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Human Neuroanatomy
Human Neuroanatomy
Second Edition

James R. Augustine

Professor Emeritus
School of Medicine
University of South Carolina
Columbia, South Carolina, USA
This edition: Copyright © 2017 John Wiley & Sons, Inc.
Published 2017 by John Wiley & Sons, Inc., Hoboken, New Jersey
Published simultaneously in Canada

First Edition: Copyright © 2008 Elsevier Inc.

Published 2008 by Academic Press, an Elsevier imprint

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Hardback ISBN: 978‐0‐4709‐6161‐2

Cover image: “Marilyn’s Brain” – MRI art by Dr. Charlotte Rae (University of Sussex). T1 weighted structural MRI images in the colors
of Warhol’s portrait of Marilyn Monroe. Figure provided by Dr. Rae.

Printed in [Printer to complete]

10 9 8 7 6 5 4 3 2 1
Contents

Prefacexiii 2.4 Fourth Week 21

About the companion websitexv 2.4.1 Formation of the Neural Tube 21
2.4.2 Rostral and Caudal Neuropores Open 22
Chapter 1 Introduction to the Nervous System 1 2.4.3 Neural Crest Cells Emerge 23
1.1 Neurons 1 2.4.4 Neural Canal – the Future Ventricular
1.1.1 Neuronal Cell Body (Soma) 2 System24
1.1.2 Axon Hillock 3 2.4.5 Neuropores Close and the Neural Tube
1.1.3 Neuronal Processes – Axons and Dendrites 3 Forms24
1.2 Classification of Neurons 4 2.4.6 Cervical Flexure Present 24
1.2.1 Neuronal Classification by Function 4 2.5 Fifth Week 24
1.2.2 Neuronal Classification by Number of Processes 4 2.5.1 Simple Tube, Complex Transformation 24
1.3 The Synapse 5 2.5.2 Five Subdivisions of the Brain Appear 24
1.3.1 Components of a Synapse 5 2.5.3 Brain Vesicles Versus Brain Regions 25
1.3.2 Neurotransmitters and Neuromodulators 5 2.6 Vulnerability of the Developing Nervous System 26
1.3.3 Neuronal Plasticity 6 2.7 Congenital Malformations of the Nervous System 27
1.3.4 The Neuropil 6 2.7.1 Spinal Dysraphism 27
1.4 Neuroglial Cells 6 2.7.2 Anencephaly 28
1.4.1 Neuroglial Cells Differ from Neurons 6 2.7.3 Microcephaly 28
1.4.2 Identification of Neuroglia 6 Further Reading 29
1.4.3 Neuroglial Function 8
1.4.4 Neuroglial Cells and Aging 9 Chapter 3 The Spinal Cord 31
1.4.5 Neuroglial Cells and Brain Tumors 9 3.1 Embryological Considerations 31
1.5 Axonal Transport 9 3.1.1 Layers of the Developing Spinal Cord 31
1.5.1 Functions of Axonal Transport 9 3.1.2 Formation of Ventral Gray Columns
1.5.2 Defective Axonal Transport 9 and Ventral Roots 32
1.6 Degeneration and Regeneration 10 3.1.3 Formation of Dorsal Gray Columns 32
1.6.1 Axon or Retrograde Reaction 10 3.1.4 Dorsal and Ventral Horns Versus Dorsal
1.6.2 Anterograde Degeneration 11 and Ventral Gray Columns 33
1.6.3 Retrograde Degeneration 11 3.1.5 Development of Neural Crest Cells 33
1.6.4 Regeneration of Peripheral Nerves 11 3.1.6 Framework of the Adult Cord
1.6.5 Regeneration and Neurotrophic Factors 13 is Present at Birth 34
1.6.6 Regeneration in the Central Nervous System 13 3.2 Gross Anatomy 34
1.7 Neural Transplantation 14 3.2.1 Spinal Cord Weight and Length 34
Further Reading 14 3.2.2 Spinal Segments, Regions, and
Enlargements34
Chapter 2 Development of the Nervous System 17 3.2.3 Spinal Segments in Each Region
2.1 First Week 19 Are of Unequal Length 34
2.1.1 Fertilization 19 3.2.4 Conus Medullaris, Filum Terminale,
2.1.2 From Two Cells to the Free Blastocyst 19 and Cauda Equina 35
2.2 Second Week 20 3.2.5 Termination of the Adult Spinal Cord 35
2.2.1 Implantation and Two Distinct Layers of Cells 20 3.2.6 Differential Rate of Growth: Vertebral
2.2.2 Primitive Streak and a Third Layer of Cells 20 Column Versus the Spinal Cord 36
2.3 Third Week 20 3.2.7 Relationship Between Spinal Segments
2.3.1 Primitive Node and Notochordal Process 20 and Vertebrae37
2.3.2 Neural Plate, Groove, Folds, and 3.3 Nuclear Groups – Gray Matter 37
Neuromeres21 3.3.1 General Arrangement of Spinal Cord Gray Matter 37
2.3.3 Three Main Divisions of the Brain 21 3.3.2 Gray Matter at Enlargement Levels 37
2.3.4 Mesencephalic Flexure Appears 21 3.3.3 Spinal Laminae 38
vi ● ● ● Contents

3.3.4 Dorsal Horn 38 6.2.2 Thermoreceptors 83

3.3.5 Intermediate Zone 38 6.2.3 Nociceptors 83
3.3.6 Ventral Horn 39 6.2.4 Chemoreceptors 83
3.4 Functional Classes of Neurons 39 6.2.5 Photoreceptors 84
3.4.1 Four Classes of Neurons in the Spinal Cord 39 6.2.6 Osmoreceptors 84
3.4.2 Somatic Afferent Versus Visceral Afferent Neurons 40 6.3 Classification of Receptors by Distribution
3.4.3 Somatic Efferent Versus Visceral Efferent Neurons 40 and Function84
3.4.4 Some Ventral Root Axons Are Sensory 40 6.3.1 Exteroceptors 84
3.5 Funiculi/Fasciculi/Tracts – White Matter 40 6.3.2 Interoceptors 84
3.6 Spinal Reflexes 41 6.3.3 Proprioceptors 84
3.7 Spinal Meninges and Related Spaces 42 6.4 Structural Classification of Receptors 84
3.7.1 Spinal Dura Mater 42 6.4.1 Free Nerve Endings 84
3.7.2 Spinal Arachnoid 43 6.4.2 Endings in Hair Follicles 85
3.7.3 Spinal Pia Mater 43 6.4.3 Terminal Endings of Nerves 85
3.8 Spinal Cord Injury 43 6.4.4 Neurotendinous Spindles 87
3.8.1 Hemisection of the Spinal Cord 43 6.4.5 Neuromuscular Spindles 87
3.8.2 Syringomyelia 44 6.5 Reflex Circuits 88
3.9 Blood Supply to the Spinal Cord 44 6.5.1 The Monosynaptic Reflex 88
Further Reading 44 6.5.2 Complex Reflexes 89
6.6 General Sensory Paths 89
Chapter 4 The Brain Stem 47 6.6.1 Classification of Sensory Paths by Function 89
4.1 External Features 47 6.7 Organization of General Sensory Paths 89
4.1.1 Medulla Oblongata 47 6.7.1 Receptors 89
4.1.2 Pons 50 6.7.2 Primary Neurons 89
4.1.3 Midbrain 50 6.7.3 Secondary Neurons 91
4.2 Cerebellum and Fourth Ventricle 50 6.7.4 Thalamic Neurons 91
4.2.1 Cerebellum 50 6.7.5 Cortical Neurons 91
4.2.2 Fourth Ventricle 52 6.7.6 Modulation of Sensory Paths 91
4.3 Organization of Brain Stem Neuronal Columns 52 Further Reading 92
4.3.1 Functional Components of the Cranial Nerves 52
4.3.2 Efferent Columns 54 Chapter 7 Paths for Pain and Temperature 95
4.3.3 Afferent Columns 54 7.1 Path for Superficial Pain and Temperature
4.4 Internal Features 54 from the Body95
4.4.1 Endogenous Substances 56 7.1.1 Modalities 95
4.4.2 Medulla Oblongata 56 7.1.2 Receptors 96
4.4.3 Pons 59 7.1.3 Primary Neurons 97
4.4.4 Midbrain 63 7.1.4 Secondary Neurons 98
Further Reading 65 7.1.5 Position of the LST in the Brain Stem 99
7.1.6 Thalamic Neurons 100
Chapter 5 The Forebrain 67 7.1.7 Cortical Neurons 100
5.1 Telencephalon 67 7.1.8 Modulation of Painful and Thermal
5.1.1 Telencephalon Medium 67 Impulses102
5.1.2 Cerebral Hemispheres 68 7.2 Path for Visceral Pain from the Body 102
5.1.3 Basal Ganglia (Basal Nuclei) 74 7.2.1 Modalities and Receptors 102
5.1.4 Rhinencephalon 77 7.2.2 Primary Neurons 103
5.2 Diencephalon 77 7.2.3 Secondary Neurons 103
5.2.1 Epithalamus 77 7.2.4 Thalamic Neurons 105
5.2.2 Thalamus 78 7.2.5 Cortical Neurons 105
5.2.3 Subthalamus 78 7.2.6 Suffering Accompanying Pain 105
5.2.4 Hypothalamus 78 7.2.7 Visceral Pain as Referred Pain 106
5.3 Cerebral White Matter 78 7.2.8 Transection of Fiber Bundles to Relieve
Further Reading 79 Intractable Pain 106
7.3 The Trigeminal Nuclear Complex 107
Chapter 6 Introduction to Ascending Sensory Paths 81 7.3.1 Organization of the Trigeminal
6.1 Receptors 81 Nuclear Complex107
6.2 Classification of Receptors by Modality 81 7.3.2 Organization of Entering Trigeminal
6.2.1 Mechanoreceptors 82 Sensory Fibers 107
 contents ● ● ● vii

