Principles of Exercise Neuroscience

Dawson J. Kidgell
Visit to download the full and correct content document:
https://textbookfull.com/product/principles-of-exercise-neuroscience-dawson-j-kidgell/
More products digital (pdf, epub, mobi) instant
download maybe you interests ...

Exercise-cognition interaction : neuroscience

perspectives 1st Edition Mcmorris

https://textbookfull.com/product/exercise-cognition-interaction-
neuroscience-perspectives-1st-edition-mcmorris/

Nutrition for sport and exercise Andrew J. Doyle

https://textbookfull.com/product/nutrition-for-sport-and-
exercise-andrew-j-doyle/

Exercise, sport, and bioanalytical chemistry :

principles and practice 1st Edition Hackney

https://textbookfull.com/product/exercise-sport-and-
bioanalytical-chemistry-principles-and-practice-1st-edition-
hackney/

Nutrition for Sport and Exercise, 4th Edition J. Andrew

Doyle

https://textbookfull.com/product/nutrition-for-sport-and-
exercise-4th-edition-j-andrew-doyle/
Principles of Managerial Finance Gitman Lawrence J

https://textbookfull.com/product/principles-of-managerial-
finance-gitman-lawrence-j/

Drugs and the Neuroscience of Behavior: An Introduction

to Psychopharmacology Adam J Prus

https://textbookfull.com/product/drugs-and-the-neuroscience-of-
behavior-an-introduction-to-psychopharmacology-adam-j-prus/

Electromagnetic Waves and Antennas, Exercise Book 1st

Edition Sophocles J. Orfanidis

https://textbookfull.com/product/electromagnetic-waves-and-
antennas-exercise-book-1st-edition-sophocles-j-orfanidis/

Principles of Chemistry A Molecular Approach Nivaldo J.

Tro

https://textbookfull.com/product/principles-of-chemistry-a-
molecular-approach-nivaldo-j-tro/

Fundamentals of Brain and Behavior : An Introduction to

Human Neuroscience 1st Edition William J. Ray

https://textbookfull.com/product/fundamentals-of-brain-and-
behavior-an-introduction-to-human-neuroscience-1st-edition-
william-j-ray/
Principles of Exercise
Neuroscience
Principles of Exercise
Neuroscience
Edited by

Dawson J. Kidgell and Alan J. Pearce

Principles of Exercise Neuroscience

Edited by Dawson J. Kidgell and Alan J. Pearce

This book first published 2020

Cambridge Scholars Publishing

Lady Stephenson Library, Newcastle upon Tyne, NE6 2PA, UK

British Library Cataloguing in Publication Data

A catalogue record for this book is available from the British Library

Copyright © 2020 by Dawson J. Kidgell and Alan J. Pearce and

contributors

All rights for this book reserved. No part of this book may be reproduced,
stored in a retrieval system, or transmitted, in any form or by any means,
electronic, mechanical, photocopying, recording or otherwise, without
the prior permission of the copyright owner.

ISBN (10): 1-5275-5813-4

ISBN (13): 978-1-5275-5813-7
 TABLE OF CONTENTS

Preface ...................................................................................................... xii

Dawson J. Kidgell, PhD and Alan J. Pearce, PhD

Acknowledgements ................................................................................. xiv

Chapter 1 .................................................................................................... 1
The Nexus between Neuroscience and the Science of Exercise
Alan J. Pearce, PhD
1. Background ....................................................................................... 1
1.1 The neuroscience of human movement........................................... 2
1.2 What is ‘motor control’? How does motor control relate within
the larger discipline of exercise science? ........................................ 4
1.3 Motor learning and skill acquisition: similarities and contrasts
to motor control ............................................................................... 6
1.4 The challenge of translating neuroscience to exercise science ....... 8
1.5 How to get the best from this book ................................................. 8
References............................................................................................. 9

Chapter 2 .................................................................................................. 10
Levels of Motor Control
Dawson J. Kidgell, PhD and Alan J. Pearce, PhD
2. Background ..................................................................................... 10
2.1 Structural arrangement of the brain contributing to movement .... 11
2.1.1 Structure of the cerebral cortex ............................................ 11
2.1.2 The brain stem ...................................................................... 12
2.2 Transmission of cortical motor signals ......................................... 13
2.3 Motor functions of the cerebellum ................................................ 13
2.3.1 Anatomical and functional organisation of the cerebellum .... 14
2.3.2 Afferent and efferent pathways of the cerebellum ............... 15
2.4 Motor functions of the spinal cord ................................................ 16
2.4.1 Organisation of the spinal cord............................................. 17
2.5 Types of reflex pathways .............................................................. 18
2.5.1 Skeletal muscle reflexes ....................................................... 19
2.6 Hierarchical organisation of motor control ................................... 21
vi Table of Contents

2.7 Motor units, fibre types, and recruitment physiology ................... 23

2.7.1 The motor unit ...................................................................... 23
2.7.2 Physiological classification and recruitment ........................ 24
2.8 Principle of motor unit recruitment ............................................... 25
2.9 Motor unit synchrony .................................................................... 26
2.9.1 Quantifying the degree of motor unit synchrony .................. 27
2.9.2 Motor unit synchronization and human motor control ......... 27
2.9.3 Motor unit synchronization in upper and lower limb
muscles .................................................................................... 28
2.10 Summary ..................................................................................... 29
References........................................................................................... 29

Chapter 3 .................................................................................................. 32
Techniques Contributing to the Understanding of Neuroscience
in Exercise
Alan J. Pearce, PhD and Dawson J. Kidgell, PhD
3. Background ..................................................................................... 32
3.1 Electroencephalography (EEG) .................................................... 33
3.2. Magnetoencephalography (MEG) ................................................ 34
3.3 Transcranial magnetic stimulation (TMS) .................................... 35
3.3.1 Single-pulse TMS ................................................................. 36
3.3.2 Paired-pulse TMS ................................................................. 38
3.4 Voluntary activation and neural drive ........................................... 40
3.4.1 Interpolated twitch and VATMS ............................................. 40
3.4.2 H-reflex ................................................................................ 42
3.4.3 V-wave ................................................................................. 44
3.5 Summary ....................................................................................... 45
References........................................................................................... 46

Chapter 4 .................................................................................................. 53
Principles of Neuroplasticity in Exercise
Dawson J. Kidgell, PhD and Ashlyn K. Frazer, PhD
4. Neuroplasticity ................................................................................ 53
4.1 Mechanisms of neuroplasticity ..................................................... 53
4.2 Short and Long-term potentiation ................................................. 54
4.2.1 NMDA receptor activation and synaptic plasticity .............. 54
4.2.2 Brain-derived neurotrophic factor and neuroplasticity ......... 55
4.3 Experimentally-induced neuroplasticity ....................................... 56
4.3.1 Transcranial direct current stimulation and neuroplasticity .... 57
4.3.2 NIBS and functional connectivity ........................................ 58
 Principles of Exercise Neuroscience vii

4.4 Is the induction of neuroplasticity via NIBS important for motor

performance? ................................................................................. 60
4.4.1 Is the magnitude of neuroplasticity and motor performance
improvement related to the BDNF polymorphism?................. 63
4.5 Is homeostatic plasticity important for motor performance? ........ 64
4.6 Use-dependent neuroplasticity ...................................................... 67
4.7 Summary ....................................................................................... 68
References........................................................................................... 69

Chapter 5 .................................................................................................. 76
Non-invasive Brain Stimulation and Exercise Performance
Shapour Jaberzadeh, PhD and Maryam Zoghi, PhD
5. Introduction ..................................................................................... 76
5.1 Transcranial Direct Current Stimulation ....................................... 77
5.2 The mechanisms behind tDCS effects during stimulation
(online effects) .............................................................................. 78
5.3 The mechanisms behind tDCS effects after the termination
of stimulation ................................................................................ 79
5.4 Montages for application of tDCS: The conventional montage .... 80
5.4.1 HD-tDCS montage ............................................................... 80
5.4.2 Other tDCS montages........................................................... 81
5.5 tDCS as a stand-alone technique ................................................... 82
5.6 tDCS as a priming technique ........................................................ 82
5.7 Halo sport tDCS device ................................................................ 82
5.8 The effects of tDCS on EP ............................................................ 83
5.9 Ethical considerations for the use of tDCS for enhancement
of EP .............................................................................................. 85
5.10 Summary ..................................................................................... 86
References........................................................................................... 86

Chapter 6 .................................................................................................. 92
Neural Control of Lengthening and Shortening Contractions
Jamie Tallent, PhD and Glyn Howatson, PhD
6. Background ..................................................................................... 92
6.1 Shortening and Lengthening Contractions .................................... 92
6.1.1 Benefits of lengthening contractions .................................... 93
6.2 Motor control of lengthening and shortening muscle contractions . 93
6.2.1 Muscle .................................................................................. 94
6.2.2 Spinal.................................................................................... 94
6.2.3 Cortico-Spinal ...................................................................... 95
6.3 Adaptations to shortening and lengthening resistance training ..... 97
viii Table of Contents

