Acute Strength Training Increases Responses

to Stimulation of Corticospinal Axons

JAMES L. NUZZO1,2, BENJAMIN K. BARRY1,2, SIMON C. GANDEVIA1,3, and JANET L. TAYLOR1,2
1
Neuroscience Research Australia, Randwick, NSW, AUSTRALIA; 2School of Medical Sciences, University of New South
Wales, Kensington, NSW, AUSTRALIA; 3Prince of Wales Clinical School, University of New South Wales, Kensington,
NSW, AUSTRALIA

ABSTRACT
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NUZZO, J. L., B. K. BARRY, S. C. GANDEVIA, and J. L. TAYLOR. Acute Strength Training Increases Responses to Stimulation of
Corticospinal Axons. Med. Sci. Sports Exerc., Vol. 48, No. 1, pp. 139–150, 2016. Purpose: Acute strength training of forearm muscles
increases resting twitch forces from motor cortex stimulation. It is unclear if such effects are spinal in origin and if they also occur with
training of larger muscles. With the use of subcortical stimulation of corticospinal axons, the current study examined if one session of
strength training of the elbow flexor muscles leads to spinal cord changes and if the type of training is important. Methods: In experiment 1,
10 subjects completed ballistic isometric training, ballistic concentric training, and no training (control) on separate days. In exper-
iment 2, 13 subjects completed ballistic isometric training and slow-ramp isometric training. Before and after training, transcranial
magnetic stimulation over the contralateral motor cortex elicited motor-evoked potentials (MEPs) in the resting biceps brachii, and
electrical stimulation of corticospinal tract axons at the cervicomedullary junction elicited cervicomedullary motor–evoked potentials
(CMEPs). Motor-evoked potential and CMEP twitch forces were also measured. Results: In experiment 1, CMEPs and CMEP twitch
forces were significantly facilitated after ballistic isometric training compared to control. In experiment 2, MEPs, MEP twitch forces,
CMEPs, and CMEP twitch forces increased for 15 to 25 min after ballistic and slow-ramp isometric training. Conclusion: Via processes
within the spinal cord, one session of strength training of the elbow flexors increases net output from motoneurons projecting to the
trained muscles. Likely mechanisms include increased efficacy of corticospinal-motoneuronal synapses or increased motoneuron
excitability. However, the rate of force generation during training is not important for inducing these changes. A concomitant increase in
motor cortical excitability is likely. These short-term changes may represent initial neural adaptations to strength training. Key Words:
BICEPS BRACHII, CERVICOMEDULLARY-EVOKED POTENTIAL, ELBOW FLEXORS, MOTONEURON, PLASTICITY,
SPINAL CORD

T
he corticospinal tract is the main descending pathway
for voluntary control of human upper-limb muscles
(19). With motor training, plastic changes occur in
this pathway, at both cortical (3,9) and spinal (35) levels.
However, the site and type of change depends on the type of
motor training performed (1,13,16).
Strength training is a form of motor training that involves
repetitive high force and/or high rate of force development
voluntary contractions. After one session of strength training
of the forearm muscles, evoked twitch forces from trans-
cranial magnetic stimulation (TMS) increase toward the train-
ing direction and in magnitude (28). Thus, strength training
Address for correspondence: James L. Nuzzo, M.S., Neuroscience Research Australia,
PO Box 1165, Randwick, NSW, Australia 2031; E-mail: j.nuzzo@neura.edu.au.
Submitted for publication April 2015.
Accepted for publication July 2015.
Supplemental digital content is available for this article. Direct URL cita-
tions appear in the printed text and are provided in the HTML and PDF
versions of this article on the journal_s Web site (www.acsm-msse.org).
0195-9131/16/4801-0139/0
MEDICINE & SCIENCE IN SPORTS & EXERCISEÒ
Copyright Ó 2015 by the American College of Sports Medicine
DOI: 10.1249/MSS.0000000000000733
alters the net excitability of corticospinal-motoneuronal pro-
jections to the muscles involved in training (28). However,
because the volley from TMS travels through corticospinal
descending paths, a change in evoked measures after training
could be cortical and/or spinal in origin. Thus, the location of
the change remains unclear.
The H-reflex can be used to elucidate activity-dependent
plasticity of the spinal cord, as changes in this measure ac-
company the acquisition and maintenance of motor skills
(35). However, the H-reflex is not easily obtained in human
upper-limb muscles (e.g., biceps brachii), which are a com-
mon focus of strength training programs. Also, the H-reflex
does not test the motoneurons through the corticospinal
APPLIED SCIENCES
path, the main path for voluntary movement. Consequently,
the H-reflex cannot assess the corticospinal-motoneuronal
synapses, one of the likely sites of modification within the
spinal cord (32). Efficacy of these synapses and excitability
of the motoneurons can be gauged by cervicomedullary
motor–evoked potentials (CMEPs) (i.e., responses to sub-
cortical stimulation of the corticospinal axons). Cervico-
medullary motor–evoked potentials have a large monosynaptic
component in biceps brachii (23), and they travel along the
same axons as motor-evoked potentials (MEP) from stimula-
tion at the motor cortex (12). Cervicomedullary motor–evoked