7.4 Path for Superficial Pain and Thermal Extremes 8.7 Certain Trigeminal Reflexes 136
from the Head108 8.7.1 “Jaw‐Closing” Reflex 136
7.4.1 Modalities and Receptors 108 8.7.2 Corneal Reflex 137
7.4.2 Primary Neurons 108 Further Reading 138
7.4.3 Secondary Neurons 110
7.4.4 Thalamic Neurons 111 Chapter 9 The Reticular Formation 141
7.5 Path for Thermal Discrimination from the Head 111 9.1 Structural Aspects 141
7.5.1 Modality and Receptors 111 9.1.1 Reticular Nuclei in the Medulla 142
7.5.2 Primary Neurons 111 9.1.2 Reticular Nuclei in the Pons 143
7.5.3 Secondary Neurons 111 9.1.3 Reticular Nuclei in the Midbrain 145
7.5.4 Thalamic Neurons 112 9.2 Ascending Reticular System 146
7.5.5 Cortical Neurons 112 9.3 Descending Reticular System 149
7.6 Somatic Afferent Components of VII, IX, and X 113 9.4 Functional Aspects of the Reticular Formation 149
7.7 Trigeminal Neuralgia 113 9.4.1 Consciousness 150
7.7.1 Causes of Trigeminal Neuralgia 113 9.4.2 Homeostatic Regulation 151
7.7.2 Methods of Treatment for Trigeminal 9.4.3 Visceral Reflexes 152
Neuralgia113 9.4.4 Motor Function 153
7.8 Glossopharyngeal Neuralgia 114 Further Reading 153
Further Reading 114
Chapter 10 The Auditory System 155
Chapter 8 Paths for Touch, Pressure, 10.1 Gross Anatomy 155
Proprioception, and Vibration 117 10.1.1 External Ear 155
8.1 Path for General Tactile Sensation from the Body 117 10.1.2 Middle Ear 155
8.1.1 Modalities and Receptors 117 10.1.3 Internal Ear 156
8.1.2 Primary Neurons 118 10.2 The Ascending Auditory Path 158
8.1.3 Secondary Neurons 118 10.2.1 Modality and Receptors 158
8.1.4 Thalamic Neurons 120 10.2.2 Primary Neurons 159
8.2 Path for Tactile Discrimination, Pressure, 10.2.3 Secondary Neurons 159
Proprioception, and Vibration from the Body 120 10.2.4 Tertiary Neurons 161
8.2.1 Modalities and Receptors 120 10.2.5 Inferior Collicular Neurons 161
8.2.2 Primary Neurons 123 10.2.6 Thalamic Neurons 161
8.2.3 Secondary Neurons 124 10.2.7 Cortical Neurons 161
8.2.4 Thalamic Neurons 126 10.2.8 Comments 164
8.2.5 Cortical Neurons 127 10.3 Descending Auditory Connections 164
8.2.6 Spinal Cord Stimulation for the 10.3.1 Electrical Stimulation of Cochlear
Relief of Pain 129 Efferents165
8.3 Path for Tactile Discrimination from the Head 130 10.3.2 Autonomic Fibers to the Cochlea 165
8.3.1 Modalities and Receptors 130 10.4 Injury to the Auditory Path 165
8.3.2 Primary Neurons 130 10.4.1 Congenital Loss of Hearing 165
8.3.3 Secondary Neurons 130 10.4.2 Decoupling of Stereocilia 165
8.3.4 Thalamic Neurons 130 10.4.3 Tinnitus 166
8.3.5 Cortical Neurons 130 10.4.4 Noise‐Induced Loss of Hearing 166
8.4 Path for General Tactile Sensation from 10.4.5 Aging and the Loss of Hearing 166
the Head131 10.4.6 Unilateral Loss of Hearing 166
8.4.1 Modalities and Receptors 131 10.4.7 Injury to the Inferior Colliculi 166
8.4.2 Primary Neurons 131 10.4.8 Unilateral Injury to the Medial
8.4.3 Secondary Neurons 132 Geniculate Body or Auditory Cortex 166
8.4.4 Thalamic Neurons 132 10.4.9 Bilateral Injury to the Primary
8.4.5 Cortical Neurons 132 Auditory Cortex167
8.5 Path for Proprioception, Pressure, and Vibration 10.4.10 Auditory Seizures – Audenes 167
from the Head133 10.5 Cochlear Implants 167
8.5.1 Modalities and Receptors 133 10.6 Auditory Brain Stem Implants 167
8.5.2 Primary Neurons 133 Further Reading 167
8.5.3 Secondary Neurons 134
8.5.4 Thalamic Neurons 134 Chapter 11 The Vestibular System 171
8.5.5 Cortical Neurons 135 11.1 Gross Anatomy 171
8.6 Trigeminal Motor Component 135 11.1.1 Internal Ear 171
viii ● ● ● Contents

11.2 The Ascending Vestibular Path 173 13.2.3 Smooth Pursuit Movements 209
11.2.1 Modalities and Receptors 173 13.2.4 Vestibular Movements 209
11.2.2 Primary Neurons 175 13.3 Extraocular Muscles 209
11.2.3 Secondary Neurons 177 13.4 Innervation of the Extraocular Muscles 210
11.2.4 Thalamic Neurons 179 13.4.1 Abducent Nucleus and Nerve 211
11.2.5 Cortical Neurons 179 13.4.2 Trochlear Nucleus and Nerve 211
11.3 Other Vestibular Connections 180 13.4.3 Oculomotor Nucleus and Nerve 213
11.3.1 Primary Vestibulocerebellar Fibers 181 13.5 Anatomical Basis of Conjugate Ocular
11.3.2 Vestibular Nuclear Projections Movements215
to the Spinal Cord 181 13.6 Medial Longitudinal Fasciculus 216
11.3.3 Vestibular Nuclear Projections 13.7 Vestibular Connections and Ocular
to Nuclei of the Extraocular Muscles 182 Movements216
11.3.4 Vestibular Nuclear Projections 13.7.1 Horizontal Ocular Movements 216
to the Reticular Formation 182 13.7.2 Doll’s Ocular Movements 216
11.3.5 Vestibular Projections to the Contralateral 13.7.3 Vertical Ocular Movements 217
Vestibular Nuclei 182 13.8 Injury to the Medial Longitudinal
11.4 The Efferent Component of the Vestibular Fasciculus218
System182 13.9 Vestibular Nystagmus 218
11.5 Afferent Projections to the Vestibular Nuclei 182 13.10 The Reticular Formation and Ocular
11.6 Vertigo 183 Movements219
11.6.1 Physiological Vertigo 183 13.11  Congenital Nystagmus 219
11.6.2 Pathological Vertigo 183 13.12 Ocular Bobbing 219
Further Reading 184 13.13 Examination of the Vestibular System 219
13.14 Visual Reflexes 221
Chapter 12 The Visual System 187 13.14.1 The Light Reflex 221
12.1 Retina 187 13.14.2 The Near Reflex 222
12.1.1 Pigmented Layer 187 13.14.3 Pupillary Dilatation 223
12.1.2 Neural Layer 187 13.14.4 The Lateral Tectotegmentospinal Tract 223
12.1.3 Other Retinal Elements 188 13.14.5 The Spinotectal Tract 223
12.1.4 Special Retinal Regions 189 13.14.6 The Afferent Pupillary Defect 225
12.1.5 Retinal Areas 190 Further Reading 225
12.1.6 Visual Fields 190
12.2 Visual Path 191 Chapter 14 The Thalamus 227
12.2.1 Receptors 191 14.1 Introduction 227
12.2.2 Primary Retinal Neurons 193 14.2 Nuclear Groups of the Thalamus 228
12.2.3 Secondary Retinal Neurons 193 14.2.1 Anterior Nuclei and the Lateral
12.2.4 Optic Nerve [Ii]194 Dorsal Nucleus229
12.2.5 Optic Chiasm 196 14.2.2 Intralaminar Nuclei 231
12.2.6 Optic Tract 197 14.2.3 Medial Nuclei 233
12.2.7 Thalamic Neurons 197 14.2.4 Median Nuclei 233
12.2.8 Optic Radiations 198 14.2.5 Metathalamic Body and Nuclei 234
12.2.9 Cortical Neurons 198 14.2.6 Posterior Nuclear Complex 235
12.3 Injuries to the Visual System 200 14.2.7 Pulvinar Nuclei and Lateral Posterior
12.3.1 Retinal Injuries 200 Nucleus235
12.3.2 Injury to the Optic Nerve 201 14.2.8 Reticular Nucleus 235
12.3.3 Injuries to the Optic Chiasm 201 14.2.9 Ventral Nuclei 236
12.3.4 Injuries to the Optic Tract 202 14.3 Injuries to the Thalamus 238
12.3.5 Injury to the Lateral Geniculate Body 202 14.4 Mapping the Human Thalamus 238
12.3.6 Injuries to the Optic Radiations 202 14.5 Stimulation of the Human Thalamus 239
12.3.7 Injuries to the Visual Cortex 203 14.6 The Thalamus as a Neurosurgical Target 239
Further Reading 204 Further Reading 240