6.4 Summary ..................................................................................... 100

References......................................................................................... 101

Chapter 7 ................................................................................................ 107

Neural Adaptations to Strength Training
Dawson J. Kidgell, PhD
7. Background ................................................................................... 107
7.1 Acute neural responses to strength training ................................ 107
7.2 Using TMS to assess the neural responses to strength training ..... 108
7.3 MEPs are acutely facilitated following a strength training
session ......................................................................................... 109
7.4 Intracortical facilitation is acutely enhanced following a strength
training session ............................................................................ 110
7.5 Why does strength training increase corticospinal excitability
and intracortical facilitation of the motor cortex? ....................... 111
7.6 Long-term neuroplastic adaptations to strength training ............. 114
7.7 Changes in strength following 2-8 weeks of strength training .... 116
7.8 Long-term strength training does not affect motor threshold or
MEP amplitude............................................................................ 117
7.9 Long-term strength training reduces motor cortex mediated
inhibition ..................................................................................... 118
7.10 Changes in spinal cord plasticity with strength training ........... 120
7.11 Changes in H-reflex and V-wave amplitude following strength
training ........................................................................................ 120
7.12 Changes in motor unit activity following strength training ...... 122
7.12.1 Single motor unit behaviour following strength training.. 123
7.12.2 Motor unit synchronization following strength training ... 124
7.13 Summary ................................................................................... 125
References......................................................................................... 125

Chapter 8 ................................................................................................ 133

Neuromuscular Responses to Fatiguing Locomotor Exercise
Callum Brownstein, PhD and Kevin Thomas, PhD
8. Background ................................................................................... 133
8.1 The role of exercise intensity and duration on neuromuscular
responses to fatiguing exercise .................................................... 135
8.2 Neuromuscular responses to “all-out” exercise .......................... 136
8.3 Neuromuscular responses to severe intensity, short-duration
exercise........................................................................................ 139
8.4 Neuromuscular responses to sustained exercise below critical
power ........................................................................................... 141
 Principles of Exercise Neuroscience ix

8.5 Neuromuscular responses to high-intensity intermittent

exercise........................................................................................ 144
8.6 The effect of exercise modality on neuromuscular responses
to locomotor exercise .................................................................. 146
8.7 Effect of exercise duration and intensity on recovery ................. 148
8.8 Origin of prolonged impairments in contractile function ............ 151
8.9 Origin of prolonged impairments in voluntary activation ........... 152
8.10 Summary ................................................................................... 153
References......................................................................................... 153

Chapter 9 ................................................................................................ 160

Sex Differences in Neuromuscular Function and Fatigability
Paul Ansdell, PhD and Stuart Goodall, PhD
9. Introduction ................................................................................... 160
9.1. A brief history of the scientific study of sex and performance .. 160
9.2. Sex differences throughout the motor pathway.......................... 161
9.2.1. Pre-motor processes .......................................................... 162
9.2.2. Intracortical and corticospinal neurons.............................. 163
9.2.3. Motor unit properties ......................................................... 164
9.2.4. Contractile apparatus ......................................................... 165
9.3. Functional neuromuscular sex differences ................................. 165
9.3.1. Maximal force production ................................................. 165
9.3.2. Force steadiness and accuracy ........................................... 166
9.3.3. Fatigability ........................................................................ 166
9.4. Female-specific neuromuscular function ................................... 169
9.4.1. The influence of hormones in vitro ................................... 169
9.4.2. In-vitro evidence ............................................................... 169
9.4.3. Functional changes across the menstrual cycle ................. 170
9.5. Summary .................................................................................... 173
9.5.1. What do we know already? ............................................... 173
9.5.2. Where do we go from here? .............................................. 174
References......................................................................................... 174

Chapter 10 .............................................................................................. 185

Motor Control Responses following Exercise-Induced Muscle Damage
Carlos Hermano Pinheiro, PhD and Alan J. Pearce, PhD
10. Background ................................................................................. 185
10.1 How does DOMS occur? .......................................................... 186
10.2 Functional outcomes affected by DOMS .................................. 188
10.3 Neuromuscular and motor control changes following DOMS .. 189
10.4 Neurophysiological studies in DOMS ...................................... 191
x Table of Contents

10.5 Summary ................................................................................... 192

References......................................................................................... 192

Chapter 11 .............................................................................................. 196

Neuromuscular Alterations and Motor Performance in Healthy Aging
Jakob Škarabot, PhD
11. Background ................................................................................. 196
11.1 Alterations in the central nervous system with advancing age .... 197
11.1.1 Alterations at the level of motor unit ................................ 197
11.1.2 The influence of synaptic inputs from spinal and
supraspinal centres ................................................................. 197
11.1.3 Reflex inputs to motoneurons........................................... 198
11.1.4 Cortical inputs to motoneurons ........................................ 199
11.2 Reduction in motor performance in healthy aging adults ......... 200
11.2.1 Motor performance during maximal contractions ............ 200
11.2.2 Control of muscle force output during submaximal tasks .. 204
11.2.3 Fatigability ....................................................................... 206
11.3 Summary ................................................................................... 208
References......................................................................................... 209

Chapter 12 .............................................................................................. 222

Cross-education
Ashlyn K. Frazer, PhD and Dawson J. Kidgell, PhD
12. Background ................................................................................. 222
12.1 Evidence of cross-education ..................................................... 222
12.2 Exercise prescription parameters .............................................. 223
12.3 Mechanisms of cross-education ................................................ 225
12.4 Interventions to enhance the cross-education effect.................. 228
12.4.1 Transcranial direct current stimulation and cross-
education................................................................................ 228
12.4.2 Electromyostimulation during cross-education ................ 229
12.4.3 Whole-body vibration (WBV) training and cross-
education ................................................................................ 230
12.5 Cross-education and neuromuscular injury ............................... 231
12.6 Summary ................................................................................... 232
References......................................................................................... 232
 Principles of Exercise Neuroscience xi

Chapter 13 .............................................................................................. 237

Using Electrophysiology to Understand Responses following Concussion
and Mild Brain Injury
Alan J. Pearce, PhD and Michael E. Buckland, MBBS PhD
13. Background ................................................................................. 237
13.1 The definition of concussion ..................................................... 238
13.2 Current standard to diagnose concussion .................................. 239
13.3 Objective measures to assess concussion .................................. 239
13.4 Electrophysiology to assess concussion .................................... 240
13.5 Electroencephalography (EEG) ERPs in concussion research .. 241
13.5.1 Visual and vestibular evoked potentials ........................... 242
13.6 Transcranial Magnetic Stimulation (TMS) EPs and concussion
research ....................................................................................... 243
13.6.1 Studies in acute concussion .............................................. 244
13.6.2 Studies in retired athletes with history of concussion
and head trauma ..................................................................... 245
13.6.3 Studies in persistent post concussion symptoms .............. 246
13.7 Combined electrophysiological modalities to assess
neurophysiology .......................................................................... 246
13.9 Summary ................................................................................... 248
References......................................................................................... 249
 PREFACE

DAWSON J. KIDGELL, PHD

AND ALAN J. PEARCE, PHD

Over the last 30 years, there has been a significant rise in research and
teaching interest that has examined the neuromuscular responses to different
types of exercise interventions. Two fields of research, neuroscience and
exercise science, have merged to provide evidence for how the nervous
system responds to exercise. An important component of exercise science,
but one area that is less recognised, although used ubiquitously through
other topics within exercise science, is the study of motor control. Motor
control is primarily concerned with the neurophysiological mechanisms that
contribute to human movement. Motor control is at the very heart of
exercise and sport science. Motor control drives how we program
appropriate exercise in health, injury and disease. Motor control is what
underpins this book we call Principles of Exercise Neuroscience.

This textbook introduces the key concepts that emphasise human motor
control and its application to exercise science and rehabilitation. The topics
covered integrate research, theory and the clinical applications of Exercise
Neuroscience that will support students, researchers and clinicians to
understand how the nervous system responds, or adapts to physical activity,
training, rehabilitation and disease. Exercise Neuroscience uses a mix of
neuromuscular physiology, electrophysiology and muscle physiology to
provide a synthesis of current knowledge and research in the field of
exercise neuroscience that specifically examines the effects of exercise
training, injury and rehabilitation on the human nervous system.

It is never easy writing a textbook like this from scratch. Indeed, this
textbook came about from the initial teaching notes to assist tertiary students
with some difficult learning concepts within the motor control subjects
taught at our respective universities. As time progressed, the lecture notes
developed into full chapters that eventually evolved into this textbook.
Further, through our international collaborative network, we were able to
draw upon international experts who were very happy to contribute and
provide their extensive knowledge and experience to develop our initial
 Principles of Exercise Neuroscience xiii

chapters into state of the art reviews. Our guest authors are leaders in their
respective areas of enquiry and provide the latest research findings in three
key areas: principles of motor control; neuroscience of exercise
performance; and clinical exercise neuroscience (injury and neuro-
rehabilitation). We sincerely thank each of them for their time and efforts
in helping us complete this book.

