,B3

...~IS4 A2

Cortical atrophy
and widening
ofsulci
Third ventricle-~.........
- - -'=4-- -- Midbrain r~
--...."""==~--Atrium and~
posterior horn
of lateral ventricle

A3

lnferior horn of
lateral ventricle
and hippocampal
formation

A5

FIGURE 1-1. MRls (traRSVetSe plane, 1-3; coronal plane, 4) from a person ~h Alzheimer disease {A) and a healthy person (B), The brain views (S) show
generalized atrophy In Alzhelmer disease.The MRls {1-4) show conic.al at!Ophy and ventricular enlargement. The MRls are T1 Images; brain tissues are shades
of gray and cerebrosplnal fluid, black. {A 1, A2, A3, Images reproduced with permission from Dr. Frank Galllard, Radlopaedla.com. M, Image courtesy of The
Dementia Research Center, Ua. lnstftute of Neurology. AS, Image courtesy of Dr. Many J de Leon [NYU Sdlool of Medicine], Dr.JerzyWeglel [lnstttutefor
Basic Researdl], and Dr. Thomas Wisniewski [NYU School of Medicine]; NIH Alzheimer's Disease Center P30 AG08051 J

Conclusion
The patient had Alzheimer disease, which is a neurodegener-
ative disease. The disease produced profound impairments of
cognition, including memory disturbances, and widespread
degeneration of the cerebral cortex.
Key neurological signs and corresponding damaged
brain structures
Brain of person with Alzheimer disease and healthy brain
No description is necessary; the amount and extent of cor-
tical atrophy is obvious in the brain of the person who had
Alzheimer disease (part AS}. Cortical atrophy is accompa-
nied by atrophy in subcortical structures as well. Because
the volume of the skull is fixed, as brain tissues decrease in
volume, there is a corresponding increase in ventricular vol-
ume. Thus, ventricular enlargement is a consequence of loss
of neural tissue.
MRls
Both the generalized cortical atrophy and ventricular enlarge-
ment can be seen on MRls of the brain. A superior-to-inferior
sequence of three images in the transverse plan (see insets}
is shown. The MRI in part 1 slices through the anterior horn
and atrium of the lateral ventricles, where enlargement is
The human nervous system carries out an enormous num-
1 ber of functions by means of many subdivisions. Indeed, the
brain's complexity has traditionally made the study of neuro-
anatomy a demanding task. This task can be greatly simplified
by approaching the study of the nervous system from the dual
perspectives of its regional and functional anatomy. Regional
neuroanatomy examines the spatial relations between brain
structures within a portion of the nervous system. Regional
neuroanatomy defines the major brain divisions as well as local,
neighborhood relationships within the divisions. In contrast,
functional neuroanatomy examines those parts of the ner-
vous system that work together to accomplish a particular task,
for example, visual perception. Functional systems are formed
by specific neural connections within and between regions of
the nervous system; connections that form complex neural
circuits. A goal of functional neuroanatomy is to develop an
understanding of the neural circuitry underlying behavior. By
knowing regional anatomy together with the functions of par-
ticular brain structures, the clinician can determine the loca-
tion of nervous system damage in a patient who has a particular
neurological impairment and, in many cases, a psychiatric
impairment. Combined knowledge of what structures do and
where they are located is essential for a complete understand-
ing of nervous system organization. The term neuroanatomy is
Chapter 1 • Organization of the Central Nervous System S
enormous. Because of the extensive cortical atrophy, the
cortical sulci are wider and filled with more cerebrospinal
fluid. Note the region around lateral sulcus and insular cor-
tex (Figure 1-1A1, where the mixture of a greater amount of
cerebrospinal fluid and thinned cortex produces a large dark
region. The inset in Figure 1-11 A illustrates the insular cortex.
The hippocampal formation is key to consolidation of short-
term to long-term memory {see Chapter 16). Its reduction in
Alzheimer disease, together with degeneration of temporal
lobe cortex, leaves a gaping hole in the temporal lobe.
Hippocampal degeneration can explain why the patient has
poor memory. Although not visible on these images, a small
nucleus on the inferior brain surface, the basal nucleus, is
severely affected early in Alzheimer disease. This nucleus con-
tains neurons that use the excitatory neurotransmitter acetyl-
choline. These neurons project widely throughout the cortex,
and with their loss, many cortical neurons are deprived of
excitatory input. This, together with the gross degeneration,
helps to explain the cognitive impairments in the patient.
These images also reveal that the brain stem is not grossly
affected. The sizes of the midbrain (parts A2-A3 and B2-B3}
and pons (parts A5 and B5} appear normal.
Reference
Brust JCM. The Practice ofNeural Science. New York, NY: McGraw-Hill;
2000.
therefore misleading because it implies that knowledge of struc-
ture is sufficient to master this discipline. Indeed, in the study of
neuroanatomy, structure and function are tightly interwoven-
so much so that they should not be separated. The interrela-
tionships between structure and function underlie functional
localization, a key principle of nervous system organization.
This chapter examines the organization of the nervous sys-
tem and the means to study it by developing the vocabulary to
describe its regional anatomy. First, the cellular constituents
of the nervous system are described briefly. Then the chapter
focuses on the major regions of the nervous system and the
functions of these regions. This background gives the reader
insight into functional localization.
Neurons and Glia Are the Two Principal Cellular
Constituents of the Nervous System
The nerve cell, or neuron, is the functional cellular unit of
the nervous system. Neuroscientists strive to understand the
myriad functions of the nervous system partly in terms of the
interconnections between neurons. The other major cellular
constituent of the nervous system is the neuroglial cell, or glia.
Glia, once thought only to provide structural and metabolic
support for neurons, are now recognized also to be important
6 Section I • The Central Nervous System

players with neurons in diverse brain functions, such as neu-
ral circuit development, learning, and the nervous system's
response to injury.
All Neurons Have a Common Morphological Plan
It is estimated that there are about 100 billion neurons in the
adult hwnan brain. Although neurons come in different shapes
and sizes, each has four morphologically specialized regions
with particular functions: dendrites, cell body, axon, and axon
terminals (Figure l-2A). Dendrites receive information from
other neurons. The "11 body contains the nucleus and cellular
organelles critical for the neuron's survival and function. The cell
body also receives information from other neurons and serves
important integrative functions. The uon conducts informa-
tion, which is encoded in the form of action potentials, to the
axon terminal. Connections between two neurons in a neural
circuit are made by the axon terminals of one and the dendrites
and cell body of the other, at the synapse (discussed below).
Despite a wide range of morphology, we can distinguish
three classes of neuron based on the configuration of their
dendrites and axons: unipolar, bipolar, and multipolar
(Figure l-2B). These neurons were drawn by the distinguished
Spanish neuroanatomist and Nobel laurate Santiago Ram6n y
Cajal at the beginning of the twentieth century. Unipolar
neurons are the simplest in shape (Figure l-2Bl). They have
no dendrites; the cell body of unipolar neurons receives and
integrates incoming information. A single axon, which origi-
nates from the cell body, gives rise to multiple processes at the
terminal. In the human nervous system, unipolar neurons are
the least common. They control exocrine gland secretions and
smooth muscle contractility.
Bipolar neurons have two processes that arise from opposite
poles of the cell body (Figure 1-282). The flow of information
in bipolar neurons is from one of the processes, which func-
tion like a dendrite, across the cell body to the other process,
which functions like an axon. A bipolar neuron subtype is a