Chapter 13 Ocular Movements and Visual Reflexes 207 Chapter 15 Lower Motor Neurons and
13.1 Ocular Movements 207 the Pyramidal System243
13.1.1 Primary Position of the Eyes 207 15.1 Regions Involved in Motor Activity 243
13.2 Conjugate Ocular Movements 207 15.2 Lower Motor Neurons 243
13.2.1 Miniature Ocular Movements 208 15.2.1 Terms Related to Motor Activity 243
13.2.2 Saccades 208 15.2.2 Lower Motor Neurons in the Spinal Cord 244
 contents ● ● ● ix

15.2.3 Activation of Motor Neurons 245 16.11 Epilogue 278

15.2.4 Lower Motor Neurons in the Brain Further Reading 279
Stem245
15.2.5 Injury to Lower Motor Neurons 246 Chapter 17 The Olfactory and Gustatory Systems 283
15.2.6 Example of a Lower Motor Neuron 17.1 The Olfactory System 283
Disorder247 17.1.1 Receptors 283
15.3 Pyramidal System 247 17.1.2 Primary Neurons 284
15.3.1 Corticospinal Component 247 17.1.3 Olfactory Fila and the Olfactory Nerve 284
15.3.2 Corticobulbar Component 252 17.1.4 Olfactory Bulb – Secondary Neurons 285
15.3.3 Clinical Neuroanatomical Correlation 255 17.1.5 Olfactory Tract 285
Further Reading 256 17.1.6 Medial Stria 285
17.1.7 Lateral Stria 285
Chapter 16 The Extrapyramidal System 17.1.8 Thalamic Neurons 288
and Cerebellum259 17.1.9 Cortical Neurons 288
16.1 Extrapyramidal System 259 17.1.10 Efferent Olfactory Connections 288
16.1.1 Extrapyramidal Motor Areas 260 17.1.11 Injuries to the Olfactory System 288
16.1.2 Basal Ganglia (Basal Nuclei) 260 17.2 The Gustatory System 290
16.1.3 Afferents to the Basal Ganglia 265 17.2.1 Receptors 290
16.1.4 Cortical–Striatal–Pallidal–Thalamo– 17.2.2 Primary Neurons 292
Cortical Circuits 266 17.2.3 Secondary Neurons 293
16.1.5 Multisynaptic Descending Paths 266 17.2.4 The Ascending Gustatory Path 293
16.1.6 Common Discharge Paths 267 17.2.5 Thalamic Neurons 293
16.1.7 Somatotopic Organization of the 17.2.6 Cortical Neurons 293
Basal Ganglia267 17.2.7 Injuries to the Gustatory System 294
16.2 Cerebellum 267 Further Reading 295
16.2.1 External Features of the Cerebellum 267
16.2.2 Cerebellar Cortex 270 Chapter 18 The Limbic System 299
16.2.3 Deep Cerebellar Nuclei 271 18.1 Historical Aspects 299
16.2.4 Cerebellar White Matter 271 18.2 Anatomy of the Limbic System 300
16.3 Input to the Cerebellum Through the 18.2.1 Olfactory System 300
Peduncles271 18.2.2 Septal Area 300
16.3.1 Inferior Cerebellar Peduncle (Icp)271 18.2.3 Mamillary Bodies of the Hypothalamus 301
16.3.2 Middle Cerebellar Peduncle (Mcp)272 18.2.4 Anterior Nuclei of the Thalamus 301
16.3.3 Superior Cerebellar Peduncle (Scp)272 18.2.5 Hippocampal Formation 301
16.4 Input to the Cerebellum 272 18.2.6 Amygdaloid Complex 303
16.4.1 Incoming Fibers to the Cerebellum 272 18.2.7 Cingulate Gyrus and Cingulum 304
16.5 Cerebellar Output 273 18.2.8 Cortical Areas 306
16.5.1 From the Fastigial Nuclei 273 18.3 Cyclic Paths of the Limbic System 306
16.5.2 From the Globose and 18.4 The Human Limbic System: A Case Study 306
Emboliform Nuclei273 18.5 Descending Limbic Paths 307
16.5.3 From the Dentate Nuclei 273 18.6 Functional Aspects of the Human Limbic
16.6 Cerebellar Circuitry 273 System307
16.7 Common Discharge Paths 273 18.6.1 Emotion 307
16.8 Cerebellar Functions 274 18.6.2 Memory 308
16.8.1 Motor Functions 274 18.7 Limbic System Disorders 308
16.8.2 Nonmotor Functions 274 18.8 Injuries to Limbic Constituents 309
16.8.3 Studies Involving the Human Cerebellum 274 18.8.1 Septal Area 309
16.8.4 Localization in the Cerebellum 274 18.8.2 Hippocampal Formation 309
16.9 Manifestations of Injuries to the Motor System 275 18.8.3 Amygdaloid Complex 309
16.9.1 Injury to the Premotor Cortex 275 18.8.4 Seizures Involving the Limbic System 309
16.9.2 Injury to the Basal Ganglia 275 18.9 Psychosurgery of the Limbic System 309
16.9.3 Injury to, or Deep Brain Stimulation 18.9.1 Drug‐Resistant Epilepsy 309
of, the Subthalamic Nucleus 276 18.9.2 Violent, Aggressive, or Restless Behaviors 310
16.9.4 Injury to the Cerebellum 277 18.9.3 Schizophrenia 310
16.9.5 Localization of Cerebellar Damage 278 18.9.4 Intractable Pain 310
16.10 Decorticate Versus Decerebrate Rigidity 278 18.9.5 Psychiatric Disorders and Abnormal
16.10.1 Decerebrate Rigidity 278 Behavior310
16.10.2 Decorticate Rigidity 278 Further Reading 310
x ● ● ● Contents

