Effects of Ageing and Tai Chi Training On Soleus H-Reflex in Olde

Southern Cross University
ePublications@SCU
Theses
2011
Effects of ageing and Tai Chi training on soleus Hreflex in older adults
Yung-Sheng Chen
Publication details
Chen, YS 2011, 'Effects of ageing and Tai Chi training on soleus H-reflex in older adults', PhD thesis, Southern Cross University,
Lismore, NSW.
Copyright YS Chen 2011
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Effects of ageing and Tai Chi training on soleus

H-reflex in older adults
Yung-Sheng Chen
Master of Science in Sport Sciences
This thesis is presented in fulfilment of the requirements for the degree of

Doctor of Philosophy at Southern Cross University
February 2011
Declaration
I certify that the work presented in this thesis is, to the best of my knowledge and belief,
original, except as acknowledged in the text, and that the material has not been
submitted, either in whole or in part, for a degree at this or any other university.
I acknowledge that I have read and understood the Universitys rules, requirements,
procedures and policy relating to my higher degree research award and to my thesis, I
certify that I have complied with the rules, requirements, procedures and policy of the
University (as they may be from time to time).
Print Name: Yung-Sheng Chen
Signature:
Date: 02/02/2011
ii
Abstract
The Hoffmann reflex (H-reflex) is used to investigate the influence of Ia afferent
projection on the spinal motoneuron activities. It has been suggested that the H-reflex is
task-dependent and demonstrates adaptations to exercise training. Much of the previous
research on the H-reflex was based on young populations. Little information is available
on the H-reflex modulation in response to exercise and training in older populations.
The objective of the research work presented in this thesis was to expand our knowledge
on the effects of ageing and Tai Chi (TC) training on the soleus (SOL) H-reflex
modulations in older adults. Four related studies have been conducted.
Study I. The aim of this study was to determine the test-retest reliability of SOL Hreflex assessment during isometric muscle contraction at various intensities and ankle
joint positions. The H-reflex was assessed when the ankle joint was placed at neutral
(0), plantarflexion 20, and dosiflexion 20 positions and during isometric
plantarflexion at 10%, 30% and 50% of the maximal voluntary contraction (MVC)
levels, on two separate days, in a group of young adults (5 males and 5 females, age
24.9 5 years). The results showed a high level of test-retested reliability (Intraclass
Correlation Coefficients, ICC) for the SOL H-reflex tested at the neutral (ICC = 0.96)
and plantarflexion (ICC = 0.92) positions, and a moderate level of ICC at the
dorsiflexion position (ICC = 0.75) during rest. The results also demonstrated that
assessing the SOL H-reflex during low intensity (10% MVC) of isometric muscle
contractions yielded more reliable test-retest outcomes (ICC = 0.92 0.95) than that
during contractions at higher intensities (30% and 50% MVC, ICC = 0.62 0.97),
regardless of ankle joint positions.
Study II. The aim of this study was to use a cross-sectional research design to compare
the effects of ankle joint position and muscle contraction intensity on SOL H-reflex gain,
iii
latency and duration between young (10 males and 10 females, age 25.1 4.0 years)
and older (10 males and 10 females, age 74.2 5.1 years) adults. The results showed
that there were significant differences of the SOL H-reflex parameters between young
and older adults under all testing conditions. However, when contraction intensity was
progressively increased, a similar down-regulation of the SOL H-reflex gain was
observed in both aged groups. This result may indicate a possible reservation of motor
function regulated by the supraspinal mechanisms in older adults.
Study III. The aim of this study was to investigate the effect of ageing on soleus (SOL)
H-reflex modulation during shortening and lengthening muscle actions. Cross-sectional
comparisons of the maximal amplitude of H-reflex (Hmax) and maximal amplitude of Mwave (Mmax) ratio were made between young (10 males and 10 females, age 24.4 4.0
years) and older (10 males and 10 females, age 73.3 5.0 years) adults during passive
movements and voluntary contractions of the muscles around the ankle joint. The Hreflex modulation during upright standing under eyes open/closed and on
stable/unstable surface conditions were also investigated in this study. The correlations
of SOL Hmax between these dynamic muscle actions and postural tasks were evaluated.
The results indicated that there were significant age-related differences of the SOL Hreflex modulation during passive and active shortening and lengthening muscle actions.
Pearson correlation analysis revealed that the SOL Hmax during both the passive and
active shortening and lengthening plantarflexions was significantly correlated with that
during the postural tasks in young adults. However, older adults only demonstrated a
significant correlation of the SOL Hmax between the passive shortening and lengthening
actions and postural tasks.
Study IV. The aim of this study was to investigate the effects of 12 weeks of TC training
on the SOL H-reflex during upright standing under different visual and somatosensory
iv
conditions in older adults. Thirty-four volunteers (17 males and 17 females, age 72.9
5.9 years) were assigned into a TC and a control group. The results demonstrated that
the SOL Hmax/Mmax ratios during upright standing under eyes open/closed and
stable/unstable surface conditions were significantly increased after the TC training in
older adults. However, the mean displacements of centre of pressure (COP) in anteriorposterior and medial-lateral directions were not significantly changed after training. The
results suggested that adaptive change of the SOL H-reflex is not related to control of
static posture after 12 weeks TC training in older adults.
This thesis demonstrated the age-related differences in the SOL H-reflex modulations
during different ankle joint positions, isometric and dynamic ankle muscle actions and
static upright standing. The adaptive change of the SOL H-reflex to 12 weeks of TC
training may provide insight into the understanding of the neural adaptation to TC
training in older adults.
List of publications
Book chapter
Chen, Y. S., & Zhou, S. (2011). H-reflex assessment as a tool for understanding motor
functions in postural control. In Posture: Types, Assessment, and Control. New
York: Nova Science Publishers (in press).
Journal publications
Chen, Y. S., Zhou, S., Cartwright, C., Crowley, Z., Baglin, R. & Wang, F. (2010). Testretest reliability of the soleus H-reflex is affected by joint positions and muscle
force levels. Journal of Electromyography and Kinesiology, 20 (5), 980-987.
Chen, Y. S. & Zhou S. (2011). Soleus H-reflex and its relation to static postural control.
Gait and Posture, 33 (2), 169-178.
Chen, Y. S., Zhou, S., & Cartwright, C. (2011). Effect of 12 weeks Tai Chi training on
soleus H-reflex and control of static posture in older adults. Archives of
Physical Medicine and Rehabilitation, 92, 886-891.
Chen, Y. S., Crowley Z, Zhou S, Cartwright C. 2011 (submitted). Effect of 12 weeks

Tai Chi training on soleus H-reflex in older adults: a pilot study. European
Journal of Applied Physiology (submitted and under review).
vi
Conference publications
Chen, Y. S., Zhou, S., Cartwright, C., Crowley, Z., Baglin, R. & Wang, F. (2009). Testretest reliability of the soleus H-reflex is affected by joint positions and force
levels. Paper presented at the 8 th Annual Conference of the Society of Chinese
Scholars on Exercise Physiology and Fitness, Baptist University, Hong Kong,
China. Abstracts Book (pp 38).
Chen, Y. S., Zhou, S., Cartwright, C. & Crowley, Z. (2010). Effects of joint position
and muscle contraction intensity on soleus H-reflex in young and older adults.
Paper presented at the 4th The Australian Association for Exercise & Sports
Science Conference, Gold Coast, Australia. Abstracts Book (pp 79).
Chen, Y. S., Zhou, S. & Cartwright, C. (2010). Effect of ageing on soleus H-reflex
modulation during shortening and lengthening muscle actions. Paper presented
at the 9 th Annual Conference of the Society of Chinese Scholars on Exercise
Physiology and Fitness, Beijing Sport University, Beijing, China.
vii
Acknowledgments
I would like to express appreciation to my principal supervisor, Professor Shi Zhou and
co-supervisor, Professor Colleen Cartwright, for their guidance to my research and
amount of time they spent on providing feedback on the thesis, that are invaluable to my
study and academic development. Their practical advice, knowledge, and comments are
very much appreciated.
Thank Robert Baglin for his generous and skilful assistance with technical support and
computer setting for data acquisition.
Also, I would like to express my gratitude to Erich Wittstock. You are always available
when I need assistance and support in the laboratory.
Thank Zac Crowley for the initial encouragement of this study.
Also extensive thanks to Li Zhang from the Library, for her helpful consultation with
the Endnote program.
Furthermore, thank Fang Wang, for her assistance during testing and data collection.
A sincere gratitude to Dr. Pedro Bezerra for his practical advice on protocol design and
technical consultation for constructing my first study.
Thank also all participants and workmates who volunteered and cooperated for this
study. I am grateful to your commitments in time and efforts in participation of my
studies.
Finally, I want to express my sincere thanks to all the members of my family for their
full support and encouragement over the time of my Ph.D. candidature.
viii
Table of Contents
Declaration ......................................................................................................................ii
Abstract. .....................................................................................................................iii
List of publications ........................................................................................................vi
Book chapter............................................................................................................vi
Journal publications.................................................................................................vi
Conference publications .........................................................................................vii
Acknowledgments........................................................................................................viii
Table of Contents...........................................................................................................ix
List of Figures ..............................................................................................................xiii
List of Tables.................................................................................................................xv
Abbreviation ................................................................................................................xvi
Units of measurement................................................................................................xviii
Chapter 1 Introduction .............................................................................................1
1.1. Introduction ...........................................................................................................2
1.2. Aims of the investigation.......................................................................................9
1.3. Research hypotheses............................................................................................11
1.4. Significance of the research.................................................................................12
1.5. Study Limitations ................................................................................................12
1.6. Delimitation .........................................................................................................13
1.7. Ethical approval ...................................................................................................13
Chapter 2 Literature review...................................................................................14
2.1. Introduction .........................................................................................................15
2.2. Validity and reliability of H-reflex measurement................................................16
2.2.1. H-wave and M-wave ....................................................................................16
2.2.2. Reliability of H-reflex ..................................................................................19
2.2.3. Methodological concerns in H-reflex assessment ........................................20
2.2.4. Limitations of H-reflex test ..........................................................................27
2.3. Control of posture................................................................................................28
2.3.1. Sensory inputs ..............................................................................................28
2.3.2. Supraspinal mechanisms ..............................................................................30
2.3.3. Spinal mechanisms .......................................................................................33
2.4. H-reflex modulation in relation to control of static posture ................................41
2.4.1. Biomechanical characteristics of standing ...................................................41
ix
2.4.2. H-reflex modulation during postural tasks ...................................................42

2.4.3. Effect of balance training on H-reflex modulation.......................................46
2.5. Effect of Tai Chi training on neuromuscular adaptations in older adults............48
2.5.1. Muscular adaptations....................................................................................49
2.5.2. Neural adaptations ........................................................................................49
2.6. Summary..............................................................................................................50
Chapter 3 General methodology ............................................................................51
3.1. Recruitment of Participants .................................................................................52
3.2. H-reflex test .........................................................................................................55
3.3. Isometric submaximal voluntary contraction ......................................................57
3.4. Passive and active dynamic muscle actions ........................................................59
3.5. Static postural tasks .............................................................................................60
3.6. Data acquisition and analysis ..............................................................................61
3.7. Statistical analyses ...............................................................................................64
Chapter 4 Test-retest reliability of the soleus H-reflex in relation to joint
positions and muscle force levels (Study I)..............................................66
4.1. Introduction .........................................................................................................67
4.2. Methods ...............................................................................................................68
4.2.1. Participants ...................................................................................................68
4.2.2. Experimental settings ...................................................................................69
4.2.3. Experimental procedures ..............................................................................70
4.2.4. Data analysis.................................................................................................72
4.2.5. Statistical analyses........................................................................................72
4.3. Results .................................................................................................................73
4.3.1. Reliability of the soleus H-reflex at different joint angles during rest .........73
4.3.2. Reliability of the soleus H-reflex during submaximal muscle contractions.74
4.3.3. Background EMG in the soleus and tibialis anterior muscles during
submaximal voluntary contractions........................................................................74
4.4. Discussion............................................................................................................80
4.5. Conclusion ...........................................................................................................84
Chapter 5 Effects of joint position and muscle contraction intensity on soleus Hreflex gain in young and older adults (Study II).....................................86
5.1. Introduction .........................................................................................................87
5.2. Methods ...............................................................................................................90
5.2.1. Participants ...................................................................................................90
x
5.2.2. Experimental settings ...................................................................................91

5.2.3. Experimental procedures ..............................................................................91
5.2.4. Data analysis.................................................................................................92
5.2.5. Statistical analyses........................................................................................93
5.3. Results .................................................................................................................94
5.3.1. The soleus H-reflex gain ..............................................................................94
5.3.2. Background EMG in the soleus and tibialis anterior muscles......................94
5.3.3. Latency of the soleus H-reflex .....................................................................95
5.3.4. Duration of the soleus H-reflex ....................................................................95
5.4. Discussion..........................................................................................................100
5.4.1. Age-related changes in the soleus H-reflex gain ........................................100
5.4.2. Age-related changes in the soleus H-reflex latency and duration ..............102
5.5. Conclusion .........................................................................................................103
Chapter 6 Effect of ageing on soleus H-reflex modulation during shortening and
lengthening muscle actions in relation to postural control (Study III)104
6.1. Introduction .......................................................................................................105
6.2. Methods .............................................................................................................107
6.2.1. Participants .................................................................................................107
6.2.2. H-reflex test ................................................................................................107
6.2.3. Passive and active dynamic muscle actions ...............................................108
6.2.4. Postural tasks ..............................................................................................109
6.2.5. Data analysis...............................................................................................109
6.2.6. Statistical analyses......................................................................................110
6.3. Results ...............................................................................................................110
6.3.1. Dynamic muscle action test........................................................................110
6.3.2. The correlation between the maximal SOL H-reflex amplitude in the
dynamic muscle action test and postural control tests..........................................111
6.4. Discussion..........................................................................................................118
6.4.1. Age-related changes in soleus H-reflex modulation during shortening and
lengthening plantarflexors actions........................................................................118
6.4.2. Correlation of soleus H-reflex modulation between dynamic plantarflexion
actions and postural tasks in young and older adults ...........................................121
6.5. Conclusion .........................................................................................................122
Chapter 7 Effect of 12 weeks Tai Chi training on soleus H-reflex and control of
static posture in older adults (Study IV) ...............................................123
xi
7.1. Introduction .......................................................................................................124

7.2. Methods .............................................................................................................127
7.2.1. Participants .................................................................................................127
7.2.2. Tai Chi training program ............................................................................128
7.2.3. H-reflex test ................................................................................................128
7.2.4. Static postural test.......................................................................................128
7.2.5. Experimental procedures ............................................................................129
7.2.6. Data analysis...............................................................................................129
7.2.7. Statistical analyses......................................................................................130
7.3. Results ...............................................................................................................130
7.3.1. Participants .................................................................................................130
7.3.2. Elimination of outlier data points ...............................................................130
7.3.3. H-reflex test ................................................................................................131
7.3.4. Centre of pressure measurement ................................................................131
7.4. Discussion..........................................................................................................136
7.4.1. Soleus H-reflex modulation........................................................................136
7.4.2. Static postural control.................................................................................137
7.4.3. The relationship between H-reflex modulation and postural control .........138
7.5. Conclusion .........................................................................................................139
Chapter 8 Summary, conclusions and implications ...........................................141
8.1. Summary of results............................................................................................142
8.2. Conclusions .......................................................................................................144
8.3. Implications of the finding and future research.................................................146
References ...................................................................................................................150
Appendix A Information sheet ...............................................................................165
Appendix B Consent form ......................................................................................175
Appendix C Call for volunteers..............................................................................182
Appendix D Health status assessment prior to exercise testing ..........................188
Appendix E Tai Chi training program..................................................................197
Appendix F Original data .......................................................................................203
xii
List of Figures
Figure 2.1: Simplified illustration of Hoffmann reflex ..................................................19
Figure 2.2: Recruitment curve ........................................................................................24
Figure 2.3: A schematic diagram of spinal networks .....................................................34
Figure 2.4: A sketch of presynaptic inhibitory process. .................................................36
Figure 2.5: Illustration of reciprocal Ia inhibitory pathway. ..........................................38
Figure 2.6: The H-reflex modulation in the soleus (SOL) and medial gastrocnemius
(MG) muscles during task-dependent body sways......................................43
Figure 3.1: Locations of the stimulation and EMG electrodes.......................................56
Figure 3.2: Participants position in the isometric voluntary contraction test................58
Figure 3.3: The standardised upright position in the static postural tasks......................61
Figure 3.4: A schematic shows the data acquisition system. .........................................63
Figure 3.5: Raw EMG trace was used to quantify the SOL H-reflex parameters. .........63
Figure 4.1: Participants leg position in the soleus H-reflex test. ..................................70
Figure 4.2: The maximal soleus H-reflex amplitude recorded during rest, and 10%, 30%
and 50% maximal voluntary contraction at the neutral, plantarflexion, and
dorsiflexion of ankle positions from a representative participant. ..............77
Figure 4.3: The soleus (A) and tibialis anterior (B) background EMG (bEMG) activity
recorded during 10%, 30% and 50% maximal voluntary contractions at the
neutral, plantarflexion and dorsiflexion positions in the Trial 1 (T1) and
Trial 2 (T2). .................................................................................................79
Figure 5.1: Typical Hmax of the soleus H-reflex recorded during rest, and muscle
contractions at 10%, 30% and 50% MVC at the neutral, plantarflexion and
dorsiflexion of ankle positions in one young (left side) and one older (right
side) participant. ..........................................................................................96
Figure 5.2: Influence of ageing on the soleus H-reflex gain during rest and submaximal
muscle contractions when the ankle joint placed at neutral, plantarflexion
and dorsiflexion positions............................................................................97
Figure 5.3: Background EMG of the soleus (A: young, C: older) and tibialis anterior (B:
young, D: older) muscles during rest and soleus contractions at 10, 30 and
50% MVC in ankle joint positions of neutral, plantarflexion and
dorsiflexion. .................................................................................................98
Figure 5.4: Latency (A) and duration (B) of maximal soleus H-reflex in the young
(filled circles) and older groups (unfilled circles). ......................................99
xiii
Figure 6.1: H-reflex and M-wave recruitment curves during passive shortening (Hreflex: ; M-wave: ) and lengthening (H-reflex: ; M-wave: ) actions,
and standing on stable surface with eyes opened (H-reflex: ; M-wave: )
from representative one young (A, B) and one older (C, D) participants,
respectively. ...............................................................................................114
Figure 6.2: Representative raw EMG traces of the maximal soleus H-reflex during
passive and active dynamic muscle activities and four different postural
conditions from one young (solid lines: A, C) and one older (dotted lines: B,
D) participants. ..........................................................................................115
Figure 6.3: The soleus Hmax to Mmax ratio during A) passive shortening and lengthening
movements, and B) voluntary shortening and lengthening muscle
contractions................................................................................................116
Figure 7.1: The SOL Hmax/Mmax ratio before and after 12-week Tai Chi training in the
Training and Control groups......................................................................133
xiv
List of Tables
Table 3.1: A summary of participants recruited in the four studies. .............................. 54
Table 3.2: Summary of the statistical analyses............................................................... 65
Table 4.1: Mean value, standard deviation (values in the brackets), intraclass correlation
coefficients (ICC), and standard error of measurement (SEM) for the maximal voluntary
contraction torque (MVC), the maximal amplitude of soleus H-reflex (Hmax), the
maximal amplitude of soleus M-wave (Mmax), and the Hmax/Mmax ratio during rest and at
three ankle joint positions in Trial 1 (T1) and Trial 2 (T2). ........................................... 75
Table 4.2: Mean value and standard deviation (in the brackets), intraclass correlation
coefficients (ICC), and standard error of measurement (SEM) for the maximal
amplitude of soleus H-reflex (Hmax) at three ankle positions during contractions at 10%,
30% and 50% maximal voluntary torque (MVC) in the Trial 1 (T1) and Trial 2 (T2).. 76
Table 6.1: Pearson correlation coefficients of the maximal soleus H-reflex amplitude
between dynamic muscle activities and postural tasks, in the young and older adult
groups. .......................................................................................................................... 117
Table 7.1: Comparison of physiological characteristics between the Tai Chi group and
Control group prior to the 12 weeks of TC training. .................................................... 134
Table 7.2: Mean values and standard deviation of the mean displacement of COP values
(anterior-posterior and medial-lateral directions) in four sensory conditions in the Tai
Chi and Control groups before and after the 12-week training or control period. ....... 135
xv
Abbreviation
A/D
Analog to digital
Ag/AgCl
Silver silver chloride
ANOVA
Analysis of variance
bEMG
Background electromyogram
C-T
Conditioning-test
CNS
Central nervous system
COM
Centre of mass
COP
Centre of pressure
COPA-P
Centre of pressure movement in anterior and posterior direction
COPM-L
Centre of pressure movement in medial and lateral direction
EEG
Electroencephalogram
EMG
Electromyogram
EPSP
Excitatory postsynaptic potential
GABA
Axo-axonal gamma-aminobutyric acids
GVS
Galvanic vestibular stimulation
Hmax
Maximal peak-to-peak amplitude of H-reflex
ICC
Intraclass coefficient correlation
IPSP
Inhibitory postsynaptic potential
ISI
Inter-stimulus interval
MEPs
Motor evoked potentials
MG
Medial gastrocnemius
Mmax
Maximal peak-to-peak amplitude of M-wave
MVC
Maximal voluntary contraction
RMANOVA
Repeated measures analysis of variance
RMS
Root mean square
SC
stable surface with eyes closed
SD
Standard deviation
SEM
Standard error of measurement
SENIAM
Surface EMG for a non-invasive assessment of muscles
SO
Stable surface with eyes open
SOL
Soleus muscle
SOT
Sensory organization test
TA
Tibialis anterior muscle

xvi
TC
Tai Chi
TES
Transcranial electrical stimulation
TMS
Transcranial magnetic stimulation
UO
Unstable surface with eyes open
UC
Unstable surface with eyes closed
xvii
Units of measurement
cm
Centimetre
Degree
/s
Degree per second
Hz
Hertz
Hour
Kg
Kilogram
kHz
Kilohertz
Microsecond
Microvolt
mA
Milliampere
mm
Millimetre
ms
Millisecond
ms.cm-1
Millisecond per centimetre
mV
Millivolt
min
Minute
N.m
Newton meter
Percentage
Second
cm2
Square centimetre
Volt
xviii
Chapter 1 Introduction
1.1. Introduction
Humans adopt an upright posture in most physical activities. During standing, this
requires stabilisation of joint positions in multiple segments of the body in order to
maintain an appropriate posture. The ability to control the vertical projection of the
centre of mass (COM) relative to the base of support is defined as postural stability
(Horak, 2006). There are two major components to determine the postural stability
during standing: neural and musculoskeletal components (Horak, 2006). Neural
component refers to receiving the sensory information from the somatosensory,
vestibular and visual systems and sensorimotor processes that occur within the central
nervous system (CNS), whilst musculoskeletal component refers to muscular actions
resulting in biomechanical changes during postural control.
During bipedal standing, the body naturally oscillates in a model of inverted pendulum
(Winter, Patla, Prince, Ishac, & Gielo-Perczak, 1998). The body sways in the anteriorposterior direction and medial-lateral directions. The kinematic action in control of
upright posture attempts to move the COM without changing the base of support (limits
of stability). When maintaining the upright posture the body sways at a mean frequency
of 1.3 Hz which corresponds to 2.6 times of postural reversion per second (Loram,
Maganaris, & Lakie, 2005b). The control of upright posture requires appropriate
strategies to maintain joint stability, mainly at ankle and hip joints (Horak, 2006). The
ankle strategy is used during standing on a stable surface with small amounts of body
sway, whereas the hip strategy is used during standing on a small base of support or an
unstable surface.
The body oscillations during standing are associated with changes in the ankle joints
within a range of 1.0-1.5 in the anterior-posterior direction (Gatev, Thomas, Kepple, &
2
Hallett, 1999). In this case, the ankle plantarflexors and dorsiflexors play an important
role in control of ankle movement and sway angle (Winter, Prince, Frank, Powell, &
Zabjek, 1996). Loram et al. (2005) utilising ultrasound imaging technique reported that
the contractile tissue of the plantarflexors is shortening during forward sway, whereas
that is lengthening during backward sway. The paradoxical muscle movements
observed in Loram et als study indicate that the plantarflexor muscles contract
concentrically, eccentrically, or even isometrically in order to maintain the position of
ankle joints.
The ankle muscle stiffness has an essential function in controlling upright posture
(Evans, Fellows, Rack, Ross, & Walters, 1983; Fitzpatrick, Gorman, Burke, &
Gandevia, 1992).Winter et al. (1998; 2001) proposed a model of stiffness control during
standing. In this model, neural mechanisms in control of ankle joint movements are
associated with neural inputs from multiple sources. The CNS regulates the tension of
ankle muscles in response to any change of ankle joint position, according to the
feedback of sensory information from the ankle joint (Winter, et al., 2001). The
feedback and feedforward controls of ankle joint movement regulated by the
supraspinal mechanisms coordinate the most joint motions in attempt to stabilising the
whole body position in the vertical alignment (Gatev et al., 1999). For example, ankle
strategy plays a crucial role to adjust standing posture during a small perturbation of
support translation in forward and backward direction. In contrast, the role of knee and
hip joint movements is minimal for postural control in this case.
Ageing is an inevitable part of the physiological process in human life. This process
results in deterioration of the neuromuscular functions and decreased physical capacity
at older ages (Nelson et al., 2007). Compared with young adults, older persons show
slower cognitive processing (Rozas, Juncos-Rabadan, & Gonzalez, 2008), motor
3
response generation (Kolev, Falkenstein, & Yordanova, 2006; Ward, 2006), nerve
conduction velocity (Rivner, Swift, & Malik, 2001), and electromechanical delay
(Mackey & Robinovitch, 2006; Zhou, Lawson, Morrison, & Fairweather, 1995), and
decreased muscle strength (Frontera et al., 2008) and power (McNeil, Vandervoort, &
Rice, 2007). As a result, older adults have less ability in performing motor tasks and
executing daily physical activities. Although the existing evidence indicates a
deterioration of neuromuscular function with ageing, older adults may adopt different
strategies in performing motor tasks as a result of compensatory adaptations (Earles,
Vardaxis, & Koceja, 2001; Scaglioni, Narici, Maffiuletti, Pensini, & Martin, 2003).
Older adults have shown an increase in postural sway during upright standing,
compared to young adults (Abrahamova & Hlavacka, 2008). The compromised postural
control is the result of age-related changes in sensorimotor (Sturnieks, St George, &
Lord, 2008) and muscular (Carrie et al., 2003) functions.
During standing, electromyographic activity (EMG) of the plantarflexors corresponds to

less than 5% of that during maximal voluntary isometric contractions (MVC). More
recently, Kouzaki and Sinohara (2010) demonstrated a significant correlation between
the coefficient of variation in the central of pressure displacement during standing and
coefficient of variation in force production during plantarflexion contractions around
5% MVC. The outcome of this study indicates that the neural control might have similar
characteristics with respect to control of ankle joint during isolated joint movement and
standing.
Motor control refers to sensorimotor performance during body movements (Latash,

Scholz, & Schoner, 2002). In the sensorimotor processes, the spinal motoneurons, such
as - and -motoneurons, are responsible for control of skeletal muscle activity. The
spinal motoneuron activity is regulated by integration of the descending inputs from the
4
supraspinal centres and the ascending inputs from the peripheral sensory receptors
(Hultborn, 2006). These neural inputs converging onto the spinal cord are major factors
to determine the firing rate and recruitment of spinal motoneurons. When a postsynaptic
membrane potential depolarizes (excitatory postsynaptic potentials, EPSP) to the
threshold level, an action potential is produced for discharge of the spinal motoneurons.
In contrast, a postsynaptic membrane potential can be reduced by inhibitory
postsynaptic potentials (IPSP), resulting in inhibition of the spinal motoneuron activity.
Variation in excitatory and inhibitory inputs can affect the recruitment of spinal
motoneurons. In human movement study, the Hoffmann reflex (H-reflex) has been
extensively used to evaluate the effectiveness of group Ia afferent inputs to excite the motoneuron during motor performance (Misiaszek, 2003).
The H-reflex is an electrically induced analogue of reflex and can be used to assess
group Ia excitatory effect on the -motoneuron activation (see detailed explanation in
Chapter 2 of this thesis). The H-reflex amplitude is influenced by neural inputs from
various afferent and efferent pathways, therefore it can be used as an indicator of the
outcomes of the motor output regulation through the supraspinal and spinal mechanisms
(Hultborn, 2006; Misiaszek, 2003). There has been evidence that the modulation of Hreflex is task-dependent (Zehr, 2002). For example, the size of SOL H-reflex has been
found to vary upon ankle joint position (Guissard & Duchateau, 2006; Hwang, 2002),
muscle contraction intensity (Butler, Yue, & Darling, 1993; Stein, Estabrooks, McGie,
Roth, & Jones, 2007), and type of dynamic muscle actions (Duclay & Martin, 2005;
Duclay, Robbe, Pousson, & Martin, 2009). These observations can help us to
understand the motor output regulation during functional tasks. However, most of the
H-reflex studies in the current literature were based on young adults and may be
inappropriate to explain the H-reflex modulation in older adults due to ageing effects
(Earles, et al., 2001; Hortobagyi, del Olmo, & Rothwell, 2006; Kido, Tanaka, & Stein,
2004).
The amplitude of H-reflex increases proportionally with increased intensity of muscle

contraction (Stein, et al., 2007). The neural mechanism underlying the H-reflex
potentiation during muscle contraction has been suggested as increased neural drive
from the supraspinal levels (Nielsen, Petersen, Deuschl, & Ballegaard, 1993). The Hreflex gain, as determined by a ratio the amplitude of H-reflex divided by the
background muscle activity, has been used to evaluate the contribution of Ia afferent
inputs to motoneuron activation during execution of motor tasks (Capaday, 1997). In the
previous studies, age-related changes in the SOL H-reflex have been reported during
10%, 20% and 30% of MVC at neutral ankle position (Angulo-Kinzler, Mynark, &
Koceja, 1998; Earles, et al., 2001). Young adults demonstrated that the SOL H-reflex
gain during submaximal voluntary muscle contractions was stronger in a lying prone
position than that at the same level of muscle contractions in a standing position. In
contrast, older adults showed no difference in the SOL H-reflex gain in the same testing
conditions (Angulo-Kinzler, et al., 1998). The difference of H-reflex gain between
young and older adults is suggested to be related to the age-related changes in
presynaptic inhibitory function (Angulo-Kinzler, et al., 1998; Earles, et al., 2001).
However, observing the H-reflex gain at the neutral ankle position may give insufficient
information in understanding the ankle strategy during postural control, because the
ankle joint position constantly changes during naturally sway of normal standing.
It has been suggested that the amplitude of SOL H-reflex increases during passive
shortening movement, compared with that during passive lengthening movement
(Nordlund, Thorstensson, & Cresswell, 2002; Pinniger, Nordlund, Steele, & Cresswell,
2001). The movement-related changes in the SOL H-reflex modulation have also been
6
observed during shortening and lengthening muscle contractions (Duclay & Martin,
2005; Duclay, et al., 2009). The differentiation of SOL H-reflex modulation during
dynamic muscle movement is mainly controlled by presynaptic inhibition via either the
central or peripheral mechanisms (Duchateau & Enoka, 2008). However, these findings
were based on young adults. It is unknown whether older adults may adopt similar
neural strategies in modulation of H-reflex during shortening and lengthening muscle
actions.
The SOL H-reflex modulation is task-dependent and is found to be related to the ability
to control upright posture (Taube, Gruber, & Gollhofer, 2008). Recently, a series of
studies conducted by Tokuno et al. (2007; 2008) suggested that the SOL H-reflex is
position- and direction-dependent during body sway in anterior-posterior direction.
They speculated that the specific characteristic of H-reflex is related to the direction of
postural sway when the triceps surae is eccentrically or concentrically contracting to
maintain upright standing. However, there is no information available to us to support
whether the SOL H-reflex modulation during upright standing is correlated to that
during dynamic muscle activities.
A very serious issue for older people is their balance control capacity. Statistically, one
in three community-dwelling older adults experiences at least one fall each year and
more than thirty percent of fallers require medical treatment after suffering fall injuries
(Sturnieks et al. 2010; Tinetti, Speechley, & Ginter, 1988). The implementation of
exercise interventions can prevent falls and reduce the risk of falling in older adults
(Sturnieks et al. 2010; Gardner, Buchner, Robertson, & Campbell, 2001). In recent
years, Tai Chi (TC) exercise has been found to be an effective exercise intervention and
widely utilised for improvement of balance control in older adults (Choi, Moon, & Song,
2005; Li et al., 2005; Lin, Hwang, Wang, Chang, & Wolf, 2006; Voukelatos, Cumming,
7
Lord, & Rissel, 2007; Zeeuwe et al., 2006). Older TC practitioners have shown better
postural control capacity than their non-TC-trained counterparts. It has been suggested
that adaptive changes after TC training in kinematic proprioception (Fong & Ng, 2006;
Li, Xu, & Hong, 2008; Tsang & Hui-Chan, 2003, 2004b), vestibular function (Tsang &
Hui-Chan, 2006; Tsang, Wong, Fu, & Hui-Chan, 2004), tactile acuity (Kerr et al., 2008)
and reaction time (Li, Xu, & Hong, 2009) are related to improvement of postural control.
Although these studies have shown evidence of neural plasticity to TC training, it is
unclear whether this neural adaptation has occurred at the spinal cord level or
supraspinal levels. The neural mechanisms at the spinal level play an important role in
regulation of spinal reflex during upright standing (Taube, Gruber, et al., 2008).
Measuring the H-reflex amplitude can provide information to understand the neural
plasticity at the spinal level (Taube, Gruber, et al., 2008). The outcome can help us to
understand whether the neural adaptations associated with TC training is relevant to the
spinal contribution.
To yield valid outcome of the H-reflex test, the method used to measure the H-reflex
needs to be reliable. In the previous studies, the test-retest reliability of H-reflex during
rest has been reported under a variety of experimental conditions (Clark, Cook, &
Ploutz-Snyder, 2007; Handcock, Williams, & Sullivan, 2001; Palmieri, Hoffman, &
Ingersoll, 2002). During static postural tasks, high reliability of SOL H-reflex test has
been reported, for example, the intraclass correlation coefficient (ICC) of 0.94 in supine
position and 0.80 in standing position was reported by Hopkins et al. (2000). Low ICC
value during upright standing is related to variability of H-reflex modulation due to
body sway. During voluntary muscle contractions, various neural mechanisms may be
involved in the SOL H-reflex modulation, depending upon the contraction intensity
(Stein, et al., 2007). However, there is no study that has reported the test-retest
reliability of H-reflex during voluntary muscle contraction. It has been demonstrated
8
that the amplitude of H-reflex is inconsistent during submaximal voluntary muscle

contractions even though under the consistent experimental conditions (Funase & Miles,
1999). Whether the variation of H-reflex is related to biological changes or a poor
reliability during repeated trials of H-reflex test has not yet been determined. This
information is important to advance current knowledge in understanding the reliability
of H-reflex test during more functional tasks.
The research focuses on the SOL H-reflex because the SOL muscle plays an important
role in maintaining upright posture during standing and the studies on the SOL H-reflex
modulation may help us to understand the neural mechanisms in postural control. In
light of the above, four studies were designed to answer the research questions. Study I
was designed to determine the test-retest reliability of SOL H-reflex during isometric
submaximal voluntary muscle contractions in neutral, plantarflexion, and dorsiflexion
positions and to establish reliable measurement of H-reflex test in our laboratory. Study
II was designed to investigate the influence of ageing on the SOL H-reflex modulation
during isometric submaximal voluntary muscle contractions in neutral, plantarflexion,
and dorsiflexion positions. Study III examined the influence of ageing on the SOL Hreflex modulation during passive and active shortening and lengthening plantarflexors
actions. Study IV investigated the SOL H-reflex adaptation and its relation to static
postural control after 12 weeks of TC training in older adults.
1.2. Aims of the investigation

The objective of the research was to advance our knowledge in understanding the effect
of ageing on the SOL H-reflex modulation during different ankle joint positions, muscle
contraction intensities, dynamic muscle actions and static postural tasks. This research
also sought to investigate the effect of 12 weeks TC training on the SOL H-reflex
modulation and control of static posture in healthy older adults.
The specific aims of the studies are stated below.
Study I: Test-retest reliability of the SOL H-reflex
The aims of this study were:
1.
To determine the test-retest reliability of SOL H-reflex at the ankle joint positions
of neutral (0), plantarflexion of 20 and dorsiflexion of 20.
2.
To determine the test-retest reliability of the SOL H-reflex during rest and
voluntary contractions at 10%, 30% and 50% of MVC levels in the abovementioned three ankle joint positions.
The details of this study are described in Chapter 4.
Study II: Effects of joint position and muscle contraction intensity on soleus Hreflex gain in young and older adults
The aim of the Study II was:
1.
To examine the age-related differences in SOL H-reflex gain during submaximal

voluntary contractions at 10%, 30% and 50% MVC when the ankle joint was
placed at neutral (0), plantarflexion of 20 and dorsiflexion of 20.
Study III: Effect of ageing on soleus H-reflex modulation during shortening and
lengthening muscle actions in relation to postural control
10
The aims of this study were:
1.
To determine age-related changes in the SOL H-reflex modulation during passive

and active shortening and lengthening muscle actions.
2.
To determine the correlation of the SOL H-reflex modulations between shortening

and lengthening muscle actions and static postural tasks in young and older adults.
Study IV: Effect of 12 weeks Tai Chi training on soleus H-reflex and static
postural control in older adults
The aims of the Study IV were:
1.
To determine the effect of 12-weeks TC training on the SOL H-reflex modulation

during static postural tasks in older adults.
2.
To determine the effect of 12-weeks TC training on static postural control in older

adults.
1.3. Research hypotheses

The following research hypotheses were tested:
1.
The test-retest reliability of the SOL H-reflex measurement is not affected by the
ankle joint position and muscle contraction intensity.
2.
Ageing affects the SOL H-reflex gain during submaximal muscle contractions at
the neutral (0), plantarflexion 20 and dosiflexion 20 positions, because the agerelated differences of neural adaptation.
11
3.
Ageing affects on the SOL H-reflex modulation during dynamic muscle actions
due to the age-related difference of neuromuscular adaptations.
4.
The maximal amplitude of SOL H-reflex between dynamic muscle actions and
static postural tasks are correlated despite the ageing effect.
5.
The maximal amplitude of SOL H-reflex is down-regulated after 12 weeks TC

training.
1.4. Significance of the research

An important aspect of postural control is that modulation of ankle muscle reflexes
plays a crucial role in maintaining upright posture. However, the characteristics of ankle
muscle reflexes during ankle joint movements and postural control is not wellunderstood in older adults. This thesis aims to improve our fundamental knowledge in
understanding motor control strategies in older adults. The studies designed and
presented in this thesis were to investigate age-related changes in reflexive function of
the soleus muscle and neural adaptation to 12-week TC training in older adults. The
age-related differences of the SOL H-reflex modulation during different modes of ankle
muscle actions as well as the contribution of spinal adaptation caused by TC training
and its relation to control of static upright posture in older adults were examined. The
outcomes provided physiological evidence that advanced current
knowledge/understanding in improving balance control and reducing the risk of falls in
old population.
1.5. Study Limitations

1.
This research was limited to investigations on the H-reflex modulation in the SOL
muscle. The outcomes may not be able to explain age-related changes in reflexive
function of other postural muscles, such as quadriceps, hamstrings or erector
12
spinae muscle groups, etc..