This is the first text devoted solely to the emerging topic of Exercise
Neuroscience. It aims to assist readers in identifying current research
findings and provide new avenues to explore the benefits of exercise on the
human nervous system. We thoroughly enjoyed writing this and we hope
that you find this text useful for your learning and professional development.

Dawson J. Kidgell, Monash University, Melbourne, Australia.

Alan J. Pearce, La Trobe University, Melbourne, Australia
June, 2020
 ACKNOWLEDGEMENTS

A text such as Principles of Exercise Neuroscience is not the effort of just

two authors. This text brings together the contributions of many research
scientists who have examined the changes that occur in the human nervous
system. We would like to especially thank Dr Eric Frazer for his assistance
in providing suggestions for revisions to this book, in particular his attention
to detail and endless editing. We would also like to acknowledge the
contributions of all the authors, in particular Professor Glyn Howatson, who
has established and leads a remarkable program of Applied Neuromuscular
Research at Northumbria University; many former PhD students from this
laboratory have contributed to this text.
 CHAPTER 1

THE NEXUS BETWEEN NEUROSCIENCE

AND THE SCIENCE OF EXERCISE

ALAN J. PEARCE, PHD

1. Background
The past thirty years have seen a significant rise in the disciplines of
neuroscience and exercise science. However, within the last ten years we
have seen these two disciplines become intersected. While neuroscience has
been an area of research for over a century, it is only in the last 20-30 years
that neuroscience has transformed from an interdisciplinary speciality under
the umbrella of neurology, psychiatry and psychology, to a stand-alone
discipline, and is now the fifth largest field of study in the sciences (Rosvall
and Bergstrom, 2010). Particularly in the last decade, with advancements in
technologies such as neuroimaging and electrophysiology, neuroscience has
captured the imagination of the wider community. In particular, the concept
of neuroplasticity, where evidence has demonstrated that the brain has the
ability to reorganise and readapt over the lifespan, has ignited a huge interest
across many unrelated fields. This has, of course, been both positive, with
advances in our understanding of the brain and nervous system, but also
negative where we are seeing the rise in what some call “neurobabble”, the
phenomenon whereby neuroscientific explanations of cognitive and motor
behaviour are more persuasive simply because they sound more technical
and authoritative (Varazzani, 2017). Despite this, continued advances in
neuroscience research will improve the lives of everyday people, as well as
contributing to improved athletic performance.

In parallel, exercise science has also emerged from the discipline of physical
education. Since the early 1990s, when the first exercise science courses
were beginning to be established in Australia, the growth in exercise science
has been so rapid that physical education can now be regarded as one of the
areas under the broader discipline of exercise science (along with exercise
physiology, strength and conditioning, biomechanics, nutrition, functional
2 Chapter 1

anatomy, skill acquisition and psychosocial determinants of health). Indeed,

many exercise science graduates will continue their learning, pursuing
postgraduate studies in education and developing courses in sport and
exercise science in secondary schools.

It should be no surprise then that the maturing disciplines of exercise science

and neuroscience have overlapped. Significant interest and growth in
research and application have occurred in the role of exercise on brain health,
neural function and neuroplasticity. Conversely, greater understanding of
clinical neurological disease has impacted on advancing exercise
programming, with studies testing the efficacy of various exercise
interventions on brain and neural integrity. Taken together, the
understanding of human motor control occurs through exploring and
investigating neural and neuromuscular function.

1.1 The neuroscience of human movement.

It is important to note that neuroscience has many different areas of research
and study. Notwithstanding animal models of neuroscience, just focusing
on the neuroscience of human nervous systems can traverse a wide
continuum of study areas (Figure 1.1).

Figure 1.1. Continuum of general areas of study within neuroscience

It is not uncommon that neuroscientists from different research areas will

not necessarily understand each other’s work. For example, there are those
who investigate the neuroscience of a single cell under physiological or
pharmacological interventions. Conversely, there are others who are
interested in neuroimaging of the healthy or diseased brain. In between, we
have various levels of neuroscientific inquiry that can overlap but are
specific sub-disciplines in their own right. Further, like exercise neuroscience
or neurophysiology, various other combined disciplines exist such as
neuroanatomy, neurochemistry, neuropsychology and the like (see Table
1.1 for a list with definitions).
 The Nexus between Neuroscience and the Science of Exercise 3

Table 1.1. List of combined areas of study within neuroscience.

Neuroanatomy Study of anatomy of neurons and neural
structures
Neurochemistry Study of the compounds that generate
physiological functioning in and between neurons
Neuroendocrinology Study of the interaction between the endocrine
and nervous systems
Neurodermatology Study of the neural structures within the skin
contributing to not only neural health but also
sensory reception.
Neurogenetics Study of the role of genetics in the development
and function of the nervous system
Neuroimmunology Study of the interaction between the immune and
nervous systems
Neurooncology Study of cancers of the brain and nervous system
Neuroopthamology Study of the interaction between the visual system
and nervous system
Neurotology Study of the neural contribution to vestibular
function
Neurophysiology Study of the functioning of the nervous system
Neuropsychology Study of the nervous systems influence in a
person’s cognition and behaviour

Of course, as this book will be focusing on aspects of exercise science and

human movement, the aim will be to synthesize and translate the growing
research output specific to human motor control. The result is that this text
narrows itself to the neural basis of human movement. Pathways, structures,
neurophysiological principles and application of these principles are
discussed but always in the context of human function and movement.

The scientific study of the neuroscience of movement is termed motor

behaviour. Motor behaviour is an umbrella term that encompasses a range
of sub-disciplines that overlap:

Motor control – the study of the underlying neurophysiological mechanisms

that contribute to movement;

Motor learning and skill acquisition – understanding the optimal

environments to practice and refine specific, purposeful movements
progressing the individual towards an accomplished level of expertise; and

Motor development – the study of movement across the lifespan; how

children develop and learn fundamental motor skills, through to
4 Chapter 1

understanding the mechanisms by which older adults decline in motor

control.

1.2 What is ‘motor control’? How does motor control

relate within the larger discipline of exercise science?
As teachers in motor control for over 15 years, this is usually the first
question that pops up from students. This is because motor control, as a term,
is rather indistinct. Indeed, as the term ‘motor control’ is becoming used
more widely, it has also been used and taken on slightly different meanings
to different professionals within the exercise and sport science domain. For
example, biomechanists discuss motor control in terms of physics being
applied on, or from within, the body during proficient performance of a
movement. Sport psychologists reference motor control in terms of actions
having an emotional or cognitive basis, for example, the use of motor
imagery to assist with rehabilitation of function or to improve on sports
skills. Physiotherapists discuss motor control in the rehabilitation setting,
with patients focusing on reducing “abnormal” movement and increasing
movement patterns that are less likely to cause injury (or re-injury).
Exercise physiologists may discuss the reduction in motor control
performance when an individual experiences fatigue. Strength and
conditioning and personal trainers need to consider motor control issues
when prescribing correct technique and resistance, to optimise strength
improvements and reduce likelihood of injury.

However, the core of motor control centres on the neuroscience that

underpins neuromuscular movement of the human body. It goes without
saying that people’s lives are filled with movement (Latash, 2008). As
undergraduates studying exercise science and related fields, human
movement is the core focus: using movement to improve strength and
muscle mass; using movement to improve skills; using movement to help
people lose weight; and using movement to assist people back to functional
daily activities.

Undergraduate study in exercise science will have motor control embedded

within course content across a number of subjects, or as a ‘stand-alone’
course. As exercise neuroscientists, we may have a bias here (who doesn’t!)
but it is quite easy to illustrate how elements of motor control permeate
throughout the curriculum of the major or core subjects areas taught in
exercise science courses (Table 1.2).
 The Nexus between Neuroscience and the Science of Exercise 5

Table 1.2. Interrelationship between motor control and other areas within
the discipline of exercise science.

Study area Relationship to Example

motor control
Anatomy (neuroanatomy) Study of the Damage to primary
structure and motor cortex leading
function of the to motor dysfunction.
brain and nervous
system.
Biomechanics/movement Study of efficient Relationship of
analysis voluntary sequential limb
movement. movement for a
specific motor task.
Exercise/sport physiology Study of The role of the brain
biochemistry and in fatiguing exercise.
physiological
systems adapting
to exercise and
sport.
Psychology of exercise and Study of Mood disorder
sport psychological influencing loss in
influences in motivation to
determining exercise.
exercise behaviour, Motor imagery to
as well as improve movement
psychological traits efficiency in sport.
required in high
performance sport.
Nutrition Study of nutrients Gut/brain interaction,
in food, how the how food intake can
body uses affect movement
nutrients, and the (interrelationship
relationship with exercise
between diet, physiology).
health, and disease.
Strength and conditioning Study of methods Neural adaptations
to induce with strength or
resistance for resistance training
training modalities.
adaptations.
6 Chapter 1

So, where does this lead us in terms of establishing what motor control
actually is? The common denominator in all these areas is the interaction of
the brain and muscle to provide meaningful (Latash, 2008) goal-directed
movement (Enoka and Stuart, 2005). Therefore, it can be argued that motor
control has a basis in neuroscience. Motor control is concerned with the
neurophysiological processes that allow movements to become more
efficient (which may also include improvement in skills, movement
efficiency, and strength). Similarly, motor control investigates how
movements can be retrained after an injury. These scenarios now have a
well-described term, neuroplasticity (Doidge, 2007), which will be
discussed later and throughout the text.