Chapter 19 The Hypothalamus 313 21.8 Parietal Lobe 347

19.1 Hypothalamic Zones (Medial to Lateral) 313 21.8.1 Primary Somatosensory Cortex (Si)348
19.2 Hypothalamic Regions (Anterior to Posterior) 315 21.8.2 Secondary Somatosensory Cortex (Sii)350
19.3 Hypothalamic Nuclei 315 21.8.3 Superior Parietal Lobule 350
19.3.1 Chiasmal Region 315 21.8.4 Inferior Parietal Lobule 352
19.3.2 Tuberal Region 319 21.8.5 Parietal Vestibular Cortex (2v) 352
19.3.3 Mamillary Region 320 21.8.6 Mirror Representation of Others’ Actions 353
19.4 Fiber Connections 321 21.8.7 Preoccipital Areas 353
19.4.1 Medial Forebrain Bundle 321 21.9 Occipital Lobe 354
19.4.2 Stria Terminalis 321 21.9.1 Primary Visual Cortex (V1) 354
19.4.3 Fornix 321 21.9.2 Secondary Visual Cortex 354
19.4.4 Diencephalic Periventricular System 321 21.10 Temporal Lobe 354
19.4.5 Dorsal Longitudinal Fasciculus 321 21.10.1 Primary Auditory Cortex (Ai)354
19.4.6 Anterior and Posterior 21.10.2 Wernicke’s Area 354
Hypothalamotegmental Tracts 322 21.10.3 Temporal Vestibular Cortex 355
19.4.7 Pallidohypothalamic Tract 322 21.10.4 Midtemporal Areas Related to Memory 356
19.4.8 Mamillothalamic Tract 322 21.10.5 Anomia 356
19.4.9 Vascular Connections 322 21.10.6 Prosopagnosia 356
19.5 Functions of the Hypothalamus 322 21.10.7 Psychomotor Seizures 356
19.5.1 Water Balance – Water Intake and Loss 322 21.11 Insula 357
19.5.2 Eating – Food Intake 322 21.12 Aphasia 358
19.5.3 Temperature Regulation 323 21.12.1 Broca’s Aphasia 358
19.5.4 Autonomic Regulation 323 21.12.2 Wernicke’s Aphasia 359
19.5.5 Emotional Expression 323 21.12.3 Conductive Aphasia 359
19.5.6 Wakefulness and Sleep – Biological Rhythms 323 21.12.4 Global Aphasia 359
19.5.7 Control of the Endocrine System 324 21.13 Alexia 360
19.5.8 Reproduction 324 21.14 Apraxia 360
Further Reading 324 21.15 Gerstmann’s Syndrome 360
21.16 Agnosia 360
Chapter 20 The Autonomic Nervous System 327 21.17 Dyslexia 360
20.1 Historical Aspects 327 Further Reading 361
20.2 Structural Aspects 328
20.2.1 Location of Autonomic Neurons of Origin 328 Chapter 22 Blood Supply to the Central Nervous
20.2.2 Manner of Distribution of Autonomic Fibers 329 System365
20.2.3 Termination of Autonomic Fibers 330 22.1 Cerebral Circulation 365
20.3 Somatic Efferents versus Visceral Efferents 331 22.2 Aortic Arch, Brachiocephalic Trunk,
20.4 Visceral Afferents 331 and Subclavian Vessels 366
20.5 Regulation of the Autonomic Nervous System 333 22.3 Vertebral–Basilar Arterial System 366
20.6 Disorders of the Autonomic Nervous System 333 22.3.1 Branches of the Vertebral Arteries 367
Further Reading 334 22.4 Blood Supply to the Spinal Cord 368
22.4.1 Extramedullary Vessels 368
Chapter 21 The Cerebral Hemispheres 337 22.4.2 Intramedullary Vessels 371
21.1 Facts and Figures 337 22.4.3 Spinal Veins 371
21.2 Cortical Neurons 338 22.5 Blood Supply to the Brain Stem and Cerebellum 372
21.3 Cortical Layers 338 22.5.1 Extrinsic or Superficial Branches 372
21.4 Cortical Columns (Microarchitecture) 343 22.5.2 Branches of the Basilar Arteries 372
21.5 Functional Aspects of the Cerebral Cortex 343 22.5.3 Intrinsic or Penetrating Branches 375
21.6 Cerebral Dominance, Lateralization, and Asymmetry 343 22.5.4 Classical Brain Stem Syndromes 377
21.7 Frontal Lobe 343 22.6 Common Carotid Artery 378
21.7.1 Primary Motor Cortex 343 22.6.1 External Carotid Artery 378
21.7.2 Premotor Cortex 344 22.6.2 Internal Carotid Artery: Cervical,
21.7.3 Supplementary Motor Area (Sma)345 Petrous, and Cavernous Parts 379
21.7.4 Cingulate Motor Areas 345 22.7 Blood Supply to the Cerebral Hemispheres 379
21.7.5 Frontal Eye Fields 345 22.7.1 Internal Carotid Artery: Cerebral Part 379
21.7.6 Broca’s Area 346 22.7.2 Branches of the Internal Carotid Artery 379
21.7.7 Prefrontal Cortex 346 22.7.3 Posterior Cerebral Artery 383
 contents ● ● ● xi

22.8 Cerebral Arterial Circle 383 23.1.4 Dural Projections 388

22.8.1 Types of Arteries Supplying the Brain 384 23.1.5 Intracranial Herniations 389
22.9 Embryological Considerations 384 23.2 Ventricular System 391
22.10 Vascular Injuries 384 23.2.1 Introduction 391
22.10.1 Brain Stem Vascular Injuries 384 23.2.2 Lateral Ventricles 391
22.10.2 Visualization of Brain Vessels 384 23.2.3 Third Ventricle 392
Further Reading 384 23.2.4 Aqueduct 393
23.2.5 Fourth Ventricle 393
Chapter 23 The Meninges, Ventricular System, 23.3 Cerebrospinal Fluid 393
and Cerebrospinal Fluid 387 Further Reading 394
23.1 The Cranial Meninges and Related Spaces 387
23.1.1 Cranial Dura Mater 387 Figure and Table References 395
23.1.2 Cranial Arachnoid 387 Index  399
23.1.3 Cranial Pia Mater 388
Preface