2.
Due to the participants schedule and the availability of the research laboratories, it
was difficult to conduct the TC training experiments at the same time of the day.
Despite a controversial finding of the SOL H-reflex modulation during circadian
rhythm was reported in young adults (Guette, Gondin, & Martin, 2005; Lagerquist,
Zehr, Baldwin, Klakowicz, & Collins, 2006), it is not known whether the H-reflex
response of older adults could be affected by circadian variations.
3.
The effects of TC training on H-reflex modulation and balance control in older

adults were limited to 12 weeks of intervention. The effects of longer-term TC
training cannot be addressed due to the limited duration of the Ph.D. candidature.
1.6. Delimitation
1.
The participants recruited for this study were healthy young and older adults. The
explanation and implication of the outcomes were based on these populations.
2.
This research examined H-reflex modulation when the body was in a steady
position. The outcomes of this study may not be applicable to H-reflex modulation
during dynamic activities (e.g., drop jumping, step climbing and walking).
1.7. Ethical approval

The research work presented in this thesis was conducted in compliance with the
Guidelines of the National Statement on Ethical Conduct in Research Involving
Humans (NHMRC). The criteria and standardisation of ethical aspects was supervised
by Southern Cross University Human Research Ethics Committee (EC00137). The
research was approved by the Human Research Ethics Committee of the University
(approval number ECN-08-092) and was performed in accordance with the Declaration
of Helsinki.
13
Chapter 2 Literature review
14
2.1. Introduction
The Hoffmann reflex, known as H-reflex, was originally described by a German
physician, Paul Hoffmann in 1910 (Hoffmann, 1910). The H-reflex test was extensively
used in the 1940s and early 1950s as a neurophysiological measurement and a noninvasive technique to examine the influence of group Ia projection on the activation of
spinal -motoneurons (Magladery, Porter, Park, & Teasdall, 1951). Between the late
1950s and mid 1980s, this test was used to investigate the inhibitory or excitatory
influences of specific pathways on the activation of -motoneurons in conditioning
reflex protocols (Hultborn, 2006). More recently, the H-reflex test has been utilised in
studies on the role of Ia afferent spinal loop in various aspects of human movement, for
example, in relation to performance of functional tasks, such as single and multiple joint
movements, postural control, and locomotion (Zehr, 2002).
The -motoneuron activity is determined by summation of excitatory and inhibitory

inputs from afferent and efferent neural pathways (Pierrot-Deseilligny & Mazevet,
2000). It has been suggested that this reflex test can be used to investigate the effects of
neural inputs from either the peripheral sensory afferents or the descending pathways on
the spinal cord circuitry during motor performance (Hultborn, 2006). Action potentials
generated by group Ia afferent inputs, as indicated by the H-reflex response, is altered
dependently with respect to changes in body orientation (Knikou & Rymer, 2003) and
joint position (Alrowayeh, Sabbahi, & Etnyre, 2005; Chen et al., 2010; Hwang, 2002).
Recent studies have shown that the modulation of H-reflex response is highly correlated
to the phase and direction of postural sway and postural stability (Earles, Koceja, &
Shively, 2000; Taube, Leukel, & Gollhofer, 2008; Tokuno, et al., 2007; Tokuno, et al.,
2008). The size of H-reflex recorded from the triceps surae decreases in association with
15
increased body sway during bipedal stance. This relationship is more obvious during
postural perturbation (Earles, et al., 2000; Taube, Leukel, et al., 2008). In contrast, the
H-reflex amplitude is down-regulated in parallel with improvement of balance control
after one session (Mynark & Koceja, 2002) and repeated sessions of balance training
(Taube, Gruber, et al., 2008).
This review aims to provide an analysis of the current literature on the relationship
between the H-reflex modulation and control of upright standing posture for a better
understanding of the physiological mechanisms involved in postural control. The review
has three major sections. The first section discusses the physiology of H-reflex test and
technical considerations for its validity and reliability. The second section addresses the
supraspinal and spinal mechanisms related to control of posture. The third section
discusses the evidence presented in the literature on the relationship between the Hreflex modulation (mainly the SOL H-reflex) and static postural control.
2.2. Validity and reliability of H-reflex measurement

2.2.1. H-wave and M-wave
The H-reflex is an electrical stimulation-induced analogue of reflex (Zehr, 2002). To

elicit the H-reflex, a percutaneous electrical stimulation is delivered to a peripheral
nerve branch which consists of mixed sensory and motor nerve fibres. When
depolarization of the nerve fibres reaches the threshold level, action potentials are
evoked and propagate on the nerve fibres in both the ascending and descending
directions. Subsequently, two types of compound muscle action potentials, M-wave and
H-wave, can be recorded from the skeletal muscle innervated by these nerve fibres.
16
The M-wave is recorded at the muscle as the result of the evoked action potential
travelling along the -motoneuron axons from the location of the electrical stimulation
to the muscle [response 1 in Figure 2.1 (Aagaard, Simonsen, Andersen, Magnusson, &
Dyhre-Poulsen, 2002)]. The M-wave amplitude reflects the number of the motoneuron axons being activated simultaneously (Tucker, et al., 2005). Under the
same stimulation intensity and testing conditions, this orthodromic action potential
should have a consistent amplitude, because it is determined by the physiological
characteristics of the efferent nerve fibres, neuromuscular junction and muscle fibres,
and is not affected by the sensory inputs and the spinal mechanisms (Frigon, Carroll,
Jones, Zehr, & Collins, 2007). However, the magnitude of M-wave may vary during
movements. The variations in the M-wave size may be caused by changes in muscle
fibre length, the distance between muscle fibres and EMG electrodes, and the spatial
relationship between the nerve and stimulation electrodes during movements (Tucker, et
al., 2005). For example, changes in joint position and level of muscle contraction can
alter action potential propagation along the sarcolemma therefore the size and duration
of M-wave response are changed (Frigon, et al., 2007).
From the location of the electric stimulation, action potentials also travel on the afferent
fibres, particularly the group Ia fibres, to the spinal motoneuron pool, that may
subsequently induce an H-wave (response 2 in Figure 2.1). If sufficient
neurotransmitters are released from the Ia afferent terminals, the -motoneurons will
respond with EPSP. Subsequently, an action potential may be generated on the
motoneuron axon if the EPSP reaches the excitation threshold. The action potential
generated by the -motoneuron propagates along the efferent motor nerve fibre to the
neuromuscular junction (response 3 in Figure 2.1), leading to an action potential on the
muscle fibres it innervates (H-wave). The H-wave recorded at the muscle is the summed
17
action potentials of different motor units, therefore its amplitude may vary depending on
the excitability of individual motoneurons in the -motoneuron pool (Misiaszek, 2003).
The sensory Ia fibres are normally larger, therefore have a lower excitation threshold
than the motor fibres (Panizza et al., 1998). At low stimulation intensities action
potentials are elicited only on the sensory Ia fibres, therefore the H-wave may be
recorded while the M-wave is absent. When the stimulation intensity is high enough to
activate the motor fibres, the evoked action potential on the -motoneuron axons will
travel in both ascending and descending directions from the location of the stimulation.
The ascending (antidromic) action potential on the -motoneuron axons (response 1* in
Figure 2.1) collides with the descending action potential from the spinal cord on the
same axons (response 3 in Figure 2.1). Thus, the H-wave may become smaller or even
disappear with increased stimulation intensity. In contrast, the descending (orthodromic)
action potential on the -motoneuron axons travels from the location of the stimulation
to the muscle (response 1 in Figure 2.1) and results in the M-wave. The amplitude of the
M-wave is getting larger with progressively increased stimulation intensity until the
maximal level (Mmax) is reached (Pierrot-Deseilligny & Mazevet, 2000). With a given
stimulation intensity, the amplitude of the H-wave is a valid indication of the Ia
excitatory effect on the target motoneurons (Pierrot-Deseilligny & Mazevet, 2000).
18
H-wave pathway
The material has been removed
M-wave pathway
Figure 2.1: Simplified illustration of Hoffmann reflex [adopted from Aaggard, et al.
(2002)]. Low intensity electrical stimulation elicits the H-wave (the second compound
EMG wave shown in the EMG amplifier), whereas high intensity electrical stimulation
elicits the M-wave (the first compound EMG wave). Because the axon diameter of the
sensory Ia fibres is relatively larger, their excitation threshold is relatively lower than
other types of fibres. Therefore, low intensity stimulation only elicits action potential on
the Ia afferent axons (response 2) but not on the efferent motor axons. The afferent
action potential induces EPSP on the spinal -motoneurons and may elicit an action
potential on the -motoneuron axons that propagates to the target muscle (response 3).
This response recorded from the muscle is the H-reflex wave. When the stimulation
intensity increases gradually to the threshold level of the -motoneuron axons, action
potential is produced on the axons at the site of stimulation and travels toward both the
muscle (response 1) and the spinal cord (response 1*). The orthodromic action potential
(response 1) propagates to the muscle that results in the M-wave, while the antidromic
action potential (response 1*) travels toward the spinal cord and may collide with the
action potential coming from the -motoneurons (response 3). The latter behaviour may
cause partial or complete cancellation of the H-wave.
2.2.2. Reliability of H-reflex
The reliability of the H-reflex test has been examined in different age populations
(Mynark, 2005), and under various experimental conditions, for instance, in different
body positions (Ali & Sabbahi, 2001; Hopkins, et al., 2000), with and over different
time periods (Clark, et al., 2007), with and without conditioning-test (C-T) stimulations
(Earles, Morris, Peng, & Koceja, 2002), and in the upper (Jaberzadeh, Scutter, Warden19
Flood, & Nazeran, 2004; Stowe, Hughes-Zahner, Stylianou, Schindler-Ivens, & Quaney,
2008) and lower (Palmieri, et al., 2002) limb muscles. In general, these investigations
have indicated that the H-reflex test is a highly reliable assessment in repeated tests. For
example, intraclass correlation coefficients (ICC) in the measurement of peak-to-peak
amplitude of the H-wave over two consecutive days have been reported as 0.99 in the
SOL, 0.99 in the peroneal, and 0.86 in the tibialis anterior (TA) muscles (Palmieri, et al.,
2002).
2.2.3. Methodological concerns in H-reflex assessment
Assessment of the H-reflex requires delivery of electrical stimulation to a peripheral

nerve and recording of action potentials (EMG) from a muscle innervated by the nerve
(Palmieri, Ingersoll, & Hoffman, 2004). The methodological considerations typically
include the configuration, placement and type of the stimulation electrodes, intensity
and duration of the stimuli, inter-stimuli interval (ISI), EMG configuration and location
of electrodes (Palmieri, et al., 2004; Pierrot-Deseilligny & Mazevet, 2000; Tucker, et al.,
2005). Furthermore, the potential effect of fatigue should also be considered in
experimental protocols that involve repeated high intensity muscle contractions or
prolonged testing time (Cresswell & Lscher, 2000; Rupp, Girard, & Perrey, 2010;
Walton, Kuchinad, Ivanova, & Garland, 2002).
Configuration and placement of stimulation electrodes
The H-reflex can be elicited using either unipolar or bipolar stimulation configuration.
The unipolar stimulation requires applying the electrical current to a location on a large
nerve trunk (Pierrot-Deseilligny & Mazevet, 2000). The cathode is placed over a mixed
peripheral nerve, for example, on the tibialis nerve (e.g. on the popliteal fossa) for
20
eliciting the H-reflex in the calf muscles, while the anode is placed on the opposite side
of the limb (e.g. on the patella) (Pierrot-Deseilligny & Mazevet, 2000).
In contrast, the bipolar stimulation is used at a more focused location for stimulation on
the nerve of interest (Palmieri, et al., 2004). Both the cathode and anode are placed on
the same nerve, for instance, on the median nerve for testing the H-reflex of the flexor
carpi radialis (Stowe, et al., 2008). In order to maintain the quality of electrical
stimulation, it is suggested that the stimulation electrodes are secured on the skin by
applying a rubber strap or elastic bandage (Capaday, 1997; Sefton, Hicks-Little, Koceja,
& Cordova, 2007).
Types of stimulation electrodes
It is known that the electrode-skin impedance is affected by the material utilised for
making the stimulation electrodes, which may influence electrical current delivered to
the tissue (Capaday, 1997). However, there is little discussion in the literature on the
optimal material of stimulation electrodes (Pierrot-Deseilligny & Mazevet, 2000;
Tucker, et al., 2005). According to a survey conducted by a European biomedical
research organization, the Surface EMG for a Non-invasive Assessment of Muscles
(SENIAM), the most common material utilised to make the electrodes is Ag/AgCl
(Hermens, Freriks, Disselhorst-Klug, & Rau, 2000). For the size of the electrode, most
laboratories have used a relatively larger electrode for the anode, in the vicinity of 20
cm2, whereas the cathode is relatively smaller, around 2 cm2, using the unipolar
configuration (Tucker, et al., 2005). Capaday (1997) suggested that it was effective to
use Ag/AgCl electrode for the cathode and a large size of metal plate made of stainless
steel or brass for the anode electrode.
21
Stimulation parameters
The optimal intensity of electrical stimulation to induce the H-reflex response is

determined individually (Brinkworth, Tuncer, Tucker, Jaberzadeh, & Turker, 2007). It
is commonly accepted that it is essential to establish an M-wave and H-wave
recruitment curve for a proper investigation of the H-reflex (Pierrot-Deseilligny &
Mazevet, 2000). To establish a complete recruitment curve, the stimulation intensity is
progressively increased from the threshold of the H-wave to a level at which there is no
more increase in the M-wave amplitude. Through this process, the intensity that elicits
threshold response and that elicits the maximal amplitude of H-reflex (Hmax) and Mmax
for each individual can be identified [see Figure 2.2 (Palmieri, et al., 2004)]. Normally,
the amplitude of the Hmax recorded in the SOL muscle is on average approximately 60%
of the amplitude of the SOL Mmax during rest (Tucker & Trker, 2004).
In general, the stimulation intensity used to elicit the Mmax is higher than that to elicit
the H-reflex and Hmax (Capaday, 1997; Tucker & Trker, 2004). The Mmax represents
the activation of all the motor axons innervating the muscle (Pierrot-Deseilligny &
Mazevet, 2000). Once the recruitment curve is established, the Mmax can be used to
normalize the smaller M-waves associated with the H-reflex. The Mmax can also be used
to evaluate the proportion of the entire motoneuron pool activated by the group Ia
afferent inputs by calculation of Hmax/Mmax ratio (Pierrot-Deseilligny & Mazevet, 2000).
In order to avoid any potential systematic changes that affect all parameters in the trials,
it is recommended to record the Mmax under the respective testing conditions as a
reference (Tucker & Turker, 2007).
There has been no common recommendation on the optimal stimulation intensity that is
normalized to the Mmax or Hmax to elicit the H-reflex. Because the M-wave is not
affected by the spinal mechanisms, stimulation intensity at a low percentage of Mmax is
22
commonly used to ensure consistency of stimulation intensity. As illustrated in Figure

2.2, the Hmax usually occurs in company with a small M-wave. However, the threshold
of sensory Ia afferent and efferent motor axons varies individually and is affected by
age (Stein & Thompson, 2006). Thus, it is suggested that the optimal stimulation should
induce the Hmax or an H-wave that is companied by an M-wave at 10 - 25% of the Mmax
throughout the experiment (Palmieri, et al., 2004).
23
Amplitude (mV)
H-reflex
M-wave
0
0
Stimulus intensity
Figure 2.2: Recruitment curve [adapted from Palmieri, et al. (2004)]. The stimulus
intensity is gradually increased from 1 to 7 (arbitrary unit). The H-reflex wave appears
first at lower intensities. The Hmax that occurs at intensity 3 is accompanied with a small
M-wave. With further increase of intensity, the size of M-wave increases accordingly
and the size of H-reflex decreases. When the maximum amplitude of M-wave is reached,
the H-reflex ultimately vanishes.
The commonly used stimulation duration is in the range of 0.5 (Capaday, 1997) to 1.0
ms (Pierrot-Deseilligny & Mazevet, 2000) in H-reflex studies, based on the large time
constant of the sensory nerve fibres (Kiernan, Lin, & Burke, 2004; Lin, Chan, PierrotDeseilligny, & Burke, 2002). Recently, Lagerquist and Collins (2008) compared the
H/M recruitment curves established by short (0.05 and 0.2 ms) and long (0.5 and 1.0 ms)
pulse durations. A left shift of the recruitment curve was demonstrated when long
duration pulses were used. The size of the H-reflex was about 30% of the size of the
Mmax during trials with 1.0 ms pulse duration, and 10% of the Mmax during trials with
0.05 ms pulse duration when the M-wave associated with the H-reflex was controlled at
5% of the Mmax. However, this study showed no significant differences in the values of
Hmax, Mmax, and Hmax/Mmax ratio despite the variations in pulse parameters.
24
The ISI is another parameter that should be considered. The size of H-reflex in response
to repeated stimuli depends upon refractory period of group Ia fibre (Capaday, 1997;
Voigt & Sinkjaer, 1998). Short ISI may diminish the amount of synaptic
neurotransmitters released to the -motoneurons, leading to a reduction of the H-reflex
amplitude, which is known as postactivation depression (Hultborn et al., 1996). It has
been reported that the effect of postactivation depression induced by preceding
electrical stimulation persists for around 8 s (Misiaszek, 2003). The optimal ISI is
therefore recommended for at least 10 s to avoid possible influence of the postactivation
depression (Palmieri, et al., 2004). Furthermore, a randomized stimulus interval is
suggested to minimise the effect of anticipation (Hultborn, et al., 1996; Mynark, 2005;
Mynark & Koceja, 2002; Schieppati, Nardone, Siliotto, & Grasso, 1995; Stein, et al.,
2007).
EMG configuration and electrode location
It is well known that the M-wave and H-reflex waveforms can be affected by recording
techniques, such as configuration and location of the EMG electrodes (Tucker, et al.,
2005; Zehr, 2002).
Bipolar configuration with the surface electrodes placed over muscle belly is most
commonly used for recording H-reflex (Pierrot-Deseilligny & Mazevet, 2000). The
advantage of bipolar configuration is that common noise signals recorded by the two
electrodes can be reduced by cancellation (Tucker, et al., 2005). However, the
disadvantage of bipolar configuration is that a potential risk of crosstalk between tested
muscle and adjacent muscles may occur if the size of the electrodes and inter-electrode
distance are too large (Beck et al., 2005). To minimise this potential problem, it is
suggested that the electrode size be around 1 cm in diameter and the optimal interelectrode distance be 2 cm for recording EMG on large muscles (Hermens, et al., 2000).
25
In contrast, the monopolar configuration consists of one active electrode placed over the
mid portion of muscle bully and one reference electrode placed over the muscle tendon
(Pierrot-Deseilligny & Mazevet, 2000). The inter-electrode distance is not a major
concern in monopolar configuration (Tucker & Turker, 2005). However, the
disadvantage is an inferior noise-to-signal ratio and the potential contribution of
crosstalk between tested muscle and adjacent muscles (Tucker, et al., 2005).
For obtaining reliable EMG recording, the location for EMG electrode placement
should be consistently determined for repeated testing. (Rainoldi, Melchiorri, & Caruso,
2004). The electrodes should not be placed over the motor point of the muscle because
it may alter the amplitude of EMG signals (Mesin, Merletti, & Rainoldi, 2009).
Hermens et al. (2000) have recommended that the proper location should be at midway
between the innervation zone and the insertion of the muscle. Zehr (2002) has
summarized the recommended electrode locations for recording H-reflex on nine upper
limb muscles and seven lower limb muscles.
Influence of fatigue
It is understood that muscle fatigue may alter motor unit recruitment and firing
frequency that can be detected by surface EMG during neuromuscular tests (Cifrek,
Medved, Tonkovi, & Ostoji, 2009). However, the potential influence of muscle
fatigue on the H-reflex is rarely addressed in the literature. This may be due to
controversial findings of the H-reflex modulation in relation to fatiguing exercise
protocols (Cresswell & Lscher, 2000; Rupp, et al., 2010; Walton, et al., 2002).
Nevertheless, caution should be taken that experimental conditions, such as high
intensity of muscle activities, long duration of experiments, and repeated application of
electrical stimulation, might lead to muscle fatigue that may affect the outcomes of Hreflex tests.
26
A recent study demonstrated that constant intensity of electrical stimulation used

throughout the experiment can lead to invalidation of the H-reflex and M-wave results if
muscle fatigue occurs (Rupp, et al., 2010). This study suggested that a re-determined
recruitment curve established by fewer numbers of electrical stimulation is helpful for
validation of the H-reflex parameters during the recovery period after an exhaustive
muscle testing protocol. However, the practicality of such a recommendation is still to
be revealed.
2.2.4. Limitations of H-reflex test
It should be noted that the H-reflex test is not a robust measurement. It could result in a
misinterpretation if the researchers neglected the physiological characteristics of the Hreflex. There are several limitations that should be emphasized when the H-reflex test is
utilised.
First, it is important to bear in mind that the spinal cord circuitry is oligosynpatic
(Pierrot-Deseilligny & Burke, 2005). Any neural input from the supraspinal and
peripheral pathways can potentially influence the H-reflex modulation through either
direct regulation of the motoneuron membrane potential or indirect regulation of
interneuronal circuitry (Misiaszek, 2003). The participants physiological and
psychological status during the experiment can affect the H-reflex results. For example,
it has been demonstrated that postural anxiety can depress the H-reflex modulation
(Sibley, Carpenter, Perry, & Frank, 2007).
Secondly, even under consistent experimental conditions, the H-reflex amplitude is

fractal and may vary temporally in time (Nozaki, Nakazawa, & Yamamoto, 1995). It is
necessary to normalize the H-reflex response to either the Mmax or controlled H-reflex
for appropriate interpretation of the H-reflex result and minimizing inter- and intra27
subject variability. To do so, it is necessary to apply extra electrical stimulations under

the respective experimental conditions that may cause discomfort to some individuals.
Thirdly, the H-reflex response may not be observed in all experimental trials for all
participants. It is possible that the H-reflex test is unsuccessful in a small proportion of a
healthy population under some experimental conditions. Unpublished observation in our
laboratory found that the SOL H-reflex response was present during passive shortening
movement but was absent during passive lengthening movement in a small number of
older adults. The mechanisms for the absence of H-reflex response during lengthening
plantarflexors muscle activity are unclear, but it is speculated to be related to quiescence
of muscle spindle activity. It has been suggested that the failure of H-reflex elicitation in
the older population might be due to age-related spinal stenosis (Egli et al., 2007).
In summary, H-reflex is a short-latency reflex which has been utilised to examine the
effects of Ia afferent inputs on the -motoneuron activation. High test-retest reliability
of H-reflex has been reported in the literature. The size of the H-wave can be affected
by recording methods and fatigue. Therefore, both technical and physiological factors
should be carefully considered for valid measurement of the H-reflex response.
2.3. Control of posture

The neural strategies involved in postural control are complex but all relate to the
interactions between sensory and motor processes at the supraspinal and spinal levels.
2.3.1. Sensory inputs
Somatosensory inputs
28
Somatosensory cue refers to proprioceptive and cutaneous sensations. Muscle spindles,

Golgi tendon organs and joint receptors are sensory sources of proprioception which
detect body displacement in relation to changes in muscle length and joint movement
(Shaffer & Harrison, 2007). The cutaneous receptors include nociceptors and thermal
receptors that detect conditions of cutaneous and subcutaneous tissues, such as pressure,
mechanical stimuli, temperature and pain (Sayenko et al., 2009; Shaffer & Harrison,
2007). Somatosensory feedback is essential in execution of motor tasks (Brooke & Zehr,
2006) including adjustment of posture during standing (Horak, Earhart, & Dietz, 2001),
particularly when visual and vestibular inputs are restricted (Winter, Allen, & Proske,
2005). For example, sensory information from the cutaneous receptors can identify
slight changes in ankle displacements (Meyer, Oddsson, & De Luca, 2004). An increase
in the afferent flow from the cutaneous receptors occurs to mediate postural reflexes
when there is an increased external perturbation (McIlroy et al., 2003). A recent study
(Sayenko, et al., 2009) demonstrated that cutaneous afferents activated from different
locations on the sole of the foot resulted in alterations of the H-reflex modulation. For
example, a conditioning stimulation applied to the metatarsal region led to inhibition of
the SOL H-reflex response, whereas a conditioning stimulation delivered to the heel
region facilitated the SOL H-reflex response.
Visual inputs
Vision provides a reliable source of sensory information for the CNS, including
contextual cues and trajectory movement of surrounding objects (Franklin, So, Burdet,
& Kawato, 2007). Through visual feedback, the supraspinal mechanisms can accurately
modify spatial orientation of the body to adapt postural behaviour and movement
(Duarte & Zatsiorsky, 2002; Wessberg & Vallbo, 1995). A recent study measuring the
H-reflex and changes of centre of pressure (COP) showed that an increase in postural
29
stability was associated with facilitation of the H-reflex amplitude when a visual target
was provided (Taube, Leukel, et al., 2008). In contrast, deprivation of visual feedback is
known to impair postural control (Hafstrm, Fransson, Karlberg, Ledin, & Magnusson,
2002; Hue et al., 2007; Van Deun et al., 2007) and to depress the H-reflex amplitude
(Earles, et al., 2000)
Vestibular inputs
The sensory receptors in the vestibular system are sensitive to any change in motion and
orientation of the head (Horak, Shupert, Dietz, & Horstmann, 1994). When the head
position changes, vestibular sensory signals are elicited by the receptors in the otoliths
organ and semicircular canals and contribute to the perception of body position for
spatial awareness. The functional roles of the vestibular system are complex but
generally related to facilitation of postural reflexes (Horak, et al., 2001). Vestibulospinal
inputs can directly affect the spinal motoneuron activation. This is shown in
experiments that use a galvanic vestibular stimulation (GVS) to elicit vestibular afferent
flows prior to an H-reflex stimulation (Kennedy, Cresswell, Chua, & Inglis, 2004). For
example, Lowrey and Bent (2009) utilised a bipolar binaural GVS 100 ms prior to
elicitation of the H-reflex response in the SOL muscle. The result showed that the SOL
H-reflex conditioned by GVS was increased about 20% compare to that without GVS.
Thus, it appears that the vestibular afferent is able to influence the SOL motoneuron
activity even in a prone position.
2.3.2. Supraspinal mechanisms
In the brain region, there are many sources of descending motor signals involving in
control of movement. This section focuses on a major contribution of cerebral motor
30
cortex, cerebellum, basal ganglia and brainstem to motor output regulation during
postural control
Cerebral motor cortex
The motor system generates appropriate motor commands after decoding and
integration of various sensory signals. The descending drive during voluntary activity is
initially executed by the motor cortex (Keller, 1993). Recent evidence with use of
transcranial magnetic stimulation (TMS) (Soto, Valls-Sole, Shanahan, & Rothwell,
2006; Taube, Gruber, et al., 2007; Taube et al., 2006; Tokuno, et al., 2008) or
electroencephalography (EEG) (Slobounov, Hallett, Cao, & Newell, 2008) has
demonstrated that the motor cortex plays an essential role during postural corrections.
For example, Solopova and colleagues (2003) reported that there was a facilitation of
motor evoked potentials (MEPs) in postural adjustment during standing on a rocking
platform. Increase in cortical excitability was observed with the size of MEPs increasing
2.2 times when the participants shifted from a stable surface to a rocking platform.
Cerebellum
The cerebellum contributes to balance and locomotion controls (Ioffe, Chernikova, &
Ustinova, 2007). Most sensory inputs transmitted to the cerebellum are carried to the
cerebellar cortex via mossy or climbing fibres. Sensory volleys from the spinal cord,
brainstem, and cerebral cortex are conveyed by mossy fibres and then reach the Purkinje
cells through granule cells and parallel fibres for ongoing sensorimotor processes.
Climbing fibres have a nature of slow discharge rate and modulate the firing pattern of
the Purkinje cells in relation to learning and error adjustment. The Purkinje cells are
responsible for organization of the cerebellar outputs. The neural processes via the
31
cerebellum are important for limb movement, control of balance and locomotion and
can become more automatic by means of postural learning processes (Ioffe, et al., 2007).
The cerebellum has a complex neural network in conjunction with many cortical,
subcortical and spinal areas. Basically, it can be divided into three different regions and
each cerebellar zone has its specific characteristics of motor functions (Morton &
Bastian, 2004). The medial zone controls the muscle tone of the antigravity muscles
during upright standing and rhythmic muscle activity during locomotion. The
intermediate zone provides an essential function in temporal and spatial adjustments of
limb movement during locomotion. The lateral zone plays a role in regulation of
walking pattern in novel tasks and association with visual feedback for gait control.
The functional role of the cerebellum involved in the H-reflex modulation has been
addressed in patients with cerebellar ataxia (Tokuda, Tako, Hayashi, & Yanagisawa,
1991) and atrophy (Mauritz, Schmitt, & Dichgans, 1981) during postural tasks. Patients
with a cerebellar lesion demonstrated a facilitation of SOL H-reflex during upright
standing in comparison with age-matched healthy counterparts (Tokuda, et al., 1991).
The potentiation of Ia afferent feedback was correlated to postural instability in the
cerebellar patients. Moreover, an earlier observation in patients with late cortical
cerebellar atrophy showed a potentiation and delay of gastrocnemius H-reflex during
bipedal standing (Mauritz, et al., 1981).
Basal ganglia and brainstem
The basal ganglia play important roles in control of automatic and voluntary movements
(Middleton & Strick, 2000; Takakusaki, Saitoh, Harada, & Kashiwayanagi, 2004). They
receive descending inputs from the cerebral cortex via cortico-basal ganglia loop and
contribute to voluntary control of movements. The basal ganglia also link to the
32
brainstem and serve a function in automatic control of postural muscle tone and
locomotion. The automatic control process in the basal ganglia-brainstem network is
dominated by the basal ganglia without motor cortical projection. When the GABAergic
projection from the basal ganglia reaches the brainstem, the pedunculopontine
tegmental nucleus (responsible for postural muscle tone) or the midbrain locomotor
region (responsible for locomotion) located in the mesopontine tegmentum are activated
for their respective function (Takakusaki, Oohinata-Sugimoto, Saitoh, & Habaguchi,
2004).
Deficits of the basal ganglia function usually accompany many movement disorders,
such as muscle weakness, rigidity, involuntary spasm and prolonged reaction time.
These movement-related changes have been commonly observed in patients with
dystonia, Parkinsons and Huntingtons diseases (Berardelli et al., 1998; Berardelli,
Rothwell, Thompson, & Hallett, 2001). The main characteristic is that these patients
have abnormality of inhibitory regulation in the main output nuclei, resulting in either
hypokinesia or hyperkinesia. In clinical investigations, a conditioning-test stimulation
with a short interval (a few milliseconds) has been utilised to estimate reciprocal
inhibition of the H-reflex in patients with dystonia (Berardelli, et al., 1998). The
existing evidence shows that dystonia reduces the degree of reciprocal inhibition and is
possibly regulated by the supraspinal mechanisms. A lack of efficiency of the spinal
reciprocal control causes strong muscle cocontractions and irrelevant muscle activities
during voluntary and involuntary movements in dystonic patients (Berardelli, et al.,
1998).
2.3.3. Spinal mechanisms
The spinal neurons integrate sensory afferent inputs and tonic drives from the
descending tracts for organizing motor functions in postural correction (Beloozerova,
33
Sirota, Orlovsky, & Deliagina, 2006; Hultborn, 2006) [Figure 2.3 (Gandevia, 2001)].
The reflex induced by group Ia afferent inputs have been described in the previous
sections. There are also other spinal mechanisms involved in sensorimotor regulation
during postural control. In this section, the structural connections and functional
importance of other inhibitory mechanisms such as presynaptic, reciprocal, recurrent
and Ib inhibitions are discussed.
Figure 2.3: A schematic diagram of spinal networks [adopted from Gandevia (2001)].
Open circles represent excitatory interneurons, whereas the filled circles represent
inhibitory interneurons. Sensory inputs from group I, II, III, and IV afferent fibres
converge toward the -motoneuron (MN). These inhibitory and excitatory interneurons
mediate the release of neurotransmitters of the -motoneurons. Factors that limit the
excitation of -motoneurons also include motor drives from descending pathways and
inhibitory adjustment from the Renshaw cells. Descending inputs can directly or
indirectly influence the inhibitory and excitatory interneurons, and motoneurons,
and Renshaw cells in the spinal circuitry. Descending inputs may selectively recruit a
particular population of interneurons and motoneurons, depending upon the state of
movement.
34
Presynaptic inhibition
Neural inputs from the group Ia (Stein, 1995), Ib (Guissard & Duchateau, 2006), II
(Aggelopoulos, Chakrabarty, & Edgley, 2008), III and IV (Iles, 1996) afferents, and the
descending tracts (Jankowska, 1992) can influence the amount of synaptic
neurotransmitters released by the nerve endings at the presynaptic level via selective
control of inhibitory interneurons. Presynaptic inhibition is a GABAergic synaptic
process, which is associated with primary afferent depolarization (Knikou, 2008). The
inhibitory interneurons release axo-axonal gamma-aminobutyric acid (GABA) that
causes depolarization of the nerve ending. The amount of the neurotransmitters released
by the nerve ending is associated with the magnitude of the action potential. A priori
depolarization will reduce the magnitude of the action potential thereby reducing the
efficacy of the synaptic transmission (Jankowska & Hammar, 2002). Presynaptic
inhibition of the Ia afferent is thought to be a crucial mechanism for mediating synaptic
efficacy between the Ia afferent fibres and -motoneurons (Stein, 1995). This neural
action causes insufficient neurotransmitters to be released to the Ia afferent terminals,
leading to a depression of motoneuron excitability (Rudomin & Schmidt, 1999) [Figure
2.4 (Widmaier, Raff, & Strang, 2008)].
35
Presynaptic
inhibition
Ia afferent
volley
Presynaptic
membrane
Synaptic
cleft
Neurotransmitters
Postsynaptic
membrane
Postsynaptic
potential
Figure 2.4: A sketch of presynaptic inhibitory process. A presynaptic synapse from last
order of interneuron can modulate the amount of neurotransmitters released from the Ia
afferent volley, leading to a change in postsynaptic membrane potential [adapted from
Widmaier, et al. (2008)].
The functional significance of presynaptic inhibition is for fine control of movement

(Zehr, 2002). For example, it has been found that presynaptic inhibition of the Ia
afferent is stronger during standing than that during lying, and during running than that
during walking (Stein, 1995). This inhibitory mechanism prevents saturation of afferent
volleys and overdrive of motoneuron activation (Pierrot-Deseilligny, 1997) and plays an
important role in sensorimotor processes for postural adjustment (Taube, Gruber, et al.,
2008).
Reciprocal inhibition
Reciprocal inhibition plays an important role in muscle coordination and joint

stabilization (Crone, Hultborn, Jespersen, & Nielsen, 1987). Reciprocal inhibition refers
to concurrent depression of motoneuron activation in antagonistic muscles, with
36
motoneuronal activation in agonistic muscles. Generally, reciprocal inhibition involves

a di-synaptic pathway and at least one Ia inhibitory interneuron (Hultborn, 2006). The Ia
inhibitory interneuron is controlled via both the supraspinal and peripheral pathways
(Hultborn, Illert, & Santini, 1976b; Leonard, Sandholdt, & McMillan, 1999; Nielsen &
Kagamihara, 1992). The corticospinal tract has monosynaptic projection to the target
motoneurons of the agonistic muscle, with a parallel projection to the Ia inhibitory
interneurons (Crone & Nielsen, 1989; Nielsen & Kagamihara, 1992; Petersen, Morita,
& Nielsen, 1998). While the motoneurons of the agonistic muscle are activated, Ia
inhibitory interneurons are activated simultaneously. The inhibitory interneurons have
synaptic connections with the motoneurons of the antagonistic muscle and cause IPSP
and therefore suppress the activation of the antagonistic muscle (Nielsen & Kagamihara,
1992). The Ia inhibitory interneurons can be activated by sensory inputs from the
muscle spindles, joint, and cutaneous receptors (Hultborn, Illert, & Santini, 1976a) and
Renshaw cells (Baret, Katz, Lamy, Penicaud, & Wargon, 2003) in the spinal segmental
pathways [Figure 2.5 (Pierrot-Deseilligny & Burke, 2005)]. Different degrees of
reciprocal Ia inhibition may be related to central regulation of the Ia inhibitory
interneurons and influence of peripheral afferents on motoneuron activation (Hwang,
Lin, & Ho, 2002).
37
Spinal cord
Cutaneous &
joint afferents
Ia
afferent
The material has
been removed
Renshaw cell
Tibialis
anterior
Ia interneuron
Soleus
Figure 2.5: Illustration of reciprocal Ia inhibitory pathway. In this diagram, open circles
represent excitatory interneurons; filled circles represent inhibitory interneurons; the
thick dash line represents Ia afferent volley; the thin dash line represents afferent inputs
from joint and cutaneous receptors, Y-shaped lines indicate excitatory synapses, and
small filled circles represent inhibitory synapses. The soleus (SOL) muscle is the
agonist, whereas the tibialis anterior muscle is the antagonist. During plantarflexion
contraction, Ia afferent input from the SOL muscle has an excitatory effect on the SOL
motoneurons. Simultaneously, Ia afferent also excites Ia inhibitory interneurons,
resulting in inhibition of antagonistic motoneurons. Joint and cutaneous afferent inputs
as well as the Renshaw cells affect the Ia interneuron that may produce an inhibitory
effect on the antagonistic muscle. It should be noted that the influence of descending
control on reciprocal inhibition is not showed in this illustration [adapted from PierrotDeseilligny & Burke (2005)].
Recurrent inhibition
Recurrent inhibition involves the Renshaw cells which are located in the ventromedial
region of the spinal ventral horns (Renshaw, 1946), and a polysynaptic pathway as
illustrated in Figure 2.3 (Gandevia, 2001; Hultborn, 2006). The Renshaw cells are
activated by orthodromic and antidromic impulses from collateral branches of the motoneuron axons and produce an inhibitory effect on the -motoneurons by releasing
inhibitory neurotransmitters (Knikou, 2008). The Renshaw cells also connect with the 38
motoneurons and Ia inhibitory interneurons for regulating inhibitory function in the

spinal circuitry (Hultborn & Pierrot-Deseilligny, 1979a). Furthermore, the degree of
recurrent inhibition can be indirectly influenced by sensory inputs from the cutaneous
and group II and III afferent fibres, leading to a presynaptic excitatory or inhibitory
effect on the - and -motoneurons (Knikou, 2008). The activity of the Renshaw cells is
also controlled by the descending drives to mediate motoneuron discharging rate and
recruitment (Baret, et al., 2003).
Recurrent inhibition serves as input-output regulator for optimal gain of muscle force
production (Katz & Pierrot-Deseilligny, 1999). It is evidenced that recurrent inhibition
is facilitated during weak muscle contraction but is depressed during strong muscle
contraction (Hultborn & Pierrot-Deseilligny, 1979b). Increase in inhibitory effect during
weak voluntary contraction would be suitable for fine movements, whereas decrease in
inhibitory effect during strong voluntary contraction would favour a large force output
(Hultborn & Pierrot-Deseilligny, 1979b).
Recurrent inhibition is also important to adjust reflex responses between the postural
muscles during standing. Barbeau et al. (2000) employed conditioning electrical
stimulation to the femoral nerve prior to recording the SOL and TA H-reflex during
postural tasks. This conditioning technique was used to investigate the functional status
of recurrent inhibition. The authors reported that recurrent inhibition was decreased in
active ankle muscles during postural tasks that required co-contraction of the synergetic
muscles. For instance, during standing with leaning backward position the quadriceps
and TA muscles were contracting to maintain appropriate posture. In this situation,
recurrent inhibition from the quadriceps to the TA did not manifest. Similarly, recurrent
inhibition from the quadriceps to the SOL muscles was reduced during a squat position
39
(the hip and knee joint angles both at 150) when these two muscle were engaged to
maintain posture.
Ib inhibition
The Ib afferent fibres are from the Golgi tendon organs and cause autogenetic inhibition
(Knikou, 2008). This pathway has synaptic connections with muscle spindle, joint and
cutaneous afferents for reflex regulation during movement (Knikou, 2008). The tendon
organs monitor the tensile force of muscle tendon during passive movements and active
muscle contractions (Shaffer & Harrison, 2007). When muscle tension increases to a
certain level, the Ib afferents may activate Ib inhibitory interneurons that cause IPSP on
the agonistic and synergistic motoneurons (Hultborn, 2006). The Ib sensory feedback is
important to prevent overstretch of contractile components and to adjust muscle
stiffness (Knikou, 2008). In humans, Ib sensory inputs from the leg muscles has a
critical function in mediating gait pattern during locomotion (Faist et al., 2006; Stephens
& Yang, 1996). For example, during the stance phase of walking, Ib sensory inputs
from the medial gastrocnemius (MG) nerve excites the SOL motoneuron activation.
Faist et al. (2006) found that during simple postural tasks (lying, sitting, or standing),
the influence of Ib inhibition was present during resting in lying and sitting positions
but was reduced during weight loading conditions (e.g. against a load or standing).
However, the functional role of Ib inhibition to postural control is still not fully
understood.
In summary, maintaining an upright stance posture is a complex behaviour that requires

sensory information from proprioceptive, visual, vestibular and cutaneous sensory
receptors, integration of sensory inputs and motor commands at the supraspinal and
spinal levels, and the execution of postural correction by the spinal motoneurons. The
spinal motoneuron activation reflects the outcome of the sensory-motor integration.
40
2.4. H-reflex modulation in relation to control of static posture