A B1 B2

B3

,.
FIGURE 1-2. Neuran1 are the functional mllular unit of the nenrau1 5)'5tem. A. A Khemiltic nerve cell is shown, illustrating tile dendrites, cell body, and
axon. Dendrltlc spines are located on the dendrites. These are sites of excitatory synapses. Inhibitory synapses are located on the shaft of the dendrites, the
cell body, and the lnltlal segment. The axon can be seen to emerge fiom the cell body. The presynaptlc termlnals of the neuron are shown synapslng on the
cell bodies of the postsynaptlc neurons. The Inset shows the spatial relations of three components of the synapse: the presynaptic axon terminal, the synaptic
d@ft, and the postsynaptlc neuron. B. Selected examples of three neuron classes: (Sf) unlpolar, (B2) bipolar, and (83) multlpolar. (A. Adapted from Kandel ER,
Schwartz JH, Jessell lM, eds. Principles ofNeural Science. 4tli ed. NewYork, NY: McGraw-Hiii, 2000. B. Reproduced wltli permission from Cajal SR. Hlstolog~du
sys~me nerveux de t'homme et des verttbrel. 2 vols. Malolne, 1909-1911 J

pseudounipolar neuron (see Figure 6-3 top). During develop-
ment the two processes of the embryonic bipolar neuron fuse
into a single process in the pseudounipolar neuron, which
bifurcates a short distance from the cell body. Many sensory
neurons, such as those that transmit information about odors
or touch to the brain, are bipolar and pseudounipolar neurons.
Multipolar neurons feature a complex array of dendrites
on the cell body and a single axon that branches extensively
(Figure l-2B3). Most of the neurons in the brain and spinal
cord are multipolar. Multipolar neurons that have long axons,
with axon terminals located in distant sites, are termed pro-
jection neurons. Projection neurons mediate communication
between regions of the nervous system and between the ner-
vous system and peripheral targets, such as striated muscle
cells. The neuron in Figure l-2B3 is a particularly complex
projection neuron. The terminals of this neuron are not shown
because they are located far from the cell body. For this type of
neuron in the human, the axon may be up to 1 m long, about
50,000 times the width of the cell body. Other multipolar neu-
rons, commonly called interneurons, have short axons that
remain in the same region of the nervous system in which the
cell body is located. Interneurons help to process neuronal
information within a local brain region.
Neurons Communicate With Each Other at Synapses
Information flow along a neuron is polarized. The dendrites and
cell body receive and integrate incoming information, which is
transmitted along the axon to the terminals. Communication of
information from one neuron to another also is polarized and
occurs at specialized sites of contact called synapses. The neu-
ron that sends information is the presynaptic neuron and the
one that receives the information is the postsynaptic neuron.
The information carried by the presynaptic neuron is most typ-
ically transduced at the synapse into a chemical signal that is
received by specialized membrane receptors on the dendrites
and cell body of the postsynaptic neuron.
The synapse consists of three distinct elements: (1) the pre-
synaptic terminal, the axon terminal of the presynaptic neuron,
(2) the synaptic cleft, the narrow intercellular space between
the neurons, and (3) the receptive membrane of the postsynap-
tic neuron. Synapses are present on dendrites, the cell body, the
initial segment of the axon, or the portion of the axon closest
to the cell body, and the presynaptic axon terminal. Synapses
located on different sites can serve different functions.
To send a message to its postsynaptic neurons, a presynap-
tic neuron releases neurotransmitter, packaged into vesicles,
into the synaptic cleft. Neurotransmitters are small molecular
weight compounds; among these are amino acids (eg, gluta-
mate, glycine, and y-aminobutyric acid [GABA]), acetylcholine,
and monoaminergic compounds such as norepinephrine and
serotonin. Larger molecules, such as peptides (eg, enkephalin
and substance P), also can function as neurotransmitters. After
release into the synaptic cleft, the neurotransmitter molecules
diffuse across the cleft and bind to receptors on the postsyn-
aptic membrane. Neurotransmitters change the permeability of
the neuronal membrane to particular ions. A neurotransmitter
can either excite the postsynaptic neuron by depolarizing it or
inhibit the neuron by hyperpolarizing it. For example, excita-
tion can be produced by a neurotransmitter that increases the
Chapter 1 • Organization of the Central Nervous System 7
flow of sodium ions across the membrane and into a neuron
(ie, depolarization), and inhibition can be produced by a neu-
rotransmitter that increases the flow of chloride ions into a
neuron (ie, hyperpolarization). Glutamate and acetylcholine
typically excite neurons, whereas GABA and glycine typically
inhibit neurons.
Many neurotransmitters, like dopamine and serotonin, have
more varied actions, exciting some neurons and inhibiting oth-
ers. Their action depends on a myriad of factors, such as the
particular membrane receptor subtype that the neurotransmit-
ter engages and whether the binding of the neurotransmitter to
the receptor leads directly to the change in ion permeability or
if the change is mediated by the actions on second messengers
and other intracellular signaling pathways (eg, G protein-
coupled receptors). For example, the dopamine receptor subtype 1
is depolarizing, whereas the type 2 receptor is hyperpolarizing;
both act through G protein-coupled mechanisms. A neuro-
transmitter can even have opposing actions on the same neuron
depending on the composition of receptor subtypes on the neu-
ron's membrane. Action through second messengers and other
intracellular signaling pathways can have short-term effects,
such as changing membrane ion permeability, or long-term
effects, such as changing gene expression. Many small mole-
cules that produce strong effects on neurons are not packaged
into vesicles; they are thought to act through diffusion. These
compounds, for example, nitric oxide, are produced in the
postsynaptic neuron and are thought to act as retrograde mes-
sengers that serve important regulatory functions on pre- and
postsynaptic neurons, including maintaining and modulating
the strength of synaptic connections. These actions are impor-
tant for learning and memory.
Although chemical synaptic transmission is the most com-
mon way of sending messages from one neuron to another,
purely electrical communication can occur between neurons.
At such electrical synapses, there is direct cytoplasmic continu-
ity between the presynaptic and postsynaptic neurons, through
gap junctions.
Glial Cells Provide Structural Support for Neurons and
Additionally Serve aBroad Set of Diverse Functions
Glial cells comprise the other major cellular constituent of
the nervous system; they outnumber neurons by about 10
to 1. Given this high ratio, the functions of glial cells must
be complex and diverse. There are two major classes of glia:
macroglia and microglia. Macroglia, of which there are four
separate types-oligodendrocytes, Schwann cells, astrocytes,
and ependymal cells-have a variety of support and nutritive
functions. Schwann cells and oligodendrocytes form the
myelin sheath around peripheral and central axons, respec-
tively {Figures 1-2A and 1-3). The myelin sheath increases
the velocity of action potential conduction. It is whitish in
appearance because it is rich in a fatty substance called myelin,
which is composed of many different kinds of myelin proteins.
Schwann cells also play important roles in organizing the for-
mation ofthe connective tissue sheaths surrounding peripheral
nerves during development and in axon regeneration following
damage in maturity. Astrocytes have important structural and
metabolic functions. For example, in the developing nervous
system, astrocytes act as scaffolds for growing axons and guides
8 Section I • The Central Nervous System