It is a great privilege to write a book on the human brain. conscious. Dr. Crosby sought to impart to me her clinically
I have studied and taught about the human brain to medical conscious, ­anatomical mindedness that hopefully is reflected
students and graduate students from an assortment of in this book.
­disciplines (biomedical science, exercise science, neurosci- The preparation of this book has come at a time when
ence, physical therapy, psychology) and also residents, there has been an enormous explosion in our knowledge
­neurologists, and neurosurgeons for some four decades. My about the nervous system. Searching Google to obtain
students have asked me thousands of questions that have information about the term “brain” results in 552 000 000
encouraged me in my own personal study, and have helped citations. If one searches PubMed for the term “brain,”
clarify my thinking about the structure and function of the some 1.6 million citations result. Therefore, keeping up
human brain. Therefore, I dedicate this book to my students with current studies of the human brain and spinal cord is
as a way of thanking them for what they have taught me. an impossible task. At the end of each chapter is a set of
I am grateful to Dr. Paul A. Young, Professor and Chairman “Further Reading” that the interested reader might want
Emeritus, Department of Anatomy and Neurobiology, Saint to consider should there be a desire to learn more about
Louis University School of Medicine, who gave me the oppor- the topics covered in that chapter or gain a different
tunity to begin my graduate studies in anatomy and served as ­perspective on a particular topic. Many of these r­ eferences
a role model to me. Dr. Young is the epitome of a dedicated relate to items in the text.
and excellent teacher and the author of an exceptional A special thank you goes to Jasna Markovac, who has
­textbook on basic clinical neuroanatomy. I am also grateful to been involved with this book in many ways from the
my distinguished colleagues Drs. Ronan O’Rahilly and beginning and enabled me to produce this edition with
­
Fabiola Müller for their many book‐related comments, sug- Wiley‐Blackwell.
gestions, and criticisms. Their studies of the embryonic It is my sincere hope that you the reader will enjoy read-
human brain are without equal. Dr. O’Rahilly has been an ing this book and that in the process you will begin to grasp
invaluable resource during the writing of this book. something of what little we do know about the structure and
It was my privilege to study with the late Dr. Elizabeth function of the human brain and spinal cord. It is my hope
C. Crosby. She was my teacher, fellow researcher, and friend. that by reading this book you will begin a lifelong study of
Dr. Crosby had a profound understanding of the human the nervous system. It is also my hope that studying the
nervous system based on her many years of study of the nervous system will lead you to do more than just write a
­comparative anatomy of the nervous system of vertebrates, book but rather make a discovery, find a cure, or actively
including humans. She had a long and distinguished career participate in some worthwhile endeavor that will relieve the
teaching medical students, residents, neurologists, and neu- suffering of those with neurological disease and give them
rosurgeons and she had many years of experience correlat- hope for a better life.
ing ­ neuroanatomy with neurology and neurosurgery in
clinical conferences and on rounds. Because of that experi- Soli Deo Gloria
ence, one could gradually see the clinicians become more James R. Augustine
anatomically minded and the anatomists more clinically Columbia, South Carolina
Other documents randomly have
different content
hand, namely in the steppes and deserts of Asia Minor, Palestine,
and Persia, and in several valleys of Arabia and India, they had a
wild horse full of promise, one which still lives, our kulan. This differs
indeed in several features from our horse, but not more than the
greyhound, the poodle, or the Newfoundland differs from the wolf or
any other wild dog, not more than the dachshund, the terrier, or the
spaniel from the jackal, not more than the pony from the Arabian
horse, or the Belgian-French cart-horse from the English racer. The
differences between our domesticated horse and the wild form which
seems to me its most probable ancestor are indeed important, but
horse and kulan seem to regard themselves as belonging to the
same blood, since they seek each other’s company.
When, on the 3rd June, 1876, we were riding through the dreary
desert steppes between the Saisan lake and the Altai—a region from
which I have drawn the main features of the above sketch—we saw
in the course of the forenoon no fewer than fifteen kulans. Among
these we observed one pair in particular. They stood on the broad
crest of a near hillock, their forms sharply defined against the blue
sky, and powerfully did they raise the desire for the chase in us and
in our companion Kirghiz. One of them made off as we appeared,
and trotted towards the mountain; the other stood quietly, and
seemed as if considering a dilemma, then raised its head once and
again, and at last came running towards us. All guns were at once in
hand; the Kirghiz slowly and carefully formed a wide semicircle with
the intention of driving the strangely stupid and inconceivably
careless creature towards us. Nearer and nearer, halting now and
then, but still steadily nearer he came, and we already looked upon
him as a sure captive. But a smile broke over the face of the Kirghiz
riding beside me; he had not only discovered the motive of the
creature’s apparently foolish behaviour, he had recognized the
animal itself. It was a Kirghiz horse, dappled like a kulan, which,
having strayed from his master’s herd, had fallen in with wild horses,
and, for lack of better company, had stayed with them. In our horses
he had recognized his kin, and had therefore forsaken his friends in
need. Having come quite near to the Kirghiz, he stopped again as if
to reflect whether he should once more yield his newly-healed back
to the galling saddle; but the first steps towards return were followed
by others, and without an attempt at flight he allowed them to halter
him, and in a few minutes he was trotting as docilely by the side of
one of the horsemen as if he had never known the free life of his
ancestors. Thus we were able to confirm by personal experience the
already accepted fact, that horse and kulan do sometimes keep
company.
The kulan is a proud, fascinating creature, full of dignity, strength,
and high spirits. He stares curiously at the horseman who
approaches him; and then, as if deriding the pursuer, trots off
leisurely, playfully lashing his flanks with his tail. If the rider spurs his
horse to full speed, the kulan takes to a gallop as easy as it is swift,
which bears him like the wind over the steppes and soon carries him
out of sight. But even when at full speed he now and again suddenly
pulls himself up, halts for a moment, jerks round with his face to the
pursuer, neighs, and then, turning, kicks his heels defiantly in the air,
and bounds off with the same ease as before. A fugitive troop always
orders itself in line, and it is beautiful to see them suddenly halt at a
signal from the leader, face round, and again take to flight.
 Fig. 18.—The Kulan (Equus hemionus).

