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 Acknowledgments vii

exploration of this field, while presenting some of the remarkable achieve-

ments of many wonderful researchers who have made this field one of the
great scientific adventures of our time.

Acknowledgments
I am most grateful to the many scientists who have helped to create the field
this book represents, and I regret that I didn’t have the space to represent more
of their great accomplishments. This revision also benefited greatly from inter-
actions with the many undergraduates who were enrolled in my course over
the past five years. Their questions and discussions motivated me to write
the Second Edition. I thank Professor Ryan Bachtell for serving as a sounding
board for some of the new material and his thoughtful input.
I thank Sydney Carroll, Editor, for her encouragement to undertake this
project and Sinauer Associates for supporting a Second Edition of this book.
The preparation of this book benefited greatly from the staff at Sinauer Associ-
ates. I would like to thank Chelsea Holabird for steering the book through the
production process, and Chris Small, Janice Holabird, and Elizabeth Morales
for their talented design ideas, book layout, and artwork.
Once again my wife, Julia A. Rudy, assumed major editorial responsibility
for the final product. Without Julie’s involvement, I cannot imagine how this
project would have been completed. Her editorial skills and commitment to
excellence are directly responsible for the book’s organizational clarity and
readability.

00_RUDY2E_FM.indd vii 10/18/13 12:43 PM

 viii About the Author

About the Author

Jerry W. Rudy is College Professor of Distinction in the Department
of Psychology and Neuroscience at the University of Colorado,
Boulder. He received his Ph.D. in psychology from the University
of Virginia in 1970, and joined the CU Boulder faculty in 1980.
The author of over 150 peer-reviewed research papers and book
chapters, Dr. Rudy has served on the editorial boards of the Journal
of Experimental Psychology: Animal Behavior Processes, Psychobiology,
Developmental Psychobiology (Editor in Chief), Behavioral Neuroscience,
Neuroscience & Biobehavioral Reviews, Learning and Memory, and
Jerry W. Rudy
Neurobiology of Learning and Memory (Associate Editor). He also
served on the governing board and as President of the International Society
for Developmental Psychobiology. He has received grant support from
the National Science Foundation, the National Institute of Mental Health,
and the National Institute of Health. Professor Rudy’s research interests
center on learning and memory processes. His research focused primarily
on understanding the complementary contributions the hippocampus
and neocortex make to learning and memory and the influence immune
products have on memory. He is currently the director of the undergraduate
neuroscience program at the University of Colorado, Boulder.

00_RUDY2E_FM.indd viii 10/18/13 12:43 PM

 Table of Contents

1 Introduction: Fundamental Concepts

and Historical Foundations 1
Learning and Memory Are Theoretical Concepts 2
Psychological and Neurobiological Approaches 3
Psychological Approach 3
Neurobiological Approach 5
Historical Influences: The Golden Age 6
Phenomena and Ideas 6
The Neuron Doctrine and Synaptic Plasticity 9
Behavioral Methods 11
Core Themes 13
Synaptic Plasticity 14
Molecules and Memory 14
Memory Systems 14
Summary 15
References 15

00_RUDY2E_FM.indd ix 10/18/13 12:43 PM

 x Contents

PART 1
Synaptic Basis of Memories 17

2 Mechanisms of Synaptic Plasticity:

Introduction 19
Two Approaches to Studying Synapses
that Support Memory 20
Simple System: The Gill Withdrawal Reflex 20
Long-Term Potentiation in the Hippocampus 24
The Conceptual Basis and Methodology of LTP 27
Understanding the Field EPSP 29
What is Synaptic Strength? 35
Inducing and Measuring LTP 36
Long-Term Depression: The Polar Opposite of LTP 39
Summary 40
References 40

3 Modifying Synapses:
Central Concepts 43
The Synapse as a Biochemical Factory 44
Postsynaptic Density 44
Other Synaptic Proteins 46
Signaling Cascades 46
First and Second Messengers 46
Protein Kinases and Phosphatases 47
Glutamate Receptors Are Critical to the Induction of LTP 48
LTP Induction Requires Both NMDA and AMPA Receptors 49
Two Events Open the NMDA Channel 51
Increasing AMPA Receptors Supports the Expression of LTP 53

00_RUDY2E_FM.indd x 10/18/13 12:43 PM

 Contents xi

An Organizing Framework: Three Principles 54

The Duration of LTP Can Vary 55
Molecular Processes Contribute to LTP Durability 56
Synapses Are Strengthened and Maintained in Stages 57
Summary 57
References 58

4 Generating and Stabilizing the Trace:

Post-Translation Processes 59
Generating the Trace 60
Dynamics of AMPA Receptor Membrane Trafficking 61
Dynamics of Actin Regulation 65
Generating the Trace: Summary 68
Stabilizing the Trace 68
Parallel Signaling Cascades Regulate Actin Dynamics 71
Cell Adhesion Molecules Help Stabilize the Trace 74
Summary 79
References 79