2.4.1. Biomechanical characteristics of standing
Change in body position in respect to vertical alignment is usually expressed as body

sway. During quiet standing on a stable surface without any external perturbation the
body oscillates in a pendulum model (Winter, et al., 1998). The ankle joints constantly
move within a range of 1.0-1.5 in the anterior-posterior direction (Gatev, et al., 1999).
The direction of body sway is inverted at a mean frequency of 1.3 Hz which
corresponds to 2.6 times of postural reversion per second (Loram, et al., 2005b).
Although postural control involves adjustments in multiple joints, it is suggested that
the ankle muscles serve as the main modulator for ankle movement and sway angle
during postural sway (Winter, et al., 2001). In response to a disturbance in posture, force
is produced to correct the shift of the COM within the base of support for maintaining
postural equilibrium (Horak, 2006). The force distribution on the ground known as the
COP can be used to reflect body sway during upright stance (Winter, et al., 1998;
Winter, et al., 2001). However, changes in COP only describe the biomechanical
behaviour of body sway as the outcomes of neural regulation. In order to examine the
neural mechanisms associated with the biomechanical changes in posture control,
several experimental methods have been frequently used, including the sensory
organization test (SOT) (Tsang & Hui-Chan, 2004a), EEG (Slobounov, et al., 2008),
transcranial electrical stimulation (TES) (Tokuno, et al., 2008), TMS (Taube, et al.,
2006), and H-reflex test (Koceja, Trimble, & Earles, 1993; Tokuno, et al., 2007; Tokuno,
et al., 2008).
41
2.4.2. H-reflex modulation during postural tasks
It has been previously found that there is a relationship between the amplitude of Hreflex and the magnitude of COP displacement (Earles, et al., 2000; Huang, Cherng,
Yang, Chen, & Hwang, 2009; Laudani, Wood, Casabona, Giuffrida, & De Vito, 2009;
Taube, Leukel, et al., 2008). The H-reflex amplitude deceases in association with
increase in mean COP displacement (Huang, et al., 2009). It is suggested that the SOL
H-reflex modulation is related to postural stability (Earles, et al., 2000; Koceja, Markus,
& Trimble, 1995). Recently, Tokuno et al. (2008) have examined the position- and
direction-dependent modulation of SOL and MG H-reflex in relation to anteriorposterior direction of body sway [see Figure 2.6 (Tokuno, et al., 2008)]. In the
direction-dependent experiment, the size of the triceps surae H-reflex was larger when
swaying in a forward direction than that when swaying in a backward direction (9-14%
difference). This result was in line with their early work that showed a greater
Hmax/Mmax ratio when the COP displacement was shifted in forward direction than that
in a backward direction (12% difference in the SOL and 23% in the MG) (Tokuno, et al.,
2007). In the position-dependent experiment, it was found that there was an additional
8.9% increase in the H-reflex amplitude when swaying further in a forward position,
compared to swaying slightly in a forward position. It can be argued that the finding
under position-dependent conditions was contrary to the relationship between the Hreflex amplitude and COP displacement found in previous studies (Earles, et al., 2000;
Huang, et al., 2009; Laudani, et al., 2009; Taube, Leukel, et al., 2008). One possible
explanation is that the SOL muscle activation in the further anterior COP position was
stronger than that in the slight anterior COP position, as indicated by baseline EMG
(Tokuno, et al., 2008). It has been shown that the SOL H-reflex response is facilitated
when the muscle activation increases (Stein, et al., 2007).
42
Figure 2.6: The H-reflex modulation in the soleus (SOL) and medial gastrocnemius
(MG) muscles during task-dependent body sways [adopted from Tokuno, et al. (2008)].
The body sways were determined by mean and standard deviation of the mean distance
in the anterior-posterior direction of COP measurement during a 90 s quiet standing.
This baseline of COP was used to compare the effects of four testing conditions in this
experiment: A: mean - 1.6 SD during forward sway; B: mean + 1.6 SD during forward
sway; C: mean + 1.6 SD during backward sway; and D: mean - 1.6 SD during backward
sway. The 1.6 SD was set to the testing conditions due to that the COP baseline signal
in addition to 1.6 SD resulted in significant difference of EMG responses in the calf
muscles during unpredictable surface perturbation. a) The H-reflex data is recorded in
different COP positions (position effect) during forward and backward body sway
(direction effect). For the position effect, the H-reflex recorded in the A and D
conditions is presented by black solid lines, whereas data recorded in the B and C
conditions is presented in gray solid lines. For the direction effect, the H-reflex value
recorded in the C and D conditions is expressed as black solid lines, whereas that in the
A and B conditions is presented as gray dash lines. b) The H-reflex amplitudes in the
mean - 1.6 SD conditions and negative direction are presented as the control of H-reflex
size. Significant facilitation of the H-reflex is observed in the mean + 1.6 SD conditions
and positive direction in the respective experimental conditions.
Influence of somatosensory inputs
It has been reported that the SOL H-reflex amplitude decreases when the postural
condition becomes more challengeable, such as standing on an unstable surface (Earles,
et al., 2000; Hoffman & Koceja, 1995). The decrease in H-reflex size observed during
postural perturbation is thought to be related to increased Ia feedback that leads to
depression of the SOL motoneuron activation (McIlroy, et al., 2003). This inhibitory
effect is suggested to prevent overdrive of autogenic excitation of motoneurons and to
43
alter saturation of motoneuron excitability for receiving descending commands from

supraspinal levels (Chalmers & Knutzen, 2002). Sefton et al. (2007) utilised a pairedstimulation technique to evaluate the possible effect of inhibitory mechanisms on the
SOL H-reflex modulation during one-leg standing under stable and unstable surface
conditions. A paired stimulation was applied to the tibial nerve with an ISI of 80 ms in
order to induce presynaptic inhibition. As expected, an increase in presynaptic
inhibition was observed under unstable surface condition as the H-reflex size was
smaller compared to that recorded under stable surface condition. Because there was no
change in the M-wave under different surface conditions, the authors speculated that the
depression of SOL H-reflex occurred at the presynaptic level rather than at the
postsynaptic level.
Influence of visual inputs
Visual inputs are known to be a predominant sensory modality over proprioceptive cues
when humans execute a variety of motor tasks (Wessberg & Vallbo, 1995). The CNS
controls voluntary movement based upon the perceptive inputs from visual tracing
(Brozovic, Gail, & Andersen, 2007). However, few studies have provided solid
evidence for the importance of visual inputs to H-reflex modulation during postural
tasks (Earles, et al., 2000; Hoffman & Koceja, 1995; Taube, Leukel, et al., 2008). Taube
et al. (2008) designed an experiment to investigate the possible influence of visual
feedback on the SOL H-reflex modulation in relation to control of upright standing
posture. Twenty-two healthy young participants were instructed to stand on stable and
unstable surfaces with two visual conditions: with and without staring at a laser pointer.
The trials with a laser pointer showed smaller COP displacements compared to the trials
without a laser pointer under the same surface condition. In contrast, the amplitude of
SOL H-reflex recorded with a laser pointer slightly increased during standing on the
44
stable surface but increased significantly during standing on the unstable surface. These
results may indicate that enhancement of visual inputs favoured stabilisation of postural
sway and facilitation of the H-reflex. Similar findings were reported in two previous
studies (Earles, et al., 2000; Hoffman & Koceja, 1995), in which there was an increase
in body sway and a decrease in the size of SOL H-reflex when visual information was
deprived.
Influence of vestibular inputs
Knikou and Rymer (2003) investigated the effect of static and dynamic changes in body
orientation on the SOL H-reflex modulation. In the static test, the participants were
placed in a supine position on a tilt table which was positioned at four forward angles
(10, 20, 40, and 60), vertical position (90), and two backward angles (-20 and 50). A baseline SOL H-reflex was recorded in the horizontal supine position. The
results revealed that the SOL H-reflex recorded at these angle positions increased to
140-180% of the baseline level. Facilitation of the SOL H-reflex was also observed
during the dynamic movement of body position in the sagittal plane at a constant
angular velocity of 1.8/s. Interestingly, the facilitation of SOL H-reflex recorded
during the dynamic movement was three times larger, while that during the static
condition was 1.6 times larger, in comparison with the baseline. The significant increase
in the SOL H-reflex in the dynamic condition compared with that in the static condition
was suggested to be a result of greater vestibulospinal inputs (Kennedy, et al., 2004).
The findings are in agreement with a previous report in that the magnitude of SOL Hreflex was increased when the body tilt was set in either forward or backward positions
in the sagittal plane, compared to the SOL H-reflex in a upright position (Chan &
Kearney, 1982). These results suggest that the input from the vestibular system has an
effect on the regulation of the spinal motoneuron activation when the body position is
45
altered. However, Trimble (1998) reported no significant changes in the SOL H-reflex
amplitude in six static positions of body tilt. The controversial findings could be related
to different methods utilised to record the SOL H-reflex in these studies, because in the
study by Trimble (1998) a strap and a traction harness were used to support the
participants head and body weight, whereas these experimental settings were not
employed in Knikou and Rymers study (2003).
2.4.3. Effect of balance training on H-reflex modulation
Change in the H-reflex has been used as an indication of functional adaptations of the
spinal motoneurons to training tasks (Taube, Gruber, et al., 2008; Wolpaw, 2007). It is
known that spinal plasticity plays a pivotal role in development of motor skills (Wolpaw,
2007). It is also known that the spinal motoneurons not only are responsible for simple
reflex interaction but also have a modulatory effect on the spinal circuitry in adaptation
to given training tasks (Wolpaw & Carp, 2006).
The H-reflex modulation can change immediately after acute balance training. Trimble
and Koceja (1994) reported that there was a down-regulation of the H-reflex of the
triceps surae after two hours of balance training. The participants practised seven sets of
balance training on an unstable balance board, then were immediately assessed for the
H-reflex when standing on a stable surface. The result indicated that the H-reflex
decreased to 26.2% of that of the pre-training test. To further support the short-term
effect of balance training, Trimble and Koceja (2001) used the identical balance training
protocol to investigate the modulation of H-reflex response in three days of training.
Two thirds of the participants (six in nine) demonstrated a downward modulation (1242% decrease) of the SOL Hmax/Mmax ratio after the balance training. Similar downregulation of H-reflex immediately after acute balance training was also found in an
older population (Mynark & Koceja, 2002). Mynark and Koceja (2002) investigated the
46
effect of balance training on the SOL H-reflex in ten young and ten older participants in
two consecutive days. The Hmax/Mmax ratio (day 1: -18.7 %; day 2: -21.0 %) and body
sway (- 10.1 % of COP measurement, only assessed in day 2) during standing both
decreased after training.
The adaptive change of SOL H-reflex is also observed after short-term exercise training.
The changes of the SOL H-reflex only occur when performing the trained tasks but not
during rest and performing non-trained tasks, indicating specificity of neural plasticity
(Schubert et al., 2008; Taube, Gruber, et al., 2007; Taube, Kullmann, et al., 2007). The
specificity of training effect on the H-reflex may be related to the changes in cortical
motoneuron excitability. Schubert et al. (2008) investigated the effect of four weeks of
balance and strength training on the spinal and corticospinal adaptations. The SOL Hreflex was used to investigate the spinal adaptation, while the SOL H-reflex conditioned
with subthreshold intensity of TMS was used to measure the corticospinal changes. A
total of thirty-seven young participants were allocated to each of the balance training,
strength training and control groups. After the training, no significant change in the
SOL Hmax/Mmax ratio was found in any of the three groups. However, the balance
training group showed a down-regulation of conditioned SOL H-reflex during postural
perturbation, whereas the strength training group demonstrated a facilitation of the
conditioned SOL H-reflex during postural perturbation. This observation was in
agreement with the researchers earlier report (Taube, Gruber, et al., 2007) in which
they demonstrated that after four weeks of balance training the SOL Hmax/Mmax ratio and
conditioned SOL H-reflex were unaltered when they were recorded at about 40 ms after
the onset of stance perturbation. In contrast, significant decrease in the SOL Hmax/Mmax
ratio and conditioned SOL H-reflex was found when it was recorded at about 120 ms
after onset of stance perturbation. The latter experimental condition was set to involve
47
the central processing mechanisms. Such evidence underscores the effect of cortical
adaptation to training on the H-reflex modulation.
It should be noted that reduction in the H-reflex amplitude associated with increase in
postural stability after balance training is a different neural behaviour to the decreased
H-reflex size during postural disturbance. The depression of H-reflex modulation during
postural disturbance is a result of spinal inhibitory regulation. This acute response is
suggested to alter the Ia excitatory effect on the target motoneurons in order to obtain
adequate descending commands for postural correction (Chalmers & Knutzen, 2002). In
contrast, down-regulation of the H-reflex found after balance trainings is a functional
status resulting from more effective regulation of motor outputs during standing (Taube,
Gruber, et al., 2008). In these training studies, the participants learned how to maintain
postural equilibrium on unstable materials (usually balance board or soft mattress).
Significant reduction of the H-reflex amplitude associated with increased body
stabilisation may be a result of altered motor efficiency after balance training.
2.5. Effect of Tai Chi training on neuromuscular adaptations in older

adults
Tai Chi (TC) is a Chinese martial art exercise, which was originally developed as a selfdefense skill (Wu, 2002). The original TC exercise is designed for fighting and consists
of more than 100 movements. Exercise intensity of the original TC routine can be quite
high and may not be suitable for general populations as a recommended exercise
intervention. Thus, modified TC routines with fewer number and slower movements
have been developed and practised by general populations (Verhagen, Immink, van der
Meulen, & Bierma-Zeinstra, 2004).
48
2.5.1. Muscular adaptations
Tai Chi is a popular exercise for promotion of muscular fitness in the older population
(Xu, Hong, & Li, 2008). In older adults, muscle strength is one of essential elements for
control of balance (Orr, 2010). Tai Chi exercise can be performed with slow and gentle
movements. Because it is weight-bearing, all TC movements require adequate level of
strength and control of lower limb muscles during unipedal and bipedal stance. It has
been shown in older adults that practice of TC exercise can improve lower limb muscle
strength, compared with their sedentary counterparts (Choi, et al., 2005; Li, et al., 2009;
Xu, et al., 2008). Xu et al. (2008) reported that TC practitioners with a minimum of 4
years practice demonstrated better muscular strength of the knee extensors and flexors
and ankle dorsiflexors than their sedentary counterparts in isokinetic muscle strength
tests. Furthermore, a 19.9% increase in the maximal concentric muscle strength of the
knee flexors after a 16-week of TC training was reported by Li et al. (2009). Significant
improvements in the isometric muscle strength of knee flexors and ankle plantarflexors
and dorsiflexors were also observed after a 12-week TC training period (Choi, et al.,
2005).
2.5.2. Neural adaptations
It has been shown in older adults that practice of TC exercise with a frequency of 3
times per week for more than 1 year can improve postural control (Fong & Ng, 2006;
Tsang & Hui-Chan, 2004b; Tsang, et al., 2004), compared with sedentary controls. It
has also been found that training 6 times per week for a 4-week period can result in a
reduction of body sway during upright standing in older adults (Tsang & Hui-Chan,
2004a). Improvement in postural control after TC training is associated with neural
adaptations in kinematic proprioception (Fong & Ng, 2006; Li, et al., 2008; Tsang &
Hui-Chan, 2003, 2004b), vestibular function (Tsang & Hui-Chan, 2006; Tsang, et al.,
49
2004) and reaction time (Fong & Ng, 2006). As indicated by the COP measurements,
older adults who practiced TC for a minium of 3 years showed better postural
performance during unipedal standing (Tsang & Hui-Chan, 2005) and bipedal standing
with alternation of visual (Wu, Zhao, Zhou, & Wei, 2002) or vestibular (Tsang & HuiChan, 2006) experimental conditions than their non-practicing counterparts.
2.6. Summary
In summary, the H-reflex test is suggested as a tool to investigate the effects of Ia
afferent inputs on the spinal motoneuron activity. The modulation of SOL H-reflex
amplitude during upright standing is found to be related to the direction and magnitude
of body sway and postural stability. Also, a down-regulation of the SOL H-reflex
response associated with increased postural stability has been observed after a short
period of balance training. However, current knowledge about the H-reflex modulation
in relation to postural control is limited to standing with simple and restricted sensory
conditions. Investigations on a single sensory condition may not be able to explain the
complex sensory interactions during postural control. Furthermore, TC has been found
to be an effective exercise intervention to improve neuromuscular functions in older
adults. Significant improvement in postural control after TC training must be associated
with neuromuscular adaptations.
50
Chapter 3 General methodology
51
The research work presented in this thesis includes four studies that were designed to
answer the specific questions and address the hypotheses, as stated in Chapter 1. Study I
was designed to determine the test-retest reliability of the SOL H-reflex during different
ankle joint positions and submaximal muscle contraction intensities in our laboratory.
Study II investigated the effects of ankle joint positions and muscle contraction
intensities on the soleus H-reflex modulation between young and older adults. Study III
examined the effect of ageing on the SOL H-reflex modulation during passive and
active shortening and lengthening ankle movements. This study also examined the
correlations between the SOL H-reflex modulation and dynamic ankle muscle actions
and static postural tasks in young and older adults. Study IV investigated the effects of
12 weeks of TC training on the SOL H-reflex modulation and control of static posture
in older adults.
This Chapter provides a description of the common methods utilised in these studies,
while the specific experimental design and methods, characteristics of the participants,
and data analysis of each study are provided in the following chapters.
3.1. Recruitment of Participants

All participants involved in the research were recruited on voluntary basis. The methods
used for participants recruitment were by word of mouth in the local community and
the media releases in local newspapers and radio broadcasts in the Northern Rivers
region, New South Wales, Australia.
To minimize the potential type I and II statistical errors, the sample size in each group
was determined by using GPower 3.0 software (Faul, Erdfelder, Lang, & Buchner,
2007). A power of 80 % and an alpha value of 0.05 in two-tailed test were used to
estimate the minimum number of participants required. It was predicted that, to
52
investigate the age-related changes in the H-reflex modulation, a minimum of 7

participants in each group were needed to minimize the type I and II errors in crosssectional comparisons (Earles, et al., 2000; Scaglioni, et al., 2003). In the intervention
study (Study IV), because there was no information regarding the effects of TC training
on the H-reflex modulation in the literature, the effect size index was calculated based
on reports that examined the effect of TC training on postural control with conflicting
sensory conditions in older adults (Tsang & Hui-Chan, 2004a). It was predicted that a
minimum of 15 participants was required in the training group to detect traininginduced changes in postural control.
An established Health Status Assessment Prior to Exercise Testing Questionnaire

(Appendix D) was used to assess health conditions of the participants and to identify
potential contraindications for participation. The inclusion criteria for participation were:
currently healthy; 20-40 years of age for the young; 65-85 years of age for the older
adult groups; and no undertaking of regular resistance training in the last six months.
Participants who were identified with contraindications or conditions described in the
following exclusion criteria were not accepted in this project. The exclusion criteria
were: 1) recent lower limb injuries; 2) neurological or lower-extremity orthopaedic
diagnoses; 3) corrected visual acuity worse than 20/100 or presence of a field defect; 4)
true vertigo; 5) acute illness; 6) the use of medication (eg. psychotrophic drugs) known
to affect sensorimotor functions (Laughton et al., 2003; Leipzig, Cumming, & Tinetti,
1999; Wolfson, Whipple, Derby, Amerman, & Nashner, 1994); and 7) waist
circumference greater than 90 cm in men and 83 cm in women (Janssen, Katzmarzyk, &
Ross, 2004; Zhu et al., 2004).
Participants who matched the criteria signed the Informed Consent form (Appendix B)
and were also verbally informed of the purposes of the studies and the experimental
53
procedures. All participants were instructed to avoid participating in strenuous exercise

within 24 hours, and to refrain from taking caffeine-containing drinks or food within 2
hours, prior to an experimental session. All experiments were carried out in the exercise
testing and biomechanics laboratories in P block, School of Health and Human Sciences,
Lismore Campus, Southern Cross University.
The participants were required to undertake a familiarization session prior to the first
formal experimental session. During the familiarization session, the participants were
introduced with the experimental procedures and devices, experienced with application
of electrical stimulation for H-reflex test and familiarised with the experimental tasks in
the respective study. Table 3.1 provides a summary of the number of participants
recruited in these studies.
Table 3.1: A summary of participants recruited in the four studies.

Group
Study I
Age (yrs)
Number of participants
Male
Female
Young
24.9 5
Study II
Young
25.1 4.8
10
10
(Cross-sectional study)
Older
74.2 5.1
10
10
Sutdy III
Young
24.4 4
10
10
(Cross-sectional study)
Older
73.3 5
10
10
Study IV
Tai Chi
72.9 4.4
11
(Longitudinal study)
Control
72.9 6.5
(Reliability Study)
Note: Data are expressed as mean SD.
54
3.2. H-reflex test

In this research, the H-reflex of SOL muscle was examined because this muscle plays
an essential role in control of upright posture during standing (Winter, et al., 2001). The
SOL H-reflex was tested in a sitting position in Study I, II and III and a standing
position in Study III and IV. The SOL H-reflex response was elicited by an electrical
impulse applied to the tibial nerve via a constant-current electrical stimulator (DS7AH,
Digitimer, Herfordshire, UK) with a single rectangular pulse of 200 s in width and
output voltage of 400 V. The current output was adjustable in the range of 0 - 999 mA.
A reusable rubber-based self-adhesive electrode with the size of 20 x 20 mm (Uni-Patch
EP 84177, MN, USA) was placed on the popliteal fossa as the cathode, while another
reusable rubber-based self-adhesive electrode with the size of 50 x 50 mm (Axelgaard
Model 895220, CA, USA) was positioned over the patella as the anode (Figure 3.1).
The location of cathode was determined where the greatest amplitude of H-reflex was
found. Because no difference of the H-reflex amplitude between the right and left legs
has been reported, all participants were measured on the right leg for the SOL H-reflex
to ensure consistency (Mezzarane & Kohn, 2002).
Electromyogram (EMG) signals from the SOL and TA muscles of the right leg were
recorded by using a Bagnoli-8 EMG system (Delsys, Boston, MA) with single
differential surface electrodes (DE 2.1, Delsys, Boston, MA). The electrode housing
contained two parallel silver bars (99.9% pure silver) of 10 mm in length and 1 mm in
diameter as the EMG sensors, spaced 10 mm apart. The electrode housing was
internally shielded and contained a pre-amplifier. A self-adhesive conductive disk
electrode (50 mm in diameter, Dermatrode, Delsys, Boston, MA) was placed over the
medial condyle of the femur bone as the reference. The electrode placement on the TA
muscle was at one thirds distance from the head of the fibula to the medial malleolus
55
and ~2 cm lateral to the tibial crest. The electrode on the SOL muscle was placed at two
thirds distance from the medial condyle of the femur to the medial malleolus and at the
central position in the medial-lateral direction of the SOL muscle border (Hermens et al.,
1999) (Figure 3.1). Skin preparation was conducted on the locations of the EMG and
stimulation electrodes. All electrode positions were carefully checked and marked with
a waterproof pen to ensure the same placements over time in repeated tests. Conductive
gel was rubbed onto the silver bars of the EMG sensors to improve electrode-skin
contacts. In addition to application of the adhesive electrode-skin interface, a surgical
adhesive tape was used to secure the electrode positions onto the skin.
To determine the maximal H-wave and M-wave in respective testing conditions, the
stimulation intensity was progressively increased with increments of 10 mA until the
Mmax was determined. The intensity for eliciting the Hmax was subsequently determined
by using 1 or 2 mA increment. Each stimulation intensity was applied four times with
10 s ISI for establishing H-wave and M-wave recruitment curve (Racinais, Gaoua, &
Grantham, 2008).
Figure 3.1: Locations of the stimulation and EMG electrodes.

56
3.3. Isometric submaximal voluntary contraction

Prior to the tests, the participants were given a 5 min warm-up exercise, which consisted
of 3 min static stretch and 2 min ankle dynamic exercises. The stretch exercises
included four static stretch activities, two for dorsiflexors and two for plantarflexors.
The participants performed each stretch exercise twice with 10 s rest interval. Each
stretch exercise was held for approximate 10 s. Afterwards, the participants moved on to
the Biodex and performed 2 min isokinetic ankle exercise at a constant angular velocity
of 60/s within their comfort levels.
A Biodex dynamometer (System 3; Biodex Medical System, Shirley, New York) was
used to perform isometric voluntary muscle contraction. Participants sat on the Biodex
chair in a semi-reclined position with the right leg fitted into the ankle attachment. The
seats back tilt was set to 55 (0 for flat). The left foot was resting on the foot-rest
attachment. The right foot was placed in the foot-plate and stabilized by Velcro straps
and the heel cup. The thigh was supported by the support arm which was fitted
proximately to the knee with the knee joint angle at 140 (180 as full extension). A
Velcro strap was applied to the thigh against the support arm. A strap over the trunk and
a strap over the hip of the testing leg were used to minimize body movement (Figure
3.2). The transverse axis of the ankle joint was carefully aligned with the rotational axis
of the dynamometer. To monitor the torque a computer screen was placed at a distance
of approximately 2 m in front of the participants at eye level.
57
Figure 3.2: Participants position in the isometric voluntary contraction test
The participants were assessed for their maximum plantarflexion strength in the neutral
0, plantarflexion 20, and dorsiflexion 20 positions. All participants performed three
successful MVC at the respective ankle placement. The duration of each contraction
was 3 s. The rest interval between contractions was 1 min in order to minimize the
potential for muscle fatigue. The criteria for successful MVC measurements were based
on the guidelines proposed by Gandevia et al. (2001). After the MVC assessment, the
participants were given 3 min rest prior to the H-reflex tests. The individual target force
at 10%, 30% and 50% MVC levels were determined by the highest MVC value found in
the respective ankle position. The participants were provided with visual feedback of
the torque production with a black horizontal line displayed on the computer screen of
the Biodex system. The location of the black line was normalised in accordance with the
target force level and appeared in the middle of the computer screen. During the
submaximal muscle contractions, participants were required to produce an adequate
level of force so that the force trace could match the black line for about 2 s.
58
3.4. Passive and active dynamic muscle actions

The Biodex dynamometer was also used to perform the dynamic muscle actions. The
speed for shortening and lengthening muscle actions was set at 60/s of constant angular
velocity. In the passive ankle movement test, the ankle joint movement was set with a
range of motion of 40. Passive shortening movement was defined as the ankle
movement from the dorsiflexion 20 to the plantarflexion 20, whereas the passive
lengthening movement was defined as the ankle movement in the opposite direction.
In the shortening contraction test, the participants ankle joint was rested at the
dorsiflexion 20 as the initial position. The participant was instructed to contract the calf
muscle statically with submaximal effort for approximately 1 s prior to each shortening
contraction in order to maintain similar thixotropic effects (Proske, 2006). When a
verbal instruction go was given, the participant immediately performed a shortening
muscle contraction with a maximal effort from the initial position to the plantarflexion
20 (terminal position). When the trial was completed at the terminal position, a 60 s
rest interval was given before the ankle position was shifted to the initial position for the
next trial.
In the lengthening contraction test, the initial and terminal ankle positions were
converted to the plantarflexion 20 and the dorsiflexion 20, respectively. The testing
procedure was identical to the shortening contraction test, except it was a lengthening
contraction.
Electrical stimulation was applied when the ankle joint passed through the neutral
position (0) during passive and active dynamic muscle actions. This setup can ensure
all evoked potentials were elicited at similar muscle length throughout the experiment
(Gerilovsky, Tsvetinov, & Trenkova, 1989).
59
3.5. Static postural tasks

When performing the postural tasks the participants were instructed to maintain bilateral
stance with feet abducted 10 and heels separated 6 cm apart on a force plate which was
mounted in the ground (type 9287, Kistler, Winterthur, Switzerland). A blank paper was
taped onto the force plate for marking the feet positions of each participant for repeated
measurement. A black colour circle of 25 mm in diameter was placed 2 m in front of the
participant at eye level as a visual target. The participant focused on the target with the
head straight and the arms resting at the side in a comfortable position (Figure 3.3).
The participants performed the static postural test with four different sensory conditions:
1) on stable surface with eyes open (SO); 2) on stable surface with eyes closed (SC); 3)
on unstable surface with eyes open (UO); and 4) on unstable surface with eyes closed
(UC). The participants stood on the force plate with and without a foam rubber mat (24
mm thickness) for testing under stable and the unstable surface conditions, respectively.
During the static postural tasks, 10 H-reflex trials were recorded from each participant.
Electric stimulation was applied to elicit the SOL H-reflex while the participant
standing on the force plate. After each stimulation, the participant was given a 10 s rest
by sitting in a chair next to the force plate. A 2 min rest interval was given between the
sensory conditions. The mean value of SOL Hmax was determined by averaging ten
repetitions under each sensory condition (Mynark, 2005).
60
Figure 3.3: The standardised upright position in the static postural tasks.
3.6. Data acquisition and analysis

The output signals from the Biodex were connected from the remote access port on the
rear of the unit, and that from the Kistler force plate were connected, to a rack-mount
analog breakout accessory (BNC-2090, National Instrument, USA) for analog-to-digital
data conversion (A/D). A custom-written LabView (National Instruments, Austin, TX)
program was used to synchronize the electrical stimulation trigger, kinematic data, and
EMG signals (Figure 3.4).
The EMG signal was filtered with a band-pass range of 20 to 450 Hz, amplified with a
gain of 1000 times, and sampled at an A/D conversion rate of 5 kHz (2 kHz in the Study
IV). A 16-bit A/D converter was used with the input range of 5V and resolution of
0.153 V (Bagnoli-8, Delsys, Boston, MA). The force plate signal was amplified via an
8-channel charge amplifier (type 9865A, Kistler, Winterthur, Switzerland) and was
sampled at an A/D conversion rate of 2 kHz.
61
The parameters of H-reflex wave and M-wave were measured off-line. The digital EMG
data was transferred to Excel datasheets (Microsoft Office, 2003 version). The peak-topeak amplitude, latency, duration of H-wave, the peak-to-peak amplitude of M-wave
and background EMG (bEMG) of the SOL and TA muscles were measured on the
Excel sheets (Figure 3.5). The bEMG was determined by using root mean square (RMS)
calculation for 100 ms duration prior to the onset of the stimulus artefact seen in the
data plots on the Excel datasheets.
In Study II, the gain, latency and duration of the H-reflex were assessed. The H-reflex
gain was a ratio as dividing the Hmax by the bEMG in the respective experimental
condition (Capaday, 1997). The latency of H-reflex response was determined as the
time elapsed between the onset of stimulus artefact and the first phase of H-wave
(Scaglioni, et al., 2003). The duration of H-reflex response was measured by the time
interval between time points where the largest positive peak and the largest negative
peak was identified (Frigon, et al., 2007).
In Study IV, static postural capacity was determined by mean displacement of the centre
of pressure (COP) in anterior-posterior (COPA-P) and medial-lateral (COPM-L) directions.
The mean distance was an average value of the total travel displacement.
62
Compatible computer
Kinematic data
Biodex
Electrical stimulator
Trigger
Force plate
Electrical stimulation
Data
acquisition
LabVIEW analog
breakout accessory
Surface EMG signals
Figure 3.4: A schematic shows the data acquisition system.
H-wave
Artefact
Background EMG
Peak-to-peak
amplitude
M-wave
Latency
2mV
30ms
Duration
Figure 3.5: Raw EMG trace was used to quantify the SOL H-reflex parameters.
63
3.7. Statistical analyses

The original data obtained from each study were checked for their normal distribution.
The Grubbs test was used to identify significant outliers in all variables prior to further
statistical analysis. This test is based on the assumption of normality and is used to
detect outliers in univariate data from a normally distributed population (Grubbs &
Beck, 1972). The outlier identification is based on the difference between the mean
value of the sample and the most extreme data, expressed in terms of the standard
deviation. The probability level was set at 0.05. The criterion for elimination of a data
was based on the difference of the suspected data point to the mean value of the group
in terms of standard deviation, ie. [the suspected single outlier mean of the sample] /
standard deviation (SD) of the sample, against the critical value set for the number of
participants. The Grubbs critical value was 2.37 SD for the control group in the Study
VI (n = 14), 2.5 SD for the older group of the dynamic test in Study III (n = 18), and
2.56 SD (n = 20) for the other groups in Studies II, III and IV. Statistical analysis was
performed on the data following elimination of significant outliers.
Table 3.2 provides a summary of the statistical measurements in each study. All
statistical analysis were performed by using SPSS for Windows (SPSS Inc, Chicago, IL,
ver. 16.0)
64
Table 3.2: Summary of the statistical analyses.

Independent variables
1. Test (1 & 2)
2. Joint position
Study I
(N, PF & DF)

3. Force level
(0, 10, 30 & 50%)
Study II
2. Hmax
3. Mmax
4. Hmax/Mmax ratio
5. SOL bEMG
2. Joint position
2. SOL bEMG
(0, 10, 30 & 50%)

1. Group (Y & O)
2. Action type (S & L)
3. Posutral tasks
(SO. SC, UO & UC)
1. ICC2.1.
2. SEM
3. 2 way RMANOVA
6. TA bEMG
1. H-reflex gain
(N, PF & DF)
Measurement
1. MVC
1. Group (Y & O)
3. Force level
Study III
Dependent variables
3. TA bEMG
4. Hmax duration
1. 2 way RMANOVA
2. 3 way RMANOVA
5. Hmax latency
1. Hmax
1. 2 way RMANOVA
2. Hmax/Mmax ratio
2. Pearson correlation
3. Hsup/Msup ratio
(Bivariate test)
1. Group (TC & C)

2. Time
Study IV
(Pre- & Post-training)

3. Postural tasks
1. Hmax/Mmax ratio
2. COPA-P
3. COPM-L
1. Independent t test
2. 2 way RMANOVA
(SO. SC, UO & UC)

Y = young group; O = older group; N = neutral; PF = plantarflexion; DF = dorsiflexion;
SO = stable surface with eyes open; SC = stable surface with eyes closed; UO =
unstable surface with eyes open; UC = Unstable surface with eyes closed; S =
shortening action; L = lengthening action; TC = Tai Chi group; C = control group;
MVC = maximal voluntary contraction; Hmax = maximal amplitude of H-reflex; Mmax =
maximal amplitude of M-wave; bEMG = background EMG; COPA-P = mean
displacement of centre of pressure in anterior-posterior direction; COPM-L = mean
displacement of centre of pressure in medial-lateral direction; ICC = intraclass
correlation coefficients; SEM = standard error of measurement; analysis of variance
with repeated measures = RMANOVA.
65
Chapter 4 Test-retest reliability of the soleus H-reflex

in relation to joint positions and muscle force levels (Study I)
66
4.1. Introduction
The H-reflex test is commonly used to investigate the excitability of the spinal motoneuron pool that may be affected by inputs from various afferent and efferent
pathways (Funase & Miles, 1999; Pierrot-Deseilligny & Mazevet, 2000). When a
method/technique is employed to assess body functional changes, it is essential to
establish its reliability and validity in repeated measurements (Weir, 2005).
The reliability of soleus (SOL) H-reflex during rest has been extensively evaluated
under a variety of experimental conditions. For example, several studies have
investigated the reliability of SOL H-reflex in lying, sitting and standing positions (Ali
& Sabbahi, 2001; Handcock, et al., 2001; Hopkins, et al., 2000). High reliability of the
SOL H-reflex measurement has been reported across different body positions as
indicated by analysis of the intraclass correlation coefficient (ICC, from 0.80 to 0.94).
The test-retest reliability of H-reflex has also been examined for populations of different
ages (Mynark, 2005), with conditioning-test (C-T) stimulations (Earles, et al., 2002),
and in upper (Jaberzadeh, et al., 2004; Stowe, et al., 2008) and lower limb muscles
(Palmieri, et al., 2002). It has been also reported that the SOL H-reflex demonstrates an
ICC of 0.93 in test-retest separated by four weeks (Clark, et al., 2007).
However, although some studies have investigated the SOL H-reflex during voluntary
muscle contractions (Butler, et al., 1993; Oya & Cresswell, 2008; Stein, et al., 2007),
there is no study found in the available literature that has examined the test-retest
reliability under such conditions. Funase and Miles (1999) reported that the amplitude
of SOL H-reflex is inconsistent during submaximal voluntary muscle contractions,
despite the fact that the electrical stimulation was controlled at a constant intensity. The
reason for the inconsistency is not known.
67
In clinical applications, the H-reflex has been used to assess spasticity, for example in
patients with spinal cord injury or after a stroke (Nielsen, Crone, & Hultborn, 2007).
The patients are tested in a particular body position. It has been shown that the H-reflex
is affected by the body position changes (Hwang, 2002). However, to our knowledge no
research has addressed the test-retest reliability of SOL H-reflex at different ankle joint
positions. Lack of consideration of body and/or ankle joint positions during the H-reflex
tests might lead to the testing results being inconsistent and incomparable for evaluation
of therapeutic treatments.
The purpose of this study was therefore two-fold: 1) to determine the test-retest
reliability of the SOL H-reflex during rest as well as during submaximal contractions at
10%, 30%, and 50% MVC; and 2) to evaluate the test-retest reliability of SOL H-reflex
at the ankle joint angles of neutral (0), plantarflexion 20 and dorsiflexion 20
respectively, in a sitting position. The soleus muscle was investigated because this
muscle plays an essential role in control of upright standing posture (Winter, et al., 2001)
and it was planned to use the SOL H-reflex test in a series of studies on neuromuscular
adaptations to exercise interventions for postural control in our laboratory.
4.2. Methods
4.2.1. Participants
Ten healthy young participants with no neuromuscular disease or injuries (5 males, 5

females, age: 24.9 5 years, height: 168.3 8.8 cm, weight: 62.7 12.3 kg)
volunteered for this study. Each participant took part in two identical experimental trials:
Test1 (T1) and Test 2 (T2), separated by 3 to 7 days. The participants were instructed to
refrain from taking any caffeine-containing substances within 2 h, and not to undertake
strenuous exercise activity within 24 h prior to the testing sessions. An established
68
health status screening questionnaire was used to identify contraindications to

participation (Appendix D), and the inclusion and exclusion criteria (section 3.1 in
Chapter 3) were used in recruitment of participants. Participants who met the inclusion
criteria and had no contraindications to the study were asked to sign the Informed
Consent form (Appendix B), went through a familiarization session and participated in
two experimental sessions. To avoid the possible influence of circadian rhythm, every
participant was required to come to the experimental sessions at a similar time of the
day. This study was approved by the Human Research Ethics Committee of Southern
Cross University and was conducted in accordance with the Declaration of Helsinki.
4.2.2. Experimental settings
used to set ankle joint positions for isometric voluntary muscle contraction. The setting
of Biodex dynamometer was presented in the Chapter 3 General Methodology, Section
3.3 (page 57).
A constant-current electrical stimulator (DS7AH, Digitimer, Herfordshire, UK) and a