Astrocyte cell body

and processes
Neuron cell body
and processes

Oligodendrocyte:
cell body and
proce88e8 forming the myelin sheath

c D
Dl D2
,/""' Oligodendrocyte

~
Myelin sheath

D3

Cytoskeletal £ilaments in axon

FIGURE 1-S. Astroc.ytu and oll9od1ndrocyhls ant the most ubiquitous types of gllal calls In the carrtnil nervous systam. Parts A and Bare hlstologlcal
sections showing examples of these cell types. A. An astrocyte {green) Is shown envelopIng a nwrcnal cell body {red}. B. Ollgodendrocytes are shown
fonning the myelin sheaths surrounding axons. A blue stain (OAPO marks nuclei in the cell bodies. The processes {green) are stained for an important
component of the rnyelln sheath, myelln basic protefn (MBP). C. Schematic drawing of an ollgodendrocy1:e and the myelln sheath that It forms around an
axon. Note the multlple wrappings of the ollgodendrocyte process around the axon. The node of Ranllfer Is not covered by the ollgodendrccyte pn>Ce$5,
forming a gap between the axon membrane and the extracellular space. Cytoskeletal fllaments are present In the axon and mitochondria are In the region
of the node of Ranvler. D. Mlcrogllal cells. Mlcroglla phagocytose proteins as well as cells In the CNS that express receptors on their membranes that enable
targeting by mlcroglla. In an anlmal model of Alzhelmer disease, a micrograph shows a mlaogllal process (red) that Is contacting a beta amylold plaque
(green; D1). Subsequent to engulfment, the plaque within the miaoglial cell is degraded and digested (D2, yellow). In a model of net1rcnal degeneration
(03), this 30 reconsttuction, from a series of micrographs, shows a microglial cell (red) contacting a neuron (green) in the spinal cord. (A, Image courtesy
of Elllsman M, Bushong E'. Unlv. <:.allfomla. San Diego. Allen NJ, 8am!$ BA. ~rosclence glla: more than Just brain glue. Notunt. 2009;457 [7230]:675-677.
8, Reproduced with permission from Lee PR, Flelds RD. Regulatlon of myelln genes lmpllcated In psychiatric disorders by functlonal activity In axons.
Front Nl!uroanat 2009;3:4. C. Adapted from Kandel ER.. Schwartz JS, Jessel! TM, eds. Ptfnctples ofNeuml Scl61Cf.'. 4th ed. New Yorlc. NY: McGraw-Hiii; 2000J 01, 02,
Courtesy of Jasmine L Pathan and Dr. Gwenn A. Garden {Department of Neurology, School of Medldne. University ofWashington, Seattle, WA); 03, Jiang Y,
Sarkar A, Amer A. Martin J. Transnaironal down-regulation of the premotor cholinergic 5)'5tem after corticospinal tract loss.J Neurosci (2018) 38(39):
8329-8344. PMIO: 30049887).
 Chapter 1 • Organization of the Central Nervous System 9

for migrating immature neurons. Many synapses are associated
with astrocyte processes that may monitor synaptic actions
and provide chemical feedback. Astrocytes also contribute to
the blood-brain barrier, which protects the vulnerable envi-
ronment of the brain from invasion of chemicals from the
periphery. which can influence neuronal firing. The last c:lass
of mac:roglia, ependymal cells. line fluid-filled cavities in the
central nervous system (see below). They play an important
role in regulating the flow of chemicals from these cavities into
the brain. Macroglia figure importantly in disease and nervous
system trauma. For example, in multiple sc:lerosis, damage to
oligodendrocytes results in the loss of the myelin sheath of
axons in particular brain regions. This, in turn, leads to impair-
ments in neural connections and functions. Astrocytes react
to inflammation after injury and neurodegenerative processes.
Microglla subserve a phagocytic or scavenger role, respond-
ing to nervous system infection or damage. They are rapidly
mobilized-they become activated (Figure 1-30)-in response
to different pathophysiologic:al conditions and trauma.
Activated microglia can destroy invading mic:roorganisms,
remove debris, and promote tissue repair. Interestingly, they
also mediate changes in neuronal properties after nervous sys-
tem damage; sometimes maladaptive changes, so they may also
hinder recovery after injury. For example, neurons often become
hyperexcitable after nervous system damage, and microglia can
be involved in this process. Microglia also play a key role in the
modification of connections between neurons, especially in the
elimination of unnecessary presynaptic connections during
development
The Nervous System Consists of Separate
Peripheral and Central Components
Neurons and glial cells of the nervous system are organized
into two anatomically separate but functionally interdepen-
dent parts: the peripheral and the central nervous systems
(Figure l-4A). The peripheral nervous system is subdivided
into somatic. autonomic. and enterlc divisions. The somatic

FIGURE 1-4. A. location of 1he central and peripheral nervous system in the body. Major peripheral nerves are shown in yellow. 8. The brain and
c.
spinal cord, viewed laterally. There are seven majordMslons cf1he central nervous system: {1) cerebral hemispheres, (2) dlencephalon. (3) mldbraln,
(4) pons, (S) cerebellum, (6) medulla, and (7) splnal cord. The mldbraln, pons, and medulla c:cmprtse the brain stem.
10 Section I • The Central Nervous System
division contains the sensory neurons that innervate the skin,
muscles, and joints. These neurons detect and, in turn, inform
the central nervous system of stimuli. This division also con-
tains the axons of motor neurons that innervate skeletal muscle,
although the cell bodies of motor neurons lie within the cen-
tral nervous system. These axons transmit signals to muscle to
regulate the force of muscle contraction. The autonomic divi-
sion contains the neurons that innervate glands and the smooth
muscle of the viscera and blood vessels (see Chapter 15). This
division, with its separate sympathetic and parasympathetic
subdivisions, regulates body functions based, in part, on infor-
mation about the body's internal state. The enteric nervous
system contains neurons that innervate the gastrointestinal sys-
tem. It functions independent of, as well as in concert with, the
autonomic nervous system.
The central nervous system consists of the spinal cord and
brain (Figure 1-4B), and the brain is further subdivided into
the medulla, pons, cerebellum, midbrain, diencephalon, and
The central nervous system develops from a specialized por-
tion of the embryonic ectoderm, the neural plate. Originally
a flattened sheet of cells, the neural plate forms a tube-like
structure-termed the neural tube-as the neurons and glial
cells proliferate. The walls of the neural tube form the neu-
ronal structure of the central nervous system. The cavity in the
neural tube forms the ventricular system.
Very early in development the rostral portion of the neural
tube forms the three hollow swellings, or vesicles, correspond-
ing to where there is an enormous proliferation of developing
neurons (Figure 1-5): (1) the prosencephalon or forebrain,
(2) the mesencephalon or midbrain, and (3) the rhomben-
cephalon or hlndbraln. The caudal portion of the neural tube
remains relatively undifferentiated and forms the spinal cord.
Two secondary vesicles emerge from the prosencephalon later
in development: the telencephalon (or cerebral hemisphere)
and the diencephalon (or thalamus and hypothalamus).
Whereas the mesencephalon remains undivided through-
out further brain development, the rhombencephalon gives
rise to the metencephalon (or pons and cerebellum) and the
myelencephalon (or medulla). The five brain vesicles and
primitive spinal cord, already identifiable by the fifth week of
fetal development give rise to the seven major divisions of
the central nervous system (see Figure 1-4).
The complex configuration of the mature brain is deter-
mined in part by how the developing brain bends, or flexes.
Flexures occur because proliferation of cells in the brain stem
and cerebral hemispheres is enormous, and the space that
the developing brain occupies in the cranium is constrained.
At the three-vesicle stage, there are two prominent flexures:
the cervical flexure, at the junction of the spinal cord and
the caudal hindbrain (or future medulla), and the cephalic
flexure, at the level of the midbrain (Figure 1-5, bottom). At
the five-vesicle stage, a third flexure becomes prominent the
pontine flexure. By birth the cervical and pontine flexures
have straightened out. The cephalic flexure, however, remains
prominent and causes the longitudinal axis oftheforebrain to
cerebral hemispheres (Figure 1-4C). Within each of the seven
central nervous system divisions resides a component ofthe ven-
tricular system, a labyrinth of fluid-filled cavities that serve var-
ious supportive functions (see Figure 1-13). Box 1-1 shows how
all of the divisions of the central nervous system and the com-
ponents of the ventricular system are present from very early in
development, from about the first month after conception.
Neuronal cell bodies and axons are not distributed uniformly
within the nervous system. In the peripheral nervous system, cell
bodies collect in peripheral ganglia and axons are contained in
peripheral nerves. In the central nervous system, neuronal cell
bodies and dendrites are located in cortical areas, which are flat-
tened sheets of cells (or laminae) located primarily on the surface
of the cerebral hemispheres, and in nuclei, which are clusters
of neurons located beneath the surface of all of the central ner-
vous system divisions. Nuclei come in various sizes and shapes;
they are commonly oval and columnar but sometimes occur in
complex three-dimensional configurations (see Figure 1-10).
deviate from that of the mid brain, hindbrain, and spinal cord
(see Figure 1-16B).
The large cavities within the cerebral vesicles develop
into the ventricular system of the brain, and the caudal cav-
ity becomes the central canal of the spinal cord (Figure 1-5).
The ventricular system contains cerebrospinal fluid, which
is produced mainly by the choroid plexus (see Chapter 3).
As the brain vesicles develop, the cavity within the cerebral
hemispheres divides into the two lateral ventricles (formerly
termed the first and second ventricles) and the third ven-
tricle (Figure 1-SB). The lateral ventricles, which develop as
outpouchings from the rostral portion of the third ventricle,
are each interconnected with the third ventricle by an inter-
ventricular fora men (of Monro) (Figure 1-5, inset). The fourth
ventricle, the most caudal ventricle, develops from the cavity
within the hindbrain. It is connected to the third ventricle by
the cerebral aqueduct {of Sylvius) and merges caudally with
the central canal (of the caudal medulla and spinal cord).
Cerebrospinal fluid normally exits from the ventricular
system into the space overlying the central nervous system's
surface through foramina in the fourth ventricle (discussed in
Chapter 3). {The central canal does not have such an aperture
for the outflow of cerebrospinal fluid.) Pathological processes
can prevent flow of cerebrospinal fluid from the ventricular
system. For example, later in development the cerebral aque-
duct becomes narrowed because of cell proliferation in the
midbrain. Its narrow diameter makes it vulnerable to the con-
stricting effects of congenital abnormalities, tumors, or swell-
ing from trauma. Occlusion can occur; however, cerebrospinal
fluid continues to be produced despite occlusion. If occlusion
occurs before the bones of the skull are fused (ie, in embryonic
development or in infancy), ventricular volume will increase,
the brain will enlarge rostral to the occlusion, and head size
will increase. This condition is called hydrocephalus. If occlu-
sion occurs after the bones of the skull are fused, ventricular
size cannot increase without increasing intracranial pressure.
This is a life-threatening condition.
 Chapter 1 • Organization of the Central Nervous System 11