5 Consolidating Synaptic Changes:

Translation and Transcription 83
The De Novo Protein Synthesis Hypothesis 84
The Genomic Signaling Hypothesis 88
Synapse-to-Nucleus Signaling 88
Soma-to-Nucleus Signaling 90
Translating Protein Requires Increased Calcium Levels 92
Extracellular and Intracellular Sources of Calcium 94
Recruiting Multiple Calcium Sources Increases LTP Duration 97
Summary 99
References 100

00_RUDY2E_FM.indd xi 10/18/13 12:43 PM

 xii Contents

6 Consolidating Synaptic Changes:

Specific Mechanisms 103
Activation of Local Protein Synthesis 104
The mTOR–TOP Pathway 104
The BDNF–TrkB Receptor Pathway 104
Synthesis of a Key New Protein: Arc 106
Arc Antisense Blocks Long-Lasting LTP 106
BDNF-TrkB Consolidation Depends on Arc 107
Arc Sustains Actin Regulation 108
Confirming the Role of Actin Regulation 108
Contribution of Single-Spine Imaging Studies 109
Distinct Actin Pools Regulate Spine Growth 111
Targeting Plasticity Products 113
Protein Degradation and LTP 116
The Ubiquitin Proteasome System 116
Protein Degradation Influences Three Phases of LTP 117
Compartment-Specific Protein Degradation 120
Summary 121
References 123

7 Maintaining the Consolidated Trace 127

PKMζ: A Promising Maintenance Molecule 129
Inhibiting PKMζ Prevents LTP Maintenance 130
PKMζ Releases and Maintains GluA2 AMPA Receptors 132
Two Key Contributions of PKMζ 134
Trapping PKMζ 134
Genetic Engineering Reveals Additional
Maintenance Molecules 136

00_RUDY2E_FM.indd xii 10/18/13 12:43 PM

 Contents xiii

Summary 137
References 138

8 Toward a Synthesis 139

Generation 139
Stabilization 141
Consolidation 142
Maintenance 143
Issues and Implications 143
The Importance of Actin Regulation 143
Synaptic Plasticity and Memory 145
Summary 148
References 149

PART 2
Molecules and Memories 151

9 Making Memories:
Conceptual Issues and Methods 153
LTP and Memory 154
Behavior and Memory 154
Test Behavior: The Window to the Memory Trace 155
The Learning–Performance Distinction 157
Dimensions of Memory Traces 158
The Concept of Memory Consolidation 159

00_RUDY2E_FM.indd xiii 10/18/13 12:43 PM

 xiv Contents

Electroconvulsive Shock and Memory Disruption 160

Memory Disruption: A Storage or Retrieval Failure? 160
Some Behavioral Test Methods for Studying Memory 161
Inhibitory Avoidance Conditioning 162
Fear Conditioning 163
Spatial Learning in a Water-Escape Task 164
Why These Three Behavioral Test Methods? 167
Methods for Manipulating Brain Function 167
Stereotaxic Surgery 167
Genetic Engineering 169
Summary 172
References 173

10 Memory Formation:
Early Stages 175
NMDA Receptors and Memory Formation 176
Pharmacological Alteration 177
Genetic Engineering 177
Cautions and Caveats 182
AMPA Receptors and Memory Formation 184
Fear Conditioning Drives GluA1 AMPA Receptors into Spines 185
Preventing AMPA Receptor Trafficking Impairs Fear
Conditioning 185
Ampakines and Cognitive Enhancement 188
NMDA and AMPA Receptors: Acquisition and Retrieval 189
CaMKII and Memory Formation 191
Preventing Autophosphorylation of CaMKII Impairs Learning 192
CaMKII and Fear Memories 193
Actin Dynamics and Memory Formation 194

00_RUDY2E_FM.indd xiv 10/18/13 12:43 PM

 Contents xv

Working and Reference Memory Depend on Glutamate

Receptors 195
An Animal Model 196
Glutamate Receptor Composition Is Critical to Working
Memory 197
Summary 199
References 200

11 Memory Consolidation 205

The Research Paradigm 207
Transcription and Enduring Memories 208
The Importance of CREB 209
Some Memory Genes 213
Summary of Transcription 215
Translation and Enduring Memories 216
The De Novo Protein Synthesis Hypothesis 216
Methodological Issues 217
Multiple Rounds of Protein Synthesis 219
Protein Degradation Processes 226
Defining the Consolidation Window 227
Summary 228
References 229

12 Memory Maintenance
and Forgetting 233
PKMζ and Memory Maintenance 234
Interfering with PKMζ Erases a Taste-Aversion Memory 234
PKMζ Strengthens New Memories and Prevents Forgetting 235