Bagnoli-8 EMG system (Delsys, Boston, MA) were used for the H-reflex test. The
procedures of H-reflex test and the method of surface EMG signal processes were
presented in the Chapter 3 General Methodology, Section 3.2 (page 55) and 3.6 (page
61). The participants testing position in Study I was illustrated in Figure 4.1.
69
Figure 4.1: Participants leg position in the soleus H-reflex test.
4.2.3. Experimental procedures
In the experimental trials, the recruitment curves of H-wave and M-wave at three ankle
positions were established at the beginning of the test. The stimulation intensity was
progressively increased with incremental steps of 10 mA from the threshold of H-reflex
to where there was no more increase in the M-wave amplitude. The exact intensity for
eliciting the Hmax was determined by using 1 or 2 mA increment (Racinais, et al., 2008).
In the recruitment curves, the Hmax and Mmax were used as resting condition for
statistical analysis and were determined in order to provide valid H-reflex data at the
respective ankle joint angles. Mean value of the H-wave amplitude was obtained by
averaging four repetitions at each testing condition.
Once the recruitment curves were established, the participants were assessed for their
maximal plantarflexion strength at the three ankle positions. Individual maximal
isometric strength was determined at the ankle joint positions of neutral (0),
plantarflexion 20 and dorsiflexion 20. All participants performed the MVC tests at
each of the respective ankle joint positions. The duration of each contraction was at
70
least 3 s. The minimum rest interval between contractions was 1 min in order to
minimize potential muscle fatigue. The criteria for successful MVC measurements were
based on the guidelines proposed by Gandevia (2001). Three successful MVCs were
recorded for each participant. The highest MVC torque of the three repetitions was used
to determine the submaximal torque levels at 10%, 30% and 50% MVC in the
subsequent H-reflex tests. After the MVC assessment, the participants were given a 3
min rest before commencing the H-reflex tests.
During the H-reflex tests, the participants were assigned to the three ankle positions in a
randomized order. The intensity of electrical stimulation used to induce the H-reflex
response was based on the stimulation intensity that induced the Hmax in the recruitment
curves (Palmieri, et al., 2004; Zehr, 2002). To minimize the potential effect of
homosynaptic post-activation depression, a minimum of 10 s resting interval was
inserted during the trials at 10% and 30%MVC (Palmieri, et al., 2004), and 30 s interval
was used during the trials at 50% MVC (Moore & Kukulka, 1991). The participants
were instructed to approach the target level of torque, as shown on the screen, as
quickly as possible. The electrical stimulus was delivered when the torque was
maintained at the required level for about 1.5 to 2 s. The three different intensities of
submaximal contractions were utilised in a randomized order. A 2 min rest interval was
offered to the participants before the ankle joint was changed to the next position. The
H-reflex test was repeated for four times under each testing condition. A total of 36
(minimum) submaximal muscle contractions were performed by each participant. The
whole experimental protocol was completed between 1.5 to 2.0 hours.
71
4.2.4. Data analysis
The digital EMG data was transferred to Excel datasheets (Microsoft Office, 2003
version) for analysis. The peak-to-peak amplitudes of H-reflex wave and M-wave were
measured offline for the respective experimental conditions.
A 100 ms bEMG before the application of percutaneous electrical stimulation was

recorded for comparison of baseline muscle activation in different motor tasks, as
suggested by Oya and Cresswell (2008). The root mean square of the EMG signal was
calculated to represent the bEMG.
4.2.5. Statistical analyses
Descriptive data of the measured variables was given as mean standard deviation (SD).
Intraclass correlation coefficients (ICC2,1, SPSS) was used to determine the test-retest
reliability over the two testing sessions. Test-retest reliability of the Hmax was analysed
for the conditions of resting, during 10%, 30% and 50% MVC, and at the three ankle
joint positions, respectively. The test-retest reliability of Mmax and Hmax/Mmax ratio at the
three different ankle positions was only assessed for the resting condition. The ranking
of ICC values followed the description in a previous study (Versino et al., 2007), with
high referring to a value which was greater than 0.75, moderate for a value between
0.4 and 0.75, and poor for a value less than 0.4. Repeated measures ANOVA
(RMANOVA) with Bonferroni adjustment was used to determine if there was any
significant difference in bEMG during submaximal muscle contractions between the
two experimental sessions. An alpha value of P < 0.05 was set for significant
differences between the means.
The measurement error was also determined by using the standard error of measurement
(SEM), which was calculated by the means of the square root of the within-subject error
72
variant (residual mean square) from the ANOVA table (Atkinson & Nevill, 1998). The
SEM was used to measure the absolute reliability, which can be used to determine the
precision of within-subject scores. A small SEM can be interpreted as a reliable
measurement. Because the ICC value is a ratio of variance for differentiation between
different subjects, it cannot be used to understand the trial-to-trial variability within
subjects. As suggested by Weir (2005), confident interpretation of test-retest reliability,
which may be affected by systematic error (eg. learning effect or fatigue), can be
exhibited when the collected data is evaluated by a statistical method with combination
of the ICC (relative reliability) and SEM (absolute reliability).
All data were analysed using SPSS for Windows (SPSS Inc, Chicago, IL, ver. 16.0).
4.3. Results
The mean, standard deviation, standard error of measurement, and intraclass correlation
coefficient of the SOL H-reflex measurements during rest and muscle contractions are
shown in Tables 4.1 and 4.2, respectively. Representative data of the SOL H-reflex
under all testing conditions from one participant is displayed in Figure 4.2, as an
example.
4.3.1. Reliability of the soleus H-reflex at different joint angles during rest
The results revealed that the test-retest reliability of SOL Hmax during rest was high at
the neutral (ICC = 0.92) and plantarflexion (ICC = 0.96) joint positions, and moderate at
the dorsiflexion (ICC = 0.75) position. The Mmax showed a high test-retest reliability at
the neutral (ICC = 0.96) and dorsiflexion (ICC = 0.89) positions, whereas it reached a
moderate level (0.75) at the plantarflexion position. The test-retest reliability of
Hmax/Mmax ratio during rest demonstrated a high level at all ankle positions
73
(plantarflexion, ICC = 0.95; neutral, 0.86 and dorsiflexion, 0.84, respectively) (Table
4.1).
4.3.2. Reliability of the soleus H-reflex during submaximal muscle contractions
The ICC values of SOL Hmax during the submaximal voluntary contractions were in the
range of 0.62 to 0.97. During the 10% MVC contractions, high ICC values were
observed at the three ankle positions (0: 0.92; 20: 0.93 and -20: 0.95). During the
30% MVC contractions, moderate ICC values were obtained across the three ankle
positions (0: 0.64; 20: 0.66 and -20: 0.62). During the 50% MVC contraction, the
highest ICC value was found at the neutral position (0.97), compared to that at the
plantarflexion (0.83) and dorsiflexion (0.79) positions (Table 4.2).
4.3.3. Background EMG in the soleus and tibialis anterior muscles during
submaximal voluntary contractions
The bEMG of the SOL and TA muscles during the 10%, 30% and 50% MVC
contractions at the three ankle positions in the two testing sessions is demonstrated in
Figure 4.3 RMANOVA revealed that there were significant differences in the bEMG
between the SOL and TA muscles at the same joint angle in the respective testing
session (P < 0.05). In the SOL muscle, the bEMG in the trials at the neutral and
dorsiflexion positions showed similar levels of activity in the T1 and T2. However, at
the plantarflexion position, the bEMG showed a quite different level of activity in the
T1 (10% MVC < 50% MVC, P < 0.05) compared to that in the T2 (10% MVC < 30%
MVC; 10% MVC < 50% MVC; 30% MVC < 50% MVC, all P < 0.05). In the TA
muscle, significant differences of bEMG between the levels of muscle activity were
observed across the three joint angles, while no significant difference in bEMG between
T1 and T2 was found under all testing conditions (Figure 4.3).
74
Table 4.1: Mean value, standard deviation (values in the brackets), intraclass correlation
coefficients (ICC), and standard error of measurement (SEM) for the maximal voluntary
contraction torque (MVC), the maximal amplitude of soleus H-reflex (Hmax), the
maximal amplitude of soleus M-wave (Mmax), and the Hmax/Mmax ratio during rest and at
three ankle joint positions in Trial 1 (T1) and Trial 2 (T2).
Neutral (0)
Plantarflexion (20)
Dorsiflexion (-20)
T1 MVC (N.m)
105.8 ( 42.5)
56.8 ( 26.1)
151.9 ( 66.1)
T2 MVC (N.m)
107.0 ( 33.6)
59.4 ( 20.9)
165.3 ( 54.3)
MVC ICC
0.96
0.98
0.97
MVC SEM
10.97
5.06
15.45
T1 Hmax (mV)
1.97 ( 0.92)
2.40 ( 0.98)
1.42 ( 0.7)
T2 Hmax (mV)
2.06 ( 0.92)
2.64 ( 0.98)
1.49 ( 0.92)
Hmax ICC
0.92
0.96
0.75
Hmax SEM
0.35
0.26
0.52
T1 Mmax (mV)
3.50 ( 1.03)
4.28 ( 1.08)
3.09 ( 1.58)
T2 Mmax (mV)
3.29 ( 1.24)
4.08 ( 0.87)
3.09 ( 1.3)
Mmax ICC
0.96
0.75
0.89
Mmax SEM
0.32
0.62
0.64
T1 Hmax/Mmax
0.57 ( 0.21)
0.57 ( 0.19)
0.6 ( 0.28)
T2 Hmax/Mmax
0.66 ( 0.28)
0.63 ( 0.20)
0.51 ( 0.26)
Hmax/Mmax ICC
0.86
0.96
0.84
Hmax/Mmax SEM
0.12
0.06
0.14
75
Table 4.2: Mean value and standard deviation (in the brackets), intraclass correlation coefficients (ICC), and standard error of measurement
(SEM) for the maximal amplitude of soleus H-reflex (Hmax) at three ankle positions during contractions at 10%, 30% and 50% maximal
voluntary torque (MVC) in the Trial 1 (T1) and Trial 2 (T2).
Neutral (0)
Plantarflexion (20)
Dorsiflexion (-20)
10%MVC
30%MVC
50%MVC
10%MVC
30%MVC
50%MVC
10%MVC
30%MVC
50%MVC
T1 Hmax (mV)
2.01
( 0.84)
1.95
( 0.8)
2.5
( 1.33)
2.06
( 0.94)
1.82
( 0.89)
1.83
( 1.54)
1.54
( 0.86)
1.72
( 0.73)
1.81
( 0.99)
T2 Hmax (mV)
2.3
( 1.06)
2.62
( 1.1)
2.4
( 1.52)
2.32
( 0.92)
2.08
( 1.25)
2.09
( 1.55)
1.47
( 0.94)
1.78
( 0.74)
2.12
( 0.75)
Hmax ICC
0.92
0.64
0.97
0.93
0.66
0.83
0.95
0.62
0.79
Hmax SEM
0.38
0.7
0.33
0.34
0.77
0.84
0.27
0.55
0.52
76
Figure 4.2: The maximal soleus H-reflex amplitude recorded during rest, and 10%, 30%
and 50% maximal voluntary contraction at the neutral, plantarflexion, and dorsiflexion
of ankle positions from a representative participant. The Trial 1 SOL H-reflex is
presented as solid black lines, whereas the Trial 2 SOL H-reflex is highlighted as thick
gray lines. Arrows indicate the stimulus artefact.
77
A
**
*
*
**
***
**
***
***
**
**
*
78
**
*
**
***
*
*
**
***
***
***
***
***
***
Figure 4.3: The soleus (A) and tibialis anterior (B) background EMG (bEMG) activity
recorded during 10%, 30% and 50% maximal voluntary contractions at the neutral,
plantarflexion and dorsiflexion positions in the Trial 1 (T1) and Trial 2 (T2). The bEMG
was recorded 100 ms prior to the delivery of electrical stimulation. Asterisks represent
the level of significant difference as *: P < 0.05, **: P < 0.01, and ***: P < 0.001.
79
4.4. Discussion
There were two novel findings in this reliability study. The first finding was that the
relative and absolute reliability of SOL H-reflex test-retest measurements was joint
angle-dependent during rest. The amplitude of SOL H-reflex was more reliable at the
neutral and plantarflexion positions than that at the dorsiflexion position. The second
major finding was that the test-retest reliability of SOL H-reflex was affected by the
intensity of voluntary muscle contractions.
Nozaki and colleagues (1995) have previously reported that the H-reflex is a
neurological response with a characteristic of considerable variability. These authors
suggested that the amplitude of H-reflex was fractal and varied in time even though the
experimental conditions were constant and controlled. The present study was designed
to examine the test-retest reliability of H-reflex in specific motor tasks. These tasks
were performed in a randomized order and the participants were tested using identical
testing protocols. Therefore the outcomes of this study were unlikely to be affected by
the time-related variability of H-reflex, if there was any, or the potential effect is
controlled by the randomization of the testing order.
The results showed that the test-retest of SOL H-reflex during rest was less reliable
when the ankle joint was placed at the dorsiflexion position (Hmax ICC = 0.75; SEM =
0.52) compared to that at the neutral (Hmax ICC = 0.92; SEM = 0.35) and plantarflexion
(Hmax ICC = 0.96; SEM = 0.26) positions. The exact mechanisms that contributed to the
differences observed at different joint angles are not known. However, the differences
may be attributable to the influences from the change in muscle length, the presynaptic
inhibition, and the corticospinal inputs. At the dorsiflexion position, the SOL muscle
was placed in a stretched position for a prolonged period. This might lead to a decrease
in the excitatory volley propagating to the -motoneurons of the synergy muscles as a
80
result of increased presynaptic inhibition (Guissard, Duchateau, & Hainaut, 2001). It

has been previously demonstrated that the position-dependent modulation of SOL Hreflex is primarily affected by the presynaptic inhibition as a result of muscle stretch and
activation of group Ia afferent (Hwang, 2002). It should be noted that other mechanisms
such as Ib inhibitory afferents may also contribute to the changes in the discharging rate
of the intrasfusal fibres when the calf muscle is stretched (Guissard & Duchateau, 2006).
It is interesting to note that higher reliability of the SOL H-reflex was observed during
rest and at lower contraction intensity (10% MVC). In contrast, lower ICC and higher
SEM values were found during contractions at 30% and 50% MVC. The inconsistency
of SOL H-reflex reliability during the submaximal muscle contractions at 30-50% MVC
is possibly due to a number of neurophysiological factors (Butler, et al., 1993; Funase &
Miles, 1999; Stein, et al., 2007).
One of the potential factors is the fluctuation of the motoneuron membrane potential
during isometric muscle contractions. It is understandable that during submaximal
voluntary contractions the pattern of motor unit recruitment and firing frequency would
vary within a group of active motor units (Nordstrom & Miles, 1991). Pasquet et al.
(2005) have shown that increase in motor unit discharging rate is a main contributor to
motor unit activity at force levels up 10% MVC, whereas increase in motor unit
recruitment is a major factor for motoneuron activation at force levels between 10% and
35% MVC. Furthermore, Nordstrom and Miles (1991) showed that motor units were
not discharging in a fixed pattern during a prolonged contraction. The net motoneuron
excitation varied as a result of modulation by supraspinal and spinal mechanisms.
Funase and Miles (1999) explained that the H-reflex response was a compound muscle
action potential from the motor units that were discharging steadily and the motor units
that were discharging inconsistently. There were also motor units whose level of
81
excitability was just below the threshold and might be activated according to additional
EPSP. This mechanism leads to varying numbers of motor units recruited in response to
group Ia afferent inputs during submaximal voluntary contractions at the time when the
electrical stimulation was applied.
Furthermore, presynaptic inhibitory mechanisms also play a critical role in the test
retest reliability of SOL H-reflex during voluntary contraction. Our findings
demonstrated that the ICC and SEM values of 10% and 30% MVC were consistent
across three ankle joint positions. It indicates that the test-retest reliability of SOL Hreflex was compromised with force levels rather than joint positions during sustained
muscle contractions at 10% and 30% MVC. However, the ICC and SEM values of 50%
MVC demonstrated the greatest variation across the three ankle positions in this
reliability study. One possible reason for these differences is that the changes in the
spinal motoneuron excitability during muscle contractions were primarily filtered by
presynaptic inhibitory mechanisms, which contributed to the regulation of the
monosynaptic transmission in the Ia pathway via either central commands (Hultborn,
Meunier, Pierrot-Deseilligny, & Shindo, 1987) or peripheral sensory feedback (Brooke
et al., 1997). This assumption may be supported by the unchanged bEMG in SOL and
TA between the two testing sessions observed in the respective testing conditions. The
unchanged SOL muscle bEMG between the T1 and T2 may indicate that the overall
level of motor unit activity was similar prior to the H-reflex records (Misiaszek, 2003).
Thus, any alteration of the H-reflex amplitude would be mediated presynaptically by
neural mechanisms via the Ia-motoneuron pathway. Based on this assumption, it is
speculated that the presynaptic inhibitory mechanism during repeated trials is one of the
primary factors that causes variation of ICC and SEM values during voluntary
contractions.
82
Co-contraction of the antagonist is also a candidate factor for inhibitory modulation of

the agonist motoneuron pool. Increase in antagonist activation may depress the
motoneuron excitability of the agonist via the reciprocal inhibitory pathway (Crone &
Nielsen, 1989). It has been suggested that during plantarflexion, co-contraction of the
TA muscle can contribute to a reciprocal inhibition of the SOL motoneuron pool,
depending upon the degree of involvement of the TA muscle (Leonard, et al., 1999). In
this reliability study, inconsistent TA muscle activity was observed that may potentially
influence the SOL H-reflex measurement to some extent.
Another factor that may have affected the variations in ICC and SEM values of SOL Hreflex is the change in muscle length. It has been reported that the size of compound
muscle action potentials can be affected by the length of the muscle (Frigon, et al., 2007)
and the spatial relationship between stimulation electrode and the nerve during
movement (Capaday, 1997). Butler et al (1993) reported that there was a variation in Mwave during different levels of submaximal voluntary contractions. This observation
might result from the two factors mentioned above. The Mmax at the respective force
levels and at the three ankle positions were not recorded in the present study. However,
it can be speculated that the changes in joint angle and contraction intensity would have
caused some changes in muscle length and the relationship between the surface
electrodes and muscle and nerve fibres that might have contributed to the variation of
the H-reflex.
In this investigation, the testretest reliability of the H-reflex during the 50% MVC
contractions was better than that during the 30% MVC contractions. The SOL and
gastrocnemius are the two main plantarflexors for force production. The motoneuron
behavior of these two muscles is different at various levels of voluntary contractions
(Tucker & Trker, 2004). Cresswell (1995) observed that the relative contribution of the
83
gastrocnemius to force production during plantarflexion increased in association with

increase in voluntary effort. It is possible that at 30% MVC both SOL and
gastrocnemius muscles were recruited alternatively, while at 50% MVC both muscles
were recruited in a more consistent way, which contributed to the larger variation
observed from the SOL at 30% MVC. However, due to lack of direct evidence on motor
unit recruitment and EMG of the gastrocnemius was not monitored in this study, this
speculation requires further confirmation.
The SOL H-reflex is frequently used to examine the excitability of motoneurons in

relation to movement patterns (e.g. muscle activation, posture, and locomotion) and
clinical interventions (e.g. therapeutic effect). For example, H-reflex test has been used
to assess the spinal motoneuron excitability in stroke patients after a certain period of
rehabilitation (Hwang, Lin, Tung, & Wang, 2004). However, an underestimation or
overestimation of therapeutic effect could occur if the selected methodology of
measurement is not reliable. As significant variability in ICC of the SOL H-reflex was
found in this study, it is suggested that careful consideration must be given to the
selection of functional testing activity, e.g. joint angles and levels of voluntary exertion
of force, in order to obtain reliable outcomes.
4.5. Conclusion
In conclusion, a high level of test-retest reliability of the SOL H-reflex, measured by
ICC and SEM, was observed when the ankle joint was placed at the neutral and 20
plantarflexion positions, while moderate ICC values and high SEM values were found
at the 20 dorsiflexion position during rest. The results of absolute and relative
reliability assessments also demonstrated that the SOL H-reflex during 10% MVC was
more reliable than that at 30% and 50% MVC. Therefore, the reliability of the SOL Hreflex appeared to be significantly affected by the ankle joint position and the level of
84
voluntary contraction. These facts should be considered in designing functional

activities for assessment of the H-reflex.
The outcomes of this study have been published in Journal of Electromyography

and Kinesiology (See detail on page vi).
85
Chapter 5 Effects of joint position and muscle

contraction intensity on soleus H-reflex gain in young and
older adults (Study II)
86
5.1. Introduction
The size of the H-reflex is frequently used to investigate the influence of group I
afferent volley on spinal motoneuron activation (Misiaszek, 2003). Changes in the Hreflex amplitude in response to given conditions have been associated with neural
strategies in performing somatic motor tasks (Hultborn, 2006). For example, it has been
shown that the amplitude of H-reflex recorded in the SOL muscle increases
proportionally with increase of contraction intensity, indicating an increased excitability
and recruitment of spinal motoneurons. The mechanism underlying this change has been
suggested as an increase in the corticospinal drives due to volitional effort (Stein, et al.,
2007).
It has been well documented that ageing affects motor functions inevitably (AnguloKinzler, et al., 1998; Earles, et al., 2001; Kido, et al., 2004). For example, older adults
demonstrate deteriorations in motor movement preparation (Sterr & Dean, 2008), motor
cortical response generation (Kolev, et al., 2006), spinal motoneuron excitability
(Scaglioni, et al., 2003), nerve conduction velocity (Rivner, et al., 2001), and
electromechanical delay (Zhou, et al., 1995), which results in slower reactions and
compromised motor performance. In addition, older adults may adopt different motor
control strategies at the spinal (Kido, et al., 2004) and cortical (Hortobagyi, et al., 2006)
levels, compared to young counterparts. The alteration of motor outputs may be a
remodelled function in compensation for the age-related deteriorations in
neuromuscular function.
H-reflex gain is a ratio determined by the input of group Ia afferent volley (H-wave)
divided by the level of background muscle activity (Capaday, 1997). The regulation of
Ia afferent inputs can be affected by neural mechanisms at the presynaptic level,
whereas the background muscle activity can reflect summation of motor outflows at the
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postsynaptic level (Capaday, 1997; Misiaszek, 2003). The ratio of H-reflex gain can be
used to estimate the contribution of Ia afferent inputs to motoneuron activation via the
Ia-motoneuron spinal loop during execution of motor tasks. It has been suggested that
the age-related changes in the H-reflex gain could be an indication of different motor
strategies adopted by young and older adults (Angulo-Kinzler, et al., 1998; Chalmers &
Knutzen, 2002). When the same level of background muscle activity was controlled in
lying supine and bipedal standing, Chalmers et al. (2002) reported that young adults
demonstrated a significant decrease of the SOL H-reflex gain from lying to standing. In
contrast, older adults demonstrated no significant change in SOL H-reflex gain from
lying to standing. Also, Angulo-Kinzler et al. (1998) demonstrated that young adults
showed an increase of SOL H-reflex gain during submaximal voluntary muscle
contractions in a lying prone position in comparison to that during the same level of
muscle contractions in a standing position, while older adults demonstrated no
significant change in the SOL H-reflex gain in the same testing conditions. The
discrepancy of the reflex gain between young and older adults is speculated to be due to
age-related changes in the spinal inhibitory functions (Earles, et al., 2001).
The latency of H-reflex response is defined as the time elapsed between the onset of
stimulus and the first phase of H-reflex response recorded from the target muscle
(Scaglioni, et al., 2003). The latency of H-reflex response is known to be influenced by
ageing (Rivner, et al., 2001; Scaglioni, et al., 2003). As the latency of H-reflex is
normalized to an individuals height for appropriate between-subject comparisons,
Scaglioni et al. (2003) reported that older adults had longer SOL H-reflex latency than
that in their young counterparts. Interestingly, it was also found that significant
difference of the SOL H-reflex response was accompanied by unchanged M-wave
latency. Because the M-wave latency was not different between the young and older
adults, it indicated that mechanisms at motoneuron axons and muscular level are not a
88
factor to affect the reflex latency. The result suggested that the slower conduction in the
reflex arc of older adults was due to age-related demyelination of sensory axons and
changes in synaptic transmission efficacy (Jankelowitz, McNulty, & Burke, 2007;
Shaffer & Harrison, 2007).
The duration of the H-wave reflects the number and synchrony of nerve fibres that are
activated in the reflex response. The Hmax duration was measured by the time interval
between time points where the largest positive peak and the largest negative peak was
identified (Frigon, et al., 2007). In the literature, very little information is available on
the H-reflex duration, particular in discussion of ageing effect. It is clear that ageing
may cause functional or structural impairement on peripheral nervous system (Doherty,
Vandervoort, & Brown, 1993; Shaffer & Harrison, 2007). However, the result presented
in Scaglioni et als study (2003) showed no signficant difference of the Hmax duration
between young and older adults. It is still unknown that unchanged Hmax duration
reported in Scaglioni et als study is related to biological factor or technical setting for
measuring the Hmax duration.
Most H-reflex studies only examined young adults under a given body position and/or
ankle joint angle. There is no information available in the current literature on the
influence of ankle joint angle and level of voluntary contraction on the SOL H-reflex
modulation in older adults. This information is particularly important to the older
population because functional characteristics of the group Ia afferents during ankle
movements can help us to understand the ankle strategy with respect to postural control
(Tokuno, et al., 2007; Tokuno, et al., 2008). To our knowledge, only two studies
reported age-related changes in the SOL H-reflex during isometric submaximal
voluntary contractions in a fixed ankle position (Angulo-Kinzler, et al., 1998; Earles, et
al., 2001). However, observations of the H-reflex at a single ankle position may not be
89
able to fully explain neural strategy in functional antigravity motor tasks in either young
or older populations. Hwang (2002) reported that the SOL H-reflex was significantly
depressed at ankle dorsiflexion position but was not significantly affected at
plantarflexion position, as compared to that at the neutral position. The possible
mechanism which contributes to depression of the SOL H-reflex at the dorsiflexion has
been speculated as an increase in presynaptic and/or postsynaptic inhibitions of the
spinal motoneurons caused by passive stretch of the muscle that activates the muscle
spindle (Guissard, et al., 2001). The effect of presynaptic inhibition mediated by
primary afferent depolarization was evidenced by a change in the H-reflex amplitude
accompanied by an unchanged size of MEPs evoked by TMS when the ankle joint was
placed at dorsiflexion of 10 (Guissard, et al., 2001). The postsynaptic inhibitory
mechanisms were demonstrated by similar reductions of H-reflex and MEP responses
when the intensity of passive stretch was increased at dorsiflexion of 20 (Guissard, et
al., 2001).
The aims of Study II were to determine: 1) the age-related differences of the SOL Hreflex gain during submaximal voluntary contractions at 10%, 30% and 50% MVC
when the ankle joint was placed at neutral 0, plantarflexion of 20, and dorsiflexion of
20 positions; and 2) the effect of ankle joint positions on the SOL H-reflex latency and
duration in young and older adults.
5.2. Methods
5.2.1. Participants
Twenty healthy young adults (10 males and 10 females) with mean age ( SD) of 25.1
4.8 years, height 168.4 8.4 cm, body mass 64.0 14.1 kg, and body mass index (BMI)
22.3 3.2, and twenty healthy older adults (10 males and 10 females) with mean age of
90
74.2 5.1 years, height 165.4 8.0 cm, body mass 74.6 12.6 kg and BMI 27.3 4.2,
volunteered for the study. A health status screening questionnaire (Appendix D) was
used to identify contraindications to participation. The volunteers who met the inclusion
criteria (Section 3.1 in Chapter 3) and had no contraindication to participation signed an
Informed Consent form (Appendix B) and undertook a familiarization session prior to
the testing session. All participants were asked to refrain from smoking and taking
caffeine-containing substances within 2 hours prior to the testing session, and not to
undertaking strenuous exercise or physical activity 24 hours prior to the testing session.
5.2.2. Experimental settings
used to set ankle joint positions and intensities of isometric voluntary muscle
contraction. The setting of Biodex dynamometer was presented in Chapter 3 General
Methodology, Section 3.3 (page 57).
A constant-current electrical stimulator (DS7AH, Digitimer, Herfordshire, UK) and a

Bagnoli-8 EMG system (Delsys, Boston, MA) were used for the H-reflex test. The
procedures of H-reflex test and the method of surface EMG signal processes were
presented in Chapter 3 General Methodology, Section 3.2 (page 55) and 3.6 (page 61).
Prior to the tests, the participants were given a 5 min warm-up exercise, which
comprised of 3 min static stretch and 2 min ankle dynamic exercise. The stretch
exercises included four static stretch activities, two for dorsiflexors and two for
plantarflexors. The participants performed each stretch exercise twice with 10 s rest
interval. Each stretch was held for approximate 10 s. Afterwards, the participants moved
91
onto the Biodex and performed 2 min isokinetic ankle exercise at a constant angular
velocity of 60/s with comfortable efforts.
The procedures of the experimental session were similar to that described in Chapter 4
(Study I) expected that it was a cross-sectional study.
Four older participants experienced muscle cramps during contractions at 50% MVC in
the plantarflexion position. The researcher assisted the participants to stretch the
plantarflexors until the symptom of muscle cramp disappeared. After appropriate
treatments, these participants were given an additional 5 min rest before continuing on
the testing.
The H-reflex and M-wave variables in all successful trials were processed by the same
researcher. The amplitudes of H-reflex wave and M-wave were measured off-line for
the respective experimental conditions. The digital EMG data was transferred to Excel
datasheets (Microsoft Office, 2003 version), and the peak-to-peak amplitude of the Hwave and M-wave was identified on the Excel sheets.
A 100 ms bEMG was recorded for comparison of baseline muscle activation in different
motor tasks (Chen, et al., 2010). The bEMG was determined by using root mean square
calculation prior to the onset of stimulus artefact seen in the data plots on the Excel
datasheets. The H-reflex gain was a ratio as dividing the Hmax by bEMG in the
respective testing condition (Capaday, 1997).
The present study utilised the normalized values for the latency with the calculation of:
Hmax latency (ms.cm-1) = Hmax latency time (ms) / body height (cm) (Scaglioni, et al.,
2003).
92
The Hmax duration was calculated by the time interval from time point of the largest
positive peak to time point of the largest negative peak in H-wave.
The latency and duration of H-reflex wave was measured during the rest condition at all
three ankle joint positions.
The data was checked for its normal distribution. If a suspicious outlier data was found,
that might be caused by other influential factors, the Grubbs test was used to confirm
and eliminate the significant outliers. The Grubbs test was used with a probability level
of 0.05 to determine any outlying value (Section 3.7 in Chapter 3). Statistical analysis
was performed on the data following elimination of the significant outliers.
Descriptive data of the measured variables was given as mean standard deviation (SD).
A three-factor [group (2) x joint angle (3) x force (4)] RMANOVA was used to
determine if there was any significant difference in the H-reflex gain and bEMG
between the SOL and TA muscles during rest and submaximal voluntary muscle
contractions in the three different ankle positions. To detect any potential effect of ankle
position, a two-factor [group (2) x joint angle (3)] RMANOVA was performed to
compare the Hmax latency and duration. When a significant interaction was found, a
post-hoc test with Bonferroni adjustment was used to locate the differences between the
mean values. An alpha value of P 0.05 was set for significant differences between the
means. All statistical analysis was performed by using SPSS for Windows (SPSS Inc,
Chicago, IL, ver. 16.0).
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5.3. Results
Representative EMG traces of the SOL H-reflex from young and older participants in
all testing conditions are illustrated in Figure 5.1.
5.3.1. The soleus H-reflex gain
For the SOL H-reflex gain, there was no significant interaction detected between group,
joint angle and force level (P = 0.35). However, significant interactions were found for
joint angle by group (P < 0.05), force level by group (P < 0.05), and joint angle by
contraction level (P < 0.05). The H-reflex gain was relatively higher during rest than
that during submaximal voluntary contractions in both age groups (P < 0.05),
independent of ankle joint positions (Figure 5.2). In addition, the H-reflex gain of the
young group was significantly higher than that of the older group in all testing
conditions (P < 0.05).
5.3.2. Background EMG in the soleus and tibialis anterior muscles
The mean values of bEMG activities in the SOL and TA muscles in all testing
conditions are illustrated in Figure 5.3. In the SOL muscle, the bEMG increased from
rest to low and high contraction intensities, with the same trend at the neutral,
plantarflexion and dorsiflexion positions in both age groups (P < 0.05) (Figure 5.3 A
and C).
In the TA muscle, the bEMG variations were different to that found in the SOL muscle
(Figure 5.3, B and D). The young group had significant differences of the TA bEMG
between the contraction levels (P < 0.05) in all three ankle positions. However, the
older group showed significant difference of the TA bEMG between all contraction
94
intensities at the neutral and plantarflexion positions (P < 0.05) but not at the
dorsiflexion (P > 0.05).
5.3.3. Latency of the soleus H-reflex
The latency and duration of the SOL H-reflex was determined during rest at all three
ankle positions. As demonstrated in Figure 5.4 A, the SOL Hmax latency was
significantly longer in the older group than that in the young group (P < 0.05). In the
young group, the latency was significantly longer at the dorsiflexion than that at the
neutral (P < 0.05) and plantarflexion positions (P < 0.05). In contrast, the older group
revealed no significant difference of the H-reflex latency between ankle joint positions
(P < 0.05).
5.3.4. Duration of the soleus H-reflex
The duration of SOL Hmax in the young group was significantly shorter than that of the
older group at all ankle positions (P < 0.05). The duration of SOL Hmax was not joint
angle-dependent in the young group, whereas in the older group significant difference
of SOL Hmax duration was found between the neutral and plantarflexion joint positions
(P < 0.05) (Figure 5.4, B).
95
Plantarflexion
Neutral
Dorsiflexion
Plantarflexion
Neutral
Dorsiflexion
Rest
10%MVC
30%MVC
50%MVC
2mV
1mV
30ms
30ms
Figure 5.1: Typical Hmax of the soleus H-reflex recorded during rest, and muscle contractions at 10%, 30% and 50% MVC at the neutral, plantarflexion
and dorsiflexion of ankle positions in one young (left side) and one older (right side) participant. The SOL background EMG as recorded 100 ms prior
to when electrical stimulation was applied is displayed above the corresponding H-reflex response. Please note the difference in Y-scale for the EMG
traces of the older participant. Arrows indicate the stimulus artefact.
96
A
*
*
*
*
*
B
*
*
*
*
*
*
Figure 5.2: Influence of ageing on the soleus H-reflex gain during rest and submaximal
muscle contractions when the ankle joint placed at neutral, plantarflexion and
dorsiflexion positions. A) Young group (N=20) demonstrated similar tendencies of
decreased SOL reflex gain at all ankle positions when contraction intensity was
progressively increased. B) Older group (N=20) also demonstrated a tendency of
decrease in SOL reflex gain. Data presented are mean values and standard deviations. *:
P < 0.05 between contraction intensities.
97
*
*
*
*
*
*
*
Figure 5.3: Background EMG of the soleus (A: young, C: older) and tibialis anterior (B:
young, D: older) muscles during rest and soleus contractions at 10, 30 and 50% MVC in
ankle joint positions of neutral, plantarflexion and dorsiflexion. The bEMG of SOL
muscle (left side) in both young and older groups increased gradually with contraction
intensity. Data presented are mean values and standard deviations. *: P < 0.05 between
contraction intensities.
98
*
*
*
**
**
**
Figure 5.4: Latency (A) and duration (B) of maximal soleus H-reflex in the young
(filled circles) and older groups (unfilled circles). Data presented are mean values and
standard deviations.*: P < 0.05, older group was greater than young group. **: P < 0.05,
between joint positions.
99
5.4. Discussion
The major findings of Study II included that the age-related change in SOL H-reflex
gain was demonstrated during different intensities of isometric submaximal voluntary
contractions and at different ankle joint positions; young and older adults showed
similar down-regulation of the SOL H-reflex gain with increased intensity of
submaximal voluntary contraction in all three ankle joint positions; the older adults
demonstrated longer latency and duration of SOL H-reflex response compared with
young adults despite changes in ankle position; the latency of SOL H-reflex was
influenced by ankle joint position in the young adults but not in the older adults.
5.4.1. Age-related changes in the soleus H-reflex gain
During voluntary muscle contraction, the effect of Ia afferent inputs on the spinal
motoneuron activity can be regulated by the descending volleys from the supraspinal
centres via mechanisms such as presynaptic inhibition (Hultborn, 2006; Misiaszek,
2003). Kawashima and colleagues (2004) have reported significant contribution of the
supraspinal mechanisms to regulate the gain of SOL stretch reflex during low intensity
of isometric voluntary muscle contractions. In order to understand the influence of the
descending drives on regulation of Ia afferent volleys in young and older adults, the
present study assessed the recruitment of SOL H-reflex gain during submaximal
contractions.
Results of Study II showed that the SOL H-reflex gain was lower in the older group
than that in the young group across all testing conditions. This finding was in agreement
with a previous observation, showing that the gain of SOL H-reflex in lying and
standing positions was higher in young adults in comparison to that in older adults
(Angulo-Kinzler, et al., 1998). However, no significant difference of the SOL bEMG
100
between the two age groups was found in all testing conditions. Therefore, the lower Hreflex gain in the older group might indicate less input from group Ia afferent volley to
the SOL motoneurons. The possible mechanisms underlying this age-related difference
in SOL H-reflex modulation may be related to several neuromuscular factors.
First, there are degenerative changes in the structure of the neuromuscular system with
ageing. These neurophysiological changes include: 1) reduction of the number of
motoneurons in the lumbosacral spinal cord (Tomlinson & Irving, 1977); 2) decrease in
the number of sensory nerve fibres (Shaffer & Harrison, 2007); 3) increase in the motor
unit innervation ratio (Doherty et al., 1993); and 4) decrease in the number of muscle
fibres (Doherty, 2003). Secondly, functional alteration in neural mechanisms is also
potentially attributable to the decrease of SOL H-reflex modulation with ageing. These
possible mechanisms include: 1) reduction of membrane excitability of sensory axons
(Kiernan, Lin, Andersen, Murray, & Bostock, 2001); 2) decrease of the efficacy of
neural transmission in the Ia-motoneuron pathway (Mynark & Koceja, 2001); 3)
changes in the spinal interneuronal activity (Earles, et al., 2001; Kido, et al., 2004) and
4) decrease in sensitivity of the muscle spindles (Shaffer & Harrison, 2007).
It was interesting to find that the young and older groups showed a similar trend of
decrease in the SOL H-reflex gain with progressive increase of contraction intensity in
three ankle joint positions. Similar findings have been reported in a previous study when
the SOL H-reflex gain was assessed during submaximal voluntary muscle contractions
up to 30% MVC in lying and standing positions (Angulo-Kinzler, et al., 1998). In
addition, Chalmers and Knutzen (2002) reported that older adults did not show a change
in the SOL H-reflex gain during lying and upright standing, whereas a down modulation
of the SOL H-reflex gain was observed during tandem stance. The later postural task in
Chalmers and Knutzens study may rely on more volitional control to maintain postural
101
stability. It seems that the descending inputs from the supraspinal level play a dominate
role in the input-output regulation of reflex gain during voluntary movement control
rather than the influence of afferent inputs from changes of body or joint position.
Although the Ia afferent transmission is impaired with ageing, it is possible that the
cortical motor functions are preserved to regulate the H-reflex gain with progressive
increase of contraction intensity in older adults.
5.4.2. Age-related changes in the soleus H-reflex latency and duration
The latency of H-reflex was significantly shorter in the young group than that in the
older group during rest across all three ankle positions (Figure 5.4 A). The results also
showed that the latency of reflex response was not significantly affected by the joint
position in the older group but was joint angle-dependent in the young group. Due to the
absence of volitional inputs from the supraspinal centres during rest condition, a
possible explanation for these differences was the effects of ageing on the peripheral
sensory system (Shaffer & Harrison, 2007). It has been suggested that older adults have
a higher threshold in sensory receptors than that in young adults (Kiernan, et al., 2001).
Demyelination of sensory axons and myelin sheath may also be attributable to the
slower propagation of afferent signals to the spinal cord in the older population (Shaffer
& Harrison, 2007). Other factors, such as segmental demyelination of motor axons,
reduction in axon diameter, change in potassium and sodium ion channel activities in
the motor axon membrane, and decline in energy metabolism in the motor axon, may
also potentially contribute to the elongated time of the H-reflex response (Jankelowitz,
et al., 2007).
There are few reports on the duration of H-reflex in the literature (Scaglioni, et al.,
2003). This might be related to some technical restrictions. For example, the type of
EMG electrodes (monopolar or bipolar configuration) and location of electrode
102
placement can result in different temporal dispersions of motor unit action potentials
(Tucker & Turker, 2005). In the present study the reflex duration in response to
different experimental conditions was examined with the same technical setting in a
cross-sectional measurement. The technical issue should not be a limitation in the
present study. In contrast to Scaglioni et als study, our result showed significant
differences of Hmax duration between young and older adults across all ankle positions.
In clinical reports, increase in duration of compound muscle action potential is related to
axonal demyelination and abnormality of muscle fibres conduction velocity due to
pathological influence (Barroso & de la Fuente, 2009). Although the exact neural
mechanism that contributed to temporal dispersion of the H-reflex in the older group
was unclear, our results demonstrated that the age-related differences in the SOL Hreflex duration were statistically significant across all three ankle joint positions.
5.5. Conclusion
In conclusion, this study was the first to address the combined effects of joint angle and
voluntary contraction intensity on the SOL H-reflex modulation in older adults. The
gain of SOL H-reflex was significantly affected by level of contraction intensity as
demonstrated by a considerable decrease of the SOL H-reflex gain associated with
increased plantarflexion intensity. The similar down-modulation of SOL H-reflex gain
in young and older adults may reflect the essential role of the supraspinal involvement
to reserve the systematic motor function during voluntary muscle activities. The
significant differences in SOL H-reflex gain, latency and duration found between young
and older adults in relation to the joint position or intensity of muscle contraction might
indicate the age-related changes in motor functions. Futher research is required to
examine the mechanisms of these differences and clinical implications of these
assessments.
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Chapter 6 Effect of ageing on soleus H-reflex

modulation during shortening and lengthening muscle actions
in relation to postural control (Study III)
104
6.1. Introduction
The H-reflex test is often used to assess neural control strategy via the Ia afferent spinal
loop (Capaday, 1997). Changes in muscle fibre length have been demonstrated to affect
the H-reflex modulation during passive (Pinniger, et al., 2001) and active muscle
actions (Duclay & Martin, 2005; Romano & Schieppati, 1987). It has been observed
that the SOL H-reflex was depressed during passive lengthening movement, compared
with that during passive shortening movement (Nordlund, et al., 2002; Pinniger, et al.,
2001). Similar modulation of the SOL H-reflex was also found during shortening and
lengthening contractions with voluntary effort at submaximal (Nordlund, et al., 2002)
and maximal (Duclay & Martin, 2005) levels. It is suggested that the movement-related
changes in the H-reflex response during shortening and lengthening muscle actions are
mainly influenced by presynaptic inhibition of Ia afferents via either the central or
peripheral mechanisms (Duchateau & Enoka, 2008). Current knowledge in respect of
the SOL H-reflex modulation during shortening and lengthening muscle actions is based
on observations in young adults. It has been demonstrated that older adults may adopt
different neural strategies to control movement in comparison with young adults (Earles,
et al., 2001; Hortobagyi, et al., 2006; Kido, et al., 2004; Scaglioni, et al., 2003).
However, whether older adults use a different neural strategy in modulation of Ia
afferent inputs during dynamic muscle actions is still unclear. The primary aim of
Study III was to investigate the effect of ageing on the SOL H-reflex modulation during
passive and active shortening and lengthening muscle actions.
During upright standing, the body sways in a pendulum model with reversals of
direction at a mean frequency of 2.6 times per second (Loram, et al., 2005b). By using
ultrasound imaging technique to record changes in muscle fascicle length, the muscle
fascicle length of the soleus muscle decreased during postural sway in a forward
105
direction (Loram, et al., 2005a). It indicates that the soleus muscle is concentrically
contracting during forward sway. A recent study by Tokuno et al. (2008) has revealed
position- and direction-dependent modulation of the SOL H-reflex during body sway in
anterior-posterior direction. The result of that study showed that the size of SOL Hreflex was larger during a sway in the anterior direction than that in the posterior
direction. This finding supported their earlier study, which demonstrated a greater
(+12%) SOL Hmax/Mmax ratio when the centre of pressure (COP) position was moving in
the anterior direction than that in the posterior direction (Tokuno, et al., 2007). These
reports indicate that the SOL H-reflex modulation is related to the direction of postural
sway when the triceps surae is eccentrically or concentrically contracting to maintain
the upright standing posture. However, currently there is no information to support
whether the SOL H-reflex modulation during upright standing is correlated to that
during dynamic muscle activities. This information can advance our knowledge in
understanding of neural control strategy with respect to balance adjustment capacity and
negotiation of postural disturbance (Mackey & Robinovitch, 2006). This is particularly
important to the older population because the ability to control posture in relation to fall
prevention is a specifically relevant issue in this population (Horak, 2006).
Recently, Duclay et al. (2005; 2008; 2009) utilised an angular velocity of 60/s and a
wide range of ankle joint movement (between plantarflexion 30 and dorsiflexion 30)
to investigate movement-related changes in the SOL H-reflex in young adults. However,
this wide range of movement may be not appropriate in testing older adults due to their
limited range of motion at the ankle joint (Gajdosik, Vander Linden, & Williams, 1999).
The present study (Study III) adopted similar methods with a modification of the joint
angle range (between plantarflexion 20 and dorsiflexion 20) to examine the agerelated changes in the SOL H-reflex modulation during dynamic ankle movements. This
range of movement was close to the maximal recovery angle of the ankle joint (16.3
106
1.5) during upright standing (Mackey & Robinovitch, 2006). The maximal recovery
angle of the ankle joint is the maximum forward lean angle where participants could
recover a stable upright stance by contraction their ankle muscles. The second aim of
Study III was to determine the correlation in the SOL H-reflex modulations between
shortening and lengthening muscle actions and postural tasks.
6.2. Methods
6.2.1. Participants
Twenty young (10 males, 10 females, age: 24.4 4 years, height: 168 6.6 cm, weight:
63.2 12.9 kg) and twenty older (10 males, 10 females, age: 73.3 5 years, height:
166.1 7 cm, weight: 74.8 12.6 kg) healthy volunteers participated in the study. An
established health status screening questionnaire (Appendix D) was used to identify any
contraindication to participation. The volunteers who met the inclusion criteria (Section
3.1 in Chapter 3) and had no contraindication to participation signed an Informed
Consent form (Appendix B) and undertook a familiarization session prior to the testing
session. All participants were instructed to refrain from taking caffeine-containing
substances and smoking within 2 hours of the testing session, and were asked not to
perform strenuous exercise activity for 24 hours before the testing session.
6.2.2. H-reflex test
The procedures of H-reflex test and the method of surface EMG signal processes were
61).
107
6.2.3. Passive and active dynamic muscle actions
used to set shortening and lengthening plantarflexors actions. Constant angular velocity
was set at 60/s. This velocity was chosen due to the high level of reproducibility of Hreflex test during dynamic muscle actions under this condition (Duclay & Martin, 2005;
Duclay, et al., 2008; Duclay, et al., 2009), and the validity of isokinetic ankle strength
measurement in older adults (Hartmann, Knols, Murer, & de Bruin, 2009), reported in
the literature. The procedures of the Biodex dynamometer setting were presented in the
Chapter 3 General Methodology, Section 3.3 (page 57).
The procedure for the passive and active dynamic muscle test were presented in the
Chapter 3 General Methodology, Section 3.4 (page 59).
Prior to the passive and active dynamic muscle test, the participants were given a 5 min
warm-up exercise, which was described in the Study III (page 57).
In the passive dynamic test, the H-wave and M-wave (H/M) recruitment curve was
established (Figure 6.1). The stimulation intensity was progressively increased, with
increments of 10 mA, from the threshold of H-reflex to where there was no more
increase in the M-wave amplitude. The intensity for eliciting the Hmax was subsequently
checked by using 2 mA increment (Racinais, et al., 2008). The mean value of SOL Hmax
and Mmax was determined by averaging four repetitions in the passive shortening or
lengthening condition and was used for data analysis. To preclude any influence of
homosynaptic post-activation depression, a minimum of 10 s ISI was used (Hultborn, et
al., 1996).
In the active dynamic test, four successful trials were recorded for each of the H-reflex
and the M-wave during the maximal shortening or lengthening contractions. The
108
stimulation intensity used for the H-reflex was that induced Hmax during the passive
movement, whereas that for the M-wave was the supra-maximal stimulation (1.5 times
of Mmax stimulation intensity) used during the passive movement.
It is understood that alteration of the spatial relationship between stimulation electrode