139

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 potentials are facilitated or depressed after periods of paired
stimulation, designed to alter synaptic efficacy through re-
peated pairs of appropriately timed transcranial magnetic and
peripheral nerve stimuli (2,32). The increase or decrease in the
CMEP was thought to reflect changes in synaptic efficacy and
was paralleled by changes in voluntary force output and sur-
face EMG of the elbow flexor muscles (32). Thus, changes at
corticospinal-motoneuronal synapses change the output from
the motoneuron pool and muscles.
Motor training may also affect the corticospinal-
motoneuronal synapses. However, only two studies have
used subcortical stimulation of corticospinal tract axons to
assess changes at a spinal level after acute motor training
(13,20). The first study was limited to three subjects (20). The
second found increased CMEPs in the first dorsal inte-
rosseous after one session of ballistic concentric training with
the index finger, but not after low-force visuomotor training
(13). Thus, ballistic training seems to facilitate corticospinal-
motoneuronal transmission via processes within the spinal
cord. However, these spinal changes have not been docu-
mented in larger muscle groups that are commonly targeted
for strength training. Therefore, this study was designed to
determine whether one session of strength training of the el-
bow flexors leads to changes in the spinal cord and if the type
of training is important for inducing such changes.
We conducted two experiments. In experiment 1, we de-
termined if ballistic training of the elbow flexors alters re-
sponses to stimulation of the corticospinal axons or motor
cortex, and if the type of ballistic training (i.e., isometric or
concentric) is important for inducing such changes. In ex-
periment 2, we determined whether evoked corticospinal
responses are influenced by the rate at which force is de-
veloped during isometric contractions (i.e., ballistic or slow-
ramp). We hypothesized that training with contractions with
a high rate of force development would produce the greatest
increase in responses to stimulation of corticospinal axons.
METHODS
Ethical Approval
The study protocol was approved by the Human Research
Ethics Committee at the University of New South Wales
and complied with the Declaration of Helsinki (2008). All
APPLIED SCIENCES
subjects provided written informed consent to participate in
the study.
Subjects
Ten subjects (age, 23.6 T 6.2 yr; 7 men) participated in
experiment 1, and 13 subjects (age, 24.6 T 7.3 yr; 7 men)
participated in experiment 2. Three subjects who participated
in experiment 1 also participated in experiment 2. Subjects
were included if they reported no contraindications to TMS
(e.g., epilepsy and metal in the body), were not currently
taking medications that may alter synaptic plasticity (e.g.,
140 Official Journal of the American College of Sports Medicine
Copyright © 2015 by the American College of Sports Medicine. Unauthorized reproduction of this article is prohibited.
antidepressants), and if CMEPs could be elicited in the right
biceps brachii without contamination from inadvertent
stimulation of cervical motor roots.
Experimental Setup
In both experiments, the subjects sat in an adjustable chair
with the right elbow, forearm, and hand resting in a pronated
position on an arm bar (Fig. 1A). The elbow was aligned with
the fulcrum of the apparatus and secured against a padded re-
straint. The wrist and distal radius were also positioned against
a padded restraint, and a strap around the wrist secured the arm
to the bar. The fingers were positioned over a metal handle,
which the subjects grasped during the training contractions.
The apparatus was designed for contractions to be
performed in the transverse plane to eliminate fatigue from
lifting and lowering the mass of the forearm. The apparatus
was also designed to accommodate both concentric and iso-
metric contractions (see ‘‘Experimental Protocol’’ for in-
structions given to the subjects on how to perform the
contractions). For concentric contractions, the arm bar was
free to pivot about the fulcrum with only inertial resistance.
Peak accelerations during these contractions were obtained
with an accelerometer (5g, ADXL 320, Analog Devices,
Norwood, MA, USA). The accelerometer was secured to the
underside of the arm bar. In addition, impact forces of the
concentric contractions were obtained from a force transducer
(Xtran, Applied Measurement, Melbourne, Australia; sam-
pling rate: 1000 Hz). This transducer also served as the me-
chanical stop at the end of the range of motion. For experiment
1, average elbow angles at the start and end of the concentric
contractions were 135- and 67-, respectively. In pilot testing,
peak acceleration during concentric contractions occurred at
approximately 120-. For isometric contractions, the arm bar
was fixed at an elbow angle of approximately 120-. Voluntary
force, rate of force development, and evoked twitch force were
obtained from a force transducer (Transducer Techniques,
Temecula, CA, USA; sampling rate: 1000 Hz).
Electromyographic activity was recorded from the right
biceps brachii and brachioradialis via surface electrodes (Ag-
AgCl). Signals were filtered, amplified (16–1000 Hz; CED
1902 amplifier; Cambridge Electronics Design, Cambridge,
United Kingdom), and sampled at 2000 Hz. All analog
signals were digitized and stored onto a personal computer
using a laboratory interface (CED Micro1401 mkII and
Spike 2 software version 6; Cambridge Electronics Design).
Transcranial Magnetic Stimulation
Magnetic stimulation (Magstim 200, Magstim, Whitland,
United Kingdom) of the motor cortex was used to obtain
MEPs from the right biceps brachii and brachioradialis.
Procedures were consistent with the guidelines for magnetic
stimulation (e.g., the subjects completed a safety screening
questionnaire to assess eligibility; measurements were ac-
quired in a quiet environment, where the subjects were
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FIGURE 1—Setup and protocols for experiments 1 and 2. A, Contractions occurred in the transverse plane for all training, with the elbow aligned
with the fulcrum of the apparatus. For ballistic and slow-ramp isometric training, force and rate of force development were measured by a force
transducer (F ). For ballistic concentric training, peak accelerations were measured by an accelerometer (a), which was secured to the underside of the
arm bar, 10.2 cm from the axis. Impact forces were obtained from a force transducer (F ). B, In experiment 1, subjects completed three testing sessions,
which differed only in the type of training performed (ballistic isometric training, ballistic concentric training, or control). In each session, two blocks
of training (12 sets of eight contractions per block) or control (17.5 min of sitting) were completed (indicated by gray boxes). Using surface EMG, MEPs
from TMS, CMEPs from electrical stimulation at the cervicomedullary junction, and Mmax from electrical stimulation of the brachial plexus at Erb’s
point were captured in the resting biceps brachii muscle before (Pre), between (Mid), and after (Post) the two blocks. The evoked responses were
acquired in sets (indicated by black arrows), with each set consisting of five CMEPs, five MEPs, and two Mmax. Evoked twitch forces were also
obtained. C, In experiment 2, subjects completed two testing sessions, which differed in the type of training (ballistic isometric training or slow-ramp
isometric training). In each session, subjects completed a control period (block 1), followed by one block of training (block 2).