00_RUDY2E_FM.indd xv 10/18/13 12:43 PM

 xvi Contents

PKMζ KO Mice Learn and Remember 235

Interpretive Caveat 236
Toward a Neurobiology of Forgetting 236
Summary 238
References 239

13 Memory Modulation Systems 241

Memory Modulation Framework 242
The Great Modulator: The Basolateral Amygdala 244
The Role of Epinephrine 246
The Epinephrine Vagus Connection 248
Norepinephrine Enhances Memories 251
Norepinephrine Enhances Glutamate Release
and Arc Translation 252
The Norepinephrine Signal in Other Storage Areas 254
The Epinephrine Liver–Glucose Connection 255
Bioenergenics and the Brain 255
Glucose Modulates Memory 255
Glucose and Aging 256
Glucose and Transcription 257
Glucocorticoids: The Other Adrenal Hormones 259
Summary 260
References 261

14 The Fate of Retrieved Memories 265

Reactivated Memory Disrupted by ECS 266
Active Trace Theory 266

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 Contents xvii

Reconsolidation Theory 268

Assessing Reconsolidation Theory 271
How Does Reactivation Destabilize the Trace? 272
Trace Restabilization and Trace Updating 275
Memory Erasure: A Potential Therapy 277
Preventing Drug Addiction Relapse 277
Eliminating Debilitating Fears 280
Stepping Back: Boundary Conditions 280
Summary 281
References 281

PART 3
Neural Systems and Memory 285

15 Memory Systems and

the Hippocampus 287
The Multiple Memory Systems Perspective 288
Example 1: Personal Facts and Emotions 288
Example 2: Personal Facts and Skills 289
The Case of Henry Molaison 290
The Episodic Memory System 292
The Animal Model Strategy 294
Studies of Patients with Selective Hippocampal Damage 296
The DNMS Paradox Resolved 298
Summary 299
References 299

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 xviii Contents

16 The Hippocampus Index

and Episodic Memory 303
Properties of Episodic Memory 304
Conscious Recollection and Contextual Information Storage 304
Automatic Capture of Episodic and Incidental Information 305
Single Episode Capture with Protection from Interference 305
Properties Summary 306
A Neural System that Supports Episodic Memory 306
The Hierarchy and the Loop 306
The MTH System 308
The Indexing Theory of Episodic Memory 309
The Hippocampus Does Not Store Content 311
Pattern Completion and Pattern Separation 311
Why Not Just Store the Memory in the Neocortex? 312
Indexing Theory and Properties of Episodic Memory 313
Evidence for the Indexing Theory 314
Animal Studies 315
Shining Light on The Index 321
Summary 323
References 324

17 The MTH System: Episodic Memory,

Semantic Memory, and Ribot’s Law 327
A Modular MTH System 329
Growing Up without the Hippocampus 329
Recognition Memory and MTH Modularity 331

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 Contents xix

The MTH System and Ribot’s Law 332

The Standard Model of Systems Consolidation 333
Challenges to the Standard Model 336
Multiple Trace Theory 339
Other Evidence Relevant to the Debate 340
Ribot’s Law Revisited: Summary 346
Summary 347
References 348

18 Actions, Habits, and the

Cortico-Striatal System 353
The Concept of Instrumental Behavior 354
Two Theories of Instrumental Behavior 356
Thorndike’s Law of Effect 356
Tolman’s Cognitive Expectancy Theory 356
Action and Habit Systems 358
With Practice, Actions Become Habits 361
A Conceptual Model for Actions and Habits 363
Action and Habit Systems Compete 364
Action Systems Are Vulnerable 365
A Cortico-Striatal System Supports Instrumental
Behavior 366
Neural Support for Actions 368
Neural Support for Habits 371
The Striatum Stores Action and Habit Memories 372
The Neural Basis of Rewarding Outcomes 374
Summary 376
References 376

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 xx Contents

19 Learning about Danger: The

Neurobiology of Fear Memories 379
The Fear System 380
The Neural Basis of Fear 382
Eliminating Dangerous Fears: Theories of Extinction 386
Neural Basis of Fear Extinction 389
The CS–noUS Neural Circuit 389
Why Fear Renews: A Role for the Hippocampus 389
Extinction Learning Depends on NMDA Receptors 391
New Insights: Extinction Can Erase Fear Memories 393
Extinction and Reconsolidation 394
Summary 395
References 396

Photo Credits 399

Glossary 401
Author Index 415
Subject Index 423

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 1 Introduction:
Fundamental Concepts
and Historical Foundations
Our uniqueness as human beings derives in large part from evolutionary
adaptations that permit experience to modify connections linking networks
of neurons in the brain. Information conveyed into the brain by our sensory
channels can leave a lasting impression on neural circuits. These networks not
only can be modified, the information contained in the modifications can be
preserved and later retrieved to influence our behavior. Our individual experi-
ences act on these networks to make us who we are. We have the ability to
learn a vast array of skills: we can become musicians, athletes, artisans, skilled
craftsmen, or cooks. Experience tunes our emotions to our environments. We
acquire food preferences and aversions. Incredibly, without intention, we also
lay down an autobiographical record of the events, times, and places in which
our experiences occur. We are connected with our past and can talk about it.
We learn and we remember.