1.3 Motor learning and skill acquisition:

similarities and contrasts to motor control.
Historically, physical education courses have focussed on the understanding
of the learning and teaching of motor skills (or motor learning). For example,
understanding the development of fundamental motor skills (FMS) such as
running, jumping, and throwing are important when teaching children these
lifelong skills. These fundamental skills are the foundation for more advanced
motor skills, particularly specific skills for sporting excellence. Teaching
motor skills effectively is ingrained within the cognitive/psychological
domain, whereby the teacher is creating an optimal learning environment.
However, with the increased interest in the study of motor development (see
section 1.4), exploration of FMS now sits within the development and
changes of skills and motor actions across the lifespan, providing motor
learning and skill acquisition the opportunity to have its own definition,
succinct from motor control and motor development.

Figure 1.2 addresses not only similarities, but also differences, between the
sub-disciplines of motor control and motor learning. Motor learning, it can
be suggested, is aimed at how people learn and acquire expertise at motor
skills. What are the environments that best optimise learning? For example,
should athletes practice the same movement action in an unchanging setting
repeatedly in a series of blocks to reinforce motor pathways? Or, should
athletes randomise the environment with the intention of learning a
particular skill but retaining the motor memory? Is it better for individuals
to learn implicitly through discovery, or have an external source (such as a
teacher or instructor) providing feedback on how their attempt was executed?
In reality, motor learning and skill acquisition are part of memory formation
and, therefore, it can be argued that this sits within the domain of cognitive
 The Nexus between Neuroscience and the Science of Exercise 7

psychology. Motor learning is creating motor memory, similar to other

forms of learning (e.g., content knowledge) that leads to memory retention.

Figure 1.2. Differences and overlap between motor control and motor learning/skill
acquisition.

Motor development focuses on the stages and underlying processes that

allow children to gain motor skills. Traditionally, within the domain of
developmental psychology, the study of motor development has taken a
‘back seat’ compared to other developmental science areas such as cognitive,
social, language, personality, perceptual and emotional development (Adolf
and Robinson, 2015). However, there has been a resurgence of interest in
studying motor development due to its advantage in direct observation and
translation to the psychological sciences. Moreover, with increased
understanding of neuroplasticity, the study of motor development allows for
the understanding of neurophysiological processes, providing a foundation
for how adults learn new motor skills and retain motor memory.

Whilst motor development is primarily focussed in children and adolescents,

it is also important to understand changes as people age, and the
consequences of these. Our society is ageing, with significant increases in
the number of people who will be aged 65 and older in the coming decades.
Understanding the neuromuscular and motor control changes with ageing is
important for current students, particularly in thinking of career opportunities
8 Chapter 1

upon graduating. As part of the Allied Health sector, exercise science

students have a unique set of skills allowing application of their
neuroscience knowledge to, for example, exercise programming for older
people, or biomechanics to understand how to prevent falls. These will be
key areas in the near future to which exercise science can make a unique
contribution (if not already).

1.4 The challenge of translating neuroscience

to exercise science
So, why does neuroscience cause such difficulty to the majority of exercise
science students? The simple reason is because the neuromuscular system
is complex. It has a number of levels contributing to meaningful movement
that work at the conscious and subconscious levels, spinal and supraspinal
levels, and between sensory interactions with the motor response. Further,
it also may not necessarily have anything to do with exercise. Motor control
is as concerned with how an individual can create a pincer grip to control a
pen during writing, as it is concerned with how a person can increase
strength without muscle hypertrophy, or how the central nervous system
controls movement as it also responds to fatiguing exercise. It is also
concerned about how control can be lost, through degeneration, or how
control is lost after a brain injury and, in some cases, how it can be regained.

With the links between neuroscience and exercise becoming closer, it is

important that exercise scientists have a competent understanding of the
brain, and how it controls the neuromuscular pathways. Moreover, it is
important that exercise scientists who may practice in related fields are able
to constructively critique new developments in the area, and to confidently
determine good neuroscience from neurobabble. With unprecedented
access to news and information on a daily basis and the rise of pop-science,
it is important that, as practitioners in the field, you will be able to advise
your clients/patients what has been demonstrated using the scientific
method, and what has not been supported (despite comprehensive spin). The
contents in this book are based upon evidence-based research; however,
each chapter aims to translate the findings allowing students to learn new
concepts confidently.

1.5 How to get the best from this book

The topics in this book have come from our teaching of motor control to
students over the last 15 years. While the topics can and do overlap, each
 The Nexus between Neuroscience and the Science of Exercise 9

chapter can be read independently and in any order. For some, all chapters
will be of interest. For others, specific chapters will be helpful for personal
and professional development. As the chapters are research based, our aim
is to guide students through each chapter with the most robust concepts, but
also some insights into future areas of enquiry. For some students, this text
may serve as a pathway to a career as an exercise science research student,
as much as it may inform students moving towards a practitioner career.
Either way, we hope that students find this book engaging, but also
challenging, in attaining future knowledge through scientific enquiry.

References
1. Adolph KE, Robinson SR. Motor Development, 2015.
2. Doidge N. The Brain That Changes Itself: Stories of Personal
Triumph from the Frontiers of Brain Science. 2007.
3. Enoka RM, Stuart DG. The contribution of neuroscience to exercise
studies. 1985.
4. Latash ML. Neurophysiological Basis of Movement. 2008.
5. Rosvall M, Bergstrom CT. Mapping change in large networks. 2010.
6. Varazzani C. The risks of ignoring the brain. 2017.
 CHAPTER 2

LEVELS OF MOTOR CONTROL

DAWSON J. KIDGELL, PHD

AND ALAN J. PEARCE, PHD

2. Background
Understanding motor control is through learning the macro and the micro
arrangement of the nervous system. In this chapter, we will focus on the
macro: structural, functional and hierarchical organisation of movement. In
chapters four and five, we will discuss how motor control occurs through
neuromodulation and neuroplastic changes. The study of neuroscience
dictates that we need to learn the structural, functional and hierarchical
anatomical arrangement of the nervous system in order for us to appreciate
how individuals improve their movement, whether it be for sport, music,
jobs requiring fine skills or, alternatively, how patients relearn and regain
movement after an injury.

The chapter will start with the basics: the major structures identified in the
central nervous system. Where appropriate, and in keeping with the
objective of this textbook, we will focus examples and discussion on how
the region contributes to movement and motor control. Bear in mind which
cognitive functions, such as attention and decision-making, can and do
influence motor output. From there, we will outline the functional
arrangement of motor control. Here, we will focus on topographic
arrangement of motor functions and, in later chapters, how their
arrangement allows for adaptation and re-organisation. Finally, we will
discuss the hierarchical levels of motor control where we will incorporate
structural and functional understanding on how movement is planned and
executed.
 Levels of Motor Control 11

2.1 Structural arrangement of the brain contributing

to movement
In simplistic terms, the brain can be divided into three parts: the cerebrum,
brain stem, and the cerebellum. Each of these structures makes important
contributions to the regulation of movement.

The cerebrum is the large ‘dome’ of the brain that is divided into right and
left cerebral hemispheres. The most superficial layer of the cerebrum is the
cerebral cortex, which is composed of tightly arranged neurons. The cortex
performs three important motor functions: (1) the organization of complex
movements; (2) the storage of learned experiences; and (3) the reception of
sensory information. For the purpose of this chapter, we will limit our
discussion to the role of the cortex in the organization of movement.
However, to appreciate this, some information about the structure of the
cerebral cortex is required.

2.1.1 Structure of the cerebral cortex

The cerebral cortex is well endowed with neurons, neuroglia, and blood
vessels. The structural organisation of the three types of cells that populate
the cortex, being pyramidal cells, stellate neurons and fusiform neurons,
enables the classification of the cortex into three types: allocortex,
mesocortex and the neocortex. The allocortex is the oldest region and is
composed of only three layers and is located in the limbic system. The
mesocortex is younger and is composed of three to six layers and is
predominantly located in the insula and cingulate gyrus. The neocortex is
the youngest region of the cortex and is composed of six layers that
comprise the bulk of the cerebral cortex (Rothwell 1994). Whilst the
cerebral cortex is structurally organised into layers, it also has organisation
through functional connections. Most corticospinal output is mediated
through pyramidal neurons and stellate (or granule) cells. The cellular
organisation of pyramidal and stellate cells within the cortex gives it a
characteristic layered or laminar appearance that can be identified as six
distinct layers:

I. Molecular layer. A layer lying immediately inferior to the pia matter and
containing very few cell bodies.

II. External granular layer. A layer of densely packed small cells

including small pyramidal and stellate cells.
12 Chapter 2

III. External pyramidal layer. A layer consisting of medium to large-sized

pyramidal cells.