As among all horses, each troop of kulans has a stallion for leader,
and he is the absolute master of them all. He leads the troop to
pasture as well as in flight, he turns boldly on carnivores which are
not superior in force, he permits no quarrelling among his followers,
and tolerates no rival, indeed no other adult male in the herd. In
every district which the kulans frequent, solitary individuals are to be
seen unaccompanied by any troop; they are stallions which have
been vanquished and driven off after furious and protracted
combats, and which must roam about alone until the next breeding
season. In September they again approach the herds, from which
the old stallion now drives off the newly-matured stallions, and a
fierce battle begins when they catch sight of an opponent. For hours
at this season they stand on the crests of steep ridges, with widely-
open nostrils raised to the wind, with their eyes on the valley before
them. As soon as the banished one sees another stallion, he rushes
down at full gallop, and fights to exhaustion with both teeth and
hoofs. Should he conquer the leader of a herd, he enters upon his
rights, and the mares follow him as they did his predecessor. After
the battles are over comes the time of wandering, for the hard winter
drives the herds from one place to another, and it is only when spring
has fully come that they return to their old quarters. Here, in the end
of May or in the beginning of June the mare brings forth her foal,
which in every respect resembles that of the domestic horse—a
somewhat awkward-looking, but very nimble and lively creature. We
had the good fortune to be able to make its acquaintance.
On climbing one of the elongated hillocks of these desert-steppes,
we suddenly saw at a short distance three old kulans and a foal,
which seemed to be only a few days old. Our Russian companion
fired a shot, and away rushed the wild horses, with their hoofs
scarce touching the ground, yet exerting their incomparable agility
almost with the ease of play, and obviously keeping themselves in
check for the sake of the foal. Down rushed all the Kirghiz and
Cossacks of our company; our attendants, carried away with the
general excitement, also gave chase; and down we also rushed. It
was a wild chase. Still playing with their strength, the wild horses
made for the distant mountains, while all the riders urged on their
steeds to their utmost, till their bellies seemed almost to touch the
ground. The desert resounded with the jubilant cries of the Kirghiz,
the thundering of their horses at full gallop, the neighing of our
slower horses which gnashed at their bridles; fluttering cloaks and
kaftans and whirling clouds of dust filled and enlivened the solitudes.
Further and further rushed the chase. Then the foal separated itself
from its older companions and fell behind; the distance increased
between it and the anxious glance which the mother repeatedly cast
back; the distance between it and our horsemen decreased; and in a
few minutes it was taken. Without resistance it gave itself up to its
pursuers; it showed no trace of the characteristic qualities of the
adults—wildness, hardly governable self-will, and inconquerable
roguishness, which often degenerates into downright spite.
Innocently it gazed at us with its large lively eyes, with apparent
pleasure it allowed us to stroke its soft skin, without resistance it
allowed itself to be led along with a halter, in child-like carelessness
it lay down beside us seeking obviously much-needed rest after the
heat of the chase. The charming creature at once captivated every
one. But who was to find a milk-mare to be its foster-mother, who
was to give it rest and care? Both were impossible, and on the
second day the lovable creature was dead. With the passion of
sportsmen would we have killed a full-grown kulan, but to see the
foal die gave us genuine sorrow.
In vain we tried to capture one of the adults; in vain we lay in
ambush beside the bound foal in hopes of inveigling its mother; in
vain we tried to effect something by driving; none of us had any luck.
As a sportsman, I left the dreary solitudes with regret, but as a
naturalist I was in the highest degree satisfied, for there I had come
to know the noblest creature of the steppes.
 THE FORESTS AND SPORT OF
SIBERIA.
Siberian scenery gives one an impression of uniformity and
monotony, which is mainly due to the fact that the whole country
consists of three zones, each more or less homogeneous within
itself, though distinct from the other two. Each of these zones
preserves its special character everywhere, and the same picture is
repeated a hundred times, satiating and blunting the senses till one
becomes almost incapable of recognizing or appreciating the charms
of any scene. Thus it is that we seldom hear anyone speak with
appreciation, much less with enthusiasm, of the scenery of this wide
region,—although it certainly deserves both—and, thus, gradually
there has become fixed in our minds an impression of Siberia which
refuses correction with an obstinacy proportionate to its falseness.
Siberia is thought of as a terrible ice-desert, without life, without
variety, without charm, as a frozen land under the curse of heaven
and of miserable exiles. But it is entirely forgotten that Siberia
includes a full third of Asia, and that a region which is almost twice
as large as the whole of Europe, which extends from the Ural to the
Pacific Ocean, from the Arctic sea to the latitude of Palermo, cannot
possibly be excessively monotonous nor uniform in all its parts. But
people usually picture only one district of Siberia, and even that in a
false light.
In truth the country is richer in variety than any one has hitherto
described it. Mountains interrupt and bound the plains, both are
brightened by flowing and standing water, the sun floods hills and
valleys with shimmering light and gleaming colour, lofty trees and
beautiful flowers adorn the whole land, and men live happily, joyous
in their homes.
Of course there are undeniable wildernesses even now in Siberia,
and these, along with the likewise real ice-deserts, the tundras, do,
to a certain extent, justify the popular impression. Dreary also are the
forests which lie between the tundra and the steppes, and form the
third zone. In them man never ventures to establish himself; on them
the industry of the settlers along their borders make relatively little
impression; within them the forces of nature hold absolute sway,
creating and destroying without interference. The flame of heaven
sets the trees ablaze, the raging winter-storm hurls them to the
ground; the forests rise and disappear without any human control,
and may in the fullest sense of the word be called primeval. Full of
mystery they attract, and at the same time inhospitably repel; inviting
they seem to the hunter, but resistant they bar his steps; rich gain
they promise the eager merchant, but postpone the fulfilment of his
wishes to the future. This girdle of forest extends, as we have
mentioned, between the steppes and the tundra. Here and there it
encroaches on both; here and there they intrude upon it. At certain
places in both the unwooded zones, a compact wood may dispute
possession with the characteristic vegetation of steppes or of tundra,
as the case may be, but such isolated woods are almost always like
islands in the sea, for whose presence there is no obvious
justification. In the steppes they are restricted to the northern slopes
of the mountains and to the valleys, in the tundra to the deepest
depressions. But in both cases they are unimportant in comparison
with the measureless extent of the forest zone, in which it is only
here and there that a stream, a lake, or a swamp interrupts the
continuity of the wilderness of trees which extends on all sides. A
conflagration may make a clearing, or, at the extreme fringe, man
may make a gap, but otherwise there is no interruption. Whole
countries, as we know them, might find space in one of these
immense forest tracts; and there are kingdoms of smaller area than
some of them. What the interior is like no one can tell, for not even
by the streams which flow from them can one penetrate far, and
even the boldest sable-hunters do not know more than a margin of at
most fifty or sixty miles.
The general impression which the Siberian forests make on the
German traveller is by no means favourable. At the apparently
boundless tracts which are wooded, he is of course astonished, but
he cannot be enthusiastic, or at least very rarely. The creative,
productive, renewing power of the North does not seem to be
adequate to balance the destructive forces. Hoary age stands side
by side with fresh youth, but somehow there is no vitality in the
combination; incomputable wealth appears in beggar’s garb; and
moribund life without any promise of vigorous rejuvenescence
inhibits any feeling of joy. Everywhere we seem to perceive the hard
struggle for existence, but nowhere are we really fascinated or
attracted by the spirit of the woods, nowhere does the interior fulfil
the expectations which the external aspect suggested. The
splendour of the primitive forests in lower latitudes is entirely and
absolutely lacking in this derelict, uncared-for woodland. The life
which stirs within them seems as if it had already fallen under the
shadow of death.[28]
True forest, full of fresh life, with continuance amid a regular
succession of changes, is rare. The devastation wrought by fire is a
much more frequent spectacle. Sooner or later a lightning-flash, or
the culpable carelessness of the Siberian, sets the forest in a blaze.
Favoured by the season and the weather the conflagration spreads
in a manner scarce conceivable. Not for hours, but for days, or even
for weeks, the destruction rages. On the mossy and turfy ground the
flames smoulder and creep further and further; the quantities of dry
and mouldy débris on the ground feed them, dry branches hanging
down to the ground, or dead trunks, still upright, lead them to the
tops of the living trees. Hissing and cracking the resinous needles
fall, and a gigantic spray of sparks rises to heaven. In a few minutes
the giant tree is dead, and the destruction spreads; the rockets which
radiate from it fall in thousands of sparks, and all around fresh
flames spring from the glowing seed. Thus every minute the fire
gains ground, and destruction spreads on all sides uncontrolled. In a
few hours square miles of the forest are ablaze. Over hundreds of
square versts steaming clouds of smoke darken the sun; slowly, but
thickly, and ever more thickly, the ashes drizzle down, and tell by day
to distant settlers, as the glow reflected in the sky proclaims by night,
that there is a fire in the forest. Affrighted animals carry terror into the
surrounding townships. Immediately after great forest-fires, bears
appear in districts where they have not been seen for years; wolves
wander over the open country in formidable troops as if it were
winter; elks, stags, roedeer and reindeer seek new homes in distant
forests; and squirrels in countless swarms hurry through wood and
plain, field and meadow, village and town. How many of the terror-
stricken beasts fall victims to the fire no one can estimate, but it has
been found that woodlands desolated by conflagration remain for
many years thereafter without fresh settlers, and that the valuable
beasts of the chase have entirely disappeared from many of these
desolated districts. The devastation is sometimes on a scale of vast
magnitude; thus, in 1870, a fire which raged for about fourteen days
destroyed a million and a quarter acres of valuable forest in the
government of Tobolsk, while clouds of smoke and showers of ashes
were borne to a distance of a thousand miles from the seat of the
conflagration.
For many years the devastated woodland remains like an immense
succession of ruins; even after a generation or two the limits of the
conflagration may be recognized and defined. The flames destroy
the life of almost all the trees, but they devour only those which were
already dry; thus stems more smoked than charred remain standing,
and even their tops may remain bereft only of their needles, young
shoots, and dry twigs. But they are dead and their destruction is in
process. Sooner or later they are bound to fall before the storm. One
after another is hurled to the ground, and one after another is robbed
of its branches, its crown, or a third or a fourth of the trunk is broken
off from the top. Across one another, at all angles, and at different
levels, thousands of these tree-corpses lie prostrate on the ground
already thickly covered with piles of débris. Some rest on their roots
and top-branches, others lean on the still upright stems of their
neighbours, and others already lie crumbling among the fallen
branches, their tops often far from their trunks, their branches
scattered all around. To the lover of the woods, those stems which
still withstand the storm have perhaps an even more doleful
appearance than those which have fallen. They stand up in
nakedness like bare masts. Only a few retain their tops, or parts of
them, for several years after the fire; but the weather-beaten twigless
branches of the crowns rather increase than lessen the
mournfulness of the picture. Gradually all the crowns sink to the
ground, and the still upright trunks become more and more rotten.
Woodpeckers attack them on all sides, chisel out nesting-holes, and
make yard-long passages leading into the tree’s heart, thus allowing
the moisture free entrance and accelerating the process of decay. In
the course of years even the largest trunk has mouldered so
completely that it is really one huge homogeneous mass of rotten
tinder which has lost all stability. Indeed, a rough shake from a man’s
hand is sufficient to make it fall into a heap of shapeless débris.
Finally, even this disappears, and there is left a treeless expanse,
broken only here and there by the last traces of a trunk.
But even here a new life begins to rise from amid the ruins. Some
years after the conflagration, the charred ground, manured by ashes
and decayed débris, begins once more to be adorned. Lichens and
mosses, ferns and heaths, and above all various berry-bearing
bushes cover the ground and the débris of the trees. These flourish
more luxuriantly here than anywhere else, and they begin to attract
animals as various as those which the flames had banished. Seeds
of birch borne by the wind germinate and become seedlings, which
gradually form, at first exclusively, a thicket as dense as if it had
sprung from man’s sowing. After some years a young undergrowth
has covered the field of the dead; after a longer interval other forest
trees gradually arise in the room of their predecessors. Every forest-
fire spares some parts of the region which it embraces; even isolated
trees may survive in the midst of the burned area, and effect the re-
sowing of the desolated tract. Sheets of water and deep gorges may
set limits to the fire, and it may even happen that the flames, leaping
over a gulley, continue their devastation on the opposite bank
without injuring the trees in the depths beneath. Moreover, individual
larch-trees which have been attacked by the fire may escape
destruction. The bases of the trunks are charred and all the needles
are shrivelled up, but often the crown bursts forth afresh, and for a
time the tree continues, though somewhat miserably, to live.
In comparison with the ravages of the flames, the devastations for
which man is directly responsible seem trivial, but in themselves they
are of no slight importance. Of forest-culture the Siberian has no
conception. The forest belongs to God, and what is His is also the
peasant’s; thus, in view of the practically infinite wealth, he never
thinks of sparing, but does what he pleases, what the needs of the
moment seem to him to demand. Every Siberian fells and roots out,
where and as he pleases, and everyone destroys infinitely more than
he really requires. For a few cones he will fell a pine, even if it be in
the prime of growth; to obtain building wood he will cut down three or
four times the quantity required, leaving the residue without a
thought, often not even using it for fuel. Already, such careless