01_RUDY2E_Chapter01.indd 1 10/18/13 11:24 AM

 2 Chapter 1

Historically, the study of learning and memory has been the domain of phi-
losophers and psychologists who have defined the relevant phenomena and
many of the important variables that influence them. Only recently have brain
scientists seriously weighed in on this topic. Armed with sophisticated meth-
ods to measure and manipulate brain processes and conceptual frameworks
to guide their application, neurobiologists have now made enormous inroads
into the mystery of how experience modifies the brain.
Consequently, an important field now exists called the neurobiology of
learning and memory. Scientists working in this field want to know how the
brain stores and retrieves information about our experiences. The goal of this
book is to present an account of some of the major accomplishments of this
field and to provide a background that will facilitate the understanding of
many of the issues and central assumptions that drive research in this field.

Learning and Memory Are Theoretical Concepts

The terms “learning” and “memory” are often used as if they are directly
observable entities, but they are not. Learning and memory are theoretical con-
cepts used to explain the fact that experience influences behavior (Figure 1.1). A
familiar example will suffice to make the point.
You have an exam tomorrow. So, over the next few hours you closet
yourself with your books and class notes. You take the test and answer the
questions to the best of your knowledge. Later you receive your grade, 90%.
Assuming that your grade would have been 50% if you had not studied,
then a reasonable person (the professor) would assume that you learned and
remembered the information needed to pass the test. The key phrase here is
“would assume.” Learning and memory were never directly observed. The
only directly observable events in this example are that (a) you spent time
with your notes and books, and (b) you took the test and performed well. That
you learned and remembered is inferred from your test performance and the
professor’s knowledge that you studied.

Learning and memory are theoretical concepts

Experience Learning–memory Behavior

observable inferred observable

Figure 1.1
Learning and memory are unobservable, inferred processes used to explain the fact
that our past experience influences our behavior.

01_RUDY2E_Chapter01.indd 2 10/18/13 11:24 AM

 Introduction 3

Larry Squire (1987) provided a useful definition of the terms learning

and memory: “Learning is the process of acquiring new information, while
memory refers to the persistence of learning in a state that can be revealed at
a later time” (p. 3). Other, more restrictive definitions have been proposed.
They usually also stipulate what learning and memory are not. For example,
a restricted definition would appropriately exclude fatigue, maturation, and
injury that might result from or be associated with experience.
Although learning and memory are theoretical concepts, neurobiologists
are motivated by the belief that they have a physical basis in the brain. A
slight modification of Squire’s definition provides a useful definition of the
field: the goal of neurobiologists working in this field is to understand how
the brain acquires, stores, and maintains representations of experience in
a state that permits the information contained in the representation to be
retrieved and influence behavior.

Psychological and Neurobiological Approaches

The study of learning and memory is the domain of both psychology and
neurobiology. It is useful to point out some fundamental differences between
the two approaches.

Psychological Approach
The general goal of psychology is to (a) derive a set of empirical principles
that describe how variation in experience influences behavior, and (b) pro-
vide a theoretical account that can explain the observed facts. The study of
memory became a science when Hermann Ebbinghaus developed the first
methods for assessing the acquisition and retention of a controlled experience.
He recognized that to study “pure memory” required a methodology that
could separate what the subject already has learned from what the
subject is now being asked to remember (Ebbinghaus, 1913). To do
this, he invented what are called nonsense syllables. A nonsense
syllable consists of a vowel placed between two consonants, such as
nuh, vag, or boc. These syllables were designed to be meaningless so
they would have to be learned without the benefit of prior knowl-
edge. Thus, for example, dog, cat, or cup would be excluded. Ebb-
inghaus made up hundreds of nonsense syllables and used them to
produce lists that were to be learned and remembered. Among the
task variables he manipulated were factors such as the number of
times a given list was presented during the memorization phase and
the interval between the learning and the test phase. Hermann Ebbinghaus

01_RUDY2E_Chapter01.indd 3 10/18/13 11:24 AM

 4 Chapter 1

Figure 1.2 100

Ebbinghaus documented the
first forgetting curve. Note that
substantial forgetting occurs in 75 nuh vag boc

Percent recalled
the first hour after learning, but
thereafter recall is fairly stable.
50

25

0
0 1 min 1h 24 h 5d 10 d 30 d
Retention interval

Ebbinghaus worked alone and was the only subject of his experiments.
He found that his test performance increased the more he practiced a given
list. He also documented the fact that retention performance was better when
he spaced the repetition of a given list than when the list was repeated with-
out inserting a break between the learning trials. He also documented the
first “forgetting curve.” As is illustrated in Figure 1.2, retention was excellent
when the test was given shortly after the learning trial, but it fell off dramati-
cally within the first hour. Remarkably, the curve stabilized thereafter.
Empirical principles such as those produced by Ebbinghaus’s experiments
led to theoretical questions about the underlying structure of the memory
(Figure 1.3). Consider Ebbinghaus’s forgetting curve. One could imagine that