IV. Internal granular layer. This layer is predominantly composed of

densely packed stellate and pyramidal cells.

V. Ganglionic layer. This layer contains large pyramidal cells (Betz cells).

VI. Multiform layer. This layer is relatively thin and mostly composed of
densely packed, spindle-shaped cells, many with axons leaving the cortex.

Layer 1 contains mainly long horizontal dendrites and axons in deeper

layers as well as thalamic afferents. The medium to large pyramidal cells in
layers III and V contain long cortico-cortical connections. In addition,
pyramidal cells in layer V also project to sub-cortical areas such as the basal
ganglia, brain stem and to the spinal cord. Layers II and IV receive afferent
inputs, with inputs from the thalamus generally terminating in layer IV.
Layer VI contains small pyramidal cells which exhibit greater morphologic
variability than the pyramidal cells located in other layers and have cortico-
cortical and cortico-thalamic projections. Layers II through VI all contain
stellate cells, some of which form excitatory connections onto pyramidal
cells and some of which form inhibitory synapses.

2.1.2 The brain stem

The brainstem is located inside the base of the skull just superior to the
spinal cord and is essentially an extension of the spinal cord. It is composed
of a series of complicated neural tracts and nuclei (groups of neurons). The
major structures of the brain stem include the medulla, pons and midbrain.
In addition to these structures, a complex neuronal network known as the
reticular formation resides in the brainstem. The brainstem performs motor
and sensory functions for the face and head (i.e., cranial nerves) along with
providing support to the body against gravity. For example, muscles of the
spinal column and the extensor muscles of the legs maintain the body
against gravity. These muscles are under the control of specific brainstem
nuclei, in particular the pontine reticular nuclei, which excite activity in the
antigravity muscles, and the medullary reticular nuclei, which inhibit the
activity of antigravity muscles. Importantly, the medullary reticular nuclei
receive collateral input from the corticospinal pathway, rubrospinal
pathway, and other motor pathways. These systems can activate the
inhibitory action of the medullary reticular nuclei and counterbalance the
signals from the pons.
 Levels of Motor Control 13

2.2 Transmission of cortical motor signals

The spinal cord is under the control of a number of neurons that descend
from the primary motor cortex (M1). The largest of these are the
corticospinal neurons that have their origins in layer V of the cerebral cortex
and extend to form the bulk of the corticospinal or pyramidal tract (Porter
1985). Although corticospinal neurons are located within six cortical
regions, the M1 has the largest concentration (Porter, 1985). Within the M1,
these corticospinal neurons are functionally organised to project to
motoneurons that control specific muscle groups (He et al. 1993), thus they
are somatotopically organised. Corticospinal neurons that arise within the
M1 descend through the internal capsule, brainstem, and medulla oblongata
and continue to descend in the dorsolateral funiculi of the spinal cord
(Alawieh et al. 2017). As the corticospinal neurons leave the M1 and
descend to the medulla, they are organised somatotopically. At the
medullary spinal junction, approximately 85-90% of the corticospinal
neurons cross the midline to form the motor pyramidal decussation
(Alawieh et al. 2017), where they continue as the dorsolateral funiculi of the
spinal cord and converge onto motoneurons within the ventral horn of the
spinal cord that innervate limb muscles (Alawieh et al. 2017). Anatomical
mapping studies reveal that the connectivity of the corticospinal tract
suggests that the remaining uncrossed ipsilateral corticospinal tract fibres
descend primarily in the dorsolateral lateral or ventral funiculi of the spinal
cord (Alawieh et al. 2017).

A small proportion of corticospinal tract fibres do not crossover at the

pyramidal decussation at the medulla; rather, they project to ipsilateral
spinal motoneurons, where they could alter the excitability of ipsilateral
pathways (Porter and Lemon 1993, Carson 2005). In the clinical
neurophysiology literature, it has been suggested that increased utilisation
of the ipsilateral pathway may provide a viable method for re-establishing
motor control of upper-limb muscles following lesions to the M1 (Alawieh
et al. 2017, see Chapter 11 for more detail). There is now good evidence for
a functional role of the corticospinal system in the control of the upper limb
and changes in its functional organisation following different forms of
exercise training (Frazer et al. 2018).

2.3 Motor functions of the cerebellum

For a long time, the cerebellum has been referred to as the silent area of the
brain mainly because electrical stimulation of the cerebellum does not
14 Chapter 2

evoke any conscious sensation or movement. However, intriguingly,

removal of the cerebellum causes highly abnormal human movement.
Although it is difficult to study the function of the cerebellum in a conscious
person, it is generally accepted that the cerebellum is vital for rapid motor
actions which include cycling, running, playing the piano and talking.
Because the cerebellum does not cause muscle contraction per se, its main
function is to assist in the sequencing of motor actions and to monitor and
correct movements whilst they are being executed. The ability to adjust
motor sequences during movements enables the M1 and other parts of the
brain to ensure that the intended movements are performed correctly.

The cerebellum continuously receives updated information about the

intended/desired sequence of muscle activity from specific motor control
centres of the cortex. It receives sensory information from peripheral
receptors such as muscle spindles and Golgi tendon organs, thus allowing
corrective adjustments to be sent to the motor system to increase or decrease
the level of muscle activation of specific muscle groups. A critical function
of the cerebellum is to assist the cerebral cortex to plan the next series of
sequential movements in advance whilst the initial movement is still being
performed. This essentially enables movements to transition smoothly from
one movement to the next. The cerebellum also learns to correct movement,
via special cerebellar nuclei (the deep cerebellar nuclei) that can adjust
motor output. The cerebellum is able to perform these functions because of
its anatomical and functional organisation.

2.3.1 Anatomical and functional organisation of the cerebellum

The cerebellum is divided into three lobes: 1) anterior lobe, 2) posterior lobe
and 3) flocculonodular lobe and two deep cerebellar fissures. The cerebellar
hemispheres are separated along the longitudinal axis by a narrow band
called the vermis. The vermis plays an important role in controlling muscle
activity of the axial body, shoulders and head. To either side of the vermis,
are cerebellar hemispheres, whereby each hemisphere divides into an
intermediate and lateral zone. The intermediate zone controls motion of
distal portions of upper and lower limbs especially the hands and feet, whilst
the lateral zone controls sequencing movements of muscles including
timing and coordination of muscle activity. The vermis and intermediate
zones contain a topographical representation of the body, similar to that
observed in the sensory cortex, motor cortex, basal ganglia, red nucleus and
the reticular formation. The topographical regions of the vermis and lateral
zones receive afferent projections from all respective parts of the body, as
well as from corresponding topographical motor areas from the cerebral
 Levels of Motor Control 15

cortex. The lateral zone of the cerebellar hemisphere is not topographically

organized, rather, it receives input signals from the cerebral cortex that
enable the cerebellum to assist the motor pathways to organise and plan
movement.

2.3.2 Afferent and efferent pathways of the cerebellum

Several neural tracts from the cerebral cortex to the cerebellum regulate
human movement. For example, the corticopontocerebellar pathway
originates from motor and premotor areas of the cerebral cortex and
somatosensory cortex and send their axons to mainly the lateral zone of the
cerebellar hemisphere. In addition, neural tracts from the brainstem, such as
the olivocerebellar tract, vestibulocerebellar tract (vestibular apparatus),
and reticulocerebellar tract, terminate predominately in the vermis. The
dorsal spinocerebellar tract transmits information mostly from muscle
spindles but also from Golgi tendon organs, tactile, and joint receptors in
the periphery to the cerebellum. Thus, it appraises the brain of the
momentary status of muscle contraction, muscle tension and limb position
and forces acting on the body surface. The other afferent pathway is the
ventral spinocerebellar tract that is mainly excited by motor signals that
arrive at the ventral horn of the spinal cord from the brain via the
corticospinal tract and from the internal motor pattern generators of the
spinal cord itself.

The main efferent pathways that exit the cerebellum include the
fastigioreticular tract and the cerebellothalamocortical tract, along with the
cerebellar nuclei. The fastigioreticular tract works in close association with
the equilibrium apparatus and brainstem nuclei to control balance and works
with the reticular formation of the brainstem to control postural stability.
The cerebellothalamocortical tract is extremely complex and passes
through many neuronal structures to finally terminate at the level of the
motor cortex, whereby it coordinates the reciprocal contractions of agonist
and antagonist muscles in the limbs; thus, it is involved in the turning ‘on’
and ‘off’ of muscle activity.