seated and fully relaxed, with eyes opened) (26). The cir-
cular coil (13.5 cm outside diameter) was positioned over
the vertex and oriented to preferentially activate the left
motor cortex. That is, anticlockwise current in the coil
resulted in a posterior-to-anterior current direction across the
left motor cortex. The optimal site for evoking biceps brachii
MEPs was found and marked on the scalp to allow accurate
placement of the coil through the experiment. The stimula-
tion intensity (range, 60%–95% of device maximum) was
that which induced the largest MEP amplitude in biceps
brachii. This resulted in MEPs of approximately 1%–2% of
the maximal compound muscle action potential (Mmax).
Twitch forces produced by the MEP were also obtained.
Cervicomedullary Junction Stimulation
Electrical stimulation (200-Ks duration, DS7AH constant
current stimulator; Digitimer, Welwyn Garden City, United
Kingdom) of the corticospinal axons at the cervicomedullary
junction was used to obtain CMEPs for the right biceps
brachii and brachioradialis. The electrodes were positioned
at the occipital grooves between the mastoid processes and
occipital bone. The cathode and anode were positioned on
the left and right sides, respectively. The stimulation inten-
sity (range, 90–185 mA) was set at that which induced a
CMEP amplitude in biceps brachii equal to approximately
15% of Mmax. Twitch forces produced by the CMEP were
also obtained.
Brachial Plexus Stimulation
Electrical stimulation (200-Ks duration, DS7AH constant
current stimulator) of the brachial plexus was delivered at
Erb’s point to obtain Mmax for the right biceps brachii and
brachioradialis. The cathode was positioned in the supracla-
vicular fossa, and the anode was positioned on the acromion.
The stimulation intensity (range, 37.5–135 mA) was set at
50% above the level necessary to produce Mmax. Twitch
forces produced by Mmax were also obtained. APPLIED SCIENCES
Experiment Protocol
Experiment 1: Ballistic isometric training, ballistic
concentric training, control. The subjects attended the
laboratory for three testing sessions, which were about a week
apart (Fig. 1B). Subjects were instructed not to participate in
strength training in the 48 h before each session. In a given
testing session, the subjects completed one of three protocols:
ballistic isometric training, ballistic concentric training, or a
session with no training (i.e., control). The subjects completed
the protocols in random order.

STRENGTH TRAINING AND CORTICOSPINAL PATH Medicine & Science in Sports & Exercised 141