Single trace Dual trace

Short-term trace
Trace strength
Trace strength

Long-term trace

Time Time
Figure 1.3
The single-trace theory explains Ebbinghaus’s forgetting curve by assuming that the
strength of a single memory trace declines monotonically as a function of time between
learning and the retention test. The dual-trace theory explains that the forgetting curve
results from two memory traces whose strength decays at different rates.

01_RUDY2E_Chapter01.indd 4 10/18/13 11:24 AM

Another random document with
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 At the end of the service the serjeant will give:—Firing party
attention. present—arms. slope—arms. volleys (with blank
cartridges)—load. present. fire. A second and third volley will be
fired. The serjeant will then give unload. fix—bayonets. slope—
arms. present—arms. slope—arms. form—fours. right. quick
—march.
The band will lead, followed by the firing party and mourners, who
will form fours as they march off without word of command.
Bayonets will be unfixed after leaving the place of burial at the
discretion of the officer in charge. The band is not to play, nor the
drums to beat, until the party is entirely clear of the burial ground.
2. Funerals of officers, warrant officers and serjeants will be
conducted in a similar manner to para. 1, the troops being detailed
for the escort according to the rank of the deceased. Officers’ swords
are reversed under the right arm, the point downwards, the hilt being
grasped with the right hand, blade uppermost, the left hand grasping
the blade behind the back.
3. Large escorts will march by companies, in fours, at the head of
the procession.
4. In the funerals of senior officers the arrangements will be made
under the supervision of the general officer commanding.
Note.—To fire three volleys over the grave the following
procedure will be adopted:—
When the men, after presenting arms, have sloped arms, the
commander will give the following commands:—
Volleys
The men will act as in the Feu-de-joie.
—Load.

Present. As in the Feu-de-joie.

Each man will at once press the trigger, and after

Fire.
a slight pause return to the position for loading.
 This will be repeated until the volleys are completed, when the
men will Unload as in Feu-de-joie.
 INDEX.
Ammunition, blank, 62
„ supply and transport of, in the field, 165
Annual course of training, auxiliary forces, 173
„ „ battalion, 179
„ „ company, 174
„ „ description of, 171
„ „ detail of, 172
„ „ general rules, 174
Assault, the, 137
Assembly, after dispersal, 65
Attack, the, 128
„ battalion in the holding, 152
„ by cavalry, meeting an, 92
„ „ or swordsmen, savage warfare, 119
„ company, general rules for, 145
„ co-operation between artillery and infantry in the, 135
„ decisive, 136
„ direction and pace in the, 143
„ distances in the, 142
„ distribution of companies in the decisive, 150
„ „ infantry in the, 128
„ duties of the several bodies in the, 131
„ feints in the, 139
„ fire in the, 133
„ firing-line and supports in the, 151
„ flank, 139
„ formations, 140
„ frontage in the, 141
Attack general rules for company in the, 145
„ „ battalion in the, 148
„ „ brigade in the, 153
„ „ division in the, 154
„ holding, 138
 „ infantry formations in regard to fire in the, 133
„ instruction for the battalion in the, 153
„ intervals in the, 143
„ local flanking fire, 140
„ machine guns in the, 144
„ nature of an offensive action, 128
„ orders, issues of, 130
„ „ for battalion in the, 149
„ practice, 85
„ preliminary measures, 129
„ rallying and redistribution, 138
„ reserve, 151
„ surprises, 140
Attack and defence, fire action, 126
„ fire discipline, 127
„ general principles, 123
„ position and duties of leaders in action, 125
„ ranges, 128
Attention, position of, 31
Auxiliary Forces, annual course of training, 173