A B
Three-vesicle stage Five-vesicle stage

r,yy Lo-1
u1 : ~ I ·~-~-~
~~\,__../~
Telencephalon' ( (I'
l ~~=:!~alon h~~phere)
( bral

2~
() (thalamus and [)r~
Diencephalon lb lnterventricular

~Rhombencephaloni
hypothalamus) ~\ I
foramen
Mesencephalon \ . Retina
(mid.brain) [ Mesencephalon 2 ~J Thir<i:
(midbrain) ~ventricle
Metencephal.on Cerebral
(hindbrain) (pons) 3a aqueduct
Myelencephalon
(medulla)

Spinal cord--
--Spinal cord
Central canal-.

Cephalic Cervical Cervical

flexure flexure flexure

FIGURE 1-5. Schemltlc lllustratlon of the three- and flw-veslcle stages of the neural tube during arty central nerwus system development.The top
portion of the figure shows dorsal vlews of the neural tube drawn without flexures. The bottom portion of the figure presents lateral views. A. Three-veslcle
stage. B. ~Ide stage. Note that the llneage of each veslcle at the five-vesicle stage Is Indicated bythe shading. The two secondary vesicles from the
fore.brain have different green shades, and the two vesicles that dertwd from the hlndbraln have different blue shades. The Inset shows the locatlon of the
lnterventrlcular foramen on one side In the flve-veslde stage. (Adapted from Kandel ER, Schwartz JH, Jessell TM, eds. Prlndples ofNeural Science. 3rd ed.
New York, NY: Mt'Graw+IHI; 1991.}

Regions of the central nervous system that contain axons have
an unwieldy number of names, the most common of which is
tract. In fresh tissue, nuclei and cortical areas appear grayish
and tracts appear whitish, hence the familiar terms gray matter
and white matter. The whitish appearance of tracts is caused by
the presence of the myelin sheath surrounding the axons (see
Figure 1-3). The gray and white matter can be distinguished in
fixed tissue using anatomical methods and in the living brain
using radiological methods (see Chapter 2, Boxes 2-1 and 2-2).
The Spinal Cord Displays the Simplest Organization
of All Seven Major Divisions
The spinal cord participates in processing sensory information
from the limbs, trunk. and many internal organs; in control-
ling body movements directly; and in regulating many visceral
functions (Figure 1-6). It also provides a conduit for the trans-
mission of both sensory information in the white matter axon
tracts that ascend to the brain and motor information in the
descending tracts.
The spinal cord consists of modules, termed seg-
ments, in which every segment has a similar basic structure
(Figure 1-6C). Each spinal cord segment contains a pair of
nerve roots (and associated rootlets) called the dorsal and
ventral roots. (The terms dorsal and ventral describe the spa-
tial relations of structures; these and other anatomical terms
are explained later in this chapter.) Dorsal roots contain only
sensory axons, which transmit sensory information into the
spinal cord. By contrast, ventral roots contain motor axons,
which transmit motor commands to muscle and other body
organs. Dorsal and ventral roots exemplify the separation of
function in the nervous system, a principle that is examined
further in subsequent chapters. These sensory and motor
axons, which are part of the peripheral nervous system,
become intermingled in the 1pinal nerves enroute to their
peripheral targets (Figure 1-6C).
12 Section I • The Central Nervous System

Ventral
surface

FIGURE 1""'- Spinal cord organization. A. Adorsal view of the central nervous system.The
horlzontal llnes over the splnal cord mark the loca'dons of the dllfefent splnal cord dMslons.
These wlll be considered In more detall In later chapters. B. A latefal vfew of the splnal cord
and the vertebral column. The dark llnes delineate the spinal cord segments. C. Surface
topography and Internal structure of a slngle spinal cord segment.