In addition to the efferent neural pathways located deep in the cerebellum,

on each side are three deep cerebellar nuclei, the dentate, interposed and
fastigial nuclei. The deep cerebellar nuclei all receive input signals from the
cerebellar cortex and the deep sensory afferent tracts to the cerebellum. The
primary source of motor output from the cerebellum is via the deep nuclear
cells. Every time an input signal is sent to the cerebellum, the afferent
signals go directly to the deep cerebellar nuclei and to the corresponding
Another random document with
no related content on Scribd:
He leaned back and clasped his hands behind his head and smiled.
“What is it?” asked Bonnie May anxiously.
“I’m afraid I couldn’t explain to you. I was just thinking about—about
certain forms of reconstruction.”
 CHAPTER XVI
MRS. THORNBURG REVEALS A SECRET

Baron shook his head slowly. He had been thinking about that
advertisement in the Times which Thornburg had answered without
any result.
“Strange,” he mused. “I won’t believe but that somebody is looking
for her—somewhere. Children like that are not dropped down and
deserted like superfluous kittens or puppies. There’s something
wrong somewhere.”
Then he remembered that Mrs. Thornburg wished to see him; that,
according to Thornburg, she had “mentioned Bonnie May.”
Possibly she knew something. At any rate, Baron felt that he ought
to call on her. It was just after the dinner-hour—of the day on which
Mrs. Baron had announced her policy of reconstruction—and the
evening was flinging a challenge to all mankind to get out of doors
and enjoy the spring air.
He took up his stick and hat and left the house.

He found Mrs. Thornburg sadly changed since he had seen her last.
She was unmistakably very ill, though the only symptoms revealed to
Baron’s inexpert eye were a pathetic thinness and pallor and a
profound lassitude.
She was alone, Thornburg having just gone out.
“It was good of you to come,” she said when Baron entered. She
spoke as if she had been expecting him. And without circumlocution
she continued: “I wanted to talk to you about the little girl. You
haven’t let anybody have her, have you?”
“No,” replied Baron. Then he added lightly: “I think we’ve changed
our minds about letting her go. It seems likely now that we’ll keep her
with us indefinitely.”
He was glad that her glance rested upon her thin, clasped hands. He
could note the effect of his statement with a steady scrutiny which
need cause him no compunction.
To his surprise she seemed quite pleased. “It makes me glad to
know that she is to be with nice people,” she said, lifting to him now
a softly grateful glance. She explained: “You see, I’m sure I’m too ill
to have her now, even if....” Her lips trembled and her eyes filled.
“But you’ll be better,” said Baron, reading her thought. Clearly she
had despaired of ever being any better. “When you’re able to have
her, she’ll be so happy to visit you. I mean Bonnie May. She’s a
wonderfully sociable little creature. If she were invited to come to see
you she would be delighted. Attentions like that—such as you would
pay to grown people—have a wonderful effect upon her.”
“Yes.... And of course some day she will be coming here to stay.”
“You mean—” Baron was surprised that his suggestion had been
received with a dully uttered, enigmatic remark, rather than gratitude
or eagerness.
“You don’t know what I mean by that?” There was regret in her tone,
reluctance in her glance—as if she knew he was not dealing
honestly and frankly by her.
“No, truly, I don’t.”
“Ah, well.... But I wanted to tell you why I was so eager to have her
when you called before. You see, I wanted to—to atone....”
She sat listlessly, lost in troubled memories, and Baron waited.
“Mr. Thornburg came to me one time, in the one moment of his
greatest need, and asked me to help him. And I failed him.”
She leaned back and closed her eyes for a moment, and Baron
thought how out of harmony she was: the ailing woman whose whole
being was in a minor key, amid surroundings which suggested only
sturdiness and well-being.
“He was always generous toward me, and patient. He was always
giving, giving, and never asking. I think I got used to that and just
took it for granted. And then one day he came home, excited, as
happy as a child ... and asked me.... It was such a little thing ... and I
refused.
“You know, he had been married when I first met him. An actress. It
didn’t last long. She got tired of the life and wanted to go back to the
stage. I think she appealed to his generosity. It would have been
easy to do that. At any rate, he allowed her to go away and take their
little girl. I can’t understand how he brought himself to let the little
daughter go, too. I have an idea he was so troubled because she
wanted to go that he didn’t realize how much the child meant to him,
or would come to mean. She was only a year old then. I never
blamed him for that episode in his life. I just concluded that the
woman was worthless. And when I married him we didn’t speak of
his other marriage—nothing in connection with it. It was just as if it
hadn’t happened. Then, after a year, or about a year, he—he made
the one request of me. The mother had offered to give him the little
girl. He wanted to bring her to me, to have her in our home.
“And that made me jealous and unhappy. I can’t explain ... or defend
myself. I could scarcely answer him when he spoke about it. And
when I didn’t answer he looked at me, and after a little a strange
expression came into his eyes. He was chilled and bewildered. He
had been so happy. He couldn’t understand. He just gave it up, and
the next day he was trying to pretend that nothing had come
between us; that I hadn’t been ungracious and cruel.
“You see, I was thinking of her child, and he was thinking of his own.
Mine was the woman’s—the narrow—point of view, and his was the
father’s. Maybe you can understand a little of what I felt. I couldn’t
have the child here in the house, while its own mother.... It would
have been like giving her a place in our home—the woman, I mean.
You can’t really separate people by putting their bodies in different
places. You see what I mean?”
“Yes,” assented Baron, “I think I see quite clearly.”
“And I was sure she was a bad woman. And I felt that if her child
were in the house, her—her real self would be here, too. Her
influence, I mean. Bodies are not everything. Sometimes they’re
even the least things of all. I was afraid that other woman’s very
presence would be here among us on the most sacred occasions: at
bedtime, to see if her child were covered up, and in the early hours
of Christmas morning, jealously looking to see what we’d given her,
and jealous of us, because we were fond of her. She would be a real
influence in the house. It couldn’t be helped.”
“But a bad woman.... Surely a bad woman would forget,” suggested
Baron.
“Well, not our kind of a woman, anyway. How could she have
deserted a man who was good to her? And how could she consent
to give up her child afterward? It might be right for her to leave her
husband; but for a mother to give up a little daughter.... No, I couldn’t
think of having here in our home a link to bind us with a woman like
that—a life out in the unknown, on the streets that are strange to us,
that are strange to all faithful, happy people.
“And then when it was too late I began to see his side of it. He was
the father just as much as she was the mother. She was his child as
much as hers—more, if he loved her more. And I began to realize
what it must be to a father to have his little daughter away from him,
perhaps not loved and provided for, possibly facing an evil future.
Oh, the night that thought came to me! And always he was so kind to
me, and patient. He did not speak of his daughter again. And I
waited.... I knew he would speak again some day, and I wanted to
grow strong enough to say to him honestly: ‘Ah, do bring her, and
she shall have love here, here in her own home’....”
She lifted her hands to her cheeks and closed her eyes. It was as if
she must shut out some of the impressions which crowded into her
mind.
Baron waited until a measure of calm came upon her. “And—he
never did?”
She opened her eyes and regarded him inquiringly.
“I mean, he never spoke of her again?”
She regarded him with a smouldering look in her eyes. Then she
leaned forward, her hands gripping the arms of her chair. “I honestly
believe you don’t know!” she whispered.
And in an instant she had taken from a little box on the table near
which she sat an envelope. She drew from it a single sheet and
passed it to Baron.
He turned a little, so that the light from the table fell upon it and read:
“Do be good to the little girl your husband has brought to you. You
ought to be, because he is her father.”
There was no name. Baron handed the sheet back to her. He was
thinking hard. “Who could have written it?” he asked.
“Of course you realize that I don’t know,” she replied. “Do you mean
to ask me what I think?”
“Well, what do you think?”
“I think her mother wrote it. I think she must have lost track of the
child, and concluded that Mr. Thornburg had taken her. I think she
must have known of my—my jealousy on that other occasion. I think
she wrote this note hoping that I would refuse to have the child in the
house if I knew who she was. It seems plain that she wants her
now.”
Baron was examining the date of the postmark on the envelope. She
saw that furrows were gathering on his forehead.
She explained: “It came some time ago. I had it with me here when
you called that first time.”
“Oh!” exclaimed Baron. “And you knew, then——”
“Yes, I knew then.”
“But you haven’t.... Mr. Thornburg....”
“I didn’t show him this. He doesn’t know. Surely you can understand.
He has acted a lie, in trying to get the little girl into the house without
telling me about her. And I can’t blame him for that, after what
happened that other time. But I can’t bear to let him know that—that I
know.”
“But don’t you see, if Bonnie May is really his daughter, and if he
weren’t afraid to tell you so, he could bring her here without any
further hinderance!”
“No, he couldn’t. Not if the mother wants her.”
Baron arose. “After all, it’s largely guesswork—conclusions reached
in the dark,” he said. “You’ve received an anonymous note. That’s all
the foundation you have for what you’ve told me. And people who
write anonymous letters....”
He reflected dubiously, and then he came to a decision.
“I’ve reason to believe,” he said, “that there is good ground for you to
reject what’s in that note.”
She leaned forward, observing him intently.
Baron was remembering the actress who had called on Thornburg;
the woman who, almost certainly, was she who had taken the child
into Thornburg’s theatre. He was recalling his question to the
manager, and the latter’s vehement, prompt response.
“You mean,” questioned Mrs. Thornburg, “that you don’t think Bonnie
May is really ... that you don’t believe it was her mother who wrote
this note?”
“It’s difficult to be quite sure of anything,” said Baron, “but I would
stake a great deal on that one thing being true—that it wasn’t Bonnie
May’s mother who wrote that anonymous note.”
 CHAPTER XVII
“A KIND OF DUEL”