Battalion advancing in review order, 196

Battalion in the attack, 148
„ distribution of companies in the decisive,
150
„ firing-line and supports, 151
„ general rules, 148
„ holding attack, 152
„ instruction, 153
„ orders, 149
„ reserve, 151
Battalion in the defence, general rules, 163
Battalion in the defence, duties of the commanding officer, 164
Battalion drill, 98
„ battalion in quarter column, 101
„ „ „ opening to column, 104
 „ „ column on the march closing to quarter
column, 106
„ „ column or quarter column advancing, in
fours, or vice versâ, 106
„ „ column changing direction, 108
Battalion formations, 100
„ general rules, 98
„ inspection of, 110
„ movements in quarter column, 103
„ movements in line or lines of company columns, 108
„ quarter column wheeling, 103
„ re-assembling, when dispersed, 109
Battalion drill, savage warfare, 113
„ „ battalion in line, 114
„ „ „ forming column or
quarter column, 116
„ „ echelon movements from line, column or quarter
column, 118
„ „ formations applicable to, 113
„ „ formation of squares, 120
„ „ general rules, 113
„ „ line advancing in column, 118
„ „ line changing position, from the halt, on a flank
company, 116
„ „ line forming column to right or left, 117
„ „ line passing obstacles or moving over broken
ground, 117
„ „ marching in line, 115
„ „ meeting attack by swordsmen, 119
„ „ quarter column or column deploying into line, 114
Battalion drill, savage warfare, quarter column on the march
forming line to a flank, 118
Battalion marching past, 192
„ review of, 189
„ training, annual course, 179
Blank ammunition, 62
Brigade in the attack, general rules, 153
 „ „ defence, general rules, 164
Brigade drill, 111
„ brigade wheeling from mass into line of battalions,
112
„ general rules, 111
„ marching in line of battalions either in column or
quarter column, 113
„ mass deploying into line of battalions and vice
versâ, 112
„ savage warfare, brigade in line advancing in
columns or quarter columns, 122
„ „ brigade forming square, 122
„ „ deployments, 122
„ „ marching in line, 122
Brigade advancing in review order, 202
„ marching past, 199
„ review of a, 196

Calls, signals, &c., 57

Casualties, skirmishing, 88
Challenging, manner of, 211
Changing direction (squad drill), 45
„ „ and forming up (squad drill), 50
„ „ as in file (squad drill), 46
„ ranks (squad drill), 49
„ „ (company drill), 76
„ step (squad drill), 39
Colour, trooping the, 213
Colours, presentation of, 220
Colours, the 188
Command, words of, how to be given, 30
Commanders, position of, in skirmishing, 88
„ instructions for, in skirmishing, 88
Commanding officer, duties of, in defence, 164
Company in the attack, general rules, 145
„ „ defence, duties of subalterns, section and squad
leaders, 162
 „ „ defence, duties of company leader, 163
„ „ „ general rules, 162
Company drill, 72
„ column changing direction, 80
„ company column forming company, 76
„ company in line forming to right or left, 78
„ company in line, or moving to a flank in fours,
forming company column, 78
„ dismissing, 83
„ drill in single rank, 81
„ fire action, 82
„ formation of fours, 76
„ forming up into line when marching, fours or file,
80
„ general rules, 72
„ inspection and telling off, 75
„ inspection in close order drill, 83
„ marching in line, 77
„ movements in line (or lines) of sections, 80
„ re-assembling when dispersed, 83
Company in line, saluting, 185
„ marching past, 185
„ sizing, 183
„ training, preliminary remarks, 67
Co-operation between artillery and infantry in the attack, 135
Counter attacks, decisive, 160
„ local, 160
Course of training, annual, auxiliary forces, 173
Course of training, annual, battalion, 179
„ „ company, 174
„ „ description of, 171
„ „ detail of, 172
„ „ general rules, 174
„ skirmishing, 54
Cover, use of, in skirmishing, 60

Defence, the, 155

 „ decisive counter attacks, 160
„ distribution of infantry for, 155
„ duties of officers commanding reserves, 159
„ entrenching a position, 157
„ fire in, 158
„ local counter attacks, 160
„ machine guns in, 158
„ occupation of the position, 156
„ position of the reserves, 159
Defence, infantry in attack and. See Attack and Defence.
Defence, the. See Company, Battalion, Brigade, Division.
Defence practice, skirmishing, 90
Defile, passage of, 53
Definitions, 16
Deployments, brigade drill, 122
Diagonal march, 49
Direction and pace in the attack, 143
Discipline, fire, 127
Dismissing, 53
Distances in the attack, 142
Distribution of companies in the decisive attack, 150
„ infantry in the attack, 128
Division in the attack, 154
„ defence, 164
Division drill, 111
Division, larger body than, marching past, 207
„ „ review of, 205
Division marching past, 204
„ review of, 202
Double march, the, 39
Dressing, when halted, 42
„ squad in two ranks, 48
„ „ with intervals, 32
Drill. See Squad, Company, Battalion, &c.
Duties of leaders in action, 125
„ officers commanding reserves, 159
„ scouts, 93
 „ skirmishers, 89
„ subalterns, section and squad leaders, 162
„ the company leader, 163
„ the commanding officer, 164
„ the several bodies in the attack, 131

Echelon movements from line, column or quarter column, 118

Entrenching tools, 169
„ a position, 157
Extending and closing, method of, 56

Feints, 139
Feu-de-joie, firing a, 221
Fire in the attack, 133
„ in the defence, 158
„ action, company, 82
„ „ in attack and defence, 126
„ discipline in attack and defence, 127
„ effect, 63
Flank attacks, 139
Flanking fire, local, 140
Formations applicable to savage warfare, 113
„ battalion drill, 100
„ in regard to fire, 133
„ in the attack, 140
„ to be practised, reviews, 188
Formation of squares, 120
„ squads, 31
„ „ in single rank, 41
„ „ in two rank, 48
„ fours, squad, 50
„ „ company, 76
Fours changing direction and forming squad, 53
Frontage in the attack, 141
Funerals, 223