and the targeted nerve during contraction can shift the H-wave and M-wave recruitment
curve from that established during rest (Capaday, 1997). Furthermore, the simulation
intensity that can induce the maximal amplitude of H-reflex during passive muscle
action might not be identical to that during maximal muscle contraction. However, it
would be difficult to establish the H/M recruitment curve and Hmax under various
conditions in one day due to the time required and the possibility of fatigue that may
affect the testing outcomes (Gandevia, 2001). Therefore, we limited our study to the Hreflex evoked by the same stimulation intensity for the purpose of comparison. This
method has been validated and utilised by Duclay et al. (2005; 2008; 2009).
6.2.4. Postural tasks
In the postural test, the H-wave and M-wave (H/M) recruitment curve was established
during standing with the SO condition (Figure 6.1). The procedures of the static postural
test were presented in the Chapter 3 General Methodology, Section 3.5 (page 60). The
force plate was not used in Study III.
All variables were measured off-line for the respective experimental conditions and
were processed by the researcher. The digital EMG data was transferred to Excel
datasheets and then all variables were measured on the Excel sheets (Microsoft Office,
2003 version).
109
the Grubbs test with the probability level of 0.05 was used to confirm and eliminate the
significant outliers (Section 3.7 in Chapter 3). Statistical analysis was conducted
following elimination of significant outliers. Descriptive data was presented as mean
standard deviation (SD). A two-factor [group (2) x action type (2)] RMANOVA was
performed to determine the main effect of age and action type, and the interaction
between these factors on the Hmax/Mmax ratio. When a significant interaction was found,
a post-hoc analysis with Bonferroni adjustment was conducted to identify where the
differences between the mean values occurred. Pearson product-movement correlation
(Bivariate test) with two-tailed test was performed to determine the relationship
between the SOL Hmax during the dynamic muscle actions and that during the postural
tasks. The correlation analyses were performed on the data within each group. An alpha
value of P 0.05 was set for significant differences between the means. All data were
analysed by using SPSS software for Windows (SPSS Inc, Chicago, IL, ver. 16.0).
6.3. Results
In the passive lengthening test, the SOL H-reflex response could not be elicited from
two older participants. Data from these two participants were excluded from data
analysis. The original EMG traces of the SOL H-reflex in all testing conditions from a
representative young and older participant, respectively, are demonstrated in Figure 6.2.
6.3.1. Dynamic muscle action test
The SOL Hmax/Mmax ratio during the passive movement demonstrated a significant
Group by Action type interaction (P = 0.001) and a main effect of Action type (P =
0.001). The post-hoc comparisons with Bonferroni adjustment revealed that the young
110
group had a higher SOL Hmax/Mmax ratio than that of the older group during the passive
lengthening action (P < 0.05). No significant difference of the SOL Hmax/Mmax ratio was
found during the passive shortening action (P > 0.05). The post-hoc comparisons with
Bonferroni adjustment also showed that the SOL Hmax/Mmax ratio during the passive
shortening movement was significantly higher than that during the passive lengthening
movement in the older group (P < 0.05; shortening: 0.40 0.22 > lengthening: 0.15
0.10). In contrast, the young group demonstrated no significant difference of the SOL
Hmax/Mmax ratio between the passive shortening and lengthening movements (0.45
0.19 and 0.48 0.20, respectively; P > 0.05).
There was no significant interaction and no significant Group main effect for the SOL
Hmax/Mmax ratio during active shortening and lengthening contractions (P > 0.05).
However, a significant main effect of Action type for the SOL Hmax/Mmax ratio during
active dynamic muscle actions was found (P = 0.001). The SOL Hmax/Mmax ratio of the
young group was significantly higher during shortening contraction than that during
lengthening contraction (P < 0.05; shortening: 0.51 0.26 > lengthening: 0.37 0.18).
There was no significant difference in the SOL Hmax/Mmax ratio found between the
shortening and lengthening muscle contractions in the older group (P > 0.05; shortening:
0.40 0.22 > lengthening: 0.36 0.18) (see Figure 6.3).
6.3.2. The correlation between the maximal SOL H-reflex amplitude in the
dynamic muscle action test and postural control tests
As illustrated in Table 6.1, there was a moderate correlation of the SOL Hmax between
the passive and active dynamic muscle actions and the four postural conditions in the
young group (P < 0.05; from r = .49 to r = .75). In contrast, the older group only
demonstrated moderate correlations between the SOL Hmax during the passive
shortening and lengthening movements and that during the postural tasks (P < 0.05,
111
from r = .54 to r = .68). The SOL Hmax of the older group during the shortening and
lengthening contractions was not significantly correlated to that recorded during the
postural conditions (P > 0.05, from r = -.15 to r = .18).
112
A
8
7
Amplitude (mV)
6
5
Shortening H-reflex
Shortening M-wave
Lengthening H-reflex
Lengthening M-wave
2
1
0
0
10
20
30 40 50 60 70 80
Stimulation intensity (mA)
90
100 110
B
6
Amplitude (mV)
5
4
Standing H-reflex
Standing M-wave
2
1
0
0
10
20
30
40
50
60
70
80
90
100 110

C
3.5
Amplitude (mV)
3
2.5
Shortening H-reflex
Shortening M-wave
Lengthening H-reflex
1.5
Lengthening M-wave
1
0.5
0
0
10 20 30 40 50 60 70 80 90 100 110 120 130 140

113
D
4
3.5
Amplitude (mV)
3
2.5
Standing H-reflex
Standing M-wave
1.5
1
0.5
0
0
10 20 30 40 50 60 70 80 90 100 110 120 130 140

Figure 6.1: H-reflex and M-wave recruitment curves during passive shortening (Hreflex: ; M-wave: ) and lengthening (H-reflex: ; M-wave: ) actions, and standing
on stable surface with eyes opened (H-reflex: ; M-wave: ) from representative one
young (A, B) and one older (C, D) participants, respectively. Please note the differences
in Y and X scales.
114
PS
PL
AS
AL
2mV
10ms
0.7mV
10ms
SO
SC
UO
UC
0.5mV
10ms
0.3mV
10ms
Figure 6.2: Representative raw EMG traces of the maximal soleus H-reflex during
passive and active dynamic muscle activities and four different postural conditions from
one young (solid lines: A, C) and one older (dotted lines: B, D) participants. Please note
the difference in Y-scale for the EMG traces. PS: passive shortening action; PL: passive
lengthening action; AS: active shortening action; AL: active lengthening action, SO:
stable surface with eyes open; SC: stable surface with eyes closed; UO: unstable surface
with eyes open; UC: unstable surface with eyes closed.
115
**
Hmax/Mmax ratio
0.8
0.6
0.4
0.2
0
Young
Older
B
1
**
Hmax/Mmax ratio
0.8
0.6
Shortening
Lengthening
0.4
0.2
0
Young
Older
Figure 6.3: The soleus Hmax to Mmax ratio during A) passive shortening and lengthening
movements, and B) voluntary shortening and lengthening muscle contractions. Data are
mean values and standard deviations. *: P < 0.05, between both groups. **: P < 0.05
between action types.
116
Table 6.1: Pearson correlation coefficients of the maximal soleus H-reflex amplitude
between dynamic muscle activities and postural tasks, in the young and older adult
groups.
Postural tasks
SO Hmax
SC Hmax
UO Hmax
UC Hmax
0.74 *
0.74 *
0.75 *
0.74 *
0.5 *
0.54 *
0.52 *
0.5 *
0.63 *
0.6 *
0.6 *
0.6 *
0.53 *
0.49 *
0.51 *
0.52 *
0.57 *
0.67 *
0.68 *
0.62 *
0.55 *
0.54 *
0.58 *
0.46
0.1
0.1
0.14
0.18
- 0.15
0.1
0.1
0.1
Young group (n = 20)

PS Hmax
PL Hmax
AS Hmax
AL Hmax
Older group (n = 18)
PS Hmax
PL Hmax
AS Hmax
AL Hmax
*:
P < 0.05, significant correlation with 2-tailed test. PS = passive shortening action, PL
= passive lengthening action, AS = active shortening action, AL = active lengthening
action, SO = stable surface with eyes open, SC = stable surface with eyes closed, UO =
unstable surface with eyes open, UC = unstable surface with eyes closed; Hmax=
maximal amplitude of SOL H-reflex.
117
6.4. Discussion
6.4.1. Age-related changes in soleus H-reflex modulation during shortening and
lengthening plantarflexors actions
Previous studies utilised a variety of experimental conditions to investigate the SOL Hreflex modulation during dynamic ankle movement in young adults. For example, in
Pinniger et al.s study (2001), a smaller range of motion for ankle joint movement
(between plantarflexion 5 and dorsiflexion 5) with slower constant angular velocities
of 2, 5 and 15/s were used, the testing was performed in lying prone body position and
the stimulation intensity was controlled at 50% of Hmax and 50% of Mmax. A study
conducted by Nordlund et al. (2002) also utilised lying prone position with an angular
velocity at 5/s to measure the SOL H-reflex during submaximal shortening and
lengthening dynamic contractions. In Duclay and Martins study (2005), although the
testing body position, constant angular velocity, contraction intensity and stimulation
intensity were similar to our study, the range of ankle motion (between plantarflexion
30 and dorsiflexion 30) was larger than that used in the current study. It has been
reported that angular velocity can alter the amplitude of SOL H-reflex during passive
(Pinniger, et al., 2001) and active lengthening actions (Duclay, et al., 2009). Also,
changes in muscle length can affect muscle spindle discharge rates and the Ia afferent
inputs. Although the H-reflex response was elicited at same angle of ankle joint, the
preceding muscle spindle activity from different ankle joint positions may lead to
different level of presynaptic and postsynaptic inhibitions of the spinal motoneurons via
peripheral and central regulations (Guissard, et al., 2001). These different experimental
settings may make the results not directly comparable, particularly in relation to older
adults. In the present study, the responses of the young and older participants were
compared under the same experimental conditions. The differences found in the SOL
118
Hmax/Mmax ratio between action types in both age groups underscore the age-related
difference in the SOL H-reflex modulation during passive and active dynamic muscle
actions.
It is interesting to find that the older group demonstrated movement-related changes in

the SOL Hmax/Mmax ratio during passive actions. It has been previously suggested that
increased presynaptic inhibition through primary afferent depolarization contributed to
the SOL H-reflex depression during passive lengthening movement as observed in
young adults (Duclay & Martin, 2005; Nordlund, et al., 2002; Pinniger, et al., 2001). It
is likely that the presynaptic inhibition of Ia afferent is the major factor that contributes
to the SOL H-reflex modulation during the lengthening movement. However, the
presynaptic inhibitory mechanisms might not be attributable factors to the reduction of
group Ia afferent volleys with ageing. Evidence suggests that older adults have less
ability to utilise the presynaptic inhibitory mechanisms to modulate the Ia afferent
inputs at the spinal level during rest (Angulo-Kinzler, et al., 1998; Chalmers & Knutzen,
2002; Earles, et al., 2001). Burke et al. (1978) observed that muscle spindle afferent
discharge was quiescent during passive shortening activity but it increased rapidly
during passive lengthening activity. It is possible that the influence of muscle spindle
output might be responsible for the significant modulation of SOL H-reflex in the older
group. However, function and structure of the intrafusal and extrafusal fibres may be
compromised with the ageing process (Shaffer & Harrison, 2007). In the current
literature, the neural control of lengthening exercise in older adults is not well
understood. Therefore, it can only be speculated that the movement-related change in
muscle spindle discharge may play a major role in the SOL H-reflex modulation in the
older group.
119
When the SOL Hmax/Mmax ratio was assessed during maximal voluntary shortening and
lengthening contractions, differences were found between young and older adults. The
SOL Hmax/Mmax ratio of the young group during the shortening contraction was
significantly higher than that during the lengthening contraction. However, this
contraction-related change of the SOL Hmax/Mmax ratio was not significant in the older
participants. Significant change in the SOL Hmax/Mmax ratio of the young group is in
agreement with the previous studies that demonstrated a depression of the SOL H-reflex
during lengthening muscle contraction at submaximal and maximal levels, in
comparison to that during shortening contraction at the same intensities (Duclay &
Martin, 2005; Nordlund, et al., 2002; Romano & Schieppati, 1987). Duclay and Martin
(2005) suggested that presynaptic inhibition controlled by descending inputs and the
homosynaptic postsynaptic depression were the two possible mechanisms underlying
the depression of SOL H-reflex during maximal lengthening contraction. In addition,
there is a difference in cortical motor outputs during shortening and lengthening
contractions (Duchateau & Enoka, 2008). Sekiguchi et al. (2003) reported that the
MEPs induced by TMS decreased during lengthening plantarflexion, compared to that
during shortening plantarflexion, although the SOL and TA bEMG values were similar
between the two contraction types. This decrease in MEPs may indicate a reduction of
the cortical descending inputs when the calf muscle voluntarily contracts in a
lengthening action.
The results of Study III showed that, in contrast to the young group, the contractionrelated motor regulation controlled by the supraspinal mechanisms was not different
between maximal shortening and lengthening contractions in the older group. However,
a potentiation of the SOL Hmax/Mmax ratio during lengthening action was observed in the
older group, from passive (0.15 0.1) to active (0.36 0.18) conditions. This
observation may indicate that the older adults used specific descending commands to
120
adjust synaptic inputs in the spinal motoneuron pool during lengthening contractions. It
has been shown that the supraspinal control of presynaptic inhibition in Ia-motoneuron
system during voluntary muscle contraction is preserved in the older population (Earles,
et al., 2001). The shift of spinal inhibitory mechanisms could lead to strong neural
modulation in the Ia afferent pathway. Therefore, it is likely that the absent modulation
of the SOL Hmax/Mmax ratio between maximal shortening and lengthening contractions
found in the older group is related to central regulation of motor output.
6.4.2. Correlation of soleus H-reflex modulation between dynamic plantarflexion

actions and postural tasks in young and older adults
The present data demonstrated that there was a significant correlation between the SOL
H-reflex responses during both the passive and active dynamic muscle actions and that
during the postural tasks in young adults. In contrast, in the older group, the significant
correlation was only found between the passive muscle movements and the postural
tasks. Several studies have reported that the H-reflex modulation is correlated to phase
and direction of postural sway and postural stability (Earles, et al., 2000; Taube, Leukel,
et al., 2008; Tokuno, et al., 2007; Tokuno, et al., 2008). This relationship is also
evidenced by alteration of H-reflex size in parallel with improvement of balance control
after short-term balance training (Taube, Gruber, et al., 2008; Taube, Kullmann, et al.,
2007). With careful control of postural direction and sway position during upright
standing, Tokuno et al (2007; 2008) speculated that the position- or direction-dependent
SOL H-reflex response was related to the shortening or lengthening activation of the
calf muscle. Recently, Laudani et al. (2009) demonstrated an exercise-induced SOL Hreflex depression associated with increased body sway in anterior-posterior direction
with repeated bouts of plantarflexion movements during upright standing. Although the
central processing mechanisms are not identical between dynamic muscle action and
121
postural control, a common characteristic is that all sensory and motor inputs finally
converge to the spinal motoneuron pool for execution of the motor response (Horak,
2006; Martin, Butler, Gandevia, & Taylor, 2008). Based on these observations, the Hreflex modulation during plantarflexion may reflect the relationship of Ia afferent
feedback between dynamic muscle activity and upright standing. It should be noted that
the functional role of the spinal neural circuitry is multifaceted in postural control.
Although this study was limited by not including measurement on COP variation, the
current results indicate a functional association of SOL H-reflex variation during
dynamic ankle movement and postural tasks.
6.5. Conclusion
In conclusion, this study was the first to demonstrate the influence of shortening and
lengthening plantarflexions on the SOL H-reflex modulation in older adults. Dissimilar
modulations of the SOL H-reflex during passive dynamic movements and voluntary
maximal dynamic contractions were found between young and older adults. These
findings suggest an age-related neural adaptation during different modes of
plantarflexion movements. Moreover, this study was the first to report a significant
correlation between the Ia excitatory effects on the SOL motoneuron activity during
dynamic ankle movements and that during upright postural conditions. The significant
correlation found between the maximal amplitude of SOL H-reflex in the dynamic
muscle actions and the postural tasks indicates a relevant characteristic of group Ia
afferent inputs in control of shortening and lengthening plantarflexions and upright
standing in young and older adults.
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Chapter 7 Effect of 12 weeks Tai Chi training on

soleus H-reflex and control of static posture in older adults
(Study IV)
123
7.1. Introduction
The ability to maintain balance and control posture is critical to older adults in relation
to prevention of falls. Poor ability to control upright posture is a risk factor to falls
during daily physical activities in the older population (Horak, 2006). It has been
reported that 30-40% of community-dwelling older adults aged over 65 years
experience at least one fall each year (Sturnieks et al. 2010; Tinetti, et al., 1988). In 30
% of these fall incidences, fallers suffer severe injuries (e.g. hip bone fractures or head
trauma) that require intensive clinical treatments (Rao, 2005). Consequently, injuries
caused by falls result in restricted mobility, functional decline and healthy
complications that can severely impact on quality of life.
Exercise training is recommended as a primary intervention to prevent falls (Sturnieks

et al. 2010). In all, balance training has been strongly recommended as an appropriate
mode of exercise to improve balance capacity in older adults who are at risk of falls
(Nelson, et al., 2007). The conventional method of balance training is to apply static
balance training on an unstable surface with or without eyes closed condition. The
participant being trained is required to maintain upright posture during bipedal standing
on a soft mattress, wobble board, ankle disc, and tilt board (DiStefano, Clark, & Padua,
2009). There are also some methods that involved use of unstable sandals (Micheli,
Ross, Blackburn, Hirth, & Guskiewicz, 2006), elastic bands with dynamic movements
(Puls & Gribble, 2007) for balance training under dynamic circumstance. DiStefano and
colleagues (2009) reported that the benefits of physiological adaptations were
manifested when the participants had undertaken balance training of at least 10 min per
session, 3 times per week, for 10 weeks training period. It is suggested that the
functional improvement in control of upright posture after balance training is associated
with neural adaptations at spinal and supraspinal levels (Taube, Gruber, et al., 2008).
124
The Hoffmann reflex (H-reflex) is frequently used to investigate the effect of group Ia
excitatory inputs on the -motoneuron activation in human movement studies. The
amplitude of H-reflex response recorded during a given motor task reflects a summation
of neural inputs converged to the target -motoneurons. The variation of H-reflex before
and after an exercise training intervention can be used as an indication of spinal
plasticity in relation to motor learning (Hultborn, 2006). The H-reflex test has been
shown to be reliable in test-retest measurements separated by several weeks, during rest
in prone (intraclass correlation coefficients, ICC=0.97) (Ali & Sabbahi, 2001), sitting
(ICC=0.92) (Hopkins, et al., 2000) or standing positions (ICC=0.80) (Chen, et al., 2010).
Tai Chi (or Tai Ji) is a Chinese martial art exercise which was originally developed as a
self-defense skill (Wu, 2002). Practicing TC regularly has been shown to improve the
neuromuscular functions, such as joint stability (Tsang & Hui-Chan, 2003, 2004b),
tactile acuity (Kerr, et al., 2008), posture control (Tsang & Hui-Chan, 2004a; Tsang, et
al., 2004), and lower limb muscle strength (Choi, et al., 2005; Li, et al., 2009). TC
movements are frequently performed within a domain of postural maintenance in either
unipedal or bipedal standing (Hong, Mao de, & Li, 2008). For older adults, the
beneficial outcomes of TC exercise in improvement of balance control have been
reported in a number of laboratory-based studies (Fong & Ng, 2006; Lelard, Doutrellot,
David, & Ahmaidi, 2010; Tsang & Hui-Chan, 2004a, 2004b, 2006). It has been found
that improvement of posture control in older adults with TC training is a result of neural
adaptation in kinematic proprioception (Fong & Ng, 2006; Li, et al., 2008; Tsang &
Hui-Chan, 2003, 2004b), vestibular function (Tsang & Hui-Chan, 2006; Tsang, et al.,
2004), and reaction time (Fong & Ng, 2006). In addition, increased lower limb muscle
strength is associated with balance improvement after TC training (Choi, et al., 2005;
Tsang & Hui-Chan, 2005). It has been suggested that muscle strength is an important
125
factor in control of balance (Orr, 2010). However, whether the neural adaptations to TC
training can be reflected in H-reflex modulation has not been examined.
There is a number of cross-sectional researches reported that older adults who have
practised TC for more than 1 year have better balance control than their non-practicing
counterparts (Tsang & Hui-Chan, 2004b, 2005, 2006; Tsang, et al., 2004). Significant
improvement of postural control in functional balance assessments has also been
observed on older adults after 16 weeks (Li, et al., 2008; Voukelatos, et al., 2007) or 12
weeks (Choi, et al., 2005; Takeshima et al., 2007) TC training, with a training frequency
of 3 times per week. In addition, significant improvement of balance performance can
be achieved after 4 week TC training if training frequency increases to 6 times per week
(Tsang & Hui-Chan, 2004a). However, a high frequency of training intervention may be
difficult to apply to some studies, which use a voluntary basis to recruit participants.
In the literature, information in relation to the effect of exercise training on H-reflex

modulation and postural control in older adults is very limited. To our knowledge, only
one study by Mynark and Koceja (2002) provides relevant information in this area. The
study showed that the SOL H-reflex amplitude was changed (18.7 to 21.0% decrease)
after a short-term balance training on two consecutive days. The changes in the SOL Hreflex were accompanied by 10.1% decrease in the mean displacement of the COP. It is
suggested that reflex regulation contributes to autogenic adjustment of muscle stiffness
in the postural muscles during upright standing (Loram, et al., 2005b). The downregulation of SOL H-reflex may be a result of functional adaptation that decreases
postural reflexive regulation due to postural improvement after balance training.
However, the effect of longer-term training on H-reflex modulation and postural control
has not been examined. Therefore it was hypothesized that the adaptations of the SOL
H-reflex is associated with an improvement in postural control after 12 weeks of TC
126
training. The hypothesis was based on the assumption that TC is a promising type of
balance exercise (Verhagen, et al., 2004). The SOL H-reflex was the focus in Study IV
because the function of SOL muscle is essential in controlling an upright standing
posture (Winter, et al., 2001). The aim of Study IV was to determine the effect of 12
weeks of TC training on the H-reflex modulation and postural control during upright
standing in older adults.
7.2. Methods
7.2.1. Participants
Healthy adults in the age range of 65 and 85 years, both males and females, were
recruited from the local community on a voluntary basis. All participants were tested
before and after the 12 weeks of TC training. A health status screening questionnaire
(Appendix D) was used to identify if there was any contraindication to participation.
Exclusion criteria included the presence of neuromuscular disorders, current lower
extremities injuries, true vertigo, acute illness, corrected visual acuity worse than 20/100
or presence of a field defect, current use of medications known to impair balance
control (e.g. benzodiazepines, anticonvulsants or antidepressants), use of a walking aid
and previous experience in TC. Thirty-four (34) community-dwelling individuals (17
males and 17 females) were accepted and assigned into either a TC training group
(11males and 9 females) or a Control group (6 males and 8 females) by their preference.
All participants signed an Informed Consent form (Appendix B) and undertook a
familiarisation session prior to the pre-training test. All participants were instructed to
refrain from taking caffeine-containing substances and smoking within two hours of the
testing session, and were also asked to avoid performing strenuous exercise or physical
activity for 24 hours before the test. The participants in the TC group were instructed
not to undertake any concurrent training during the 12-week period, whereas the
127
participants in the control group were asked to maintain their regular daily physical
activities during the same period. All participants were tested before and after the 12
weeks of TC training.
7.2.2. Tai Chi training program
The TC training program was delivered by a qualified TC instructor. The participants

practiced TC exercise three times per week for 12 weeks. Each training session
consisted of 20 minutes warm-up exercises and 40 minutes TC practice. The warm-up
exercises consisted of 22 modified TC activities. The first six weeks was a learning
period. The participants were taught the 13-form Yang style TC (Appendix E)
(Verhagen, et al., 2004). In the following six weeks, the participants repeatedly
practiced the sequence at a slow pace and the individuals performance was closely
monitored by the instructor.
The procedures of H-reflex test and the method of surface EMG signal processes were
61).
7.2.4. Static postural test
The procedures of static postural test and the method of COP signal processes were
61).
128
The participants were standing barefoot on the floor while repeated stimuli were
delivered to SOL to establish the H-wave and M-wave recruitment curve. The
stimulation intensity was increased progressively with increments of 10 mA until there
was no further increase in the Mmax. The intensity for eliciting the Hmax was
subsequently checked by using 2 mA increments (Racinais, et al., 2008). The mean
values of SOL Hmax and Mmax were determined by averaging four trials. Once the
recruitment curves were constructed, the participants moved to the force plate for the Hreflex and static postural tests.
During the main test, the participants were required to maintain an upright stance for at
least 10 s in each trial. Electrical stimulation was applied to record the SOL H-reflex
followed by 10 s COP measurements. A 10 s rest interval was given between trials. The
mean values of SOL Hmax and COP variables were determined by averaging 10 trials
under each sensory condition, as recommended by Mynark (2005). Three minutes rest
interval was given between sensory conditions. The whole experimental protocol was
completed within 1.5 hours.
The parameters of focus included the Hmax/Mmax ratio and the mean displacement of
COPA-P and COPM-L. All parameters were measured off-line for the respective
experimental condition and were processed by the same researcher. The digital EMG
data was transferred to Excel datasheets and then all values of the variables were
determined on the Excel sheets (Microsoft Office, 2003 version). The data files were
coded by a serial number, therefore the researcher was blind to the participants
Intervention or Control status during the analysis.
129

the Grubbs test with the probability level of 0.05 was used to confirm and eliminate the
significant outliers (Section 3.7 in Chapter 3). Statistical analysis was conducted after
elimination of significant outliers, if any (SPSS Inc, Chicago, IL, ver. 16.0). Descriptive
data of the measured variables was presented as mean standard deviation (SD). An
independent t test was used to compare age, height, body weight and BMI between the
two groups in the baseline measurement. A two-way [Time (2, pre vs post tests) x
Condition (4 sensory conditions)] RMANOVA with Group as the between-subject
factor was performed to compare the Hmax/Mmax ratio and mean displacement of COPA-P
and COPM-L. When a significant main effect or an interaction was found, a post-hoc
analysis with Bonferroni adjustment was conducted to identify the differences between
the mean values. An alpha value of P 0.05 was set for significant differences between
the means.
7.3. Results
7.3.1. Participants
The physical profiles of age, height, body weight, and BMI of the TC and control
groups are shown in Table 7.1. No significant differences were found between the two
groups in all comparisons prior to training.
7.3.2. Elimination of outlier data points
In the pre-training test, a significant outlier of the SOL Hmax/Mmax ratio was found in the
respective sensory condition in the training group. In the post-training test, there was a
significant outlier of the SOL Hmax/Mmax ratio in the SO condition in the control group.
130
For the mean displacement of COP measurement, a significant outlier of the COP A-P
was identified in the SO, UO, and UC conditions from one control participant. These
data points were excluded from further statistical analysis.
The results revealed no significant Condition or Group main effect and no significant
interaction for the SOL Hmax/Mmax ratio. However, a significant main effect of Time
(pre-post 12 weeks) was observed (P = 0.048). Significant differences were found in the
SOL Hmax/Mmax ratio between the pre-training and post-training tests in the TC group in
all of the four sensory conditions (P < 0.05), whereas no significant difference was
found in the Control group. The SOL Hmax/Mmax ratio of the training group increased by
20%, 31%, 26% and 28 % from the baseline measurement, in the SO, SC, UO, and UC
sensory conditions, respectively. In contrast, the SOL Hmax/Mmax ratio of the Control
group did not show any significant change (by -4%, -4%, 0.4%, and 0.3%) in the
respective sensory condition after the 12-week period (see Figure 7.1).
7.3.4. Centre of pressure measurement
There was no significant Time or Group main effect and no significant interaction for
the mean displacement of COPA-P and COPM-L. However, there was a significant main
effect of Condition for the COPA-P (P = 0.001) and COPM-L (P = 0.001) measurements
(Table 7.2). In the pre-training test, statistically significant differences of the COPA-P
and COPM-L were found between all sensory conditions, except the COPA-P and COPM-L
of SC vs UO comparisons in the Control group (P = 0.108 and P = 0.068, respectively).
In the post-training test, significant differences of the COPA-P and COPM-L were also
found between all sensory conditions, except the COPA-P and COPM-L of SO vs SC
comparisons in the TC group (P = 0.113 and P = 0.069, respectively) and the COPA-P of
131
SC vs UO (P = 0.317) and COPM-L of SO vs SC and UO vs UC comparisons in the

control group (P = 0.49 and P = 0.181, respectively) (Table 7.2).
132
0.6
SC Hmax/Mmax ratio
SO Hmax/Mmax ratio
0.6
0.4
0.2
*
0.4
0.2
0
Tai Chi
Control
Tai Chi
Control
0.6
UC Hmax/Mmax ratio
UO Hmax/Mmax ratio
0.6
*
0.4
0.2
*
0.4
0.2
0
Tai Chi
Control
Tai Chi
Control
Pre-training test
Post-training test
Figure 7.1: The SOL Hmax/Mmax ratio before and after 12-week Tai Chi training in the
Training and Control groups. A) stable surface with eyes open; B) stable surface with
eyes closed; C) unstable surface with eyes open and D) unstable surface with eyes
closed. *: P < 0.05, significant difference between the pre-and post-training tests.
133
Table 7.1: Comparison of physiological characteristics between the Tai Chi group and
Control group prior to the 12 weeks of TC training.
Tai Chi group (N = 20)
Control group (N = 14)
P value
72.9 4.4
72.9 6.5
.982
11M 9F
6M 8F
.501
Height (cm)
168.3 9.6
163.8 6.3
.139
Body Weight (kg)
75.5 15.3
70.5 8.9 kg
.288
26.5 3.7
26.2 2.5
.817
Age (years)
Gender (male/female)
BMI (kg/m )
Note: Data are expressed as mean SD.
134
Table 7.2: Mean values and standard deviation of the mean displacement of COP values
(anterior-posterior and medial-lateral directions) in four sensory conditions in the Tai
Chi and Control groups before and after the 12-week training or control period.
Tai Chi group (N = 20)
Control group (N = 14)
COPA-P (mm)
COPM-L (mm)
COPA-P (mm)
COPM-L (mm)
Pre-training
3.02 ( 0.9) #
1.76 ( 0.64) #
3.1 ( 0.8) #
1.77 ( 0.68) #
Post-training
3.03 ( 0.84) #
1.87 ( 0.63) #
2.67 ( 0.41) #
1.65 ( 0.55) #
Pre-training
3.54 ( 1.18) *#
2.26 ( 1.05) *#
3.68 ( 0.8) *#
2.22 ( 0.86) *#
Post-training
3.44 ( 0.95) #
2.07 ( 0.77) #
3.39 ( 0.9) *#
1.98 ( 0.73) #
Pre-training
4.11 ( 1.18) *#
2.77 ( 0.96) *#
4.31 ( 1.01) *#
2.72 ( 0.81) *#
Post-training
4.28 ( 1.52) *#
2.63 ( 0.85) *#
3.74 ( 0.69) *#
2.7 ( 0.83) *
Pre-training
4.96 ( 1.5) *
3.65 ( 1.4) *
5.14 ( 1.3) *
3.72 ( 1.22) *
Post-training
5.07 ( 1.79) *
3.76 ( 1.5) *
4.24 ( 0.79) *
3.16 ( 0.88) *
SO condition
SC condition
UO condition
UC condition
Note: No significant pre to post training differences were found in all variables. No
significant differences of the COP measurement in anterior-posterior and medial-lateral
directions were found between SO and SC conditions in the post-training test in the
training group. For the control group, the COPA-P and COPM-L between SC and UO
conditions were not significantly different in the pre-training test. No significant
differences were also found in the COPA-P between SC and UO conditions and the
COPM-L in SO vs SC and UO vs UC comparisons in the post-training test. SO = stable
surface with eyes open, SC = stable surface with eyes closed, UO = unstable surface
with eyes open, and UC = unstable surface with eyes close.
*: P < 0.05, differ significantly to the SO condition.
: P < 0.05, differ significantly to the SC condition.
: P < 0.05, differ significantly to the UO condition.
#: P < 0.05, differ significantly to the UC condition.
135
7.4. Discussion
7.4.1. Soleus H-reflex modulation
It has been suggested that the SOL H-reflex response is training-specific to a given
exercise intervention. For example, down-regulation of the SOL H-reflex was found
after balance training (Taube, Gruber, et al., 2007; Taube, Kullmann, et al., 2007;
Trimble & Koceja, 1994, 2001), whereas the SOL H-reflex was up-regulated after
strength training (Aagaard, et al., 2002; Taube, Kullmann, et al., 2007). The adaptation
of H-reflex to a specific training may be primarily attributable to the neural mechanisms
at the spinal cord level and possibly also related to the influences from the supraspinal
levels on the spinal inhibitory mechanisms (Schubert, et al., 2008; Taube, Gruber, et al.,
2008).
In the literature, the up-regulated H-reflex modulation was reported in association with
improvement of muscle strength after resistance training (Aagaard, et al., 2002;
Holtermann, Roeleveld, Engstrom, & Sand, 2007). Aagaard et al. (2002) demonstrated
that, through 14 weeks of resistance training in young adults, the SOL Hmax/Mmax ratio
was increased during maximal voluntary muscle contraction and was associated with
increases in the size of V-wave response (volitional response from the supraspinal level)
and plantarflexion muscle strength. In another study, a significant increase in the SOL
Hmax/Mmax ratio was also reported during a postural perturbation task after six weeks of
strength training in young athletes (Taube, Kullmann, et al., 2007). Conversely,
Scaglioni et al. (2002) reported that older adults showed significant improvement in
plantarflexion muscle strength but no change in H-reflex after 16 weeks of resistance
training. However, in that study the SOL H-reflex was assessed during rest in lying
prone position. These findings suggested that the increased H-reflex modulation may be
due to changes in the spinal inhibitory mechanisms (Aagaard, et al., 2002; Taube,
136
Kullmann, et al., 2007) and increased descending drives from the supraspinal levels
(Aagaard, et al., 2002), while the absence of H-reflex alteration during rest might
indicate the adaptations only occur to the specific motor tasks.
In Study IV, an increase in the SOL Hmax/Mmax ratio found in the TC group may indicate
an adaptive change of the spinal mechanisms after 12-week training. As mentioned
above, the up-regulation of SOL H-reflex is possibly due to an increase in neural
activation from the supraspinal levels. Because TC exercise is performed under weightbearing condition and most of the manoeuvres rely on knee and ankle movements
during unipedal or bipedal stance, increase in lower limb muscle strength has been
reported after participation in TC training for a period of 12 weeks (Choi, et al., 2005),
16 weeks (Li, et al., 2009), or more than three years (Tsang & Hui-Chan, 2005),
compared with sedentary controls. Thus, it appears that TC and resistance training may
both contribute to a similar training effect on neuromuscular adaptation.
7.4.2. Static postural control
A primary benefit of TC exercise is that it reduces the age-related deterioration in the

neuromuscular functions and improves postural control (Choi, et al., 2005; Lin, et al.,
2006; Takeshima et al., 2007). However, the results of Study IV showed no significant
changes in the mean displacement of COPA-P and COPM-L of the post-training test,
compared with that of the pre-training test in any of the four sensory conditions or in
either group. The absence of postural improvement in the current study may be related
to insufficient length of the training period. Our finding is in line with the reports that
showed no significant modification of static (Lelard, et al., 2010) and dynamic postural
control (Fong & Ng, 2006) after three months of TC practice. However, beneficial
effects on balance control have been reported after TC training for more than one year
(Fong & Ng, 2006; Tsang & Hui-Chan, 2004b; Tsang, et al., 2004). Older adults who
137
had practiced TC for three years or longer showed better postural performance during
unipedal standing (Tsang & Hui-Chan, 2005) and bipedal standing with alteration of
visual (Wu, et al., 2002) or vestibular (Tsang & Hui-Chan, 2006) condition than their
non-practicing counterparts.
7.4.3. The relationship between H-reflex modulation and postural control
The H-reflex modulation appears to be task-dependent. It has been shown that the
relationship between the H-reflex modulation and COP values during upright standing
under different visual and/or somatosensory conditions is reciprocal (Earles, et al., 2000;
Taube, Leukel, et al., 2008). For example, the SOL H-reflex amplitude is larger and the
COP value is smaller during upright standing under SO condition, compared to those
under UO condition. The major mechanism underlying the SOL H-reflex depression
during standing with UO condition has been suggested to be an increased presynaptic
inhibition that prevents oversaturation of the motoneurons (Chalmers & Knutzen, 2002).
This adaptation to challenging postural conditions acts as an important modulator in
order to receive more descending inputs for postural adjustment. However, this
relationship is altered after balance training as the SOL H-reflex is down-regulated in
parallel with decrease in COP values. It is suggested that the adaptive change of Hreflex is related to alterations of the spinal inhibitory mechanisms with a possible
involvement of descending control (Taube, Gruber, et al., 2008). The reduction of SOL
H-reflex after balance training is speculated to be the result of improved effectiveness of
motor output regulation. In Study IV, the SOL H-reflex increased during the postural
tasks with or without challenging conditions after 12 weeks TC training. The findings of
increased SOL H-reflex and unchanged COP, contrary to previous reports, may indicate
different effects between TC and balance training.
138
Balance training that was used in some studies included unipedal or bipedal standing
exercises on an unstable surface. The ability to control static upright posture is
improved after repeated exposure to the unstable surface (DiStefano, et al., 2009). In
contrast, TC exercise consists of a series of manoeuvres that are performed in a
continuous manner on a stable surface. Tai Chi practitioners practice all movements in
unipedal or bipedal support pattern and stepping in various directions (Hong, et al.,
2008). All manoeuvres are frequently performed in dynamic circumstances. Tsang et al.
(2005) reported that significant differences were found between TC practitioners with a
minimum of three years of training and healthy controls in dynamic postural control
tasks but not in statistic postural control tasks. Therefore, it appears that the H-reflex
adaptation demonstrates a task-dependent specificity. Future studies are needed to
confirm this speculation.
It should be noted that the volunteer participants in this study were assigned to the TC
and Control groups by their preferences. The participants were recruited from the same
local population and no significant differences were found between the two groups in
their age, height, body mass or health conditions. Although their preferences for group
allocation were mostly based on personal convenience rather than their physical
capability, the potential bias caused by this convenience sample could not be
determined, and is therefore a limitation of this study.
7.5. Conclusion
An increase of the SOL Hmax/Mmax ratio during static postural tasks was observed after
12 weeks of TC training in the older adults, under four sensory conditions. However,
the training-induced alteration in H-reflex was not accompanied by improvement of
performance in the static postural control tasks.
139
The outcomes of this study have been published in Archives of Physical Medicine
and Rehabilitation (See detail on page vi).
140
Chapter 8 Summary, conclusions and implications
141
8.1. Summary of results