The Brain Stem and Cerebellum Regulate Body
Functions and Movements
The next three divisions-medulla. pons. and midbrain-
comprise the brain stem (Figure 1-7). The brain stem hu
three general functions. First, it receives sensory information
from cranial structures and controls the muscles of the head.
These functions are similar to those of the spinal cord. Cranial
nerves, the sensory and motor nerve roots that enter and exit
the brain stem, are parts of the peripheral nervous system and
are analogous to the spinal nerves (Figure 1-7). Second, simi-
lar to the spinal cord, the brain stem is a conduit for informa-
tion flow because ascending sensory and descending motor
tracts travel through it Finally, nuclei in the brain stem inte-
grate diverse information from a variety of sources for arousal.
behavioral responses to the environment, and other higher
brain functions.
In addition to these three general functions, the various
divisions of the brain stem each subserve spec:ifi.c sensory and
motor functions. For example. portions of the medulla par-
ticipate in essential blood pressure and respiratory regulatory
mechanisms. Indeed, damage to these parts of the brain is
almost always life threatening. Parts of the pons and miclbrain
play a key role in the control ofeye movement.
The principal functions of the cerebellum are to regulate
eye and limb movements and to maintain posture and balance
(Figure 1-8). Limb movements become poorly coordinated
when the cerebellum is damaged. In addition, parts of the cer-
ebellum play a role in higher brain functions, including lan-
guage, cognition, and emotion (Chapter 13).
Another random document with
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luminosity of radio-active minerals. A shining world, a world glowing
eternally with cold white light, a luminous, titanic sphere that rushed
through the darkness of infinite space like some pale, gigantic moon.
And upon the surface of the glowing plains beneath us rose dense
and twisted masses of dark, leafless vegetation, distorted tree-
growths and tangles of low shrubs that were all of deepest black in
color, springing out of that glowing soil and twisting blackly and
grotesquely above its feeble light, stretching away over plain and hill
and valley like the monstrous landscape of some undreamed-of hell!
And now, as our ship slanted down across the surface of the glowing
sphere, there gleamed ahead a deepening of that glow, a
concentration of that feeble light which grew stronger as we raced on
toward it. And it was a city! A city whose mighty buildings were each
a truncated pyramid in shape, towering into the air for thousands
upon thousands of feet, a city whose every building and street and
square glowed with the same faint white light as the ground upon
which they stood, a metropolis out of nightmare, the darkness of
which was dispelled only by the light of its own great glowing
structures and streets. Far away stretched the mass of those
structures, a luminous mass which covered square mile upon square
mile of the surface of this glowing world, and far beyond them there
lifted into the dusky air the shining towers and pyramids of still other
cities.
We straightened, trembling, turning toward each other with white
faces. And then, before any could speak, Dal Nara had whirled to the
window and uttered a hoarse shout. "Look!" she cried, and pointed
down and outward toward the titanic, glowing buildings of the city
ahead; for from their truncated summits were rising suddenly a
swarm of long black shapes, a horde of long black cones which were
racing straight up toward us.
I shouted an order to the pilot, and instantly our ship was turning and
slanting sharply upward, while around us our cruisers sped up with
us. Then, from beneath, there sped up toward us a shining little
cylinder of metal which struck a cruiser racing beside our own. It
exploded instantly into a great flare of blinding light, enveloping the
cruiser it had struck, and then the light had vanished, while with it
had vanished the ship it had enveloped. And from the cones beneath
and beyond there leapt toward us other of the metal cylinders,
striking our ships now by the dozens, flaring and vanishing with them
in great, silent explosions of light.
"Etheric bombs!" I cried. "And our ship is the only battle-cruiser—the
rest have no weapons!"
I turned, cried another order, and in obedience to it our own cruiser
halted suddenly and then dipped downward, racing straight into the
ascending swarm of attacking cones. Down we flashed, down, down,
and toward us sprang a score of the metal cylinders, grazing along
our sides. And then, from the sides of our own downward-swooping
ship there sprang out brilliant shafts of green light, the deadly de-
cohesion ray of the ships of the Federation Fleet. It struck a score of
the cones beneath and they flamed with green light for an instant
and then flew into pieces, spilling downward in a great shower of tiny
fragments as the cohesion of their particles was destroyed by the
deadly ray. And now our cruiser had crashed down through the
swarm of them and was driving down toward the luminous plain
below, then turning and racing sharply upward again while from all
the air around us the black cones swarmed to the attack.
Up, up, we sped, and now I saw that our blow had been struck in
vain, for the last of our ships above were vanishing beneath the
flares of the etheric bombs. One only of our cruisers remained,
racing up toward the zenith in headlong flight with a score of the
great cones in hot pursuit. A moment only I glimpsed this, and then
we had turned once more and were again diving down upon the
attacking cones, while all around us the etheric bombs filled the air
with the silent, exploding flares. Again as we swooped downward our
green rays cut paths of annihilation across the swarming cones
beneath; and then I heard a cry from Hurus Hol, whirled to the
window and glimpsed above us a single great cone that was diving
headlong down toward us in a resistless, ramming swoop. I shouted
to the pilot, sprang to the controls, but was too late to ward off that
deadly blow. There was a great crash at the rear of our cruiser; it
spun dizzily for a moment in midair, and then was tumbling crazily
downward like a falling stone toward the glowing plain a score of
miles below.

3
I think now that our cruiser's mad downward plunge must have
lasted for minutes, at least, yet at the time it seemed over in a single
instant. I have a confused memory of the bridgeroom spinning about
us as we whirled down, of myself throwing back the controls with a
last, instinctive action, and then there came a ripping, rending crash,
a violent shock, and I was flung into a corner of the room with terrific
force.
Dazed by the swift action of the last few minutes I lay there
motionless for a space of seconds, then scrambled to my feet. Hurus
Hol and Dal Nara were staggering up likewise, the latter hastening at
once down into the cruiser's hull, but Nal Jak, the wheelman, lay
motionless against the wall, stunned by the shock. Our first act was
to bring him back to consciousness by a few rough first-aid
measures, and then we straightened and gazed about us.
Apparently our cruiser's keel was resting upon the ground, but was
tilted over at a sharp angle, as the slant of the room's floor attested.
Through the broad windows we could see that around our prostrate
ship lay a thick, screening grove of black tree-growths which we had
glimpsed from above, and into which we had crashed in our mad
plunge downward. As I was later to learn, it was only the shock-
absorbing qualities of the vegetation into which we had fallen, and
my own last-minute rush to the controls, which had slowed our fall
enough to save us from annihilation.
There was a buzz of excited voices from the crew in the hull beneath
us, and then I turned at a sudden exclamation from Hurus Hol, to
find him pointing up through the observation-windows in the
bridgeroom's ceiling. I glanced up, then shrank back. For high above
were circling a score or more of the long black cones which had
attacked us, and which were apparently surveying the landscape for
some clue to our fate. I gave a sharp catch of indrawn breath as they
dropped lower toward us, and we crouched with pounding hearts
while they dropped nearer. Then we uttered simultaneous sighs of
relief as the long shapes above suddenly drove back up toward the
zenith, apparently certain of our annihilation, massing and wheeling
and then speeding back toward the glowing city from which they had
risen to attack us.
We rose to our feet again, and as we did so the door clicked open to
admit Dal Nara. She was a bruised, disheveled figure, like the rest of
us, but there was something like a grin on her face.
"That cone that rammed us shattered two of our rear vibration-
projectors," she announced, "but that was all the damage. And
outside of one man with a broken shoulder the crew is all right."
"Good!" I exclaimed. "It won't take long to replace the broken
projectors."
She nodded. "I ordered them to put in two of the spares," she
explained. "But what then?"
I considered for a moment. "None of our other cruisers escaped, did
they?" I asked.
Dal Nara slowly shook her head. "I don't think so," she said. "Nearly
all of them were destroyed in the first few minutes. I saw Ship 16
racing up in an effort to escape, heading back toward the Galaxy, but
there were cones hot after it and it couldn't have got away."
The quiet voice of Hurus Hol broke in upon us. "Then we alone can
take back word to the Federation of what is happening here," he
said. His eyes suddenly flamed. "Two things we know," he
exclaimed. "We know that this dark star's curving path through
space, which will bring it so fatally near to our own sun in passing, is
a path contrary to all the laws of astronomical science. And we know
now, too, that upon this dark-star world, in those glowing cities
yonder, live beings of some sort who possess, apparently, immense
intelligence and power."
My eyes met his. "You mean——" I began, but he interrupted swiftly.
"I mean that in my belief the answer to this riddle lies in that glowing
city yonder, and that it is there we must go to find that answer."
"But how?" I asked. "If we take the cruiser near it they'll sight us and
annihilate us."
"There is another way," said Hurus Hol. "We can leave the cruiser
and its crew hidden here, and approach the city on foot—get as near
to it as possible—learn what we can about it."
I think that we all gasped at that suggestion, but as I quickly revolved
it in my mind I saw that it was, in reality, our only chance to secure
any information of value to take back to the Federation. So we
adopted the idea without further discussion and swiftly laid our plans
for the venture. At first it was our plan for only us three to go, but at
Dal Nara's insistence we included the pilot in our party, the more
quickly because I knew her to be resourceful and quick-witted.

Two hours we spent in sleep, at the suggestion of Hurus Hol, then

ate a hasty meal and looked to our weapons, small projectors of the
de-cohesion ray similar to the great ray-tubes of the cruiser. Already
the ship's two shattered vibration-projectors had been replaced by
spares, and our last order was for the crew and under-officers to
await our return without moving beyond the ship in any event. Then
the cruiser's hull-door snapped open and we four stepped outside,
ready for our venture.
The sandy ground upon which we stood glowed with the feeble white
light which seemed to emanate from all rock and soil on this strange
world, a weird light which beat upward upon us instead of down. And
in this light the twisted, alien forms of the leafless trees around us
writhed upward into the dusky air, their smooth black branches
tangling and intertwining far above our heads. As we paused there
Hurus Hol reached down for a glowing pebble, which he examined
intently for a moment.
"Radio-active," he commented. "All this glowing rock and soil." Then
he straightened, glanced around, and led the way unhesitatingly
through the thicket of black forest into which our ship had fallen.
Silently we followed him, in single file, across the shining soil and
beneath the distorted arches of the twisted trees, until at last we
emerged from the thicket and found ourselves upon the open
expanse of the glowing plain. It was a weird landscape which met
our eyes, a landscape of glowing plains and shallow valleys patched
here and there with the sprawling thickets of black forest, a pale,
luminous world whose faint light beat feebly upward into the dusky,
twilight skies above. In the distance, perhaps two miles ahead, a
glow of deeper light flung up against the hovering dusk from the
massed buildings of the luminous city, and toward this we tramped
steadily onward, over the shining plains and gullies and once over a
swift little brook whose waters glowed as they raced like torrents of
rushing light. Within an hour we had drawn to within a distance of
five hundred feet from the outermost of the city's pyramidal buildings,
and crouched in a little clump of dark tree-growths, gazing
fascinatedly toward it.
 "It was a weird landscape that met their eyes."