General principles, attack and defence, 123

General remarks, skirmishing, 84
General rules, annual course of training, 174
„ ceremonial, 181
„ attack practice, 85
„ battalion drill, 98
„ brigade and division drill, 111
„ company drill, 72
„ savage warfare, 113
„ skirmishing, 54
„ company in attack, 145
„ battalion „ , 148
„ brigade „ , 153
„ division „ , 154
„ company in defence, 162
„ battalion „ , 163
„ brigade and division in defence, 164
Guards—
General rules, 209;
Guards of honour, 213;
Relieving or posting, 210;
Relieving and posting sentries, and marching reliefs, 210;
Sentries challenging, 211;
Trooping the Colour, 213;
Turning out at night, 212;

Halt, the, 37
Holding attacks, 138
„ attack, battalion, 152

Infantry formations in regard to fire, 133

Inspection of skirmishers, 65
„ in skirmishing, 95
„ of, and telling off the company, 75
„ in close order drill, 83
„ of a battalion, 110
Instruction of the recruit, method of, 29
„ for battalion in attack, 153
Instructions for commanders in attack, 88
„ training of the recruit, 25
Intervals in the attack, 143
„ skirmishing, 64
Issue of orders, the attack, 130

Length of pace and time in marching, 35

Line advancing in column, 118
„ battalion in, 114
„ brigade marching in, 122
„ brigade advancing in columns or quarter columns, 122
„ changing front, 116
„ company forming to right or left, 78
„ „ company column, 78
„ forming column or quarter column, 116
„ „ to the right or left, 117
„ -firing and supports, 151
„ marching in, company, 77
„ marching in, 115
„ of sections, movements in, 80
„ of company columns, movements in, 108
„ of battalions, brigade wheeling from mass into, 112
„ „ mass deploying into, 112
„ „ marching in, 113
Line passing obstacles or moving over broken ground, 117
„ quarter column or column deploying into, 114
Local counter attacks, 160
„ flanking fire, 140

Machine guns in the attack, 144

„ „ defence, 158
March, double, 39
„ diagonal, 43
„ „ (two ranks), 49
„ turning on the, 40
Marching as in file, 46
„ „ forming up into squad, 47
 „ in squad and file, 49
Marching past, company, 185
„ battalion, 192
„ brigade, 199
„ division, 204
„ larger body of troops than a division, 207
Marching, position in, 36
„ quick, 37
„ squad, 42
„ with arms, 47
Marking time, 38
Method of extending and closing, 56
„ instruction, special training of non-commissioned
officers, 179
„ „ of the recruit, 29
Movements in line of sections, 80
„ quarter column, 103
„ line or lines of company columns, 108

Non-commissioned officers, special training of, 179

Obstacles, 53
Occupation of a position, 156
Orders, battalion in attack, 149
„ issue of, in the attack, 130
„ reporting and passing, 64

Pace and time in marching, 35

Passage of a defile, 53
Passing orders, 64
Plates, key to, 15
Preliminary measures for the attack, 129
„ remarks, training of the recruit, 21
„ „ company training, 67
„ training, skirmishing, 55
Presentation of Colours, 220

Rallying and redistribution, 138

Ranges, 128
Reconnaissance practice, 91
Recruit, method of instruction of, 29
„ training of the, 21
Reinforcing, 64
Relieving or posting a guard, 210
„ and posting sentries and marching reliefs, 210
Remarks, general, on skirmishing, 84
„ preliminary, on the training of the recruit, 21
„ „ on company training, 67
Reporting and passing orders, 64
Reserve, the, 151
„ position of, in the defence, 159
„ duties of officers commanding the, 159
Retirements, 91
Review of a battalion, 189
„ of a brigade, 196
„ of a division, 202
„ of any larger body of troops than a division, 205
„ Royal, 208
„ order, battalion advancing in, 196
Review order, brigade advancing in, 202
Rules. See General Rules.