procedure has entailed serious consequences. The woods in the
neighbourhood of townships, and here and there even those near
the highways, are worked out, and appear scarce better than those
which the fire has devastated; and still the work of destruction goes
on. It is only since 1875 that there have been forest-officers in
Western Siberia, and even they give their attention rather to the
exploitation than to the renewal of the woods.
Even where neither man nor fire has ravaged them the forests
present an appearance essentially different from ours—an
appearance of complete, absolutely uncontrolled naturalness. It is
but rarely, however, that this attracts us. At first, perhaps, we are
impressed by seeing at one glance all stages of growth and decay;
but the dead soon becomes more conspicuous than the living, and
this depresses instead of stimulating.[29] In forests thus left in their
natural state, thick growth alternates with clearing, tall trees with
mere thicket, hoary senility with vigorous youth. Mouldering trees
stand or lean, hang or lie everywhere. From the remains of fallen
stems young shoots sprout; gigantic corpses bar the way within the
thickets. Willows and aspens, which, with the birch, are the most
abundant foliage-trees of Western Siberia, appear at times in
irreproachable perfection, and at times as if they had been
persistently hindered from full growth. Stems thicker than a man’s
waist bear tangled crowns of small size, on which, year after year,
fresh twigs break forth without being able to grow into branches;
other apparently aged trees remain not more than bushes; and
others, broken across the middle, have their split, cracked, and
twisted upper parts connected to the trunk only by the splintered
bark. Rarely does one get a complete picture; everything looks as if
it were going to ruin, and could advance only in decay.
Yet this sketch is not true of all the woods in this vast region; there
are indeed woodlands, especially in the south of the zone, on which
the eye rests with satisfaction. Locality, situation, soil, and other
conditions are sometimes alike propitious and combine to produce
pleasing results. The growth of the individual trees becomes
vigorous, and the general composition of the wood changes; the
undergrowth, which is luxuriant everywhere, becomes diversified in
the most unexpected manner. Gladly one welcomes each new
species of tree or bush which reduces the marked poverty of species
in these forests, but even from the richest tracts many trees are
awanting which we rarely miss in Europe at the same latitude. It
must be confessed that the forests of Siberia are uniform and
monotonous, like the steppes, and like the tundra.
In the river-valleys of the forest zone the uniformity is perhaps most
conspicuous. Here the willows predominate, forming often extensive
woods by the banks and on the islands, almost to the complete
exclusion of other trees. Over wide stretches willows alone form the
woods of the valley, and in many places the trees rise to a stately
height, yet even then without often gaining in impressiveness or
charm. For the isolated willow-tree is not more, but rather less
picturesque than the willow bushes; its crown is always thin and
irregular, it is not close-set but loose and open, in fact almost
scraggy. On frequent repetition it becomes wearisome. When the
willows stand, as is usual, close beside one another, they form a
dense thicket, and then, even more than the isolated tree, they lack
character, for all the stems rise like posts and all the crowns fuse into
a close, straight-contoured mass of foliage, suggestive of a clipped
hedge, in which the individual trees are entirely merged. As pleasing
additions to such monotonous woods we welcome the sprinkling of
poplars, the silver poplar in the south, the aspen in the north, both of
them giving some animation to the willows. In the valley of the
stream too, but only in those places which are not subject to
regularly-recurring floods, the birch appears in addition to the trees
already noticed; indeed birch-covered tracts occur with some
constancy as connecting links between the willow-woods and the
pine-forests. But it is only in the south of the zone that the birch
attains its full size and vigour; it is as unresisting a victim to the
flames as the most resinous pine, and is therefore incapable of
greatly affecting the general aspect of the forest. More or less
unmixed birch woods bound the forest zone to the south, and
sometimes intrude far into the steppes, yet it is but rarely that they
form thick, compact, well-established stretches of timber; and they
are, when one sets foot in them, disappointing.
On the other hand, the pine-forests which cover all the regions
between the river-courses often fascinate and satisfy the traveller
from the west. If the tundra has not gained upon them or begun to
make its desolating mark, they consist in the main of vigorous pines
and Norway spruce firs, the pichta or Siberian silver fir, the cembra
pine, and more rarely larches. Among these there are aspens and
willows, with occasional mountain-ash and bird-cherry, while birches
often appear in as great vigour as in woods which consist exclusively
of this accommodating tree. The pichta and the cembra pine are the
characteristic trees of all West Siberian pine-forests, and vie with
one another in beauty and vigour of growth. The pichta is a
particularly beautiful tree. Nearly related to our silver fir, and
representing it in all East Russian and West Siberian woodlands,
even from a distance it catches the eye, standing out impressively
from among all the other conifers. From the silver fir and from the
Norway spruce fir the pichta is distinguished by the stateliness of its
slender conical crown and by the rich, delicate, bright green needles.
Almost always it overtops the other trees of the forest; usually,
indeed, the topmost third is above the crowns of its neighbours, thus
effectively breaking the sky-line of the forest and giving an individual
character to certain regions. The cembra or stone pine, which
flourishes especially in the south of the forest zone, though it also
occurs far to the north, has round, smooth, usually compact tops
which contrast well with those of the other pines and firs; and it also
contributes not a little to the external adornment of the forest,
towards making it seem more attractive than it is. Pines and spruce
firs are nowhere absent, but they do not flourish everywhere as they
do in the mountains of Central Germany; towards the north they sink
rapidly into crippled senility. And so is it also with the larches, whose
true home is Siberia; it is only in the south of the forest zone,
especially on the mountains, that they attain the stately height of
those in our country.
The above-named species include almost all those which occur
regularly in the woodlands of Western Siberia. There seems to be a
complete absence of oak and beech, elm and ash, lime and maple,
silver fir and yew, hornbeam and black poplar. On the other hand,
there are many kinds of bushes and shrubs in abundance
everywhere. Even in the north the undergrowth of the forests is
surprisingly rich and luxuriant. Currants and raspberries flourish to a
latitude of 58°, a species of woodbine occurs up to 67°; juniper, white
alder, sallow, crowberry, bilberry, cranberry, and cloudberries
increase rather than decrease as one goes north; and even on the
margins of the tundra, where dwarf-birches and marsh-andromedas,
mosses, and cowberries insinuate themselves into the interior of the
woods, the ground is still everywhere thickly covered, for the mosses
thrive the more luxuriantly the poorer the woods become. The
steppes also contribute to enrich the woods, for in the south of the
forest zone, not only most of the steppe-bushes and shrubs, but also
various herbs and flowers, enter or fringe the forest. Thus certain
wooded stretches of this border-land become natural parks, which in
spring and early summer display a surprising splendour of blossom.
As an instance of a forest glorious in such charms, I may mention
that region known as “Taiga,”[30] which lies between the towns
Schlangenberg and Salain, in the domain of Altai. In the broad tract
which this beautiful forest covers there is a most pleasing succession
of long ridges and rounded hills, valleys, troughs, and basins. One
hill rises beyond and above another, and everywhere one sees a
sky-line of forest. Pines and pichta firs, aspen and willow, mountain-
ash and bird-cherry, are in the majority among the high trees, and
are mingled in most pleasing contrasts of bright and dark colour, of
light and shade. The soft lines of the foliage trees are pleasingly
broken by the conical summits of the pichta firs which overtop them.
The two species of Siberian pea-tree, guelder-rose and woodbine,
wild rose and currant are combined in the brightly blooming
undergrowth; Umbellifers as tall as a man, especially hemlock;
spiræa; ferns such as maidenhair, larkspur and foxglove, bluebell
and hellebore all shooting up in unparalleled luxuriance, weave a
gay carpet, from which the wild hops climb and twine up to the tall
trees. It is as if the art of the landscape gardener had been
intelligently exercised, as if man had fashioned the whole with an
eye to scenic effect.
In the south, the forests show their greatest beauty in spring, in the
north, in autumn. By the first days of September the leaves of the
foliage trees begin to turn yellow here, and by the middle of the
month the north Siberian forest is more brilliant than any of ours.
From the darkest green to the most flaming red, through green and
light green, light yellow and orange yellow, pale red and carmine, all
the shades of colour are represented. The dark Norway spruce firs
and pichta firs are followed by the cembra pines and larches; and
next in order come the few birches which are not yet yellowed. The
white alders display all gradations from dark to light green and to
greenish yellow; the aspen leaves are bright cinnabar red, the
mountain-ash and the bird-cherry are carmine. So rich and yet so
harmonious is the mingling of all these colours that sense and
sentiment are satisfied to the full.
Such are the pictures which the woodlands of Western Siberia
display to the traveller. But all the sketches we have attempted to
give have been taken from within a narrow fringe. To penetrate
further into the primeval forests, in summer at least, seems to the
western traveller absolutely impossible. On the slopes of the
mountains he is hindered by thickets and masses of débris, on
highland and plain alike by prostrate trees and a tangle of bushes, in
the hollows and valleys by standing and flowing water, by brooks and
swamps. Wide-spread talus from the rocks, blocks and boulders
rolled into heaps and layers form barriers on all the hills; lichens and
mosses form a web over the rocks, and treacherously conceal the
numerous gaps and clefts between them; a young undergrowth is
rooted between and upon the old possessors of the soil; and the old
trees as well as the young increase the risk of attempting to traverse
these regions. On the low ground the obstacles which the forests
present are hardly less formidable. Literally impenetrable thickets
such as exist in the virgin forests of equatorial countries there are
none, but there are obstacles enough. The prostrated trunks are all
the more troublesome because most of them lie, not on the
untrodden path, but at an inconvenient height above it; they are
turnpikes in the most unpleasant sense of the word. Sometimes it is
possible to climb over them or to creep under them; but equally often
neither is possible, and one is forced to make a circuit, which is the
more unwelcome, since, without constant reference to the compass,
it is only too easy to stray from the intended direction. Real clearings
are met with but rarely, and if one tries to walk across them, deep
holes and pools full of mud and decayed débris soon show that here
also the greatest caution is necessary. If the traveller trusts to one of
the many cattle-tracks, which, in the south of the forest-zone, lead
from every village to the forest, and penetrate into it for some
distance, even then sooner or later he finds himself at fault. It is
impossible to tell, or even to guess, whither such a path leads, for it
intersects hundreds of others, and runs through tangled brushwood,
through tall grass concealing unpleasant débris of trees, through
moss and marsh; in short, they are not paths for human foot. Thus,
though there are not everywhere insuperable obstacles, one meets
everywhere and continuously with hindrances so numerous and so
vexatious, that even where the plague of mosquitoes is not
intolerable, the traveller is apt to return much sooner than he had
intended. Only in winter, when hard frost has covered all the pools,
bogs, and swamps with a trustworthy crust, when deep snow has
smoothed off most of the roughnesses, and is itself coated with a
hard layer of ice, only then are the forests accessible to the hunter
equipped with snow-shoes, and accompanied by weather-hardened
dogs; only then can even the natives think of making long
expeditions.
Siberian forests are dumb and dead, “dead to the point of
starvation”, as Middendorf most justly says. The silence which reigns
within them is a positive torture. When the pairing of the black-cock
is past one may hear the song of the fieldfare and the black-throated
thrush, the warbling of the white-throat, the linnet, and the pine
grosbeak, the melody of the wood-wren, and the call of the cuckoo,
but hardly ever all these voices at once. The trilling call of the
greenshank and redshank becomes a song, the chattering of the
magpie gains a new charm, even the cawing of the hooded crow and
the raven seem cheerful, and the call of a woodpecker or a titmouse
most refreshing. The silence expresses the desolateness of the
woods. He who hopes to be able to lead in them a joyous
sportsman’s life will be bitterly disappointed. Doubtless all the
immense woods of this region have more tenants, especially birds
and mammals, than we are at first inclined to believe, but these
animals are so unequally distributed over the immeasurable area,
and probably also wander so widely, that we can arrive at no
standard for estimating their numbers. Miles and miles are, or
appear to be for a time at least, so lifeless and desolate that
naturalist and sportsman alike are almost driven to despair, their
expectations are so continually disappointed. All the reports of even
experienced observers who have sojourned there leave one still in
the dark. Districts which seem to combine all the conditions
necessary for the vigorous and comfortable life of certain species of
animals, shelter, to all appearance, not a single pair, not even a
male, naturally fond of roving. In such woods, far from human
settlements, and to some extent beyond the limits of human traffic,
one cannot but hope at length to fall in with the species which should
frequent such places, but the hope is as fallacious as the supposition
that one is more likely to meet with them in the heart of the forest
than on its outskirts. The fact is that the regions under man’s
influence, which he has modified, and to some extent cultivated,
seem often to exhibit a more abundant and diverse life than the
interior of the forest-wilderness. Wherever man has founded stable
settlements, rooted out trees, and laid out fields and pasture-lands,
there gradually arises a greater diversity of animal life than is to be
seen in the vast untouched regions which remain in their original
monotony. It seems as if many animals find suitable localities for
settlement only after the ground is brought under cultivation. Of
course the fact that certain animals are more abundant in the
neighbourhood of man, where they are ruthlessly hunted, than they
are in the inaccessible forest, where danger scarce threatens them,
implies a gradual reinforcement from without. At certain seasons at
least there must be migrations of more or less considerable extent,
and in these most of the West Siberian animals take part. All the
observations hitherto made corroborate this view.