The scene before us was one of unequaled interest and activity.

Over the masses of huge, shining buildings were flitting great
swarms of the long black cones, moving from roof to roof, while in
the shining streets below them moved other hordes of active figures,
the people of the city. And as our eyes took in these latter I think that
we all felt something of horror, in spite of all the alien forms which we
were familiar with in the thronging worlds of the Galaxy.
For in these creatures was no single point of resemblance to
anything human, nothing which the appalled intelligence could seize
upon as familiar. Imagine an upright cone of black flesh, several feet
in diameter and three or more in height, supported by a dozen or
more smooth long tentacles which branched from its lower end—
supple, boneless octopus-arms which held the cone-body upright
and which served both as arms and legs. And near the top of that
cone trunk were the only features, the twin tiny orifices which were
the ears and a single round and red-rimmed white eye, set between
them. Thus were these beings in appearance, black tentacle-
creatures, moving in unending swirling throngs through streets and
squares and buildings of their glowing city.
Helplessly we stared upon them, from our place of concealment. To
venture into sight, I knew, would be to court swift death. I turned to
Hurus Hol, then started as there came from the city ahead a low,
waxing sound-note, a deep, powerful tone of immense volume which
sounded out over the city like the blast of a deep-pitched horn.
Another note joined it, and another, until it seemed that a score of
mighty horns were calling across the city, and then they died away.
But as we looked now we saw that the shining streets were
emptying, suddenly, that the moving swarms of black tentacle-
creatures were passing into the pyramidal buildings, that the cones
above were slanting down toward the roofs and coming to rest.
Within a space of minutes the streets seemed entirely empty and
deserted, and the only sign of activity over all the city was the
hovering of a few cones that still moved restlessly above it.
Astounded, we watched, and then the explanation came suddenly to
me.
"It's their sleep-period!" I cried. "Their night! These things must rest,
must sleep, like any living thing, and as there's no night on this
glowing world those horn-notes must signal the beginning of their
sleep-period."
Hurus Hol was on his feet, his eyes suddenly kindling. "It's a chance
in a thousand to get inside the city!" he exclaimed.
The next moment we were out of the shelter of our concealing trees
and were racing across the stretch of ground which separated us
from the city. And five minutes later we were standing in the empty,
glowing streets, hugging closely the mighty sloping walls of the huge
buildings along it.
At once Hurus Hol led the way directly down the street toward the
heart of the city, and as we hastened on beside him he answered to
my question, "We must get to the city's center. There's something
there which I glimpsed from our ship, and if it's what I think——"
He had broken into a run, now, and as we raced together down the
bare length of the great, shining avenue, I, for one, had an
unreassuring presentiment of what would happen should the huge
buildings around us disgorge their occupants before we could get out
of the city. Then Hurus Hol had suddenly stopped short, and at a
motion from him we shrank swiftly behind the corner of a pyramid's
slanting walls. Across the street ahead of us were passing a half-
dozen of the tentacle-creatures, gliding smoothly toward the open
door of one of the great pyramids. A moment we crouched, holding
our breath, and then the things had passed inside the building and
the door had slid shut behind them. At once we leapt out and
hastened on.
We were approaching the heart of the city, I judged, and ahead the
broad, shining street we followed seemed to end in a great open
space of some sort. As we sped toward it, between the towering
luminous lines of buildings, a faint droning sound came to our ears
from ahead, waxing louder as we hastened on. The clear space
ahead was looming larger, nearer, now, and then as we raced past
the last great building on the street's length we burst suddenly into
view of the opening ahead and stepped, staring dumfoundedly
toward it.
It was no open plaza or square, but a pit—a shallow, circular pit not
more than a hundred feet in depth but all of a mile in diameter, and
we stood at the rim or edge of it. The floor was smooth and flat, and
upon that floor there lay a grouped mass of hundreds of half-globes
or hemispheres, each fifty feet in diameter, which were resting upon
their flat bases with their curving sides uppermost. Each of these
hemispheres was shining with light, but it was very different light
from the feeble glow of the buildings and streets around us, an
intensely brilliant blue radiance which was all but blinding to our
eyes. From these massed, radiant hemispheres came the loud
droning we had heard, and now we saw, at the pit's farther edge, a
cylindrical little room or structure of metal which was supported
several hundred feet above the pit's floor by a single slender shaft of
smooth round metal, like a great bird-cage. And toward this cage-
structure Hurus Hol was pointing now, his eyes flashing.
"It's the switch-board of the thing!" he cried. "And these brilliant
hemispheres—the unheard-of space-path of this dark star—it's all
clear now! All——"
He broke off, suddenly, as Nal Jak sprang back, uttering a cry and
pointing upward. For the moment we had forgotten the hovering
cones above the city, and now one of them was slanting swiftly
downward, straight toward us.
We turned, ran back, and the next moment an etheric bomb crashed
down upon the spot where we had stood, exploding silently in a
great flare of light. Another bomb fell and flared, nearer, and then I
turned with sudden fierce anger and aimed the little ray-projector in
my hand at the hovering cone above. The brilliant little beam cut
across the dark shape; the black cone hovered still for a moment,
then crashed down into the street to destruction. But now, from
above and beyond, other cones were slanting swiftly down toward
us, while from the pyramidal buildings beside us hordes of the black
tentacle-creatures were pouring out in answer to the alarm.
In a solid, resistless swarm they rushed upon us. I heard a yell of
defiance from Dal Nara, beside me, the hiss of our rays as they
clove through the black masses in terrible destruction, and then they
were upon us. A single moment we whirled about in a wild mêlée of
men and cone-creatures, of striking human arms and coiling
tentacles; then there was a shout of warning from one of my friends,
something hard descended upon my head with crushing force, and
all went black before me.
 4