Saluting, company in line, 185

Savage warfare, formations applicable to—
Battalion in line, 114
„ forming column or quarter column, 116
Echelon movements from line, column or quarter column, 118
Formation of squares, 120
General rules, 113
Line advancing in column, 118
„ changing front from the halt on a flank company, 116
„ forming column to the right or left, or vice versâ, 117
„ passing obstacles, or moving over broken ground, 117
Marching in line, 115
Meeting an attack by cavalry or swordsmen, 119
 Quarter column or column deploying into line, 114
„ on the march forming line to a flank, 118
Section leaders, duties of, 162
Sentries, relieving and posting, 210
„ challenging, 211
Shelter trench exercise, 95
Single rank, company drill, 81
Skirmishing—
Assembly after dispersal, 65
Blank ammunition, 62
Course of training, 54
Cover, 60
Fire effect, 63
General rules, 54
Inspection of skirmishers, 65
Intervals, 64
Method of extending and closing, 56
Practical skirmishing, 59
Preliminary training, 55
Reinforcing, 64
Reporting and passing orders, 64
Signals, whistle blasts, calls, 57
Company training—
Casualties, 88
Commanders, instructions for, 88
„ position of, 88
Defence practice, 90
General remarks, 84
„ rules, 85
Meeting attack by cavalry, 92
Reconnaissance practice, 91
Retirements, 91
Scouting, 93
Scouts, duties of, 93
„ training of, 94
Shelter trench exercise, 95
Skirmishers, duties of, 89
 Skirmishing, inspection in, 95
Squad drill—
Method of instruction of the recruit, 29
Words of command, 30
With intervals—
Dressing, 32
Formation, 31
Position of attention, 31
Saluting, 34
Standing at ease, 32
Turnings, 33
Marching—
Changing step, 39
Double march, 39
Drum and pace stick, 36
Halt, 37
Length of pace and time, 35
Marking time, 38
Position in marching, 36
Quick march, 37
Side step, 40
Stepping back, 38
„ short, 39
„ out, 37
Turning on the march, 40
Single rank—
Changing direction, 45
„ as in file, 46
Diagonal march, 43
Dressing when halted, 42
Formation, 41
Marching as in file, forming up into squad, 47
„ „ 46
„ in squad, 42
„ with arms, 47
Turnings, 42
In two ranks—
 Changing direction and forming up, 50
„ ranks, 49
Diagonal march, 49
Dismissing with or without arms, 53
Dressing, 48
Formation, 48
„ of fours, 50
Fours changing direction and forming squad, 53
Marching in squad and in file, 49
Obstacles, 53
Passage of a defile, 53
Squad leaders, duties of, 162
Square, brigade forming, 122
„ forming, 120
Subalterns, duties of, company in defence, 162
Surprises, 140
Supply of ammunition in the field, 165
Syllabus of training (recruit), 21

Training, course of, skirmishing, 54

„ preliminary, skirmishing, 55
„ of scouts, 94
„ of non-commissioned officers, 179
„ annual course. See Annual Course.
„ recruits, 21
„ syllabus of recruits, 22
Transport of ammunition in the field, 165
Turnings. See Squad Drill.
LONDON:
PRINTED FOR HIS MAJESTY’S STATIONERY OFFICE,
BY HARRISON AND SONS, ST. MARTIN’S LANE,
PRINTERS IN ORDINARY TO HIS MAJESTY.

(Wt. w 27691 150,000 2 | 05—H & S 5749)

 FOOTNOTES:
[1] In marching order without the rifle the arms will be retained
as in the position of Attention.
[2] The number of paces to which men are to extend is to be
communicated by word of mouth.
[3] This signal denotes extension from the centre. If the
extension is to be made to the right, finish the signal by pointing
to the right. If the extension is to be made to the left, finish the
signal by pointing to the left.
[4] This signal denotes “Close on the centre.” If it is desired to
close on the right, finish the signal by pointing to the right. If the
close is to be on the left point to the left.
Note.—All signals should be made with whichever arm will
show most clearly what is meant.
[5] Rifle regiments will drill in close order at the Trail. On the
command quick (or double) march, the men will Trail arms
without further word of command, and will halt and stand at ease
on the word or signal Halt.
[6] When a company without arms is inspected, section
commanders will close up the rear rank on completion of
inspection of their sections, giving the command close ranks.
march.
[7] When constructing trenches by night, tracing tapes are used
to show the line of the proposed trench.
[8] With tripod equipment, the machine gun detachment will
include two pack animals, each of which will carry 7 boxes of
ammunition (250 rounds to the box).
[9] Recruits, as soon as their training as such is completed, are
to take their place in the ranks irrespective of the season of the
year.
[10] In the following sections dealing with ceremonial, including
guard duties, the command to slope arms and fix bayonets will
not apply to men of Rifle Regiments, who will remain at the Order
 when halted, will present arms from the Order without fixing
bayonets, and will march past at the Trail.
[11] The term “corps,” when used with reference to troops, is
intended to designate troops which are not allotted to any
particular field unit.
[12] In coming to the Engage from the Slope, seize the rifle
with the right hand at the small and come to the Engage.

Transcriber’s Notes:

1. Obvious printers’, punctuation and spelling errors have been corrected

silently.

2. Where hyphenation is in doubt, it has been retained as in the original.

3. Some hyphenated and non-hyphenated versions of the same words have

been retained as in the original.
 *** END OF THE PROJECT GUTENBERG EBOOK INFANTRY
TRAINING, 1905 ***

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