The research work presented in this thesis provided quantitative evidence on the SOL
H-reflex modulation during different modes of muscle actions and static postural tasks
in older adults. The current results demonstrated the age-related differences in H-reflex
modulation that may indicate the differences of motor strategies in control of the SOL
muscle in older adults during various plantarflexors activities (e.g. ankle joint positions,
submaximal isometric voluntary muscle contractions, passive and active lengthening
and shortening muscle actions) and upright standing with eyes open or closed on a
stable or unstable surface. In addition, adaptation of the SOL H-reflex to 12 weeks TC
training in healthy older adults was investigated. The major outcomes of this thesis are
summarised below.
Study I
The reliability of SOL H-reflex measurement was affected by the level of muscle
contraction and the ankle joint position. Study I demonstrated a relatively high testretest reliability of the SOL H-reflex measurement during rest in the neutral ankle
position. The values of ICC were higher at the neural 0 and plantarflexion 20 than that
at the dorsiflexion 20 during rest. High reliability of test-retest H-reflex measurement
was also found during 10% MVC than that during 30% and 50% MVC.
Study II
The results of Study II showed that there is ageing effect of the SOL H-reflex during
submaximal isometric muscle contractions. The SOL H-reflex gain in the young adults
was relatively higher than that in the older adults during 10%, 30% and 50% MVC
contractions at the neutral, plantarflexion and dorsiflexion joint positions. Moreover, the
SOL H-reflex gain was significantly affected by the intensity of isometric submaximal
142
voluntary contraction. It is interesting to note that young and older groups together
showed a similar downward regulation of the SOL H-reflex gain when the intensity of
plantarflexors contraction progressively increased (10% MVC > 30% MVC > 50%
MVC) at the three ankle positions. This study also showed age-related changes in the
SOL H-reflex latency and duration in the three different ankle joint placements. The
latency of H-reflex was not influenced by ankle joint position in older adults, whereas it
was joint angle-dependent in young adults. There was a significant difference in the
SOL H-reflex duration between the young and older adults. The significant differences
of the SOL H-reflex results found between the young and older groups might reflect the
ageing effect on neural control of the SOL muscle.
Study III
The research in Study III was the first to demonstrate age-related differences in the SOL
H-reflex modulation during passive and active shortening and lengthening muscle
actions. The young group showed a significant difference of the SOL Hmax/Mmax ratio
during maximal shortening and lengthening muscle contractions, whereas the older
group demonstrated a significant difference of the SOL Hmax/Mmax ratio during passive
shortening and lengthening muscle movements. In the older group, a facilitation of the
SOL Hmax/Mmax ratio from passive to active lengthening muscle action was observed. In
addition, the results also revealed a significant correlation between the amplitude of
SOL Hmax during passive and active dynamic plantarflexors movements and that during
upright standing with eyes open/closed on a stable/unstable surface in young group. In
contrast, in the older group, the significant correlation was only found in comparison
between passive dynamic muscle action and postural tasks but not in comparison
between active dynamic muscle action and postural tasks.
Study IV
143
It is interesting to note that the results of SOL H-reflex and COP measurements after 12
weeks TC training were counter-intuitive to our hypotheses. An increase in the SOL
Hmax/Mmax ratio during bipedal standing with eyes open/closed on a stable/unstable
surface was observed after 12 weeks of TC training in the older adults. In the current
study, the SOL Hmax/Mmax ratio in the training group was facilitated with a range of
120% - 131% increase after TC training. However, the training-induced alteration in the
H-reflex was not accompanied by improvement of performance in the COPA-P and
COPM-L. The SOL H-reflex adaptation to TC training was not related to the ability to
control static upright posture when 12-week training was completed.
8.2. Conclusions
The research work presented in this thesis investigated the effect of ageing on the SOL
H-reflex modulation during different modes of ankle joint actions and during
maintenance of an upright posture under different visual and somatosensory conditions.
The adaptation of the SOL H-reflex to 12 weeks of TC training in older adults was also
examined in this thesis. The major conclusions from this series of studies are listed
below.
1) The conclusion of Study I was that testretest reliability of the SOL H-reflex was
apparently affected by the intensity of voluntary contraction and ankle joint position
selected in the experiment. A number of neural mechanisms, such as changes in motor
unit recruitment and firing frequency, presynaptic inhibition and reciprocal inhibition,
are potentially influencing repeated measurement of SOL H-reflex. To obtain reliable
outcome of the SOL H-reflex in the repeated measurements, the level of background
muscle activity and selection of ankle joint angle should be carefully considered. This
reliability study set a foundation for further studies on the H-reflex modulation in our
laboratory.
144
2) This study was the first to demonstrate the age-related differences of the SOL Hreflex gain during isometric voluntary muscle contractions with intensities of 10%, 30%
and 50% MVC at the neutral 0, plantarflexion 20 and dorsiflexion 20. The findings
support our hypothesis that ageing affects the spinal reflexive function during
submaximal voluntary muscle contractions. However, the young and older groups
together showed a similar downward regulation of the SOL H-reflex gain when the
intensity of plantarflexors contraction progressively increased. This observation
indicated a preservation of cortical motor function for regulation of SOL H-reflex gain
with progressive increase of contraction intensity in older adults. In addition, the results
of Study II also revealed age-related changes in the SOL H-reflex latency and duration
and the H-reflex latency was not joint angle-dependent in older adults. In physiological
point of view, factors underlying these changes are multiple but elongation of reflex
response and increase in H-reflex duration may result in slow reaction to adjust posture
during standing.
3) There were age-related differences in the SOL H-reflex modulation during passive
and active shortening and lengthening actions. This age-related difference in the SOL
H-reflex modulation during passive and active dynamic ankle movements was a novel
finding. The modulation of group Ia projection in older adults was affected by the mode
of muscle action (shortening and lengthening) during passive movement but not during
active contraction, while in young adults movement-related change of the SOL H-reflex
was observed during active contraction but not during passive movement. During
lengthening muscle action, increase in the SOL Hmax/Mmax ratio from passive to active
conditions was observed in older adults. This finding suggests a cortical contribution to
H-reflex modulation during dynamic muscle contraction in older adults. Furthermore,
the positive correlation found between the maximal amplitude of SOL H-reflex in
dynamic muscle actions and static postural tasks indicated the influence of group Ia
145
afferent inputs in control of shortening and lengthening plantarflexions and upright

standing in young and older adults. This correlation possibly indicated a similarity of
motor output regulation in the Ia-motoneuron pathway during dynamic plantarflexions
and upright standing. In older adults, the SOL H-reflex during static postural control is
significantly correlated to that during passive dynamic muscle action rather than active
dynamic muscle action may implicate an essential role of involuntary control in
regulating reflexive function during standing. The results have indicated the role of
group Ia afferent feedback on the SOL muscle activation in relation to static postural
control during bipedal standing in humans.
4) The increase of the SOL Hmax/Mmax ratio during static postural tasks after 12 weeks
of TC training in the older adults indicates a possibility of neural adaptation at the
supraspinal levels, similar to previous observations in resistance training (Aagaard, et al.,
2002; Taube, Kullmann, et al., 2007). However, the neural adaptation to 12 weeks of
TC training was not associated with balance improvement. It is unclear what
functionality is related to this adaptive change in the current study. Based on
observations in previous findings, benefit of balance improvement has been reported
after more than one year TC training. The training period engaged in the current study is
insufficient to see significant improvement of postural control. It is possible to assume
that functional improvement in postural control follows up physiological adaptation
during long-term TC practice. Nevertheless, the findings suggest a task-dependent
specificity of H-reflex adaptation to exercise training intervention in older adults.
8.3. Implications of the finding and future research

The findings of this thesis was first to demonstrate the age-related differences of motor
output regulation in control of the SOL muscle during various motor tasks. Furthermore,
the neural adaptation to 12 weeks TC training in older adults was revealed as the result
146
showing the potentiation of SOL Hmax/Mmax ratio after training. It is suggested that the
SOL muscle plays a dominant role in control of upright posture (Gatev, et al., 1999).
Understanding the motor functions of this antigravity muscle during different modes of
ankle movements can advance current knowledge of neural strategy in controlling ankle
muscles during postural control. This is particularly important for the older population
since improving balance control has become a major topic of health promotion in the
gerontological area (Horak, 2006; Tinetti, et al., 1988). Despite the literature regarding
the effect of aging on reflex function are little, this thesis provides fundamental
evidence to advance current knowledge in this context. The outcomes of this series of
studies will have implications to the H-reflex test in ageing study and clinical practice.
In clinical practice, the H-reflex measurement is usually used to test peripheral motor
function. Clinical practitioners often used the testing result to evaluate therapeutic
treatments in patients with neurological disorders. However, misjudgment of therapeutic
evaluation may occur if the standard of testing condition was not claimed in repeated
measurement. The results of Study I indicated that the ankle joint position and
background muscle activity may affect the reliability of the test outcomes. This may
help our current knowledge to establish a standard instruction for testing the H-reflex in
clinical purpose. However, it remains a question whether the results of this reliability
study on healthy populations would be applicable to neurological patients. The
availability of H-reflex measurement to populations with different pathologies needs to
be established in future studies.
The results from Studies II and III revealed the age-related differences in the SOL Hreflex modulation during different modes of isolated ankle movements. These results
indicated older adults adopted different neural strategy to control plantarflexors during
isomeric, shortening and lengthening muscle contractions, compared with young adults.
147
Similar downward regulation of the SOL H-reflex gain found between young and older
groups in Study II and a potentiation of the SOL Hmax/Mmax ratio during active
lengthening action in older group in Study III may implicate a preservation of cortical
motor function in older adults. However, the relevant supraspinal mechanisms
underlying these age-related changes in neuromuscular adaptation could not be fully
explained in Studies II and III. Future studies utilising TMS and EEG are required to
answer this question.
The second purpose of Study III was to investigate correlation of the maximal SOL Hreflex amplitude between dynamic ankle actions and static postural tasks. Tokuno et al.
(2008) reported greater H-reflex amplitudes of triceps surae during a further forward
body sway position than a slight forward body sway position. In this case, the triceps
surae assumes to perform in a manner of shortening contraction as a result of increasing
in the H-reflex amplitude (i.e. increase of motor output). If this assumption is true,
understanding relationship of the SOL H-reflex between dynamic muscle actions and
bipedal standing can offer meaningful information regarding ankle strategy in postural
control. This is particularly important to older adults because information about this
matter is not available in the literature. Indeed, the finding was limited by lack of
biomechanical measurements for muscle lengthening and kinematic movement in the
experiment. The functional implication of study III was restricted by many limitations
of research design. However, the significant correlation of H-reflex amplitude between
dynamic muscle actions and postural control found in the present study may open a
window to explore the scientific question in this stream. Future studies should
implement supplementary measurements (e.g. ultrasound scanner) for appreciating
better interpretation of neuromuscular function in this research area.
148
The results of Study IV provide novel evidence on the spinal plasticity to TC training
intervention. The underpinning of quantitative evidence in relation to the adaptation of
SOL H-reflex to 12-week TC training in older adults was elucidated. It has been
speculated that the potentiation of the SOL H-reflex after TC training is relevant to
cortical adaptation. However, the cortical contributions to TC training are still unclear.
Knowledge in this field is important because older adults tend to utilise the supraspinal
mechanisms to control movements (Chalmers & Knutzen, 2002; Earles, et al., 2001).
Future studies need to determine the extent of neural contribution from the supraspinal
mechanisms after TC training. Relevant information in this aspect can help us to
develop proper exercise interventions for balance improvement in older adults.
149
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164
Appendix A
Information sheet
165
INFORMATION SHEET
Name of Project: Effects of joint positions and force levels on muscle stretch reflex
in young and older adults
You are invited to participate in our research that examines the influence of
ageing on muscle stretch reflex under various conditions. The research requires two
groups of healthy volunteers: young adult group (20 to 40 years of age) and older adult
group (65 to 85 years of age). Both male and female, who are not currently undertaking
regular resistance training, may participate in the study. The research will be conducted
by researchers at Department of Exercise Science and Sport Management, Southern
Cross University, Lismore Campus. If you have met these criteria and are interested in
participating in this project, or want to know more about it before making the decision,
you are welcome to discuss with us.
What is the research about
The research aims to examine the effects of ageing on neuromuscular function
with a particular focus on the muscle stretch reflex. This reflex plays critical role in
maintenance posture. However, little information is available on the factors that may
affect this reflex in older adults. This study will investigate the effects of various muscle
contraction levels and ankle joint angles on the reflex in the calf muscles in a sitting
position. We hope to find some differences between young and older adults that allow a
better understanding of the effect of ageing on the reflex and other neuromuscular
function, and development of better exercise programs for prevention of falls for older
populations.
What will be involved
Participants are invited to visit the laboratory at least two times. The first visit
will be a familiarization trial and the second visit is for a formal experimental trial. Prior
to participation, we would like to know whether there is any contraindication to
participation in muscle strength and reflex testing. The researcher will interview the
potential participants for their health status. The participants who show no
contraindications will be given a familiarization trial during which further detail of the
experimental procedure will be explained.
During the formal experiment trial, participants muscle strength will be assessed and
muscle stretch reflex of the soleus (calf) muscle will be measured while the participant
sits on a testing chair with the right foot strapped to a dynamometer. A couple of
electrodes are placed on the calf muscle for recording muscle responses. The soleus
stretch reflex is evoked by giving an electrical pulse to the calf during a muscle
contraction. Each muscle contraction will be held for approximately 3 seconds. Several
trials are performed under each of the above-mentioned conditions.
166
Each testing session would require approximately two hours of time to complete.
Prior to each testing session, it would be appreciated if the participants do not smoke, or
take caffeine-containing drinks or food within 2 hours, and do not have strenuous
exercise within 24 hours.
Possible Discomforts and Risks
Performing muscle contractions with maximal effort may have some potential
risks, including injuries to muscle/tendon, and increased blood pressure. Electrical
stimulation on muscles may cause an unfamiliar sensation. The stimulation only lasts
for a fraction of a second, and will cause no harm to the body systems.
Responsibilities of the Researcher
The researchers will take all necessary measures in an attempt to minimise the
above-mentioned potential risks. In respect of privacy, any information that is obtained
in connection with this study and that can be identified with the participants will remain
confidential and will be disclosed only with their permission. The original data
collected will have to be retained for at least seven years by the researcher as required
by the University. The data with personal identity will then be destroyed in a way that
the privacy of the participants is protected.
Responsibilities of the Participants
It is important that the participants disclose all current health conditions that may
be considered as contraindications to perform such maximal and submaximal muscle
contractions, and discuss with the researcher if there is any concern.
Freedom of Consent
Your participation in this research is totally voluntary. It is your decision on
participation. You are free to withdraw your consent and to discontinue participation at
any time. However, we would appreciate you letting us know your decision.
Inquiries
If you have any questions, we expect you to ask us. If you have any additional
questions at any time please talk to the researchers, Mr. Yung Sheng Chen or Mr. Zac
Crowley, Dr Shi Zhou, who will be happy to answer any queries you may have.
Mr. Yung Sheng Chen, Mr. Zac Crowley, Dr Shi Zhou
School of Health and Human Sciences
P.O. Box 157, Lismore, NSW 2480
Work phone number: (02) 66203759 (Mr. Chen and Mr. Crowley), (02) 66203991 (Dr
Zhou)
Email: y.chen.33@scu.edu.au, z.crowl.10@scu.edu.au, shi.zhou@scu.edu.au
The ethical aspects of this study have been approved by the Southern Cross University
Human Research Ethics Committee (HREC). The Approval Number is ECN-08-092. If
you have any complaints or reservations about any ethical aspect of your participation
in this research, you may contact the Committee through the Ethics Complaints Officer:
Ms Sue Kelly
Ethics Complaints Officer and Secretary
HREC
167
PO Box 157
Lismore, NSW, 2480
Telephone (02) 6626-9139 or fax (02) 6626-9145
Email: sue.kelly@scu.edu.au
All complaints, in the first instance, should be in writing to the above address. All
complaints are investigated fully and according to due process under National
Statement on Ethical Conduct in Research Involving Humans and this University. Any
complaint you make will be treated in confidence and investigated, and you will be
informed of the outcome.
168
INFORMATION SHEET
Name of Project: Effect of ageing on the soleus H-reflex modulation during
shortening and lengthening muscle actions
The research aims to examine the influence of ageing and long-term physical
activity on neuromuscular function with a focus on muscle stretch reflex during static
and dynamic exercises. The research requires participation of healthy individuals, both
male and female, who are either in the age range of 20 to 40 years who are not
participating in regular physical activities, or 65 to 85 years who are currently either
undertaking or not undertaking regular physical activity. If you have met these criteria
and are interested in participating in this project, or want to know more about it before
making the decision, you are welcome to discuss with us.
It has been demonstrated that decline in neuromuscular function is a common
phenomenon when the human age increases. Maintaining a moderate level of physical
activity has shown to be beneficial to older adults. The aim of this study is to investigate
whether there is a difference between young and older adults in the muscle reflex, and
between older adults who participate in regular exercise and who do not. The outcomes
of the study will allow us to have a better understanding of the effects of aging on
neuromuscular function and to find better ways in improving balance and fall
prevention, particularly for older populations.
Participants are required to visit the laboratory at least two times. The first visit
will be a familiarization trial and the second visit will be the experimental trial. Prior to
participation, we would like to know whether there is any contraindication to
participation. The researcher will interview the potential participants and use a
questionnaire for checking the medical history and current health status. The
participants who show no contraindications will be given a familiarization trial during
which further detail of the experimental procedure will be explained. All of the testing
sessions will be conducted in the Exercise Science Laboratories in P Block, Southern
Cross University, Lismore Campus.
The muscle stretch reflex is examined while the participant sits on a testing chair
with the right foot strapped to a dynamometer. A couple of electrodes are placed on the
calf muscle for recording muscle responses. The muscle stretch reflex is evoked by
giving an electrical pulse to the calf during a muscle contraction. Each muscle
contraction will be held for approximately 3 seconds. The reflex test will also be
performed while in a standing position. Several trials are performed under the abovementioned conditions.
169
Ability in postural control will be tested while standing on a force plate under
four conditions: on firm or soft surface and with eyes open or closed.
take caffeine containing drinks or food within 2 hours, and do not have strenuous
for a fraction of a second and will cause no harm to the body systems.
Responsibilities of the Participant
It is important that the participants disclose all current conditions that may be
considered as contraindications to perform such maximal and submaximal muscle
Freedom of Consent
Inquiries
Zhou)
170
Ms Sue Kelly
HREC
PO Box 157
Lismore, NSW, 2480
Telephone (02) 6626-9139 or fax (02) 6626-9145
171
INFORMATION SHEET
Name of Project: Effect of Tai Chi training on muscle stretch reflex in older adults
You are invited to participate in a research that examines the effects of 12 weeks
Tai Chi training on the muscle stretch reflex modulation in relation to postural stability
in older adults. The research requires participation of healthy individuals, both male and
female, who are in the age range of 65 to 85 years, and have no experience of
undertaking Tai Chi training. If you have met these criteria and are interested in
participating in this project, or want to know more about it before making the decision,
you are welcome to discuss with us.
Ageing may affect individuals ability in control of balance and increase in the
chance of fall injuries. Muscle stretch reflex plays an important role in control of
balance. It has been shown that Tai Chi exercise has beneficial effects in improving
balance control in older adults. However, whether the improvement is related to changes
in muscle stretch reflex is unknown. The aim of this study is to evaluate the effect of Tai
Chi training on reflex modulation in older adults. The outcomes of the study will allow
us to have a better understanding of the effect of Tai Chi training on postural control
that may help us to find better ways in improving balance and preventing falls in older
populations.
40 volunteers, in the age range of 65-85 years, both males and females, will be
randomly allocated into a control and a Tai Chi training group. The control group will
maintain their normal daily activities for 12 weeks, while the Tai Chi group will
participate in training, 3 sessions per week for 12 weeks.
Prior to the training, participants will attend a familiarization trial followed by a
pre training testing trial to establish the baseline data. The same tests will be repeated
after 12 weeks of training or control period. During the first visit, the researcher will
interview the potential participants to find out if there is any contraindication to
participation. The participants who show no contraindications will be given a
familiarization trial during which further detail of the experimental procedure will be
explained.
The muscle stretch reflex is examined while the participant sits on a testing chair
with the right foot strapped to a dynamometer. A couple of electrodes are placed on the
calf muscle for recording muscle responses. The muscle stretch reflex is evoked by
giving an electrical pulse to the calf during a muscle contraction. Each muscle
contraction will be held for approximately 3 seconds. The reflex test will also be
performed while in a standing position. Several trials are performed under the abovementioned conditions.
172
Ability in postural control will be tested while standing on a force plate under
four conditions: on firm or soft surface and with eyes open or closed.
for a fraction of a second, and will cause no harm to the body systems.
Responsibilities of the Participant
It is important that the participants disclose all current conditions that may be
considered as contraindications to perform such maximal and submaximal muscle
Freedom of Consent
Inquiries
Zhou)
173
Ms Sue Kelly
HREC
PO Box 157
Lismore, NSW, 2480
Telephone (02) 6626-9139 or fax (02) 6626-9145
174
Appendix B
Consent form
175
CONSENT FORM
(This consent form is based on the National Statement on Ethical Conduct in Human Research (National Statement/NS)
Title of research project:

Effects of body position and vision on muscle stretch reflex in young and older
adults
NOTE: This consent form will remain with the Southern Cross University researchers
for their records.
Tick the box that applies, sign and date and give to the researcher
I agree to take part in this research project at Southern Cross University
as specified above.
Yes
No
I have been provided with information at my level of comprehension

about the purpose, methods, demands, risks, inconveniences and
possible outcomes of this research. I understand this information.
Yes
No
I understand that my participation is voluntary.
Yes
No
I can choose not to participate in part or all of this research at any time,
without consequence.
Yes
No
I understand that any information that may identify me, will be

de-identified at the time of analysis of any data. Therefore, I, or
any information I have provided cannot be linked to my person/
company. (Privacy Act 1988 Cth)
Yes
No
I understand that neither my name nor any identifying information will

be disclosed or published, except with my permission.
Yes
No
I give permission for identifying information to be published

or disclosed.
Yes
No
I understand that all information gathered in this research is confidential.

It is kept securely and confidentially for 7 years at the University (unless
there are special circumstances, that have been explained to me.
Yes
No
I am aware that I can contact the Supervisor or other researchers at any

time with any queries.
Yes
No
176
I understand that the ethical aspects of this research have been

approved by the SCU Human Research Ethics Committee.
Yes
If I have concerns about the ethical conduct of this research, I understand that I can
contact the SCU Ethics Complaints Officer. All inquiries are confidential and
should be in writing, in the first instance, to the following:
Ethics Complaints Officer
PO Box 157
Lismore NSW 2480
Participants name:
Participants signature:
Date:
Contact: Tel:
Email:
177
Yes
No
No
CONSENT FORM

Effects of ageing on the soleus H-reflex modulation during shortening and
lengthening muscle actions
for their records.
as specified above.
Yes
No

Yes
No
Yes
No
Yes
No

Yes
No

Yes
No

or disclosed.
Yes
No

Yes
No

Yes
No

178
Yes
PO Box 157
Lismore NSW 2480
Participants name:
Date:
Contact: Tel:
Email:
179
Yes
No
No
CONSENT FORM

Effect of Tai Chi training on muscle stretch reflex in older adults
for their records.
as specified above.
Yes
No

Yes
No
Yes
No
Yes
No

Yes
No

Yes
No

or disclosed.
Yes
No

Yes
No

Yes
No

Yes
No
180
PO Box 157
Lismore NSW 2480
Participants name:
Date:
Contact: Tel:
Email:
181
Yes
No
Appendix C
Call for volunteers
182
CALL FOR VOLUNTEERS

FOR A RESEARCH ON NEUROMUSCULAR FUNCTION
Name of Project:
Effects of joint positions and force levels on muscle stretch reflex in young and
older adults
Purpose:
The research aims to examine the effects of ageing on neuromuscular function with a
particular focus on the muscle stretch reflex. We hope to find some differences between
young and older adults that allow a better understanding of the effect of ageing on the
reflex and other neuromuscular function, and development of better exercise programs
for prevention of falls for older populations.
Volunteers sought:
The research requires two groups of healthy volunteers: 20 healthy young adults (20 to
40 years of age) and 20 healthy older adults (65 to 85 years of age). Both male and
female, who are not currently undertaking regular resistance training, may participate in
the study.
What will be involved:
Participants are invited to visit the laboratory at least two times. The first visit will be a
familiarization trial and the second visit is for a formal experimental trial.
During the formal experiment trial, participant's muscle strength will be assessed and
stretch reflex is evoked by giving a percutaneous electrical pulse to the calf during a
muscle contraction. Each muscle contraction will be held for approximately 3 seconds.
Several trials are performed under each of the above-mentioned conditions.
Each testing session would require approximately 1.5 to 2 hours of time to complete.
Further information:
If you have any questions, we expect you to ask us. If you have any additional questions
at any time please talk to the researchers, Mr. Yung Sheng Chen, Mr. Zac Crowley, or
Dr Shi Zhou, who will be happy to answer any queries you may have.
183
Work phone number: (02) 66203759 (Mr. Chen and Mr. Crowley); (02) 66203991 (Dr
Zhou) Email: y.chen.33@scu.edu.au, z.crowl.10@scu.edu.au, shi.zhou@scu.edu.au
184
CALL FOR VOLUNTEERS

Name of Project:
Effect of ageing on the soleus H-reflex modulation during shortening and
lengthening muscle actions
Purpose:
The research aims to examine the effects of ageing on neuromuscular function with a
particular focus on the muscle stretch reflex. We hope to find some differences between
young and older adults that allow a better understanding of the effect of ageing on the
reflex and other neuromuscular function, and development of better exercise programs
for prevention of falls for older populations.
Volunteers sought:
The research requires two groups of healthy volunteers: 20 healthy young adults (20 to
40 years of age), 20 sedentary older adults (65 to 85 years of age), and 20 active older
adults (65 to 85 years of age). Both male and female, who are not currently undertaking
regular resistance training, may participate in the study.
Participants are invited to visit the laboratory at least two times. The first visit will be a
familiarization trial and the second visit is for a formal experimental trial.
During the formal experiment trial, participants muscle strength will be assessed and
stretch reflex is evoked by giving a percutaneous electrical pulse to the calf during a
muscle contraction. Each muscle contraction will be held for approximately 3 seconds.
Several trials are performed under each of the above-mentioned conditions.
Ability in postural control will be tested while standing on a force plate under four
conditions: on firm or soft surface and with eyes open or closed.
Each testing session would require approximately 1.5 to 2 hours of time to complete.
Crowley, or Dr Shi Zhou, who will be happy to answer any queries you may have.
School and Health and Human Sciences
Zhou),
Email: y.chen.33@scu.edu.au, z.crowl.10@scu.edu.au, shi.zhou@scu.edu.au or, come to
their offices in P Block (P 1.48 and P1.50), Southern Cross University, Lismore Campus.
185
CALL FOR VOLUNTEERS

Name of Project:
Effect of Tai Chi training on muscle stretch reflex in older adults
Purpose:
The aim of this study is to evaluate the effect of Tai Chi training on reflex modulation in
older adults. The outcomes of the study will allow us to have a better understanding of
the effect of Tai Chi training on postural control that may help us to find better ways in
improving balance and preventing falls in older populations.
Volunteers sought:
40 volunteers, in the age range of 65-85 years, both males and females, will be
randomly allocated into a control and a Tai Chi training group. The control group will
maintain their normal daily activities for 12 weeks, while the Tai Chi group will
participate in training, 3 sessions per week for 12 weeks.
Prior to the training, participants will attend a familiarization trial followed by a pre
training testing trial to establish the baseline data. The same tests will be repeated after
12 weeks of training or control period. During the first visit, the researcher will
interview the potential participants to find out if there is any contraindication to
participation. The participants who show no contraindications will be given a
familiarization trial during which further detail of the experimental procedure will be
explained.
During the formal experiment trial, participants ability in postural control will be tested
while standing on a force plate under four conditions: on firm or soft surface and with
eyes open or closed. Muscle stretch reflex of the soleus (calf) muscle will be measured
while the participant sits on a testing chair with the right foot strapped to a
dynamometer. A couple of electrodes are placed on the calf muscle for recording
muscle responses. The soleus stretch reflex is evoked by giving a percutaneous
electrical pulse to the calf. Several trials are performed under each of the abovementioned conditions.
Each testing session would require approximately 1.5 hours of time to complete. Prior
to each testing session, it would be appreciated if the participants do not smoke, or take
caffeine-containing drinks or food within 2 hours, and do not have strenuous exercise
within 24 hours.
Crowley, or Dr Shi Zhou, who will be happy to answer any queries you may have.
School and Health and Human Sciences
Zhou),
186
Email: y.chen.33@scu.edu.au, z.crowl.10@scu.edu.au, shi.zhou@scu.edu.au or, come to

their offices in P Block (P 1.48 and P1.50), Southern Cross University, Lismore Campus.
187
Appendix D
Health status assessment prior to
exercise testing
188
HEALTH STATUS ASSESSMENT PRIOR TO EXERCISE TESTING

(Adapted from ASMF)
_____________________________________________________________________________________
__
This form is used as a pre-participation health and risk factor screening device and should be completed
prior to the commencement of an exercise test or exercise program.
The information obtained in this screening will be kept as CONFIDENTIAL. Only the responsible staff
member and the medical practitioner may access to the information.
Sections (1) to (9) should be completed by the client before seeing a medical practitioner.
Section (10) should be completed by the staff.
Sections (11) and (12) should be completed by a medical practitioner.
_____________________________________________________________________________________
___
Clients Surname (Mr., Mrs., Ms.) : ______________________________________
Given Names: _______________________________________________________
Date of Birth: ________________________________________________________
Address: ____________________________________________________________
_________________________________________________ Postcode: __________
Contact Telephone: (Home) ____________________ (Work/Mobile) ____________________
Reasons for participating in exercise testing or training:
______________________________________________________________________
______________________________________________________________________
189
(1) FAMILY MEDICAL HISTORY.

Has any near relative brother (B), sister (S), father (F), mother (M), grandparents (GP) suffered:
Please tick the appropriate column
No
Yes
Relation
Age
Remarks/Details
Apoplexy (stroke)
Congenital heart trouble
Rheumatic heart disease
Heart operation
Angina
Heart attack
Sudden death
High blood pressure
High cholesterol
'Hardening of arteries'
Asthma
Lung disorder
Bronchitis, emphysema
Hay fever
Diabetes
Gout
Arthritis
Epilepsy
190
(2) PAST MEDICAL HISTORY

Have you suffered any of the following conditions at any time:
(Please tick the appropriate column)
No
Yes
Details
Rheumatic or scarlet fever

Heart trouble or murmur
Heart palpitation
High blood pressure
Heart attack
Chest pain/Angina
Stroke
Disease of arteries or veins
Undue limiting shortness of breath with
exercise
Fainting or blackout
Loss of consciousness or fainting with exercise
Epilepsy
Lung or bronchial disease
Asthma
Hay fever
Anaemia
Diabetes
Thyroid disease
Arthritis, rheumatism or gout spondylitis,
disc trouble or back injury
Serious accident or injury
Surgical operation
Congenital abnormality
Other serious illness (or conditions that may
affect exercise)
For female only:
Having normal/regular periods
191
(3) SOCIAL HISTORY

Marital Status: _________________________ Number of Children: _________________
Occupation: ______________________________________________________________
(4) HABITS
Please tick and fill in your usual habit of consumption of the following substances:
Alcoholic beverage:
( ) Nil
( ) Every day: number of drinks per day: _________________
( ) Irregular: number of drinks per week: _________________
Smoking:
( ) Never smoked
( ) Gave up ______ years ago
( ) Now smoke
Cigarettes:
______________ per day
Pipe:
______________ per day
Cigars:
______________ per day
(5) The poliovirus eradication campaign is almost complete but risks exist for people who have been
to West Africa, Central Africa, Ethiopia, India, Pakistan, Afghanistan and Eastern Turkey.
Have you been to any of the listed areas in the previous 6 months?
_____________________________________________________________________________
(6) DIETARY SUPPLEMENTS

Please tick if you regularly take any of the following:
( ) Vitamins:
Type: ____________________________________________
( ) Minerals:
Type: ____________________________________________
( ) Iron:
Type: ____________________________________________
( ) Other compounds: _____________________________________________________
192
(7) PRESENT MEDICAL CONDITIONS

This section should be completed at the INITIAL EXAMINATION and also at examination
IMMEDIATELY PRIOR TO THE EXERCISE TEST.
Are you currently suffering or have you in the recent past suffered any of the following conditions (Please
tick the appropriate column):
Initial Exam
Yes
No
Second Exam
Yes
No
Cough
Stuffy nose or sore throat
Tonsillitis, glandular fever
Hepatitis
Diarrhoea/vomiting
Headaches
Shortness of breath
Pain in chest, left arm or neck at rest, or during
physical activities
Heart palpitations
Cramp in legs
Abnormal loss of blood
Insomnia
Indigestion or constipation
Swollen, stiff or painful joints
Backache
Sports injury or other injury
Other symptom or illness, or surgery
Any deterioration in training or competitive
performance
Any other conditions that may contraindicate to exercise
or affect exercise capacity
For female only:
Currently in pregnancy
If yes, provide details
193
(8) CURRENT MEDICATION

This section should also be checked IMMEDIATELY PRIOR TO THE EXERCISE TEST.
State the name and dosage of any drugs or medicines that you are taking regularly:
Drug
Dose
Time of last dose
(9) EXERCISE OR PHYSICAL ACTIVITY HISTORY

Present exercise and/or physical activities (activities may include walking to work, gardening, bike
riding, swimming, weights training, and sports):
Type of activity
Intensity (rate 1-5

gentle to intense)
Hours per day
_________________
______________
______________
______________
_________________
______________
______________
______________
_________________
______________
______________
______________
_________________
______________
______________
______________
_________________
______________
______________
______________
Days per week
I attest that the information provided by me in completing this form is to the best of my knowledge a true
and accurate reflection of my current health status. In the event that I display symptoms of illness at any
point during my participation in this exercise test I will advise the testing staff immediately.
Signed: ________________________________
Name: _________________________
Date: __________________________
(10) BODY MASS, HEIGHT AND BODY MASS INDEX (to be completed by the staff)
Body Mass (kg) : _______________________________ Height (m) : ___________________________

Body Mass Index (kg/m2) : _______________________
194
(11) PHYSICAL EXAMINATION

This section should be completed by a medical practitioner.
_____________________________________________________________________________________
(a) General appearance including glands, and lymph nodes
_____________________________________________________________________________________
(b) Cardiovascular system
(i)
Peripheral vessel and pulses
(ii)
Neck veins
(iii)
Apex beat position
(iv)
Heart sounds
(v)
Resting heart rate
(vi)
Blood Pressure: Lying: ___________ mmHg, Standing: ___________ mmHg
(vii)
12 leads ECG (a copy of resting ECG should be on file).
_____________________________________________________________________________________
(c) Respiratory system
_____________________________________________________________________________________
(d) Abdomen
_____________________________________________________________________________________
(e) Nervous system
_____________________________________________________________________________________
(f) Fundi
_____________________________________________________________________________________
(g) Locomotor system
_____________________________________________________________________________________
(h) Varicose veins
_____________________________________________________________________________________
(i) Urinalysis:
Protein: ______________________
Sugar: _______________________
Specific Gravity: _______________
Other results: _________________
_____________________________________________________________________________________
(j) Blood lipid profile:
TC: _________________________
LDL: ________________________
Triglyceride: __________________
Other results: __________________
_____________________________________________________________________________________
(k) Other examinations:
_____________________________________________________________________________________
195
(12) MEDICAL PRACTITIONER'S SUMMARY:

(a) Comments (detail any significant abnormalities or reservations):
___________________________________________________________________________
___________________________________________________________________________
___________________________________________________________________________
(b) Recommendations:
The medical practitioner should underline and initial the appropriate clause:
(i)
Fit to undergo maximal exercise tests.
(ii)
Not fit to undergo maximal exercise test, but may undergo

submaximal test without special precautions.
(iii)
Not fit to undergo maximal exercise test but may undergo submaximal test with
following precaution:
Precaution: _________________________________________________
___________________________________________________________
(iv)
Not fit to undergo any exercise test.
Signed: ________________________________
Name: _________________________
Date: __________________________
Contact Telephone Number: (Work) ____________________ (A/H) ______________________
196
Appendix E
Tai Chi training program
197
Warm-up exerciseises
1. Supporting the sky with both hands
2. Drawing a bow to each side
3. Holding up a single hand
4. Looking back like a cow
5. Lowering the head and hips removes
6. Touching the feet with both hands
7. Clentching the fists
8. Shaking the body wards
9. Open your hands and turn your head
10. Backstroke
11. Turn the wheel
12. Clean the mirror
13. Picking fruit
14. Swing your arms and turn your head
15. Twist your body and let your arms swing
16. Reaching down and leaning back
17. Turn to look at the moon
18. Interlace fingers behind your neck and squeeze your
elbows together
19. Fingers exercises
20. Eyes exercises
21. Kicks
22. Massage
198
12 weeks Tai Chi program

Period
Week 1
Learning Period
Week 2
Week 3
Week 4
Week 5
Yang style Tai Chi

1. Beginning of Tai Chi
2. Grasp birds tail
3. Single whip
4. Raise hands and step up
5. White stork displays its wings
6. Brush knee and twist left
7. Play guitar
8. Brush knee and twist left, right, and left
9. Play guitar
10. Brush knee and twist left
11. Deflect sideways, parry, step forward and punch
Week 6
12. Apparent counter and closure
Practice Period
13. Cross hands
Week 7 ~ Week 12
13 slow forms of Yang style Tai Chi
199
Instructions of 13 slow forms

1) Beginning of Tai Chi
Check your position.
Raise your hands to shoulder level.
Turn you palms and pull them down.
2) Grasp Birds Tail

Bend your knees.
Raise your right arm and turn your right foot.
Raise your left arm and step your left foot.
Right arm forms a circle and step your right foot.
Left arm follows right arm (like holding a football).
Turn your palms and pull them to the left.
Left hand forms a circle and touch your right wrist.
Press your hands forward, separate, pull back and push.
3) Single Whip
Turn your body and turn your right foot.
Keep your hands at eye level and pull them to the left.
Put your hands down and pull them to the right.
Right hand forms a bird head and look to the right.
Step your left foot and turn your left palm out.
4) Raise Hands and Step Up
Open your right palm and raise your hands to the centre.
Raise your right foot to the centre (right heel touches the
floor)
200
5) White Stork Displays Its Wings

Turn your body to the left (left hand up, right hand down).
Step your right foot (right hand up, left hand down).
Point your left foot and turn your right palm up.
6) Brush Knee and Twist Left

Right hand forms a circle to the back of your right ear.
Left hand follows right hand and step your left foot.
Left hand brushes left knee and right hand pushes to the
front.
7) Play Guitar
Step your back foot in and raise your hands to the centre (left
heel touches the floor).
8) Brush Knee and Twist: Left, Right, and Left

front.
Left hand forms a circle to the back of your left ear.
Right hand follows left hand and step your right foot.
Right hand brushes right knee and left hand pushes to the
front.
front.
201
9) Play Guitar
Step your back foot in and raise your hands to the centre (left
heel touches the floor).
10) Brush Knee and Twist Left

Left hand brushes left knee and right hand pushes to the front.
11) Deflect Sideways, Parry, Step Forward and Punch

Right hand makes a fist and turn your left foot.
Both hands form a circle and twist your right foot to the front.
Step your left foot and withdraw your right fist to the right.
Your right fist strikes out and your left palm goes under your
right upper arm.
12) Apparent Counter and Closure
Open your right palm and withdraw to the right.
Turn your palms and push.
13) Cross Hands