Faint light was filtering through my eyelids when I came back to

consciousness. As I opened them I sat weakly up, then fell back.
Dazedly I gazed about me. I was lying in a small, square room lit
only by its own glowing walls and floor and ceiling, a room whose
one side slanted steeply upward and inward, pierced by a small
barred window that was the only opening. Opposite me I discerned a
low door of metal bars, or grating, beyond which lay a long, glowing-
walled corridor. Then all these things were suddenly blotted out by
the anxious face of Hurus Hol, bending down toward me.
"You're awake!" he exclaimed, his face alight. "You know me, Ran
Rarak?"
For answer I struggled again to a sitting position, aided by the arm of
Dal Nara, who had appeared beside me. I felt strangely weak,
exhausted, my head throbbing with racing fires.
"Where are we?" I asked, at last. "The fight in the city—I remember
that—but where are we now? And where's Nal Jak?"
The eyes of my two friends met and glanced away, while I looked
anxiously toward them. Then Hurus Hol spoke slowly.
"We are imprisoned in this little room in one of the great pyramids of
the glowing city," he said. "And in this room you have lain for weeks,
Ran Rarak."
"Weeks?" I gasped, and he nodded. "It's been almost ten weeks
since we were captured there in the city outside," he said, "and for all
that time you've lain here out of your head from that blow you
received, sometimes delirious and raving, sometimes completely
unconscious. And in all that time this dark star, this world, has been
plunging on through space toward our Galaxy, and our sun, and the
theft and doom of that sun. Ten more days and it passes our sun,
stealing it from the Galaxy. And I, who have learned at last what
forces are behind it all, lie prisoned here.
"It was after we four were brought to this cell, after our capture, that I
was summoned before our captors, before a council of those strange
tentacle-creatures which was made up, I think, of their own
scientists. They examined me, my clothing, all about me, then
sought to communicate with me. They do not speak—
communicating with each other by telepathy—but they strove to
enter into communication with me by a projection of pictures on a
smooth wall, pictures of their dark star world, pictures of our own
Galaxy, our own sun—picture after picture, until at last I began to
understand the drift of them, the history and the purpose of these
strange beings and their stranger world.
"For ages, I learned, for countless eons, their mighty sun had flashed
through the infinities of space, alone except for its numerous planets
upon which had risen these races of tentacle-creatures. Their sun
was flaming with life, then, and on their circling planets they had
attained to immense science, immense power, as their system rolled
on, a single wandering star, through the depths of uncharted space.
But as the slow eons passed, the mighty sun began to cool, and their
planets to grow colder and colder. At last it had cooled so far that to
revive its dying fires they dislodged one of their own planets from its
orbit and sent it crashing into their sun, feeding its waning flames.
And when more centuries had passed and it was again cooling they
followed the same course, sending another planet into it, and so on
through the ages, staving off the death of their sun by sacrificing
their worlds, until at last but one planet was left to them. And still
their sun was cooling, darkening, dying.
"For further ages, though, they managed to preserve a precarious
existence on their single planet by means of artificial heat-
production, until at last their great sun had cooled and solidified to
such a point that life was possible upon its dark, dead surface. That
surface, because of the solidified radio-active elements in it, shone
always with pale light, and to it the races of the tentacle-creatures
now moved. By means of great air-current projectors they
transferred the atmosphere of their planet to the dark star itself and
then cast loose their planet to wander off into space by itself, for its
orbit had become erratic and they feared that it would crash into their
own great dark star world, about which it had revolved. But on the
warm, shining surface of the great dark star they now spread out and
multiplied, raising their cities from its glowing rock and clinging to its
surface as it hurtled on and on and on through the dark infinities of
trackless space.
"But at last, after further ages of such existence, the tentacle-races
saw that again they were menaced with extinction, since in
obedience to the inexorable laws of nature their dark star was
cooling still further, the molten fires at its center which warmed its
surface gradually dying down, while that surface became colder and
colder. In a little while, they knew, the fires at its center would be
completely dead, and their great world would be a bitter, frozen
waste, unless they devised some plan by which to keep warm its
surface.
"At this moment their astronomers came forward with the
announcement that their dark-star world, plunging on through empty
space, would soon pass a great star-cluster or Galaxy of suns at a
distance of some fifteen billion miles. They could not invade the
worlds of this Galaxy, they knew, for they had discovered that upon
those worlds lived countless trillions of intelligent inhabitants who
would be able to repel their own invasion, if they attempted it. There
was but one expedient left, therefore, and that was to attempt to jerk
a sun out of this Galaxy as they passed by it, to steal a star from it to
take out with them into space, which would revolve around their own
mighty dark world and supply it with the heat they needed.
"The sun which they fixed on to steal was one at the Galaxy's very
edge, our own sun. If they passed this at fifteen billion miles, as their
course then would cause them to do, they could do nothing. But if
they could change their dark star's course, could curve inward to
pass this sun at some three billion miles instead of fifteen, then the
powerful gravitational grip of their own gigantic world would grasp
this sun and carry it out with it into space. The sun's planets, too,
would be carried out, but these they planned to crash into the fires of
the sun itself, to increase its size and splendor. All that was needed,
therefore, was some method of curving their world's course inward,
and for this they had recourse to the great gravity-condensers which
they had already used to shift their own planets.
"You know that it is gravitational force alone which keeps the suns
and planets to their courses, and you know that the gravitational
force of any body, sun or planet, is radiated out from it in all
directions, tending to pull all things toward that body. In the same
way there is radiated outward perpetually from the Galaxy the
combined attractive gravitational force of all its swarming suns, and a
tiny fraction of this outward-radiating force, of course, struck the dark
star, pulling it weakly toward the Galaxy. If more of that outward-
radiating force could strike the dark star, it would be pulled toward
the Galaxy with more power, would be pulled nearer toward the
Galaxy's edge, as it passed.
"It was just that which their gravity-condenser accomplished. In a low
pit at the heart of one of their cities—this city, in fact—they placed
the condenser, a mass of brilliant hemispherical ray-attracters which
caused more of the Galaxy's outward-shooting attractive force to fall
upon the dark star, which condensed and concentrated that radiating
force upon the dark star, thereby pulling the dark star inward toward
the Galaxy's edge in a great curve. When they reached a distance of
three billion miles from the Galaxy's edge they planned to turn off the
great condenser, and their dark star would then shoot past the
Galaxy's edge, jerking out our sun with it, from that edge, by its own
terrific gravitational grip. If the condenser were turned off before they
came that close, however, they would pass the sun at a distance too
far to pull it out with them, and would then speed on out into space
alone, toward the freezing of their world and their own extinction. For
that reason the condenser, and the great cage-switch of the
condenser, were guarded always by hovering cones, to prevent its
being turned off before the right moment.
"Since then they have kept the great gravity-condenser in unceasing
operation, and their dark star has swept in toward the Galaxy's edge
in a great curve. Back in our own solar system I saw and understood
what would be the result of that inward curve, and so we came here
—and were captured. And in those weeks since we were captured,
while you have lain here unconscious and raving, this dark star has
been plunging nearer and nearer toward our Galaxy and toward our
sun. Ten more days and it passes that sun, carrying it out with it into
the darkness of boundless space, unless the great condenser is
turned off before then. Ten more days, and we lie here, powerless to
warn any of what forces work toward the doom of our sun!"

There was a long silence when Hurus Hol's voice had ceased—a
whispering, brain-crushing silence which I broke at last with a single
question.
"But Nal Jak——?" I asked, and the faces of my two companions
became suddenly strange, while Dal Nara turned away. At last Hurus
Hol spoke.
"It was after the tentacle-scientists had examined me," he said
gently, "that they brought Nal Jak down to examine. I think that they
spared me for the time being because of my apparently greater
knowledge, but Nal Jak they—vivisected."
There was a longer hush than before, one in which the brave, quiet
figure of the wheelman, a companion in all my service with the fleet,
seemed to rise before my suddenly blurring eyes. Then abruptly I
swung down from the narrow bunk on which I lay, clutched dizzily at
my companions for support, and walked unsteadily to the square,
barred little window. Outside and beneath me lay the city of the dark-
star people, a mighty mass of pyramidal, glowing buildings, streets
thronged with their dark, gliding figures, above them the swarms of
the racing cones. From our little window the glowing wall of the great
pyramid which held us slanted steeply down for fully five hundred
feet, and upward above us for twice that distance. And as I raised
my eyes upward I saw, clear and bright above, a great, far-flung field
of stars—the stars of our own Galaxy toward which this world was
plunging. And burning out clearest among these the star that was
nearest of all, the shining yellow star that was our own sun.
I think now that it was the sight of that yellow star, largening steadily
as our dark star swept on toward it, which filled us with such utter
despair in the hours, the days, that followed. Out beyond the city our
cruiser lay hidden in the black forest, we knew, and could we escape
we might yet carry word back to the Federation of what was at hand,
but escape was impossible. And so, through the long days, days
measurable only by our own time-dials, we waxed deeper into an
apathy of dull despair.
Rapidly my strength came back to me, though the strange food
supplied us once a day by our captors was almost uneatable. But as
the days fled by, my spirits sank lower and lower, and less and less
we spoke to each other as the doom of our sun approached, the only
change in any thing around us being the moment each twenty-four
hours when the signal-horns called across the city, summoning the
hordes in its streets to their four-hour sleep-period. At last, though,
we woke suddenly to realization of the fact that nine days had
passed since my awakening, and that upon the next day the dark
star would be plunging past the burning yellow star above us and
jerking it into its grip. Then, at last, all our apathy dropped from us,
and we raged against the walls of our cells with insensate fury. And
then, with startling abruptness, came the means of our deliverance.