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 For the ballistic isometric and concentric protocols, the
subjects performed two blocks of training. Each block
consisted of 96 voluntary contractions (12 sets of eight) of
the elbow flexor muscles. Every 4 s, a brief auditory signal
indicated when to perform each contraction. Sets were sep-
arated by a 60-s rest period.
The subjects received instruction and feedback through-
out training. For ballistic isometric and concentric training,
the subjects were instructed to produce ‘‘quick bursts’’ of
force by briefly contracting ‘‘as hard and fast as possible.’’
During ballistic isometric training, the subjects received
visual feedback of peak rate of force development. Similarly,
during ballistic concentric training, the subjects received vi-
sual feedback of peak acceleration as well as peak impact
force. To encourage the subjects to maximize performance
throughout training, target cursors were placed on the monitor.
For the control condition, the subjects were seated with the
right arm resting on the arm bar for the same time that was
required to complete the training (i.e., 17.5 min per block).
Measurements of motor pathway excitability were obtained
from sets of stimuli that occurred before, between, and after
the two blocks of training. Each set of stimuli consisted of five
CMEPs, five MEPs, and two Mmax. Sets of stimuli occurred
before block 1 (Pre); 30 s (Mid-0), 5 min (Mid-5), and 10 min
(Mid-10) after block 1; and 30 s (Post-0) and 5 (Post-5),
10 (Post-10), 15 (Post-15), 20 (Post-20), 25 (Post-25), and
30 (Post-30) min after block 2. All stimulation occurred with
the subject at rest (i.e., in a seated position with the right arm
resting on the arm bar). Electromyography was monitored to
ensure that the subjects were relaxed before the stimulation
being delivered. Individual stimuli were given 10 s apart.
Experiment 2: Ballistic isometric training, slow
isometric training. The subjects attended the laboratory
for two testing sessions, approximately a week apart (Fig. 1C).
In a given session, the subjects completed one of two protocols:
ballistic isometric training or slow-ramp isometric training.
The subjects completed the two protocols in random order.
The protocol for experiment 2 was based on the findings
from experiment 1. In the control session from experiment 1,
CMEPs were depressed and then plateaued after sitting for
17.5 min (see ‘‘Discussion’’). Additionally, in experiment 1,
the greatest facilitation of the CMEPs occurred after the first
block of ballistic isometric training. Therefore, experiment 2
was designed so that the subjects had a control period of
17.5 min of sitting, before performing one block of training
APPLIED SCIENCES
of the elbow flexors. Within this block of training, the number
of sets, contractions per set, and rest between sets were the
same as described for experiment 1.
The ballistic isometric contractions in experiment 2 were
similar to those in experiment 1, with the exception that the
subjects were required to ramp down the force over 2 s after
the initial ballistic burst. This was done in an effort to match
the force-time integrals of the ballistic isometric and the
slow-ramp isometric contractions. For slow-ramp isometric
contractions, the subjects were required, over a period of 2 s,
to gradually increase force up to a target of 75% of maximal
142 Official Journal of the American College of Sports Medicine
Copyright © 2015 by the American College of Sports Medicine. Unauthorized reproduction of this article is prohibited.
voluntary contraction (MVC). This target was used as an-
other means to match the slow-ramp and ballistic isometric
contractions, as pilot data and results from experiment 1
revealed that 75% MVC was the approximate peak force
achieved during ballistic isometric contractions. To assist
subjects in achieving the appropriate rate and amplitude of
force production, a target force was displayed on the moni-
tor. Subjects were then required to track this target force
with a line that represented their actual force. For both the
ballistic and slow-ramp isometric training, the subjects re-
ceived a 2-s auditory signal to inform them when to start and
end each contraction. Contractions were performed every
4 s. The rate of force development and shape of the EMG
profiles were purposely different between the ballistic iso-
metric and slow-ramp isometric contractions, but we attempted
to equalize them for peak force, impulse (force  time), biceps
brachii EMG root mean square (RMS) amplitude, and inte-