Turn your body and turn your left foot.
Both arms form a circle and cross hands (your right foot steps
in)
202
Appendix F
Original data
203
Study I
MVC (N.m)
Test 1
Test 2
Participants
20
-20
20
-20
90.0
170.0
240.0
89.0
166.0
244.0
67.0
130.0
214.0
69.0
126.0
202.0
82.0
127.0
163.0
74.0
117.0
171.0
54.0
115.0
143.0
52.0
92.0
193.0
38.1
54.9
61.6
51.3
88.3
113.8
43.3
93.5
173.2
47.0
103.6
170.8
22.9
50.7
74.6
29.0
64.0
95.7
42.4
86.0
105.8
54.4
88.5
114.7
98.0
163.5
242.6
91.3
153.4
237.4
10
30.6
66.9
101.2
36.9
70.8
111.0
Mean
56.8
105.8
151.9
59.4
107.0
165.3
SD
26.1
42.5
66.1
20.9
33.6
54.3
204
Plantarflexion
Hmax (mV)
Test 1
Test 2
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
2.10
3.33
3.63
4.75
3.13
4.07
4.23
4.46
3.10
3.32
2.48
0.51
2.93
2.69
0.82
0.41
4.61
2.97
2.12
4.17
4.68
3.00
3.58
4.46
1.16
1.15
1.31
1.33
1.26
1.79
1.59
1.81
2.39
1.66
1.36
1.35
2.71
2.08
0.87
1.14
1.36
1.75
1.08
0.56
1.55
2.70
3.39
3.39
2.22
2.02
1.41
0.69
1.90
1.71
1.92
1.84
1.97
2.38
2.50
2.40
2.22
2.59
1.94
0.51
2.25
1.46
1.84
2.06
2.73
1.95
1.74
2.34
10
2.88
0.51
0.51
0.49
3.27
0.61
0.69
0.59
Mean
2.40
2.06
1.82
1.83
2.64
2.32
2.08
2.09
SD
0.98
0.94
0.89
1.54
0.98
0.92
1.25
1.55
205
Neutral
Hmax (mV)
Test 1
Test 2
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
1.08
1.11
1.39
4.28
2.39
2.36
3.47
4.77
2.26
2.29
2.43
2.26
2.57
2.54
3.30
1.62
3.59
3.12
2.61
4.01
3.40
4.14
3.56
4.32
1.01
0.57
1.23
1.66
0.75
0.61
0.56
1.05
2.28
2.59
1.58
1.71
2.37
2.61
1.59
1.13
0.59
1.32
3.09
4.57
0.48
1.41
3.29
4.07
2.08
1.98
2.15
1.69
1.54
1.42
1.57
1.54
3.01
2.86
2.68
2.64
3.01
3.06
2.93
3.22
1.64
1.58
0.44
1.37
1.88
1.55
2.04
1.27
10
2.17
2.71
1.88
0.85
2.17
3.30
3.84
1.04
Mean
1.97
2.01
1.95
2.50
2.06
2.30
2.62
2.40
SD
0.92
0.84
0.80
1.33
0.92
1.06
1.10
1.52
206
Dorsiflexion
Hmax (mV)
Test 1
Test 2
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
1.78
1.19
1.99
2.59
1.10
1.20
0.84
1.45
1.46
1.71
2.36
3.05
1.33
0.79
1.24
3.23
0.83
2.43
2.25
2.62
2.81
2.43
2.16
2.23
0.83
0.31
0.49
0.54
0.38
0.79
1.57
1.50
1.89
1.80
1.36
1.14
1.60
1.73
2.34
1.64
0.28
0.46
1.36
0.96
0.34
0.23
1.18
2.37
1.48
1.17
1.68
1.83
1.19
1.23
1.41
1.73
2.75
3.17
2.83
2.36
3.14
3.43
3.21
2.58
1.07
1.28
0.76
0.35
1.08
0.92
1.35
1.15
10
1.83
1.89
2.14
2.66
1.96
1.96
2.48
3.31
Mean
1.42
1.54
1.72
1.81
1.49
1.47
1.78
2.12
SD
0.70
0.86
0.73
0.99
0.92
0.94
0.74
0.75
207
Mmax (mV)
Test 1
Test 2
Participants
20
-20
20
-20
3.92
2.56
1.71
4.26
2.46
2.72
4.41
4.01
3.80
4.49
4.09
3.89
6.77
5.23
6.73
4.46
5.18
4.63
5.02
4.79
4.34
5.19
5.48
5.44
4.93
3.95
2.90
5.49
3.36
2.55
4.24
2.43
1.87
3.76
2.11
1.77
3.61
3.02
2.20
2.87
2.18
1.72
3.35
3.94
3.39
3.05
3.23
3.92
3.19
2.39
1.72
3.94
2.48
1.98
10
3.41
2.65
2.18
3.35
2.35
2.26
Mean
4.28
3.50
3.09
4.09
3.29
3.09
SD
1.08
1.03
1.58
0.87
1.24
1.30
208
Plantarflexion
SOL bEMG (mV)

Test 1
Participants
Rest
10%
30%
Test 2
50%
Rest
10%
30%
50%
0.0047 0.0271 0.0234 0.0326 0.0026 0.0161 0.0259 0.0377
0.0019 0.0143 0.0453 0.0488 0.0024 0.0172 0.0189 0.0231
0.0024 0.0215 0.0384 0.0381 0.0104 0.0216 0.0286 0.0383
0.0033 0.0141 0.0148 0.0155 0.0022 0.0331 0.0308 0.0352
0.0022 0.0138 0.0217 0.0217 0.0129 0.0141 0.0268 0.0302
0.0047 0.0098 0.0184 0.0284 0.0024 0.0141 0.0213 0.0335
0.0016 0.0109 0.0174 0.0176 0.0044 0.0141 0.0168 0.0251
0.0027 0.0190 0.0316 0.0361 0.0025 0.0157 0.0187 0.0242
0.0023 0.0051 0.0063 0.0064 0.0056 0.0080 0.0154 0.0124
10
0.0024 0.0146 0.0184 0.0186 0.0023 0.0225 0.0267 0.0261
Mean
0.0028 0.0150 0.0236 0.0264 0.0048 0.0177 0.0230 0.0286
SD
0.0011 0.0062 0.0117 0.0127 0.0039 0.0068 0.0054 0.0080
209
Neutral
SOL bEMG (mV)

Test 1
Test 2
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
0.0048
0.0177
0.0337
0.0433
0.0025
0.0194
0.0368
0.0615
0.0019
0.0124
0.0326
0.0661
0.0023
0.0101
0.0348
0.0408
0.0025
0.0128
0.0273
0.0402
0.0101
0.0234
0.0322
0.0352
0.0031
0.0173
0.0263
0.0300
0.0022
0.0336
0.0347
0.0391
0.0022
0.0193
0.0369
0.0345
0.0136
0.0221
0.0465
0.0371
0.0042
0.0265
0.0386
0.0492
0.0025
0.0239
0.0387
0.0492
0.0017
0.0254
0.0162
0.0185
0.0043
0.0186
0.0613
0.0487
0.0027
0.0155
0.0359
0.0354
0.0024
0.0159
0.0187
0.0232
0.0023
0.0074
0.0139
0.0143
0.0060
0.0104
0.0158
0.0216
10
0.0025
0.0304
0.0302
0.0258
0.0024
0.0279
0.0560
0.0473
Mean
0.0028
0.0185
0.0292
0.0357
0.0048
0.0205
0.0375
0.0404
SD
0.0010
0.0071
0.0085
0.0152
0.0040
0.0074
0.0144
0.0122
210
Dorsiflexion
SOL bEMG (mV)

Test 1
Test 2
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
0.0046
0.0161
0.0250
0.0450
0.0026
0.0265
0.0282
0.0498
0.0018
0.0187
0.0361
0.0568
0.0024
0.0125
0.0271
0.0418
0.0024
0.0145
0.0202
0.0252
0.0110
0.0177
0.0232
0.0335
0.0032
0.0138
0.0248
0.0289
0.0021
0.0314
0.0407
0.0560
0.0022
0.0302
0.0221
0.0219
0.0129
0.0256
0.0414
0.0528
0.0040
0.0149
0.0367
0.0416
0.0024
0.0151
0.0315
0.0443
0.0017
0.0204
0.0242
0.0389
0.0043
0.0130
0.0315
0.0501
0.0027
0.0192
0.0311
0.0463
0.0025
0.0141
0.0201
0.0265
0.0022
0.0074
0.0132
0.0150
0.0055
0.0110
0.0190
0.0273
10
0.0025
0.0192
0.0330
0.0502
0.0023
0.0288
0.0513
0.0733
Mean
0.0027
0.0174
0.0266
0.0370
0.0048
0.0196
0.0314
0.0455
SD
0.0009
0.0059
0.0075
0.0136
0.0039
0.0076
0.0103
0.0142
211
Plantarflexion
TA bEMG (mV)
Test 1
Participants
Rest
10%
30%
Test 2
50%
Rest
10%
30%
50%
0.0033 0.0048 0.0071 0.0083 0.0032 0.0062 0.0089 0.0102
0.0034 0.0049 0.0070 0.0096 0.0096 0.0073 0.0075 0.0096
0.0242 0.0326 0.0341 0.0338 0.0039 0.0062 0.0099 0.0139
0.0051 0.0327 0.0341 0.0330 0.0265 0.0064 0.0080 0.0095
0.0116 0.0140 0.0146 0.0151 0.0142 0.0157 0.0180 0.0200
0.0069 0.0073 0.0087 0.0104 0.0030 0.0038 0.0049 0.0081
0.0055 0.0065 0.0073 0.0086 0.0054 0.0064 0.0080 0.0098
0.0085 0.0091 0.0129 0.0174 0.0150 0.0177 0.0198 0.0210
0.0030 0.0034 0.0034 0.0040 0.0047 0.0063 0.0086 0.0095
10
0.0031 0.0034 0.0039 0.0042 0.0054 0.0098 0.0132 0.0142
Mean
0.0075 0.0119 0.0133 0.0144 0.0091 0.0086 0.0107 0.0126
SD
0.0065 0.0114 0.0115 0.0108 0.0075 0.0045 0.0048 0.0046
212
Neutral
TA bEMG (mV)
Test 1
Test 2
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
0.0033
0.0116
0.0135
0.0274
0.0032
0.0115
0.0139
0.0402
0.0040
0.0089
0.0134
0.0358
0.0102
0.0091
0.0137
0.0266
0.0244
0.0223
0.0332
0.0399
0.0038
0.0128
0.0162
0.0221
0.0057
0.0247
0.0360
0.0395
0.0299
0.0126
0.0133
0.0202
0.0116
0.0140
0.0214
0.0252
0.0146
0.0181
0.0225
0.0286
0.0070
0.0162
0.0169
0.0296
0.0032
0.0148
0.0141
0.0256
0.0057
0.0185
0.0103
0.0135
0.0054
0.0122
0.0219
0.0319
0.0079
0.0111
0.0175
0.0239
0.0140
0.0146
0.0187
0.0214
0.0031
0.0056
0.0062
0.0102
0.0047
0.0082
0.0088
0.0145
10
0.0031
0.0172
0.0103
0.0159
0.0060
0.0187
0.0225
0.0310
Mean
0.0076
0.0150
0.0179
0.0261
0.0095
0.0133
0.0165
0.0262
SD
0.0065
0.0060
0.0098
0.0105
0.0083
0.0034
0.0047
0.0072
213
Dorsiflexion
TA bEMG (mV)
Test 1
Test 2
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
0.0074
0.0037
0.0049
0.0074
0.0034
0.0046
0.0061
0.0076
0.0028
0.0043
0.0055
0.0077
0.0103
0.0070
0.0072
0.0101
0.0257
0.0298
0.0320
0.0320
0.0040
0.0053
0.0064
0.0088
0.0042
0.0302
0.0418
0.0414
0.0267
0.0043
0.0055
0.0073
0.0114
0.0127
0.0115
0.0121
0.0145
0.0161
0.0168
0.0171
0.0125
0.0073
0.0083
0.0094
0.0053
0.0030
0.0034
0.0044
0.0060
0.0070
0.0066
0.0071
0.0055
0.0061
0.0064
0.0080
0.0376
0.0079
0.0093
0.0135
0.0265
0.0177
0.0185
0.0188
0.0029
0.0031
0.0033
0.0036
0.0047
0.0056
0.0066
0.0090
10
0.0031
0.0033
0.0034
0.0036
0.0071
0.0082
0.0098
0.0122
Mean
0.0114
0.0109
0.0127
0.0138
0.0108
0.0078
0.0087
0.0103
SD
0.0116
0.0105
0.0132
0.0127
0.0090
0.0050
0.0050
0.0045
214
Study II
Plantarflexion
H-reflex gain (%)

Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
667.01
150.23
180.90
136.82
241.53
146.96
57.71
15.07
337.68
231.64
54.70
10.51
61.71
22.18
10.26
7.35
449.19
138.49
124.99
116.57
67.83
29.58
29.87
17.81
51.03
54.10
51.78
51.25
114.50
38.61
34.04
7.04
480.06
425.19
309.97
193.85
92.69
36.16
18.27
8.39
209.57
147.38
32.31
37.66
25.91
22.90
18.89
22.73
420.66
241.98
135.50
84.50
167.41
48.53
25.96
13.68
355.51
191.05
202.00
135.28
140.28
26.91
20.32
20.56
242.25
106.49
44.69
19.14
158.34
103.62
43.52
32.28
10
488.38
245.28
113.10
188.80
265.92
58.71
42.08
36.11
11
285.79
27.25
26.03
22.44
29.59
15.84
15.59
9.37
12
1236.15
40.06
110.33
107.89
151.18
19.89
20.65
18.45
13
367.96
331.66
160.37
119.81
39.54
30.83
21.33
20.96
14
325.87
178.98
110.38
73.94
256.52
31.34
23.11
27.36
15
149.61
44.00
63.35
53.36
254.70
93.42
195.91
145.31
16
69.71
105.27
97.82
78.50
73.72
26.19
46.27
31.81
17
681.33
255.46
141.63
79.16
70.22
32.52
37.16
29.70
18
177.51
70.80
69.82
19.08
147.71
91.59
50.33
47.93
19
260.48
89.89
59.38
24.60
17.81
18.99
12.90
10.04
20
1219.22
194.82
133.95
108.95
284.68
76.57
36.77
12.51
Mean
423.75
163.50
111.15
83.10
133.09
48.57
38.05
26.72
SD
322.46
104.24
68.34
54.98
88.22
35.57
39.46
29.99
215
Neutral
H-reflex gain (%)

Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
499.11
133.22
103.17
110.25
140.45
145.53
56.13
14.39
245.92
183.63
74.31
34.23
43.10
29.26
15.04
5.32
336.38
177.03
110.58
122.61
68.00
83.04
45.66
21.70
29.97
18.16
16.20
26.79
73.93
64.22
41.26
42.05
491.75
82.97
190.48
164.58
106.64
51.41
23.35
15.00
174.16
118.37
34.27
30.43
20.12
14.42
22.56
16.00
713.47
107.85
69.48
53.66
117.61
40.39
9.54
21.51
112.09
75.93
53.74
83.59
134.95
34.71
23.71
13.28
211.86
127.52
59.74
47.67
90.14
96.71
55.14
54.02
10
314.01
149.87
129.01
59.07
141.78
42.96
41.96
48.57
11
177.50
118.04
68.59
21.96
39.94
54.15
13.85
10.22
12
656.09
64.34
79.28
99.04
123.31
82.36
37.04
27.69
13
258.81
201.87
151.54
85.75
27.90
13.53
23.87
17.46
14
221.39
175.76
134.50
99.21
168.92
44.32
29.30
24.26
15
124.79
102.84
36.77
39.12
214.91
35.99
60.35
23.93
16
59.62
80.95
89.80
51.73
81.48
48.29
23.72
27.30
17
722.00
136.71
75.30
78.51
116.19
21.83
21.14
34.99
18
178.19
77.71
72.19
19.29
111.29
83.91
47.46
40.03
19
295.75
98.08
48.70
12.85
20.80
24.72
18.47
14.03
20
1210.68 136.27
143.01
86.42
279.64
77.28
41.92
39.92
Mean
351.68
118.36
87.03
66.34
106.05
54.45
32.57
25.58
SD
289.34
45.74
44.55
40.11
65.78
32.63
15.30
13.54
216
Dorsiflexion
H-reflex gain (%)

Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
238.28
75.34
33.59
32.13
274.91
132.28
41.85
19.73
209.99
91.44
65.54
53.66
28.12
8.98
11.92
7.06
255.24
136.92
93.11
66.77
24.50
58.42
37.11
32.40
17.54
25.04
38.45
26.86
17.02
30.32
31.75
25.29
371.23
161.03
59.79
22.01
52.49
29.45
24.15
14.85
124.35
67.63
56.49
31.16
8.05
6.96
9.54
8.58
686.39
212.16
76.94
56.66
28.56
37.28
15.95
5.96
77.85
13.24
37.46
47.28
84.40
30.66
18.17
13.52
146.42
60.84
53.92
39.47
61.98
65.67
59.47
39.54
10
194.54
83.62
70.80
42.07
151.26
9.34
6.21
6.34
11
116.69
68.23
48.21
45.12
40.27
45.88
33.80
20.52
12
384.57
68.59
23.56
4.49
75.36
57.46
28.79
18.86
13
185.68
124.53
74.49
34.24
23.33
32.00
25.34
22.46
14
186.13
140.48
91.99
54.15
80.74
36.42
40.94
22.25
15
73.71
95.54
90.79
40.89
53.51
41.95
35.12
28.70
16
66.25
43.60
24.36
39.08
41.46
31.82
9.98
15.00
17
550.63
144.96
58.17
55.94
64.05
28.64
36.93
27.65
18
142.84
76.45
45.65
11.41
111.62
60.88
44.07
42.30
19
260.08
102.42
68.16
42.43
20.17
15.03
15.59
13.72
20
989.86
186.07
111.35
73.70
126.68
71.07
41.54
26.00
Mean
263.91
98.91
61.14
40.97
68.42
41.53
28.41
20.54
SD
237.82
52.19
24.12
17.15
62.01
28.47
14.27
10.39
217
Plantarflexion
SOL bEMG (V)

Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
4.69
27.06
23.40
32.60
9.46
14.75
22.75
34.75
9.18
14.32
45.31
48.82
4.69
24.59
31.23
36.92
10.41
21.63
28.61
38.30
10.41
49.91
71.34
70.00
24.73
33.15
30.77
35.22
9.18
24.34
29.35
45.46
4.76
5.75
10.96
17.48
4.76
24.96
34.39
49.08
12.94
14.10
26.84
30.19
12.94
56.00
59.46
62.02
4.68
9.83
18.42
28.38
4.68
23.40
31.47
38.71
4.37
14.11
16.80
25.06
4.37
29.04
55.15
53.95
9.18
19.00
31.58
36.09
5.60
12.60
27.42
39.63
10
5.60
7.97
15.36
12.42
4.16
20.69
32.88
38.70
11
11.45
22.55
26.69
26.13
13.67
11.48
22.59
28.76
12
4.16
29.21
54.36
58.63
8.84
56.65
59.39
74.09
13
13.67
14.99
28.96
34.36
13.12
33.93
42.98
43.69
14
8.84
18.92
31.20
31.15
7.05
55.23
79.47
64.11
15
13.12
28.52
33.94
37.95
5.62
8.24
9.54
12.40
16
18.56
19.95
27.85
28.96
9.58
37.98
40.54
64.76
17
7.05
14.56
24.00
40.13
5.89
21.69
25.27
25.59
18
5.89
33.39
27.39
36.40
7.32
18.95
38.51
41.84
19
7.32
12.38
20.69
35.62
5.31
14.41
19.79
16.48
20
5.31
28.63
44.35
59.32
5.94
26.74
32.23
41.56
Mean
9.29
19.50
28.37
34.66
7.63
28.28
38.29
44.13
SD
5.34
8.29
10.48
11.49
3.44
16.91
19.93
16.50
218
Neutral
SOL bEMG (V)

Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
4.79
17.74
33.67
43.25
15.98
14.79
23.73
43.28
9.18
12.45
32.64
66.14
4.79
11.06
22.36
30.47
10.11
23.37
32.18
35.24
10.11
16.06
39.14
57.63
24.93
33.64
34.66
39.06
9.18
15.55
26.01
36.16
4.69
24.08
16.25
20.01
4.69
11.87
26.83
40.15
13.60
22.07
46.51
37.14
13.60
34.85
33.31
40.97
4.22
26.50
38.64
49.17
4.22
17.45
29.68
35.13
4.31
18.56
61.29
48.68
4.31
22.07
29.52
39.95
9.83
15.50
35.93
35.43
5.99
10.31
20.06
21.21
10
5.99
10.37
15.83
21.58
4.68
17.57
23.00
26.20
11
12.21
27.92
55.96
47.29
13.45
6.65
19.05
21.03
12
4.68
30.27
58.71
68.77
10.15
14.48
30.68
39.24
13
13.45
20.95
29.90
55.67
14.70
23.28
38.82
39.25
14
10.15
16.31
24.53
28.47
6.98
34.05
59.13
87.18
15
14.70
20.89
31.04
42.03
5.36
14.87
17.88
24.64
16
20.98
32.77
30.08
41.65
9.45
18.60
41.41
49.45
17
6.98
31.39
66.87
72.14
5.87
25.17
31.58
34.81
18
5.87
18.39
29.31
50.03
7.26
16.61
36.72
49.57
19
7.26
17.21
20.89
51.48
5.18
7.08
13.72
20.84
20
5.18
51.99
47.51
85.38
6.51
21.33
39.78
47.90
Mean
9.65
23.62
37.12
46.93
8.12
17.69
30.12
39.25
SD
5.70
9.42
14.62
16.66
4.21
8.02
10.53
16.47
219
Dorsiflexion
SOL bEMG (V)

Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
4.60
16.14
25.02
45.04
8.53
17.58
32.89
38.24
6.93
18.72
36.08
56.83
4.60
11.36
17.47
21.77
11.00
17.74
23.22
33.47
11.00
15.46
25.98
42.62
21.61
31.41
40.74
55.97
9.18
13.04
20.41
34.45
4.71
13.31
25.55
60.16
4.71
18.28
19.90
23.63
12.88
25.55
41.38
52.76
12.88
28.29
36.68
38.98
4.00
14.95
36.73
41.64
4.00
13.28
24.55
36.75
4.31
13.04
31.48
50.12
4.31
22.64
31.24
34.14
10.10
19.17
31.11
46.34
5.54
11.86
15.92
23.50
10
5.54
11.03
19.04
27.35
4.31
13.89
19.95
27.00
11
16.79
28.76
51.34
73.26
12.02
7.50
14.70
20.39
12
4.31
21.32
44.33
70.95
9.92
17.71
34.70
39.85
13
12.02
16.20
26.92
41.24
14.76
19.59
24.63
29.40
14
9.92
16.87
31.12
58.70
8.30
25.02
30.67
57.78
15
14.76
16.54
23.97
32.98
5.91
12.18
18.96
20.34
16
16.09
23.63
53.36
86.98
9.72
20.19
42.51
47.29
17
8.30
30.85
57.64
100.44
5.28
20.97
24.13
29.40
18
5.28
15.12
24.77
43.19
7.25
11.68
21.18
29.16
19
7.25
12.94
31.24
40.39
5.30
5.88
12.64
17.07
20
5.30
29.24
66.50
118.66
6.10
17.59
34.83
49.79
Mean
9.29
19.63
36.08
56.82
7.68
16.20
25.20
33.08
SD
5.06
6.43
12.94
23.33
3.55
5.62
7.15
10.36
220
Plantarflexion
TA bEMG (V)
Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
3.34
4.84
7.09
8.29
4.62
6.03
8.35
9.91
3.44
4.92
7.03
9.63
3.34
5.50
7.40
10.47
3.91
6.15
9.88
13.91
3.91
13.36
16.79
16.00
5.13
6.43
8.00
9.51
3.44
12.36
10.00
12.78
13.43
12.11
12.77
15.19
13.43
3.84
4.61
5.57
14.19
15.65
18.03
20.02
14.19
10.16
10.83
10.70
6.94
7.26
8.69
10.40
6.94
10.27
11.83
13.65
5.38
6.45
8.00
9.76
5.38
7.74
9.94
11.60
3.44
9.14
12.92
17.38
4.67
7.62
8.80
9.26
10
4.67
6.28
8.57
9.55
5.26
5.03
6.21
7.33
11
5.41
9.77
13.21
14.16
13.69
7.57
11.13
12.89
12
5.26
6.07
10.26
12.14
7.62
6.77
9.87
14.07
13
13.69
12.79
12.87
15.42
6.58
7.80
10.31
10.38
14
7.62
7.84
9.96
14.18
8.10
5.91
7.39
9.07
15
6.58
8.33
10.40
11.72
7.69
9.54
11.72
17.44
16
7.64
7.90
9.65
9.92
27.50
27.96
29.68
26.95
17
8.10
7.84
9.79
12.98
4.99
16.48
25.27
18.80
18
4.99
5.69
7.09
9.53
3.96
8.59
9.14
10.65
19
3.96
4.22
4.96
6.64
4.05
5.73
7.07
7.57
20
4.05
7.04
10.07
14.17
4.17
9.71
12.07
13.08
Mean
6.56
7.84
9.96
12.22
7.68
9.40
11.42
12.41
SD
3.43
2.88
2.92
3.33
3.70
2.71
2.85
3.15
221
Neutral
TA bEMG (V)
Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
3.33
11.60
13.51
27.45
6.74
11.62
12.51
30.40
3.99
8.93
13.45
35.77
3.33
18.24
32.25
34.40
3.78
12.84
16.22
22.14
3.78
12.14
12.24
35.61
5.75
12.58
13.27
20.20
3.44
17.88
22.14
21.47
14.23
17.14
13.12
16.20
14.23
7.68
10.56
22.20
14.56
18.09
22.46
28.58
14.56
22.65
15.74
25.94
7.01
16.16
16.87
29.61
7.01
13.67
15.18
25.15
5.40
12.16
21.89
31.89
5.40
14.31
11.30
23.46
13.99
11.12
17.46
23.90
4.67
8.98
10.80
16.86
10
4.67
8.23
8.75
14.51
5.70
4.16
10.24
17.80
11
5.96
18.71
22.46
30.96
15.42
6.55
17.14
16.15
12
5.70
19.70
22.92
36.07
8.29
8.92
10.83
20.00
13
15.42
16.38
18.09
30.65
6.63
14.80
15.37
25.42
14
8.29
10.25
12.10
18.49
8.17
20.83
17.53
49.02
15
6.63
13.90
14.92
24.71
6.92
16.70
18.78
24.04
16
7.53
18.86
12.64
25.93
24.88
22.54
31.51
41.21
17
8.17
21.21
23.41
49.82
5.68
21.04
22.06
26.07
18
5.68
11.04
12.06
30.73
4.01
11.84
15.84
29.32
19
4.01
10.93
9.23
30.70
4.16
5.63
6.79
13.50
20
4.16
27.10
20.65
56.70
4.30
12.84
15.99
29.73
Mean
7.41
14.85
16.27
29.25
7.87
13.65
16.24
26.39
SD
3.93
4.79
4.69
10.21
4.00
5.39
5.83
8.63
222
Dorsiflexion
TA bEMG (V)
Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
7.39
3.70
4.85
7.37
35.11
5.08
5.88
6.66
2.84
4.25
5.53
7.75
7.39
30.44
50.10
49.82
3.96
5.29
6.42
8.82
3.96
6.28
6.62
10.23
4.16
4.35
5.55
7.34
3.44
19.81
15.47
6.21
16.58
10.61
10.76
12.61
16.58
3.93
3.53
3.80
14.50
16.06
16.78
17.08
14.50
9.05
9.40
8.65
12.49
7.34
8.32
9.37
12.49
10.96
11.28
11.63
5.49
6.10
6.36
8.01
5.49
7.06
8.14
8.83
26.52
7.86
9.33
13.49
4.73
6.92
7.41
9.34
10
4.73
5.56
6.59
9.01
5.18
3.95
4.20
4.99
11
7.10
8.22
9.79
12.23
11.82
6.55
51.13
97.15
12
5.18
5.43
6.26
9.05
7.98
3.40
3.84
4.93
13
11.82
13.07
13.24
13.35
6.44
5.12
6.08
6.86
14
7.98
7.02
7.78
8.99
7.28
29.40
10.86
7.20
15
6.44
5.86
6.14
9.71
6.69
13.65
22.35
25.17
16
6.76
6.58
7.18
9.24
27.03
27.45
27.61
28.53
17
7.28
7.84
8.15
9.51
4.66
16.65
17.03
25.93
18
4.66
5.21
6.03
6.79
4.03
7.90
7.96
8.37
19
4.03
3.99
4.27
4.95
4.18
3.42
4.13
5.51
20
4.18
4.43
5.74
10.45
48.74
5.96
6.20
8.30
Mean
8.20
6.94
7.75
9.76
11.89
11.15
13.96
16.91
SD
5.73
3.15
3.04
2.80
11.95
8.89
14.05
22.03
223
H-reflex latency (ms.cm-1)

Young group
Older group
Participants
20
-20
20
-20
0.210
0.218
0.221
0.242
0.242
0.251
0.218
0.221
0.227
0.250
0.229
0.270
0.206
0.207
0.212
0.247
0.247
0.253
0.205
0.216
0.227
0.261
0.263
0.257
0.213
0.211
0.217
0.291
0.290
0.302
0.218
0.222
0.228
0.243
0.276
0.248
0.226
0.220
0.236
0.238
0.233
0.235
0.204
0.196
0.199
0.243
0.225
0.241
0.219
0.224
0.232
0.248
0.272
0.226
10
0.209
0.204
0.225
0.239
0.195
0.219
11
0.207
0.204
0.214
0.229
0.241
0.260
12
0.208
0.209
0.219
0.251
0.256
0.290
13
0.206
0.207
0.226
0.240
0.237
0.246
14
0.203
0.200
0.192
0.251
0.244
0.282
15
0.208
0.209
0.213
0.233
0.239
0.247
16
0.232
0.232
0.240
0.262
0.286
0.293
17
0.208
0.213
0.232
0.240
0.232
0.239
18
0.240
0.243
0.250
0.224
0.229
0.237
19
0.217
0.219
0.225
0.252
0.257
0.277
20
0.219
0.225
0.238
0.204
0.200
0.210
Mean
0.214
0.215
0.224
0.244
0.245
0.254
SD
0.010
0.011
0.014
0.015
0.023
0.023
224
H-reflex duration (ms)

Young group
Older group
Participants
20
-20
20
-20
9.75
10.25
11.00
13.75
16.00
13.50
13.50
14.25
13.75
19.50
22.00
10.00
10.50
10.25
10.00
16.25
15.25
17.75
18.75
10.00
10.25
13.00
15.00
16.00
11.75
13.75
14.00
16.00
17.75
15.00
12.50
13.50
13.00
18.50
22.00
17.75
12.00
11.50
12.25
12.75
9.00
10.50
13.25
20.00
26.25
17.50
30.75
30.00
14.00
12.50
12.50
16.25
35.25
44.00
10
13.25
12.25
13.00
15.25
26.50
39.00
11
10.25
11.50
12.00
16.00
19.00
13.00
12
8.00
8.75
10.00
20.00
23.75
31.25
13
10.25
10.00
20.50
10.00
16.50
13.50
14
9.50
10.00
22.50
12.00
22.25
36.25
15
9.50
9.75
17.25
13.75
13.50
11.75
16
10.75
11.00
13.00
18.25
32.75
31.25
17
10.50
10.00
10.25
16.25
13.50
14.00
18
19.75
18.75
18.75
12.75
23.25
22.50
19
12.75
11.75
15.75
11.00
10.50
20.50
20
13.75
13.00
14.00
12.60
11.40
12.05
Mean
12.21
12.14
14.50
15.07
19.80
20.98
SD
2.94
2.90
4.46
2.84
6.87
11.48
225
Study II (after the Grubbs test)
Plantarflexion
H-reflex gain (%)

Young group
10%
30%
Older group
Participants
Rest
10%
30%
50%
667.01
150.23 180.90 136.82 241.53
57.71
15.07
337.68
231.64
10.51
61.71
22.18
10.26
7.35
449.19
138.49 124.99 116.57
67.83
29.58
29.87
17.81
51.03
54.10
51.78
51.25
114.50
38.61
34.04
7.04
480.06
425.19
193.85
92.69
36.16
18.27
8.39
209.57
147.38
32.31
37.66
25.91
22.90
18.89
22.73
420.66
241.98 135.50
84.50
167.41
48.53
25.96
13.68
355.51
191.05 202.00 135.28 140.28
26.91
20.32
20.56
242.25
106.49
158.34 103.62
43.52
32.28
10
488.38
245.28 113.10 188.80 265.92
58.71
42.08
36.11
11
285.79
27.25
26.03
29.59
15.84
15.59
9.37
12
1236.15
40.06
110.33 107.89 151.18
19.89
20.65
18.45
13
367.96
331.66 160.37 119.81
39.54
30.83
21.33
20.96
14
325.87
178.98 110.38
73.94
256.52
31.34
23.11
27.36
15
149.61
44.00
63.35
53.36
254.70
93.42
16
69.71
105.27
97.82
78.50
73.72
26.19
46.27
31.81
17
681.33
255.46 141.63
79.16
70.22
32.52
37.16
29.70
18
177.51
70.80
69.82
19.08
147.71
91.59
50.33
47.93
19
260.48
89.89
59.38
24.60
17.81
18.99
12.90
10.04
20
1219.22 194.82 133.95 108.95 284.68
76.57
36.77
12.51
Mean
423.75
163.50 100.69
83.10
133.09
43.39
29.74
20.48
SD
322.46
104.24
54.98
88.22
27.74
13.66
11.27
54.70
44.69
51.17
50%
19.14
22.44
Rest
226
Neutral
H-reflex gain (%)

Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
499.11
133.22
103.17
110.25
140.45
56.13
14.39
245.92
183.63
74.31
34.23
43.10
29.26
15.04
5.32
336.38
177.03
110.58
122.61
68.00
83.04
45.66
21.70
29.97
18.16
16.20
26.79
73.93
64.22
41.26
42.05
491.75
82.97
190.48
164.58
106.64
51.41
23.35
15.00
174.16
118.37
34.27
30.43
20.12
14.42
22.56
16.00
713.47
107.85
69.48
53.66
117.61
40.39
9.54
21.51
112.09
75.93
53.74
83.59
134.95
34.71
23.71
13.28
211.86
127.52
59.74
47.67
90.14
96.71
55.14
54.02
10
314.01
149.87
129.01
59.07
141.78
42.96
41.96
48.57
11
177.50
118.04
68.59
21.96
39.94
54.15
13.85
10.22
12
656.09
64.34
79.28
99.04
123.31
82.36
37.04
27.69
13
258.81
201.87
151.54
85.75
27.90
13.53
23.87
17.46
14
221.39
175.76
134.50
99.21
168.92
44.32
29.30
24.26
15
124.79
102.84
36.77
39.12
214.91
35.99
60.35
23.93
16
59.62
80.95
89.80
51.73
81.48
48.29
23.72
27.30
17
722.00
136.71
75.30
78.51
116.19
21.83
21.14
34.99
18
178.19
77.71
72.19
19.29
111.29
83.91
47.46
40.03
19
295.75
98.08
48.70
12.85
20.80
24.72
18.47
14.03
20
136.27
143.01
86.42
77.28
41.92
39.92
Mean
306.47
118.36
87.03
66.34
96.92
49.66
32.57
25.58
SD
212.65
45.74
44.55
40.11
52.97
25.28
15.30
13.54
227
Dorsiflexion
H-reflex gain (%)

Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
238.28
75.34
33.59
32.13
41.85
19.73
209.99
91.44
65.54
53.66
28.12
8.98
11.92
7.06
255.24
136.92
93.11
66.77
24.50
58.42
37.11
32.40
17.54
25.04
38.45
26.86
17.02
30.32
31.75
25.29
371.23
161.03
59.79
22.01
52.49
29.45
24.15
14.85
124.35
67.63
56.49
31.16
8.05
6.96
9.54
8.58
686.39
212.16
76.94
56.66
28.56
37.28
15.95
5.96
77.85
13.24
37.46
47.28
84.40
30.66
18.17
13.52
146.42
60.84
53.92
39.47
61.98
65.67
59.47
39.54
10
194.54
83.62
70.80
42.07
151.26
9.34
6.21
6.34
11
116.69
68.23
48.21
45.12
40.27
45.88
33.80
20.52
12
384.57
68.59
23.56
4.49
75.36
57.46
28.79
18.86
13
185.68
124.53
74.49
34.24
23.33
32.00
25.34
22.46
14
186.13
140.48
91.99
54.15
80.74
36.42
40.94
22.25
15
73.71
95.54
90.79
40.89
53.51
41.95
35.12
28.70
16
66.25
43.60
24.36
39.08
41.46
31.82
9.98
15.00
17
550.63
144.96
58.17
55.94
64.05
28.64
36.93
27.65
18
142.84
76.45
45.65
11.41
111.62
60.88
44.07
42.30
19
260.08
102.42
68.16
42.43
20.17
15.03
15.59
13.72
20
186.07
111.35
73.70
126.68
71.07
41.54
26.00
Mean
225.71
98.91
61.14
40.97
57.56
36.75
28.41
20.54
SD
169.94
52.19
24.12
17.15
39.57
19.34
14.27
10.39
228
Plantarflexion
SOL bEMG (V)

Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
4.69
27.06
23.40
32.60
9.46
14.75
22.75
34.75
9.18
14.32
45.31
48.82
4.69
24.59
31.23
36.92
10.41
21.63
28.61
38.30
10.41
49.91
71.34
70.00
33.15
30.77
35.22
9.18
24.34
29.35
45.46
4.76
5.75
10.96
17.48
4.76
24.96
34.39
49.08
12.94
14.10
26.84
30.19
12.94
56.00
59.46
62.02
4.68
9.83
18.42
28.38
4.68
23.40
31.47
38.71
4.37
14.11
16.80
25.06
4.37
29.04
55.15
53.95
9.18
19.00
31.58
36.09
5.60
12.60
27.42
39.63
10
5.60
7.97
15.36
12.42
4.16
20.69
32.88
38.70
11
11.45
22.55
26.69
26.13
13.67
11.48
22.59
28.76
12
4.16
29.21
54.36
58.63
8.84
56.65
59.39
74.09
13
13.67
14.99
28.96
34.36
13.12
33.93
42.98
43.69
14
8.84
18.92
31.20
31.15
7.05
55.23
79.47
64.11
15
13.12
28.52
33.94
37.95
5.62
8.24
9.54
12.40
16
18.56
19.95
27.85
28.96
9.58
37.98
40.54
64.76
17
7.05
14.56
24.00
40.13
5.89
21.69
25.27
25.59
18
5.89
33.39
27.39
36.40
7.32
18.95
38.51
41.84
19
7.32
12.38
20.69
35.62
5.31
14.41
19.79
16.48
20
5.31
28.63
44.35
59.32
5.94
26.74
32.23
41.56
Mean
8.48
19.50
28.37
34.66
7.63
28.28
38.29
44.13
SD
4.02
8.29
10.48
11.49
3.44
16.91
19.93
16.50
229
Neutral
SOL bEMG (V)

Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
4.79
17.74
33.67
43.25
15.98
14.79
23.73
43.28
9.18
12.45
32.64
66.14
4.79
11.06
22.36
30.47
10.11
23.37
32.18
35.24
10.11
16.06
39.14
57.63
33.64
34.66
39.06
9.18
15.55
26.01
36.16
4.69
24.08
16.25
20.01
4.69
11.87
26.83
40.15
13.60
22.07
46.51
37.14
13.60
34.85
33.31
40.97
4.22
26.50
38.64
49.17
4.22
17.45
29.68
35.13
4.31
18.56
61.29
48.68
4.31
22.07
29.52
39.95
9.83
15.50
35.93
35.43
5.99
10.31
20.06
21.21
10
5.99
10.37
15.83
21.58
4.68
17.57
23.00
26.20
11
12.21
27.92
55.96
47.29
13.45
6.65
19.05
21.03
12
4.68
30.27
58.71
68.77
10.15
14.48
30.68
39.24
13
13.45
20.95
29.90
55.67
14.70
23.28
38.82
39.25
14
10.15
16.31
24.53
28.47
6.98
34.05
15
14.70
20.89
31.04
42.03
5.36
14.87
17.88
24.64
16
20.98
32.77
30.08
41.65
9.45
18.60
41.41
49.45
17
6.98
31.39
66.87
72.14
5.87
25.17
31.58
34.81
18
5.87
18.39
29.31
50.03
7.26
16.61
36.72
49.57
19
7.26
17.21
20.89
51.48
5.18
7.08
13.72
20.84
20
5.18
47.51
85.38
6.51
21.33
39.78
47.90
Mean
8.85
22.13
37.12
46.93
8.12
17.69
28.59
36.73
SD
4.54
6.83
14.62
16.66
4.21
8.02
8.21
10.61
230
Dorsiflexion
SOL bEMG (V)

Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
4.60
16.14
25.02
45.04
8.53
17.58
32.89
38.24
6.93
18.72
36.08
56.83
4.60
11.36
17.47
21.77
11.00
17.74
23.22
33.47
11.00
15.46
25.98
42.62
21.61
31.41
40.74
55.97
9.18
13.04
20.41
34.45
4.71
13.31
25.55
60.16
4.71
18.28
19.90
23.63
12.88
25.55
41.38
52.76
12.88
28.29
36.68
38.98
4.00
14.95
36.73
41.64
4.00
13.28
24.55
36.75
4.31
13.04
31.48
50.12
4.31
22.64
31.24
34.14
10.10
19.17
31.11
46.34
5.54
11.86
15.92
23.50
10
5.54
11.03
19.04
27.35
4.31
13.89
19.95
27.00
11
16.79
28.76
51.34
73.26
12.02
7.50
14.70
20.39
12
4.31
21.32
44.33
70.95
9.92
17.71
34.70
39.85
13
12.02
16.20
26.92
41.24
14.76
19.59
24.63
29.40
14
9.92
16.87
31.12
58.70
8.30
25.02
30.67
57.78
15
14.76
16.54
23.97
32.98
5.91
12.18
18.96
20.34
16
16.09
23.63
53.36
86.98
9.72
20.19
42.51
47.29
17
8.30
30.85
57.64
100.44
5.28
20.97
24.13
29.40
18
5.28
15.12
24.77
43.19
7.25
11.68
21.18
29.16
19
7.25
12.94
31.24
40.39
5.30
5.88
12.64
17.07
20
5.30
29.24
66.50
6.10
17.59
34.83
49.79
Mean
9.29
19.63
36.08
53.57
7.68
16.20
25.20
33.08
SD
5.06
6.43
12.94
18.73
3.55
5.62
7.15
10.36
231
Plantarflexion
TA bEMG (V)
Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
3.34
4.84
7.09
8.29
4.62
6.03
8.35
9.91
3.44
4.92
7.03
9.63
3.34
5.50
7.40
10.47
3.91
6.15
9.88
13.91
3.91
13.36
16.79
16.00
5.13
6.43
8.00
9.51
3.44
12.36
10.00
12.78
13.43
12.11
12.77
15.19
13.43
3.84
4.61
5.57
14.19
20.02
14.19
10.16
10.83
10.70
6.94
7.26
8.69
10.40
6.94
10.27
11.83
13.65
5.38
6.45
8.00
9.76
5.38
7.74
9.94
11.60
3.44
9.14
12.92
17.38
4.67
7.62
8.80
9.26
10
4.67
6.28
8.57
9.55
5.26
5.03
6.21
7.33
11
5.41
9.77
13.21
14.16
13.69
7.57
11.13
12.89
12
5.26
6.07
10.26
12.14
7.62
6.77
9.87
14.07
13
13.69
12.79
12.87
15.42
6.58
7.80
10.31
10.38
14
7.62
7.84
9.96
14.18
8.10
5.91
7.39
9.07
15
6.58
8.33
10.40
11.72
7.69
9.54
11.72
17.44
16
7.64
7.90
9.65
9.92
26.95
17
8.10
7.84
9.79
12.98
4.99
18.80
18
4.99
5.69
7.09
9.53
3.96
8.59
9.14
10.65
19
3.96
4.22
4.96
6.64
4.05
5.73
7.07
7.57
20
4.05
7.04
10.07
14.17
4.17
9.71
12.07
13.08
Mean
6.56
7.42
9.54
12.22
6.63
7.97
9.64
12.41
SD
3.43
2.28
2.29
3.33
3.49
2.55
2.73
3.15
232
Neutral
TA bEMG (V)
Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
3.33
11.60
13.51
27.45
6.74
11.62
12.51
30.40
3.99
8.93
13.45
35.77
3.33
18.24
34.40
3.78
12.84
16.22
22.14
3.78
12.14
12.24
35.61
5.75
12.58
13.27
20.20
3.44
17.88
22.14
21.47
14.23
17.14
13.12
16.20
14.23
7.68
10.56
22.20
14.56
18.09
22.46
28.58
14.56
22.65
15.74
25.94
7.01
16.16
16.87
29.61
7.01
13.67
15.18
25.15
5.40
12.16
21.89
31.89
5.40
14.31
11.30
23.46
13.99
11.12
17.46
23.90
4.67
8.98
10.80
16.86
10
4.67
8.23
8.75
14.51
5.70
4.16
10.24
17.80
11
5.96
18.71
22.46
30.96
15.42
6.55
17.14
16.15
12
5.70
19.70
22.92
36.07
8.29
8.92
10.83
20.00
13
15.42
16.38
18.09
30.65
6.63
14.80
15.37
25.42
14
8.29
10.25
12.10
18.49
8.17
20.83
17.53
15
6.63
13.90
14.92
24.71
6.92
16.70
18.78
24.04
16
7.53
18.86
12.64
25.93
22.54
41.21
17
8.17
21.21
23.41
49.82
5.68
21.04
22.06
26.07
18
5.68
11.04
12.06
30.73
4.01
11.84
15.84
29.32
19
4.01
10.93
9.23
30.70
4.16
5.63
6.79
13.50
20
4.16
27.10
20.65
4.30
12.84
15.99
29.73
Mean
7.41
14.85
16.27
27.80
6.97
13.65
14.50
25.20
SD
3.93
4.79
4.69
8.13
3.77
5.39
4.17
7.12
233
Dorsiflexion
TA bEMG (V)
Young group
Older group
Participants
Rest
10%
30%
50%
Rest
10%
30%
50%
7.39
3.70
4.85
7.37
5.08
5.88
6.66
2.84
4.25
5.53
7.75
7.39
30.44
50.10
49.82
3.96
5.29
6.42
8.82
3.96
6.28
6.62
10.23
4.16
4.35
5.55
7.34
3.44
19.81
15.47
6.21
16.58
10.61
10.76
12.61
16.58
3.93
3.53
3.80
14.50
17.08
14.50
9.05
9.40
8.65
12.49
7.34
8.32
9.37
12.49
10.96
11.28
11.63
5.49
6.10
6.36
8.01
5.49
7.06
8.14
8.83
7.86
9.33
13.49
4.73
6.92
7.41
9.34
10
4.73
5.56
6.59
9.01
5.18
3.95
4.20
4.99
11
7.10
8.22
9.79
12.23
11.82
6.55
51.13
12
5.18
5.43
6.26
9.05
7.98
3.40
3.84
4.93
13
11.82
13.07
13.24
13.35
6.44
5.12
6.08
6.86
14
7.98
7.02
7.78
8.99
7.28
29.40
10.86
7.20
15
6.44
5.86
6.14
9.71
6.69
13.65
22.35
25.17
16
6.76
6.58
7.18
9.24
27.03
27.45
27.61
28.53
17
7.28
7.84
8.15
9.51
4.66
16.65
17.03
25.93
18
4.66
5.21
6.03
6.79
4.03
7.90
7.96
8.37
19
4.03
3.99
4.27
4.95
4.18
3.42
4.13
5.51
20
4.18
4.43
5.74
10.45
5.96
6.20
8.30
Mean
7.24
6.46
7.28
9.76
8.55
11.15
13.96
12.68
SD
3.87
2.37
2.23
2.80
6.02
8.89
15.49
11.64
234
H-reflex latency (ms.cm-1)

Young group
Older group
Participants
20
-20
20
-20
0.210
0.218
0.221
0.242
0.242
0.251
0.218
0.221
0.227
0.250
0.229
0.270
0.206
0.207
0.212
0.247
0.247
0.253
0.205
0.216
0.227
0.261
0.263
0.257
0.213
0.211
0.217
0.290
0.302
0.218
0.222
0.228
0.243
0.276
0.248
0.226
0.220
0.236
0.238
0.233
0.235
0.204
0.196
0.199
0.243
0.225
0.241
0.219
0.224
0.232
0.248
0.272
0.226
10
0.209
0.204
0.225
0.239
0.195
0.219
11
0.207
0.204
0.214
0.229
0.241
0.260
12
0.208
0.209
0.219
0.251
0.256
0.290
13
0.206
0.207
0.226
0.240
0.237
0.246
14
0.203
0.200
0.192
0.251
0.244
0.282
15
0.208
0.209
0.213
0.233
0.239
0.247
16
0.232
0.232
0.240
0.262
0.286
0.293
17
0.208
0.213
0.232
0.240
0.232
0.239
18
0.243
0.250
0.224
0.229
0.237
19
0.217
0.219
0.225
0.252
0.257
0.277
20
0.219
0.225
0.238
0.204
0.200
0.210
Mean
0.212
0.215
0.224
0.242
0.245
0.254
SD
0.008
0.011
0.014
0.013
0.023
0.023
235
H-reflex duration (ms)

Young group
Older group
Participants
20
-20
20
-20
9.75
10.25
11.00
13.75
16.00
13.50
13.50
14.25
13.75
19.50
22.00
10.00
10.50
10.25
10.00
16.25
15.25
17.75
18.75
10.00
10.25
13.00
15.00
16.00
11.75
13.75
14.00
16.00
17.75
15.00
12.50
13.50
13.00
18.50
22.00
17.75
12.00
11.50
12.25
12.75
9.00
10.50
13.25
17.50
30.75
30.00
14.00
12.50
12.50
16.25
35.25
44.00
10
13.25
12.25
13.00
15.25
26.50
39.00
11
10.25
11.50
12.00
16.00
19.00
13.00
12
8.00
8.75
10.00
20.00
23.75
31.25
13
10.25
10.00
20.50
10.00
16.50
13.50
14
9.50
10.00
22.50
12.00
22.25
36.25
15
9.50
9.75
17.25
13.75
13.50
11.75
16
10.75
11.00
13.00
18.25
32.75
31.25
17
10.50
10.00
10.25
16.25
13.50
14.00
18
18.75
18.75
12.75
23.25
22.50
19
12.75
11.75
15.75
11.00
10.50
20.50
20
13.75
13.00
14.00
12.60
11.40
12.05
Mean
11.82
11.72
13.88
15.07
19.80
20.98
SD
2.40
2.30
3.59
2.84
6.87
11.48
236
Study III
Hmax/Mmax ratio
Young group
Older group
Participants
PS
PL
AS
AL
PS
PL
AS
AL
0.96
0.93
0.71
0.21
0.62
0.20
0.37
0.24
0.43
0.72
0.54
0.39
0.21
0.15
0.63
0.49
0.49
0.63
0.07
0.12
0.14
0.05
0.36
0.33
0.12
0.09
0.48
0.32
0.06
0.02
0.75
0.48
0.21
0.28
0.46
0.45
0.31
0.19
0.26
0.33
0.39
0.34
0.24
0.25
0.25
0.17
0.59
0.47
0.49
0.38
0.19
0.28
0.14
0.03
0.16
0.29
0.30
0.28
0.73
0.29
0.29
0.04
0.18
0.07
0.67
0.78
0.80
0.50
0.32
0.15
0.09
0.07
10
0.22
0.25
0.42
0.29
0.90
0.52
0.23
0.36
11
0.62
0.73
1.05
0.75
0.21
0.09
0.45
0.24
12
0.55
0.67
0.33
0.52
0.67
0.33
0.39
0.32
13
0.10
0.23
0.51
0.48
0.52
0.26
0.16
0.23
14
0.49
0.57
0.55
0.57
0.58
0.21
0.95
0.66
15
0.74
0.50
0.86
0.81
0.33
0.27
0.48
0.42
16
0.57
0.70
0.66
0.59
0.62
0.26
0.52
0.26
17
0.44
0.33
0.12
0.04
0.44
0.13
0.63
0.61
18
0.75
0.37
0.21
0.29
0.50
0.04
0.17
0.24
19
0.48
0.36
0.58
0.25
20
0.54
0.47
0.61
0.52
Mean
0.48
0.48
0.51
0.40
0.40
0.17
0.41
0.34
SD
0.19
0.20
0.26
0.20
0.22
0.13
0.24
0.16
237
Hmax (mV)
Young group
Older group
Participants
PS
PL
AS
AL
PS
PL
AS
AL
3.19
2.94
3.68
1.16
1.79
0.57
1.58
1.24
2.13
3.58
4.24
3.39
0.36
0.25
1.59
1.36
3.31
4.23
0.69
1.20
0.39
0.13
1.39
1.35
0.36
0.28
2.26
1.72
0.05
0.02
0.89
0.81
0.92
1.24
2.94
3.26
0.76
0.46
0.87
1.29
1.58
1.37
1.65
1.76
0.36
0.24
1.42
1.17
2.53
1.78
1.59
2.08
0.23
0.05
0.41
0.76
1.06
0.92
3.43
1.52
0.59
0.07
0.53
0.21
1.79
2.24
3.75
2.66
0.32
0.14
0.15
0.11
10
1.61
1.82
4.18
2.84
2.03
1.11
0.71
1.03
11
2.20
2.93
5.72
4.55
0.44
0.18
1.40
0.76
12
2.97
3.03
2.62
3.93
0.88
0.36
0.74
0.57
13
0.40
0.85
2.91
2.60
0.70
0.33
0.32
0.42
14
3.98
4.76
5.54
5.67
0.59
0.22
1.20
0.82
15
5.53
3.62
8.30
7.64
0.41
0.37
0.77
0.74
16
2.58
3.14
4.88
4.26
1.51
0.62
1.94
0.96
17
2.05
1.40
0.83
0.27
1.07
0.30
2.36
2.18
18
2.34
1.10
1.06
1.55
2.20
0.16
1.13
1.63
19
1.45
1.05
2.82
1.34
20
3.46
2.92
5.52
4.93
Mean
2.27
2.26
3.43
2.92
0.81
0.31
1.08
0.97
SD
1.27
1.28
1.98
1.82
0.65
0.26
0.59
0.51
238
Hmax (mV)
Young group
Older group
Participants
SO
SC
UO
UC
SO
SC
UO
UC
3.17
3.39
3.15
3.17
1.98
1.98
2.04
2.14
2.14
2.02
2.08
1.97
0.91
0.65
0.35
0.68
1.97
1.85
1.73
1.71
0.64
0.77
0.61
0.84
0.35
0.40
0.45
0.43
0.16
0.17
0.13
0.11
0.91
0.60
0.68
0.66
0.48
0.58
0.45
0.44
1.71
1.66
1.49
1.39
0.68
0.64
0.45
0.44
2.64
2.67
2.68
2.65
0.13
0.12
0.06
0.06
1.43
1.22
1.18
1.24
0.84
0.73
0.59
0.73
1.84
1.80
1.76
1.73
0.46
0.39
0.35
0.38
10
3.50
3.77
3.49
3.95
1.40
1.29
1.22
1.05
11
2.30
2.34
1.76
1.67
0.88
0.61
0.77
0.88
12
1.90
1.97
1.87
1.98
0.72
0.66
0.67
0.64
13
0.78
0.59
0.73
0.62
1.59
1.46
1.13
1.27
14
2.64
2.88
2.47
2.68
0.80
0.76
0.84
0.80
15
5.62
4.94
5.03
5.41
0.59
0.60
0.64
0.63
16
2.63
2.59
2.65
2.40
0.76
0.68
0.73
0.81
17
1.42
1.49
1.39
1.22
0.44
0.44
0.46
0.42
18
1.58
1.58
1.63
1.62
0.85
1.09
0.85
0.99
19
3.44
2.96
2.55
2.55
0.37
0.20
0.27
0.17
20
2.31
2.19
2.08
1.97
0.12
0.10
0.12
0.11
Mean
2.21
2.15
2.04
2.05
0.74
0.70
0.64
0.68
SD
1.17
1.11
1.06
1.18
0.47
0.47
0.46
0.48
239
Study III (after the Grubbs test)
Hmax/Mmax ratio
Young group
Older group
Participants
PS
PL
AS
AL
PS
PL
AS
AL
0.93
0.71
0.21
0.62
0.20
0.31
0.29
0.43
0.72
0.54
0.39
0.21
0.15
0.57
0.54
0.49
0.63
0.07
0.12
0.14
0.05
0.34
0.35
0.12
0.09
0.48
0.32
0.06
0.02
0.53
0.68
0.21
0.28
0.46
0.45
0.31
0.19
0.22
0.38
0.39
0.34
0.24
0.25
0.25
0.17
0.57
0.49
0.49
0.38
0.19
0.28
0.14
0.03
0.15
0.29
0.30
0.28
0.73
0.29
0.29
0.04
0.17
0.07
0.67
0.78
0.80
0.50
0.32
0.15
0.10
0.06
10
0.22
0.25
0.42
0.29
0.90
0.25
0.33
11
0.62
0.73
1.05
0.75
0.21
0.09
0.45
0.24
12
0.55
0.67
0.33
0.52
0.67
0.33
0.41
0.30
13
0.10
0.23
0.51
0.48
0.52
0.26
0.17
0.22
14
0.49
0.57
0.55
0.57
0.58
0.21
0.96
0.65
15
0.74
0.50
0.86
0.33
0.27
0.43
0.46
16
0.57
0.70
0.66
0.59
0.62
0.26
0.53
0.26
17
0.44
0.33
0.12
0.04
0.44
0.13
0.66
0.58
18
0.75
0.37
0.21
0.29
0.50
0.04
19
0.48
0.36
0.58
0.25
20
0.54
0.47
0.61
0.52
Mean
0.45
0.48
0.51
0.37
0.40
0.15
0.40
0.36
SD
0.19
0.20
0.26
0.18
0.22
0.10
0.22
0.18
240
Hmax (mV)
Young group
Older group
Participants
PS
PL
AS
AL
PS
PL
AS
AL
3.19
2.94
3.68
1.16
1.79
0.57
1.58
1.24
2.13
3.58
4.24
3.39
0.36
0.25
1.59
1.36
3.31
4.23
0.69
1.20
0.39
0.13
1.39
1.35
0.36
0.28
2.26
1.72
0.05
0.02
0.89
0.81
0.92
1.24
2.94
3.26
0.76
0.46
0.87
1.29
1.58
1.37
1.65
1.76
0.36
0.24
1.42
1.17
2.53
1.78
1.59
2.08
0.23
0.05
0.41
0.76
1.06
0.92
3.43
1.52
0.59
0.07
0.53
0.21
1.79
2.24
3.75
2.66
0.32
0.14
0.15
0.11
10
1.61
1.82
4.18
2.84
2.03
0.71
1.03
11
2.20
2.93
5.72
4.55
0.44
0.18
1.40
0.76
12
2.97
3.03
2.62
3.93
0.88
0.36
0.74
0.57
13
0.40
0.85
2.91
2.60
0.70
0.33
0.32
0.42
14
3.98
4.76
5.54
5.67
0.59
0.22
1.20
0.82
15
3.62
8.30
0.41
0.37
0.77
0.74
16
2.58
3.14
4.88
4.26
1.51
0.62
1.94
0.96
17
2.05
1.40
0.83
0.27
1.07
0.30
2.36
2.18
18
2.34
1.10
1.06
1.55
2.20
0.16
1.13
1.63
19
1.45
1.05
2.82
1.34
20
3.46
2.92
5.52
4.93
Mean
2.10
2.26
3.43
2.67
0.81
0.26
1.08
0.97
SD
1.01
1.28
1.98
1.47
0.65
0.17
0.59
0.51
241
Hmax (mV)
Young group
Older group
Participants
SO
SC
UO
UC
SO
SC
UO
UC
3.17
3.39
3.15
3.17
2.14
2.02
2.08
1.97
0.91
0.65
0.35
0.68
1.97
1.85
1.73
1.71
0.64
0.77
0.61
0.84
0.35
0.40
0.45
0.43
0.16
0.17
0.13
0.11
0.91
0.60
0.68
0.66
0.48
0.58
0.45
0.44
1.71
1.66
1.49
1.39
0.68
0.64
0.45
0.44
2.64
2.67
2.68
2.65
0.13
0.12
0.06
0.06
1.43
1.22
1.18
1.24
0.84
0.73
0.59
0.73
1.84
1.80
1.76
1.73
0.46
0.39
0.35
0.38
10
3.50
3.77
3.49
3.95
1.40
1.29
1.22
1.05
11
2.30
2.34
1.76
1.67
0.88
0.61
0.77
0.88
12
1.90
1.97
1.87
1.98
0.72
0.66
0.67
0.64
13
0.78
0.59
0.73
0.62
1.59
1.46
1.13
1.27
14
2.64
2.88
2.47
2.68
0.80
0.76
0.84
0.80
15
4.94
0.59
0.60
0.64
0.63
16
2.63
2.59
2.65
2.40
0.76
0.68
0.73
0.81
17
1.42
1.49
1.39
1.22
0.44
0.44
0.46
0.42
18
1.58
1.58
1.63
1.62
0.85
1.09
0.85
0.99
19
3.44
2.96
2.55
2.55
0.37
0.20
0.27
0.17
20
2.31
2.19
2.08
1.97
0.12
0.10
0.12
0.11
Mean
2.03
2.15
1.88
1.87
0.68
0.63
0.56
0.60
SD
0.84
1.11
0.78
0.86
0.38
0.37
0.32
0.34
242
Study IV
Pre-training test
Hmax/Mmax ratio
Tai Chi group
Control group
Participants
SO
SC
UO
UC
SO
SC
UO
UC
0.72
0.71
0.63
0.60
0.46
0.42
0.45
0.49
0.33
0.33
0.30
0.27
0.31
0.21
0.19
0.24
0.36
0.32
0.33
0.30
0.56
0.56
0.57
0.60
0.31
0.25
0.27
0.29
0.31
0.32
0.26
0.26
0.49
0.45
0.41
0.41
0.21
0.21
0.22
0.20
0.25
0.22
0.19
0.14
0.15
0.14
0.07
0.07
0.22
0.18
0.23
0.21
0.40
0.34
0.34
0.29
0.35
0.24
0.19
0.21
0.06
0.04
0.05
0.04
0.10
0.11
0.09
0.09
0.35
0.36
0.19
0.24
10
0.24
0.21
0.21
0.28
0.57
0.63
0.61
0.53
11
0.42
0.38
0.35
0.36
0.47
0.51
0.49
0.43
12
0.10
0.10
0.09
0.20
0.24
0.18
0.24
0.22
13
0.19
0.21
0.19
0.19
0.40
0.36
0.35
0.32
14
0.13
0.10
0.07
0.10
0.19
0.22
0.18
0.21
15
0.27
0.25
0.24
0.25
16
0.36
0.29
0.37
0.22
17
0.40
0.40
0.32
0.29
18
0.09
0.12
0.09
0.10
19
0.39
0.33
0.31
0.39
20
0.51
0.47
0.40
0.36
Mean
0.31
0.28
0.26
0.26
0.33
0.32
0.30
0.30
SD
0.16
0.15
0.14
0.12
0.15
0.17
0.17
0.16
243
Pre-training test
Mean displacement of COPA-P (mm)
Tai Chi group
Control group
Participants
SO
SC
UO
UC
SO
SC
UO
UC
2.96
4.10
4.93
6.35
2.32
2.91
3.24
3.69
2.43
3.24
2.99
4.44
2.05
2.53
4.12
4.48
3.88
6.31
5.47
7.84
3.84
3.46
5.27
6.83
1.81
2.36
2.61
3.77
3.06
2.62
4.48
3.77
3.76
3.18
4.22
3.63
2.47
4.30
3.96
5.50
2.61
2.47
2.84
3.19
3.15
3.36
3.69
5.38
2.91
2.65
3.60
3.61
4.89
5.33
6.53
7.91
2.07
2.88
4.04
5.53
2.60
3.20
2.86
3.68
4.57
5.81
6.60
4.60
2.45
3.57
3.66
5.68
10
2.31
2.38
2.70
3.12
3.07
4.25
3.45
5.40
11
3.66
4.36
4.81
5.18
2.40
3.17
4.25
3.46
12
1.84
2.27
2.73
3.40
4.22
4.52
5.83
6.25
13
4.21
4.35
5.61
6.84
3.39
3.92
4.74
4.67
14
3.39
3.56
4.69
5.29
3.42
4.30
4.18
5.20
15
2.38
3.31
3.90
4.72
16
2.70
2.72
4.01
4.91
18
4.64
1.88
4.70
2.01
8.18
2.58
7.35
2.86
19
3.72
3.98
4.42
6.86
20
2.72
4.08
5.32
5.69
Mean
3.02
3.54
4.31
4.96
3.10
3.68
4.30
5.14
SD
0.90
1.18
1.47
1.50
0.81
0.80
1.01
1.30
17
244
Pre-training test
Mean displacement of COPM-L (mm)
Tai Chi group
Control group
Participants
SO
SC
UO
UC
SO
SC
UO
UC
1.65
2.82
2.08
4.41
1.16
1.30
2.19
2.60
1.77
1.85
2.34
2.17
1.23
1.49
2.49
3.32
2.36
3.53
4.52
6.24
2.88
2.90
4.06
5.16
1.46
2.11
1.78
2.53
2.18
2.05
2.77
3.02
2.04
1.94
2.30
2.79
1.94
2.85
2.76
3.59
1.28
1.22
1.69
2.72
2.49
2.68
2.82
4.39
1.03
1.74
2.51
2.69
2.81
3.94
3.72
5.69
1.41
1.81
2.49
3.76
0.75
0.94
1.05
1.46
2.82
4.21
4.10
5.49
1.82
2.90
3.09
4.78
10
1.62
1.24
1.85
2.37
1.08
1.51
2.62
4.16
11
2.87
3.10
3.28
3.54
1.03
1.57
1.36
2.03
12
1.13
1.18
2.22
2.45
1.90
3.13
2.74
4.92
13
3.05
4.69
4.64
7.08
1.36
1.69
3.26
3.54
14
2.18
2.65
3.42
3.90
2.10
2.19
3.22
3.40
15
1.57
2.21
2.37
2.87
16
1.25
2.26
3.07
3.67
17
1.49
1.53
4.03
4.01
18
0.97
0.94
1.36
1.89
19
1.08
1.09
2.17
3.49
20
2.16
3.01
3.10
4.90
Mean
1.76
2.26
2.77
3.65
1.77
2.22
2.72
3.72
SD
0.63
1.05
0.96
1.40
0.68
0.86
0.81
1.22
245
Post-training test
Hmax/Mmax ratio
Tai Chi group
Control group
Participants
SO
SC
UO
UC
SO
SC
UO
UC
0.55
0.59
0.53
0.57
0.52
0.57
0.49
0.45
0.35
0.36
0.39
0.41
0.31
0.25
0.24
0.26
0.28
0.23
0.16
0.20
0.52
0.45
0.43
0.44
0.26
0.28
0.22
0.24
0.38
0.22
0.29
0.33
0.57
0.53
0.50
0.50
0.40
0.39
0.33
0.36
0.35
0.35
0.22
0.17
0.30
0.36
0.29
0.25
0.33
0.30
0.33
0.29
0.40
0.36
0.35
0.32
0.65
0.63
0.54
0.58
0.05
0.05
0.05
0.06
0.15
0.14
0.12
0.13
0.46
0.34
0.32
0.33
10
0.24
0.24
0.21
0.19
0.48
0.52
0.47
0.35
11
0.45
0.40
0.36
0.36
0.22
0.16
0.15
0.13
12
0.19
0.23
0.17
0.16
0.17
0.15
0.15
0.19
13
0.20
0.21
0.23
0.24
0.29
0.27
0.34
0.34
14
0.09
0.07
0.08
0.11
0.20
0.21
0.20
0.20
15
0.44
0.50
0.40
0.43
16
0.40
0.36
0.42
0.39
17
0.32
0.34
0.33
0.29
18
0.11
0.11
0.13
0.12
19
0.45
0.49
0.35
0.36
20
0.56
0.49
0.40
0.48
Mean
0.35
0.34
0.30
0.31
0.33
0.31
0.29
0.29
SD
0.16
0.16
0.14
0.15
0.14
0.15
0.13
0.11
246
Post-training test
Tai Chi group
Control group
Participants
SO
SC
UO
UC
SO
SC
UO
UC
2.64
4.07
3.55
5.08
2.90
2.61
2.77
3.99
2.78
3.88
3.89
4.83
2.64
3.06
3.97
4.34
4.50
6.53
7.03
8.67
2.79
2.81
3.16
3.63
2.67
2.59
2.95
3.86
2.15
3.21
3.48
3.61
2.86
4.40
4.37
5.12
2.78
3.12
4.35
4.44
2.28
2.89
3.03
2.90
2.50
2.78
4.71
4.17
2.81
3.01
3.63
3.78
4.11
5.04
6.56
8.97
1.91
2.10
2.88
3.17
2.43
3.79
3.02
3.67
4.15
3.00
4.30
5.02
3.00
4.03
3.51
4.43
10
2.34
2.85
2.71
3.27
3.04
3.14
3.88
4.74
11
3.40
4.03
4.49
6.29
2.04
2.48
3.03
3.14
12
1.75
1.87
2.33
2.40
2.71
4.59
5.01
6.01
13
3.64
5.06
4.77
6.58
2.24
2.11
3.51
3.59
14
3.82
4.58
5.76
5.97
3.50
4.70
4.15
5.34
15
3.03
3.27
3.85
4.10
16
3.16
3.06
5.61
7.65
17
3.69
4.02
6.07
6.84
18
2.26
2.15
2.47
2.84
19
4.27
4.17
4.97
6.49
20
3.25
3.71
4.25
5.75
Mean
3.06
3.56
4.15
5.03
2.77
3.39
3.94
4.58
SD
0.78
1.12
1.28
1.74
0.55
0.90
1.00
1.47
247
Post-training test
Tai Chi group
Control group
Participants
SO
SC
UO
UC
SO
SC
UO
UC
1.44
1.99
1.66
3.97
1.99
1.40
2.35
3.28
2.03
2.29
2.55
3.05
2.17
1.95
2.86
2.62
3.17
4.51
4.55
7.32
1.40
1.30
1.93
2.65
1.99
1.73
2.23
2.57
1.24
2.08
2.13
2.71
1.90
3.37
2.61
4.67
1.68
1.98
2.64
3.63
1.50
1.55
1.89
2.36
1.88
2.67
3.78
3.73
1.67
1.64
2.38
2.92
2.32
2.02
3.22
3.93
1.58
1.45
1.84
2.07
0.73
0.92
1.25
1.37
1.91
2.02
2.31
3.28
2.41
3.28
3.45
3.72
10
1.63
1.97
2.41
3.52
1.16
1.66
2.49
2.89
11
2.56
3.03
3.24
3.68
1.22
1.67
2.03
2.42
12
1.21
1.37
1.82
2.25
1.56
3.04
3.28
5.13
13
3.23
3.62
3.98
6.63
0.95
1.04
2.08
2.90
14
2.67
2.88
3.26
5.24
2.39
2.72
4.31
3.27
15
1.63
1.71
2.27
2.35
16
1.60
2.12
2.84
5.05
17
1.80
2.26
4.76
6.01
18
1.09
0.90
1.36
1.28
19
0.88
1.31
2.02
2.93
20
2.40
2.03
3.32
4.37
Mean
1.87
2.19
2.66
3.78
1.65
1.98
2.70
3.16
SD
0.63
0.89
0.93
1.62
0.55
0.73
0.83
0.88
248
Study IV (after the Grubbs test)

Pre-training test
Hmax/Mmax ratio
Tai Chi group
Control group
Participants
SO
SC
UO
UC
SO
SC
UO
UC
0.46
0.42
0.45
0.49
0.33
0.33
0.30
0.27
0.31
0.21
0.19
0.24
0.36
0.32
0.33
0.30
0.56
0.56
0.57
0.60
0.31
0.25
0.27
0.29
0.31
0.32
0.26
0.26
0.49
0.45
0.41
0.41
0.21
0.21
0.22
0.20
0.25
0.22
0.19
0.14
0.15
0.14
0.07
0.07
0.22
0.18
0.23
0.21
0.40
0.34
0.34
0.29
0.35
0.24
0.19
0.21
0.06
0.04
0.05
0.04
0.10
0.11
0.09
0.09
0.35
0.36
0.19
0.24
10
0.24
0.21
0.21
0.28
0.57
0.63
0.61
0.53
11
0.42
0.38
0.35
0.36
0.47
0.51
0.49
0.43
12
0.10
0.10
0.09
0.20
0.24
0.18
0.24
0.22
13
0.19
0.21
0.19
0.19
0.40
0.36
0.35
0.32
14
0.13
0.10
0.07
0.10
0.19
0.22
0.18
0.21
15
0.27
0.25
0.24
0.25
16
0.36
0.29
0.37
0.22
17
0.40
0.40
0.32
0.29
18
0.09
0.12
0.09
0.10
19
0.39
0.33
0.31
0.39
20
0.51
0.47
0.40
0.36
Mean
0.29
0.26
0.24
0.25
0.33
0.32
0.30
0.30
SD
0.13
0.11
0.11
0.11
0.15
0.17
0.17
0.16
249
Pre-training test
Tai Chi group
Control group
Participants
SO
SC
UO
UC
SO
SC
UO
UC
2.96
4.10
4.93
6.35
2.32
2.91
3.24
3.69
2.43
3.24
2.99
4.44
2.05
2.53
4.12
4.48
3.88
6.31
5.47
7.84
3.84
3.46
5.27
6.83
1.81
2.36
2.61
3.77
3.06
2.62
4.48
3.77
3.76
3.18
4.22
3.63
2.47
4.30
3.96
5.50
2.61
2.47
2.84
3.19
3.15
3.36
3.69
5.38
2.91
2.65
3.60
3.61
4.89
5.33
6.53
7.91
2.07
2.88
4.04
5.53
2.60
3.20
2.86
3.68
4.57
5.81
6.60
4.60
2.45
3.57
3.66
5.68
10
2.31
2.38
2.70
3.12
3.07
4.25
3.45
5.40
11
3.66
4.36
4.81
5.18
2.40
3.17
4.25
3.46
12
1.84
2.27
2.73
3.40
4.22
4.52
5.83
6.25
13
4.21
4.35
5.61
6.84
3.39
3.92
4.74
4.67
14
3.39
3.56
4.69
5.29
3.42
4.30
4.18
5.20
15
2.38
3.31
3.90
4.72
16
2.70
2.72
4.01
4.91
18
4.64
1.88
4.70
2.01
2.58
7.35
2.86
19
3.72
3.98
4.42
6.86
20
2.72
4.08
5.32
5.69
Mean
3.02
3.54
4.11
4.96
3.10
3.68
4.30
5.14
SD
0.90
1.18
1.18
1.50
0.81
0.80
1.01
1.30
17
250
Pre-training test
Tai Chi group
Control group
Participants
SO
SC
UO
UC
SO
SC
UO
UC
1.65
2.82
2.08
4.41
1.16
1.30
2.19
2.60
1.77
1.85
2.34
2.17
1.23
1.49
2.49
3.32
2.36
3.53
4.52
6.24
2.88
2.90
4.06
5.16
1.46
2.11
1.78
2.53
2.18
2.05
2.77
3.02
2.04
1.94
2.30
2.79
1.94
2.85
2.76
3.59
1.28
1.22
1.69
2.72
2.49
2.68
2.82
4.39
1.03
1.74
2.51
2.69
2.81
3.94
3.72
5.69
1.41
1.81
2.49
3.76
0.75
0.94
1.05
1.46
2.82
4.21
4.10
5.49
1.82
2.90
3.09
4.78
10
1.62
1.24
1.85
2.37
1.08
1.51
2.62
4.16
11
2.87
3.10
3.28
3.54
1.03
1.57
1.36
2.03
12
1.13
1.18
2.22
2.45
1.90
3.13
2.74
4.92
13
3.05
4.69
4.64
7.08
1.36
1.69
3.26
3.54
14
2.18
2.65
3.42
3.90
2.10
2.19
3.22
3.40
15
1.57
2.21
2.37
2.87
16
1.25
2.26
3.07
3.67
17
1.49
1.53
4.03
4.01
18
0.97
0.94
1.36
1.89
19
1.08
1.09
2.17
3.49
20
2.16
3.01
3.10
4.90
Mean
1.76
2.26
2.77
3.65
1.77
2.22
2.72
3.72
SD
0.63
1.05
0.96
1.40
0.68
0.86
0.81
1.22
251
Post-training test
Hmax/Mmax ratio
Tai Chi group
Control group
Participants
SO
SC
UO
UC
SO
SC
UO
UC
0.55
0.59
0.53
0.57
0.52
0.57
0.49
0.45
0.35
0.36
0.39
0.41
0.31
0.25
0.24
0.26
0.28
0.23
0.16
0.20
0.52
0.45
0.43
0.44
0.26
0.28
0.22
0.24
0.38
0.22
0.29
0.33
0.57
0.53
0.50
0.50
0.40
0.39
0.33
0.36
0.35
0.35
0.22
0.17
0.30
0.36
0.29
0.25
0.33
0.30
0.33
0.29
0.40
0.36
0.35
0.32
0.65
0.63
0.54
0.58
0.05
0.05
0.05
0.06
0.15
0.14
0.12
0.13
0.46
0.34
0.32
0.33
10
0.24
0.24
0.21
0.19
0.48
0.52
0.47
0.35
11
0.45
0.40
0.36
0.36
0.22
0.16
0.15
0.13
12
0.19
0.23
0.17
0.16
0.17
0.15
0.15
0.19
13
0.20
0.21
0.23
0.24
0.29
0.27
0.34
0.34
14
0.09
0.07
0.08
0.11
0.20
0.21
0.20
0.20
15
0.44
0.50
0.40
0.43
16
0.40
0.36
0.42
0.39
17
0.32
0.34
0.33
0.29
18
0.11
0.11
0.13
0.12
19
0.45
0.49
0.35
0.36
20
0.56
0.49
0.40
0.48
Mean
0.35
0.34
0.30
0.31
0.33
0.31
0.29
0.29
SD
0.16
0.16
0.14
0.15
0.14
0.15
0.13
0.11
252
Post-training test
Tai Chi group
Control group
Participants
SO
SC
UO
UC
SO
SC
UO
UC
2.64
4.07
3.55
5.08
2.90
2.61
2.77
3.99
2.78
3.88
3.89
4.83
2.64
3.06
3.97
4.34
4.50
7.03
8.67
2.79
2.81
3.16
3.63
2.67
2.59
2.95
3.86
2.15
3.21
3.48
3.61
2.86
4.40
4.37
5.12
2.78
3.12
4.35
4.44
2.28
2.89
3.03
2.90
2.50
2.78
4.71
4.17
2.81
3.01
3.63
3.78
5.04
1.91
2.10
2.88
3.17
2.43
3.79
3.02
3.67
4.15
3.00
4.30
5.02
3.00
4.03
3.51
4.43
10
2.34
2.85
2.71
3.27
3.04
3.14
3.88
4.74
11
3.40
4.03
4.49
6.29
2.04
2.48
3.03
3.14
12
1.75
1.87
2.33
2.40
2.71
4.59
5.01
6.01
13
3.64
5.06
4.77
6.58
2.24
2.11
3.51
3.59
14
3.82
4.58
5.76
5.97
3.50
4.70
4.15
5.34
15
3.03
3.27
3.85
4.10
16
3.16
3.06
5.61
7.65
17
3.69
4.02
6.07
6.84
18
2.26
2.15
2.47
2.84
19
4.27
4.17
4.97
6.49
20
3.25
3.71
4.25
5.75
Mean
3.06
3.44
4.15
5.03
2.67
3.39
3.74
4.24
SD
0.78
0.95
1.28
1.74
0.41
0.90
0.69
0.79
253
Post-training test
Tai Chi group
Control group
Participants
SO
SC
UO
UC
SO
SC
UO
UC
1.44
1.99
1.66
3.97
1.99
1.40
2.35
3.28
2.03
2.29
2.55
3.05
2.17
1.95
2.86
2.62
3.17
4.55
7.32
1.40
1.30
1.93
2.65
1.99
1.73
2.23
2.57
1.24
2.08
2.13
2.71
1.90
3.37
2.61
4.67
1.68
1.98
2.64
3.63
1.50
1.55
1.89
2.36
1.88
2.67
3.78
3.73
1.67
1.64
2.38
2.92
2.32
2.02
3.22
3.93
1.58
1.45
1.84
2.07
0.73
0.92
1.25
1.37
1.91
2.02
2.31
3.28
2.41
3.28
3.45
3.72
10
1.63
1.97
2.41
3.52
1.16
1.66
2.49
2.89
11
2.56
3.03
3.24
3.68
1.22
1.67
2.03
2.42
12
1.21
1.37
1.82
2.25
1.56
3.04
3.28
5.13
13
3.23
3.62
3.98
6.63
0.95
1.04
2.08
2.90
14
2.67
2.88
3.26
5.24
2.39
2.72
4.31
3.27
15
1.63
1.71
2.27
2.35
16
1.60
2.12
2.84
5.05
17
1.80
2.26
4.76
6.01
18
1.09
0.90
1.36
1.28
19
0.88
1.31
2.02
2.93
20
2.40
2.03
3.32
4.37
Mean
1.87
1.98
2.66
3.78
1.65
1.98
2.70
3.16
SD
0.63
0.73
0.93
1.62
0.55
0.73
0.83
0.88
254

Effects of Ageing and Tai Chi Training On Soleus H-Reflex in Olde

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Effects of Ageing and Tai Chi Training On Soleus H-Reflex in Olde

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Southern Cross University

Effects of ageing and Tai Chi training on soleus

Master of Science in Sport Sciences

This thesis is presented in fulfilment of the requirements for the degree of

Print Name: Yung-Sheng Chen

Chen, Y. S., Crowley Z, Zhou S, Cartwright C. 2011 (submitted). Effect of 12 weeks

Thank Zac Crowley for the initial encouragement of this study.

2.4.2. H-reflex modulation during postural tasks ...................................................42

5.2.2. Experimental settings ...................................................................................91

7.1. Introduction .......................................................................................................124

Silver silver chloride

Central nervous system

Centre of pressure movement in anterior and posterior direction

Centre of pressure movement in medial and lateral direction

Excitatory postsynaptic potential

Axo-axonal gamma-aminobutyric acids

Galvanic vestibular stimulation

Maximal peak-to-peak amplitude of H-reflex

Intraclass coefficient correlation

Inhibitory postsynaptic potential

Motor evoked potentials

Maximal peak-to-peak amplitude of M-wave

Maximal voluntary contraction

Repeated measures analysis of variance

Root mean square

stable surface with eyes closed

Standard error of measurement

Surface EMG for a non-invasive assessment of muscles

Stable surface with eyes open

Sensory organization test

Tibialis anterior muscle

Transcranial electrical stimulation

Transcranial magnetic stimulation

Unstable surface with eyes open

Unstable surface with eyes closed

Degree per second

Millisecond per centimetre

During standing, electromyographic activity (EMG) of the plantarflexors corresponds to

Motor control refers to sensorimotor performance during body movements (Latash,

The amplitude of H-reflex increases proportionally with increased intensity of muscle

that the amplitude of H-reflex is inconsistent during submaximal voluntary muscle

1.2. Aims of the investigation

The specific aims of the studies are stated below.

Study I: Test-retest reliability of the SOL H-reflex

The aims of this study were:

The details of this study are described in Chapter 4.

The aim of the Study II was:

To examine the age-related differences in SOL H-reflex gain during submaximal

The details of this study are described in Chapter 5.

The aims of this study were:

To determine age-related changes in the SOL H-reflex modulation during passive

To determine the correlation of the SOL H-reflex modulations between shortening

The details of this study are described in Chapter 6.

The aims of the Study IV were:

To determine the effect of 12-weeks TC training on the SOL H-reflex modulation

To determine the effect of 12-weeks TC training on static postural control in older

The details of this study are described in Chapter 7.

1.3. Research hypotheses

The maximal amplitude of SOL H-reflex is down-regulated after 12 weeks TC

1.4. Significance of the research

1.5. Study Limitations

spinae muscle groups, etc..

The effects of TC training on H-reflex modulation and balance control in older