For hours there had been a busy clanging of tools and machines
somewhere in the great building above us, and numbers of the
tentacle-creatures had been passing our barred door carrying tools
and instruments toward some work being carried out overhead. We
had come to pay but little attention to them, in time, but as one
passed there came a sudden rattle and clang from outside, and
turning to the door we saw that one of the passing creatures had
dropped a thick coil of slender metal chain upon the floor and had
passed on without noticing his loss.
In an instant we were at the door and reaching through its bars
toward the coil, but though we each strained our arms in turn toward
it the thing lay a few tantalizing inches beyond our grasp. A moment
we surveyed it, baffled, fearing the return at any moment of the
creature who had dropped it, and then Dal Nara, with a sudden
inspiration, lay flat upon the floor, thrusting her leg out through the
grating. In a moment she had caught the coil with her foot, and in
another moment we had it inside, examining it.
We found that though it was as slender as my smallest finger the
chain was of incredible strength, and when we roughly estimated the
extent of its thick-coiled length we discovered that it would be more
than long enough to reach from our window to the street below. At
once, therefore, we secreted the thing in a corner of the room and
impatiently awaited the sleep-period, when we could work without
fear of interruption.
At last, after what seemed measureless hours of waiting, the great
horns blared forth across the city outside, and swiftly its streets
emptied, the sounds in our building quieting until all was silence,
except for the humming of a few watchful cones above the great
condenser, and the deep droning of the condenser itself in the
distance. At once we set to work at the bars of our window.
Frantically we chipped at the rock at the base of one of the metal
bars, using the few odd bits of metal at our command, but at the end
of two hours had done no more than scratch away a bare inch of the
glowing stone. Another hour and we had laid bare from the rock the
lower end of the bar, but now we knew that within minutes the sleep-
period of the city outside would be ending, and into its streets would
be swarming its gliding throngs, making impossible all attempts at
escape. Furiously we worked, dripping now with sweat, until at last
when our time-dials showed that less than half an hour remained to
us I gave over the chipping at the rock and wrapped our chain firmly
around the lower end of the bar we had loosened. Then stepping
back into the cell and bracing ourselves against the wall below the
window, we pulled backward with all our strength.
A tense moment we strained thus, the thick bar holding fast, and
then abruptly it gave and fell from its socket in the wall to the floor,
with a loud, ringing clang. We lay in a heap on the floor, panting and
listening for any sound of alarm, then rose and swiftly fastened the
chain's end to one of the remaining bars. The chain itself we
dropped out of the window, watching it uncoil its length down the
mighty building's glowing side until its end trailed on the empty
glowing street far below. At once I motioned Hurus Hol to the
window, and in a moment he had squeezed through its bars and was
sliding slowly down the chain, hand under hand. Before he was ten
feet down Dal Nara was out and creeping downward likewise, and
then I too squeezed through the window and followed them,
downward, the three of us crawling down the chain along the huge
building's steeply sloping side like three flies.
I was ten feet down from the window, now, twenty feet, and glanced
down toward the glowing, empty street, five hundred feet below, and
seeming five thousand. Then, at a sudden sound from above me, I
looked sharply up, and as I did so the most sickening sensation of
fear I had ever experienced swept over me. For at the window we
had just left, twenty feet above me, one of the tentacle-creatures was
leaning out, brought to our cell, I doubted not, by the metal bar's
ringing fall, his white, red-rimmed eye turned full upon me.
I heard sighs of horror from my two companions beneath me, and for
a single moment we hung motionless along the chain's length,
swinging along the huge pyramid's glowing side at a height of
hundreds of feet above the shining streets below. Then the creature
raised one of its tentacles, a metal tool in its grasp, which he brought
down in a sharp blow on the chain at the window's edge. Again he
repeated the blow, and again.
He was cutting the chain!

5
For a space of seconds I hung motionless there, and then as the tool
in the grasp of the creature above came down on the chain in
another sharp blow the sound galvanized me into sudden action.
"Slide on down!" I cried. They didn't, however, but followed me up
the chain, though Dal Nara and I alone came to grips with the
horrible dead-star creature. I gripped the links with frantic hands,
pulling myself upward toward the window and the creature at the
window, twenty feet above me.
Three times the tool in his hand came down upon the chain while I
struggled up toward him, and each time I expected the strand to
sever and send us down to death, but the hard metal withstood the
blows for the moment, and before he could strike at it again I was up
to the level of the window and reaching up toward him.
As I did so, swift black tentacles thrust out and gripped Dal Nara and
me, while another of the snaky arms swept up with the tool in its
grasp for a blow on my head. Before it could fall, though, I had
reached out with my right hand, holding to the chain with my left, and
had grasped the body of the thing inside the window, pulling him
outside before he had time to resist. As I did so my own hold slipped
a little, so that we hung a few feet below the window, both clinging to
the slender chain and both striking futilely at each other, he with the
metal tool and I with my clenched fist.
A moment we hung there, swaying hundreds of feet above the
luminous stone street, and then the creature's tentacles coiled swiftly
around my neck, tightening, choking me. Hanging precariously to our
slender strand with one hand I struck out blindly with the other, but
felt consciousness leaving me as that remorseless grip tightened.
Then with a last effort I gripped the chain firmly with both hands,
doubled my feet under me, and kicked out with all my strength. The
kick caught the cone-body of my opponent squarely, tearing him
loose from his own hold on the chain, and then there was a sudden
wrench at my neck and I was free of him, while beneath Dal Nara
and I glimpsed his dark body whirling down toward the street below,
twisting and turning in its fall along the building's slanting side and
then crashing finally down upon the smooth, shining street below,
where it lay a black little huddled mass.
Hanging there I looked down, panting, and saw that Hurus Hol had
reached the chain's bottom and was standing in the empty street,
awaiting us. Glancing up I saw that the blows of the creature I had
fought had half severed one of the links above me, but there was no
time to readjust it; so with a prayer that it might hold a few moments
longer Dal Nara and I began our slipping, sliding progress
downward.
The sharp links tore our hands cruelly as we slid downward, and
once it seemed to me that the chain gave a little beneath our weight.
Apprehensively I looked upward, then down to where Hurus Hol was
waving encouragement. Down, down we slid, not daring to look
beneath again, not knowing how near we might be to the bottom.
Then there was another slight give in the chain, a sudden grating
catch, and abruptly the weakened link above snapped and we
dropped headlong downward—ten feet into the arms of Hurus Hol.
A moment we sprawled in a little heap there on the glowing street
and then staggered to our feet. "Out of the city!" cried Hurus Hol.
"We could never get to the condenser-switch on foot—but in the
cruiser there's a chance. And we have but a few minutes now before
the sleep-period ends!"
Down the broad street we ran, now, through squares and avenues of
glowing, mighty pyramids, crouching down once as the ever-
hovering cones swept by above, and then racing on. At any moment,
I knew, the great horns might blare across the city, bringing its
swarming thousands into its streets, and our only chance was to win
free of it before that happened. At last we were speeding down the
street by which we had entered the city, and before us lay that
street's end, with beyond it the vista of black forest and glowing plain
over which we had come. And now we were racing over that glowing
plain, a quarter-mile, a half, a mile....
Abruptly from far behind came the calling, crescendo notes of the
mighty horns, marking the sleep-period's end, bringing back into the
streets the city's tentacle-people. It could be but moments now, we
knew, before our escape was discovered, and as we panted on at
our highest speed we listened for the sounding of the alarm behind
us.
It came! When we had drawn to within a half-mile of the black forest
where our cruiser lay hidden, another great tumult of horn-notes
burst out over the glowing city behind, high and shrill and raging. And
glancing back we saw swarms of the black cones rising from the