grated rectified biceps brachii EMG.
Cervicomedullary motor–evoked potentials, MEPs, and
Mmax were obtained in sets as described for experiment 1.
However, in experiment 2, the ‘‘Mid’’ measures occurred
after the control period of sitting, and the ‘‘Post’’ measures
occurred after one block of training. Additionally, MVCs of
the elbow flexors were performed after the Mid measures
but before the one block of training. These MVCs were used
to establish the 75% MVC target for the slow-ramp iso-
metric training.
Data Analysis
In experiment 1, peak rate of force development and peak
acceleration provided measures of performance in the bal-
listic isometric and ballistic concentric training sessions,
respectively. Additionally, RMS amplitude of biceps brachii
EMG was measured across 80 ms before the peak rate of
force development or peak acceleration. The start of this
time segment typically coincided with the beginning of bi-
ceps brachii EMG burst at the start of the contraction. For
the performance and EMG measures, each of the eight
contractions in a training set were analyzed, and an average
was computed for each set. Raw EMG data were used for
comparisons between training sets within a given training
condition, and normalized EMG data (%MVC) were used
for comparisons between training conditions.
For MEPs, CMEPs, and Mmax, the areas of the waveforms
were measured. Biceps brachii and brachioradialis MEPs
and CMEPs were then normalized to the Mmax closest in
time, to account for any peripheral changes. For each set of
stimuli, the five CMEPs were averaged, as were the five
MEPs and the two Mmax. The exception to this is the Pre
value, which represents the average of three sets of stimuli
performed before block 1. Additionally, the amplitudes of
the evoked twitch forces were measured as the difference
between mean prestimulus force (over 100 ms) and the peak
flexion force in the 200 ms after the stimulus. However, the
twitch forces were obtained incidentally. They were only
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available for the control and isometric training days per the
experimental setup on those days. Twitch forces were not
available for the concentric training day.
In experiment 2, the peak rate of force development served
as the measure of performance for the ballistic isometric
training, whereas the area difference between the target force
line and the subject_s actual force line provided the measure of
performance for the slow-ramp isometric training. Peak force
and impulse under the force–time curve were also measured.
Additionally, RMS amplitude and area under the rectified
EMG signal for biceps brachii were measured across the entire
contraction duration. The analysis of the evoked responses
was the same as for experiment 1.
Statistical Analysis
Statistical analyses were conducted with SPSS version
21 (IBM, Armonk, NY, USA). For experiment 1, one-way
repeated-measures ANOVAs were used to assess changes in
performance and EMG data in training sets 1, 12, and 24
(i.e., the sets closest in time to the evoked potential mea-
surements). A dependent t-test was used to compare the
MVC-normalized EMG data between the two training con-
ditions. Two-way repeated-measures ANOVAs (factors:
training condition and time) were used to assess changes in
the evoked potentials and twitch forces. For this analysis,
MEPs and CMEPs were normalized to Mmax and then nor-
malized to Pre, whereas Mmax and twitch forces evoked by
the MEP, CMEP, and Mmax stimuli were just normalized to
Pre. The Pre time point was not included in the analysis.
Mauchly’s test was used to assess sphericity of the repeated
measures. In cases where sphericity of the repeated mea-
sures was violated, Huynh–Feldt corrections were applied.
For post hoc analyses, a Bonferroni correction was applied
to account for multiple comparisons. Statistical significance
was set at the 0.05 alpha level.
For experiment 2, a dependent t-test was used to assess
changes in performance variables in training sets 1 and 12. A
dependent t-test was also used to compare the peak rate of
force development, peak force, impulse under the force-time
curve, and MVC-normalized EMG between the different
training conditions. Two-way repeated-measures ANOVAs
(factors: training condition and time) were used to assess
changes in the evoked potentials and twitch forces. For this
analysis, MEPs and CMEPs were normalized to Mmax,
whereas the twitch forces evoked by the MEP and CMEP
were not. Planned contrasts were used to compare values at
all time points to Pre.