That night in his attic room Baron arrived, by perfectly logical

reasoning, at two conclusions, each of which was precisely the
opposite of the other.
The first of these conclusions was that he had a perfect right to
shape Bonnie May’s future according to his own inclination. The
second was that he had no right at all to do such a thing.
He arrived at the first conclusion in this manner:
He had made an honest effort to locate any person or persons
having a legal and just claim on the child, and he had failed. If the
Thornburgs had any claim upon her, it was not his fault that they had
bungled their affairs until they were unwilling to make their claim
public.
Therefore he had a right to have and to hold Bonnie May, and to
regard her, if not as his own, at least as a permanent member of the
household.
His second and contrary opinion began to shape itself when he
recalled the picture of Mrs. Thornburg, helpless and despairing,
greatly desiring the presence of the child in her own home in order
that she might complete a great moral victory over herself.
A man couldn’t oppose his claims and advantages to a need like
that!
Besides—it was borne in upon Baron more and more strongly—
there was a very serious question as to the child’s best interests.
She was an actress, born and bred, and some day she would surely
hear the call of the theatre. Not in the near future certainly. Baron
couldn’t bear to associate children and the stage. But in a few
years....
And if she were ever to return to the profession which was her
birthright, it was Thornburg she would need, and not the Barons.
Moreover, Thornburg was a wealthy man, and childless. He was now
ready to take the child into his home as his own. There could be only
one outcome to such an arrangement—an outcome wholly in Bonnie
May’s favor.
Therefore, his—Baron’s—right to keep the child was of the shakiest
possible nature.
And having reached these two conclusions, dwelling now upon the
one and now upon the other, Baron extinguished his light and went
to bed.

In the morning at about seven o’clock, while he was standing before

the glass with a military hair-brush in his hand, his problem was
solved for him in a flash. He stood with the brush suspended in air. A
light leaped into his eyes.
“How simple!” he exclaimed. “The very way out of it. The only way.”

At three o’clock that afternoon he entered Thornburg’s private office,

after having taken the precaution of ascertaining (1st) that Thornburg
had returned from luncheon in a fairly good humor, and (2d) that the
manager was alone.
“You know I had a little talk with Mrs. Thornburg about Bonnie May
last night,” he began, when Thornburg had thrust a chair toward him.
He was assuming his most casual manner, primarily because it
suited his present purpose, and also because he had not failed to
note that Thornburg’s face had darkened slightly at sight of him.
“Yes, I know.” The manager glanced at his desk as if he were a very
busy man.
“I felt the least bit—up a tree, as the fellow said, after I had talked to
her,” continued Baron. “You know I want to—to be decent about
things.”
“Of course,” agreed the manager, giving part of his attention to the
papers which were strewn about his desk. “And I suppose the child
is a good deal of a burden——”
He glanced up, and Baron wondered why a man shouldn’t be able to
keep the light of triumph out of his eyes when he really tried to.
“Not at all!” he interrupted blandly.
“——or that you are sure she will be, when the novelty of having her
about wears off.” He squared about sharply, with the air of a man
who means to do something handsome. “I’m still ready to take her, if
you decide that you’d like to give her up. Of course, I don’t know how
soon I might change my mind. In case Mrs. Thornburg loses interest,
I’d be through with the case, naturally.”
He turned to his desk again and examined a letter which came
uppermost, frowning and pursing his lips as if he were giving it deep
consideration.
Baron did not wholly succeed in repressing a smile. “All wrong,” he
said amiably. “The Greeks must have borne gifts to you before now,
Thornburg. No, I’m not tired of her. I’m not likely to be, either. Why,
she’s like a tonic. Sense? You wouldn’t believe it. She’s forever
surprising you by taking some familiar old idea and making you really
see it for the first time. She can stay at our house until the roof falls
in, if she only will—though of course I don’t hope she’d be willing to.
But don’t think there’s any question of our getting tired of her. She’s
not that kind. I might add, neither are we.”
Much to his amazement Thornburg sprang to his feet excitedly.
“I don’t know what you’re getting at!” he exclaimed. “If you’ve got
anything to say, why not say it and be done with it?”
“I don’t know what you’re getting at!” he exclaimed. “If
you’ve got anything to say, why not say it and be done
 with it?”
Baron arose, too. He thought he was justified in feeling offended. “I
think,” he said quietly, “I haven’t got anything to say, after all.” He
managed to keep his voice and eyes under control. These
proclaimed no unfriendliness. But his lips had become somewhat
rigid.
“But you did have,” retorted Thornburg. He sat down again and
produced a handkerchief with which he wiped his face and neck
nervously. “Come, don’t pay any attention to my bad manners. You
know I’ve got a thousand things to worry me.”
“Yes, I know. I’m really trying to help—or I had the thought of helping.
You—you make it a bit difficult.”
“There was something about the little girl,” said Thornburg.
“Yes. As to her—status. Chapter I—the inquiry for her, and our little
flurry—seems to be completed.”
“They probably didn’t care about her very much.”
“Well—possibly. At any rate, we seem to have come to a full stop for
the time being. And I’ve been thinking about the future. I ought to tell
you that after my talk with Mrs. Thornburg, the case didn’t seem
quite so simple as it had seemed.”
Thornburg, clasping his knee in his hands, was bending upon the
floor a gaze darkened by labored thought.
“I’ve begun to feel a kind of moral responsibility. At first I thought only
of my own point of view. My family’s, I mean. Our interests and
pleasures. But you see there’s also something to be said from the
standpoint of our—our guest. I wouldn’t want to lessen her chances
of future happiness. I wouldn’t want to have my way altogether and
then find out after a while that it had been the wrong way. I never
realized before how much the people of the stage are born and not
made. That’s the gist of the matter. There will come a time when
nothing in the world is going to keep Bonnie May off the stage.
That’s my conviction now.”
“They say children do inherit—” interposed Thornburg.
“The question of her future stumps you a bit. It’s not as if she were
like any other little girl I ever heard of. It’s like this: I’d like to have a
skylark in a cage, if it would sing for me. But I’d never be able to
forget that its right place was in the sky. You see what I mean. I don’t
want to be wholly responsible for keeping Bonnie May—out of the
sky.”
“Well?”
“My ideas aren’t exactly definite. But I want her to be free. I want her
to have a part in working things out the way she wants them.”
“That’s good sense. Turn her over to me, then.”
“That’s not the idea at all. I think up to a certain point it may be good
for her to experience the—the gentle tyrannies which are part of her
life with us. On the other hand, if she becomes identified with you (I
don’t know just what other word to use), and you get to be fond of
her, why then in a material sense.... Oh, I don’t like the tone of that at
all. But you’ll get the idea, and take it for granted that what I’m trying
to get at is that I don’t want to stand in Bonnie May’s light.”
Baron tried to join the manager in the latter’s impatient laugh. “You’ll
have to excuse my denseness,” said Thornburg. “I get your meaning
as easy as I can see into a pocket. The way it sounds to me is that
you’re sure you want to keep her, and that you’re just as sure that
you don’t want to keep her.”
“That’s nearly it,” admitted Baron, flushing slightly. “Suppose I say
that I want to keep her a part of the time, and that I’d like you to keep
her the other part. Suppose I offer to share her with you: to
encourage her to visit Mrs. Thornburg a day at a time—days at a
time—a week at a time. Suppose we take her on a kind of
partnership basis. No unfair influence; no special inducements.
Suppose I make it plain to her that you and Mrs. Thornburg are her
real friends, and that you will be glad to have her come as often as
she likes, and stay as long as she likes.”
Thornburg’s eyes were beginning to brighten.
“Would you,” added Baron, “do the same thing by us? I mean, would
you encourage her to come to us when she felt like it, and see that
she had the chance to go as freely as she came?”
Thornburg’s flushed face was all good-nature now. The little barriers
which he had kept between his visitor and himself fell away
completely.
“A kind of duel between us,” he elaborated, “to see which of us has
the best attractions to offer?”
“Well—yes, you might put it that way, I suppose. That’s a theatrical
phrase, I believe. Perhaps it wouldn’t have occurred to me. At any
rate, the plan I’ve outlined would give her a chance to do a little
deciding on her own account. It would give her a chance to give her
affections to those who win them. It would place some of the
responsibility for her future on her own shoulders. And whatever
conclusions she came to I’d be willing to bank on.”
“That,” declared Thornburg with enthusiasm, “is what I call the
proposition of a first-class sport.” He extended his hand to Baron.
“You stick to your part of the bargain and I’ll play fair to the letter.”
He would have shown Baron out of the office, then. He had a taste
for suitable climaxes, too. But Baron lingered, chiefly because he
didn’t like the prospect of an almost mischievous conflict which the
manager seemed to welcome and to anticipate.
“She can be loyal to us all,” he said, “if she’s encouraged in being.”
At the sound of his own words he fell to thinking.
No, she wouldn’t need to be encouraged. She would be loyal without
that. There was nothing to fear on that score at all.
He looked up rather whimsically. “Well, I’ll tell her,” he said.
“You’ll tell her——”
“That she has been invited to visit Mr. and Mrs. Thornburg, and
make herself quite at home.”
 CHAPTER XVIII
MRS. BARON TAKES UP THE GAUNTLET