Fig. 19.—Reindeer Flocking to Drink.

The only stationary animals, in the usual sense of the word, seem to
be certain mountain species and those which hibernate in caves and
holes; all the rest migrate more or less regularly. At the pairing and
nesting time all the West Siberian animals segregate themselves,
except those which are gregarious during the breeding season. Later
on, the parents and their young combine with their fellows in herds or
flocks, which, impelled by the need for more readily obtainable food,
and perhaps also driven off by the plague of mosquitoes, set out on
their wanderings together. Localities rich in fodder attract the
herbivorous creatures, which are the first to arrive, but on their heels
come others, and finally their enemies. Thus certain parts of the
woodland are depopulated and others are peopled, and there occur
actual blocks in the migratory stream, which must be the more
striking in contrast to the usual desolateness and emptiness of the
forest. The scenes of old conflagrations are favourite rendezvous,
for, on the fertilized soil, berry-bearing bushes of various sorts have
sprung up and attained luxuriant growth. Here, in autumn, the
Siberian herbivores find a rich harvest, and not only they, but wolves
and foxes, martens and gluttons, sables and bears, primarily
attracted by the collected herds, may be seen banqueting, devouring
the berries with evident pleasure. The different animals thus brought
together seem to remain for a time in a certain correlation. The
herbivores, as observant sportsmen have noticed, keep with
unmistakable constancy to the berries; and the carnivores follow
closely in their tracks.
These migrations explain how it is that during certain years some of
the woods are filled with all kinds of beasts of the chase, while during
other years they are entirely forsaken. The traveller from the west,
who journeys in late winter or early spring in Western Siberia,
beholds with astonishment a flock of three to five hundred black-
game rise in crowded flight from the highway through the forest, and
learns with not less astonishment a little later that the same or even
more favourable woods are but sparsely stocked with these birds. In
summer he searches in the most suitable localities for the hazel-
grouse, and is discouraged because his search is continually futile;
in autumn he is pleasantly surprised to see, in the same places,
abundance of the same game.
So peculiar are the conditions due to the monotonous uniformity of
wide stretches, that the huntsman who will make sure of his booty
must be very familiar with them; indeed, even the most skilful and
experienced sportsman is always and everywhere in the
measureless forests at the mercy of chance. Whatever be the game
he pursues, he never can predict where he will find it. Yesterday the
goddess of the chase was kind to extravagance; to-day she refuses
him every aid. There is no lack of game, but the huntsman who had
to live on what he shot would starve. A sportsman’s life, such as is
possible in other latitudes, is inconceivable in Western Siberia; the
profit to be derived from the forest chase is inconsiderable. Some
animals, for example the beaver,[31] seem already to have been
exterminated; and others, especially the much-prized sable, have
withdrawn from the inhabited districts into the interior of the forest.
Everywhere in Siberia one hears the common complaint, that game
becomes scarcer every year; and it is certain that from one
decennium to another the diminution is perceptible. For this man is
not wholly responsible; the forest-fires and the devastating
epidemics which now and then break out are probably as much, if
not more, to blame. At the same time, no Siberian ever realizes that
a temporary sparing of the game is the first condition of its
preservation. Sportsman-like hunting is unknown; the most varied
means are used to kill as many animals as possible. Gun and rifle
are mere accessories; pitfalls and nets, spring-guns and poison are
the most important agents employed by natives and immigrants
alike.
“Game” to the Siberian means every animal which he can in any way
use after its death, the elk and the flying squirrel, the tiger and the
weasel, the capercaillie and the magpie. What the superstition of one
race spares falls as a booty to the other; animals whose flesh the
Russians despise are delicacies to the Mongolian palate. Ostiaks
and Samoyedes take young foxes, martens, bears, owls, swans,
geese, and other creatures, treat them tenderly as long as they are
young, care for them sedulously until the fur or plumage is fully
developed, and then kill them, eating the flesh and selling the skin.
The number of skins brought from Siberia to the markets there and
in Europe is computed in millions: the number used in the country
itself is much smaller, but still very considerable. The quantity of
furred, and especially of feathered game, which is transported to a
distance in a frozen state, also mounts up to many hundreds of
thousands. Along with the furs of mammals the skins of certain birds
are at present much exported, especially those of swans, geese,