Having decided upon what he conceived to be an admirable plan of

action, Baron was unwilling to believe that he ought to be in any
hurry to execute his plan.
For the time being Bonnie May was getting along very well indeed. In
fact, Baron made a point of looking into this matter with a good deal
of thoroughness, from a somewhat new angle, and he was greatly
pleased by what he discovered.
Little by little the child had become habituated to the home
atmosphere. This, of course, was due largely to the fact that the
other members of the family had become habituated to having her
about. They no longer felt constrained to utter pleasant nothings, or
to hold their tongues, because of her presence. When they forgot her
“strangeness,” she ceased to be strange.
She obediently and even intelligently attended when Mrs. Baron
gave her her lesson on the piano.
“Though I think,” she confided to Baron on one occasion, “I could get
hold of the high places without going through all the funny business
she seems to regard so highly.”
Baron spoke in defense of the “funny business,” and presently she
agreed with him.
The guest’s wardrobe had been made gloriously complete, and in
this relationship another pleasant development was to be noted.
Bonnie May had been painfully accustomed to the use of trunks.
Now she made the acquaintance of bureau drawers, and her delight
was unbounded. She spent hours in arranging her things. She won
Flora’s genuine applause by her skill and taste in this matter.
Flora bought her a hat.
She looked at it in a queerly detached manner for an instant. “Oh, a
hat,” she commented. She might have been repeating a word
spoken by a travel-lecturer, describing some interesting place which
did not seem to concern her. It appeared that she never had owned
a hat.
She put it on before the glass. “Oh!” she cried. She thrust impulsive
arms about Flora’s neck and hugged her.
Flora enjoyed that experience so much that she bought another hat
which she described as “unmade.” Ribbons of gay colors, and white
lace, and little silk flowers of various hues, came with it, and the child
was given these materials to experiment with as she pleased. Flora
gave advice, and was ready with assistance.
Again the result was interesting. Bonnie May experienced a joy
which was rapt, almost tremulous in quality. A desert-bred bird,
coming upon an oasis, might have regarded its surroundings with the
same incredulous rapture.
Baron’s room became hers permanently, and here she developed a
keen delight in “housekeeping.” Here also she received Mrs. Baron
and Flora as guests, and amazed them by her performance of the
part of hostess.
“I call it nonsense,” declared Mrs. Baron to Flora, after the two had
paid a formal call. But her face was flushed with happiness and her
voice was unwontedly soft.
“Not nonsense,” responded Flora; “it’s just happiness.”
She spent whole afternoons with Mrs. Shepard in the kitchen and
dining-room. She learned how to bake little cakes.
It became her duty—by her own request—to set the table, and upon
this task she expended the most earnest thought.
Baron commented upon this on one occasion. “Ah, you’re not an
artist, after all. You’re a Gretchen,” he said.
“But everything about the table is so pretty and nice,” she
responded. “It’s as elegant as a table in a play, and ever so much
more sensible. You know something always happens when you sit
down to a table on the stage. A servant comes in and says: ‘Beg
pardon, mum, but there’s a gentleman—he says he’s your uncle
from Green Bay’—and then everybody gets up in a hurry, because
the uncle is supposed to believe his niece has a lot of children he’s
been helping to support, when she hasn’t got any at all. Or
something like that.”
In brief, there were a hundred accumulating evidences to prove that
Bonnie May in the Baron household was the right individual in the
right place.
It is true that Mrs. Baron did not forget how Thornburg had called on
a certain night to take the child away, and how she had given him to
understand—she supposed—that she would expect him back on the
same errand some other time. And Baron could not free his mind of
the fact that he had voluntarily entered into a compact by which his
guest must sooner or later be lost to the household at least a part of
the time.
But these were matters which were not discussed in the family.
A week passed—two weeks, and Baron hadn’t seen Thornburg or
communicated with him. One day in June the thermometer shot up in
real midsummer fashion, and the audiences in most of the theatres
were such that all the shrewd managers became listless and absent-
minded. The “regular season” was over.
Thornburg closed his theatre and turned his attention to a summer
resort where there was an opportunity to launch an al fresco
entertainment scheme. “Everybody was leaving town.” There
remained only the uncounted thousands for whom some lighter form
of entertainment must be provided.
The flight of time, the inevitable march of events, brought to Baron a
realization of the fact that there was a promise he must keep. And so
one day, during an hour in the attic, he spoke to Bonnie May.
She didn’t seem to pay any attention at all to his preliminary words. It
slowly dawned upon her that what Baron was saying concerned her
in a special way.
“... people you will be interested in, I am sure,” Baron was saying.
“Thornburg, the name is.” He glanced at her; but the name had
made no impression. “Mrs. Thornburg is not very strong, and a
cheerful visit ought to be just the thing to help her. Mr. Thornburg is a
theatrical man. Why, it was his theatre I met you in. They have a
beautiful home.”
“Oh, that makes me think,” was all the reply he received. “What
became of the man who had a play?”
“Eh—a play?”
“You remember—when I first came. He had the first act and read it to
you in the library, and I had to go to bed.”
“Oh—Baggot. He’s probably forgotten all about it by this time. Or
writing another that he’ll never finish.”
She shook her head, unconvinced. “He was so enthusiastic,” she
objected.
So for the time being there was an end to the discussion of her visit
to the Thornburgs.

Another week passed, and then Baron had an extraordinarily busy

day.
In the forenoon came a letter from one of the dramatic editors for
whom Baron had done special work occasionally.
“They are launching some sort of a dramatic stock enterprise out at
Fairyland to-night,” the letter ran, “and I’m hoping you can do it for
me. Thornburg is managing it. I don’t hope it will be much as a
dramatic proposition, but you might be able to get some readable
impressions. Please let me know.”
A later mail brought a communication from Thornburg.
The sight of the manager’s signature brought Baron up with a jerk—
but he was reassured by the first few lines. Thornburg wasn’t
charging him with bad faith. Instead, he was enclosing an order for
an unlimited number of seats for the Fairyland opening.
“I understand,” ran a pencilled line by way of postscript and
explanation, “that you are to represent the Times to-night.”
Also there was a letter from Baggot. Baggot’s play had reached a
stage where it needed Baron’s inspection. The budding playwright
asked no questions. He merely declared his intention of calling that
night.
Baron went up into the attic to look at the morning paper. He wanted
to know what they were doing out at Fairyland, and who was doing it.
And while he noted one impressive name after another, he was
arrested by an altogether amazing sound down in his mother’s
sitting-room. Mrs. Baron had been giving Bonnie May her music
lesson, and now, the lesson done, she was singing for her pupil.
The thin old voice faltered on some of the notes, but the words came
clear enough:
“... She’s all the world to me,
And for bonnie Annie Laurie....”
Baron smiled and shook his head.
“What was it,” he mused, “about a plan of reconstruction?”
Then he went down-stairs to telephone his acceptance to the man
on the Times.
Baggot he completely forgot.

When Baron entered the dining-room at dinner-time that evening

Flora looked at him with mild surprise.
“All dressed up and nowhere to go,” said she.
“But there is somewhere to go. I’m going to write up the Fairyland
opening. Would you like to go with me?”
“No, thank you.”
It was clearly understood that Baron’s question had been put in a
spirit of jest. It was understood that Flora and her kind did not go to
the Fairylands—and their kind.
But Bonnie May failed to grasp the situation.
“What’s Fairyland?” she inquired.
“A large enclosure occupied entirely by mad people, and with a
theatre in one corner.”
She ignored the reference to mad people. “Oh! a theatre. What are
they playing?”
“A piece called ‘The Second Mrs. Tanqueray,’” said Baron.
She sat up, swiftly erect, and clasped her hands. “How fine!” was her
comment. “Do you think you could take me?”
“I should say not!” Baron responded without thinking. His unthinking
refusal was a result of the habitual Baron attitude. But as he
regarded her thoughtfully, and noted the puzzled inquiry in her
glance, he couldn’t quite understand why he had been so emphatic,
so confident of being right. “It’s not a play a little girl would care for,”
he added, now on the defensive.
She smiled indulgently. “The idea! I mean, anybody would be
interested in it.”
“What’s it about?” challenged Baron.
“A lady who died because they were unkind to her—even the people
who loved her. It’s about a lot of snobs and a—a human being.” She
spoke with feeling. She sensed the fact that again she was being
required to stand alone.
Baron frowned. “How in the world did you find out anything about a
play like that?”