APPLIED SCIENCES

FIGURE 2—Raw traces and group data (mean T SEM) of performance measures in experiment 1. A, Raw performance and biceps brachii EMG
traces for one subject during ballistic isometric training. A 55% increase in peak rate of force development (RFD) from sets 1 to 24 is indicated by the
black arrows. B, Raw performance and biceps brachii EMG traces for the same subject during ballistic concentric training. A 28% increase in peak
acceleration from sets 1 to 24 is indicated by the black arrows. In the raw acceleration traces, the vertical lines at the end represent the moment of
impact with the force transducer. C, Group data for peak RFD and biceps brachii EMG RMS amplitude during ballistic isometric training. Each
square represents the average of the eight contractions for that training set. D, Group data for peak acceleration and biceps brachii EMG RMS
amplitude during ballistic concentric training.

STRENGTH TRAINING AND CORTICOSPINAL PATH Medicine & Science in Sports & Exercised 143

Copyright © 2015 by the American College of Sports Medicine. Unauthorized reproduction of this article is prohibited.
 Data in the text are reported as mean T SD, and data in
the figures are shown as mean T SEM. For illustration, data
from experiment 2 are normalized to Pre, although statistical
analyses were performed on the data that were not normalized.
RESULTS
Experiment 1: Ballistic Isometric Training, Ballistic
Concentric Training, Control
Training performance. Figure 2A shows an individual
subject_s raw traces for rate of force development and biceps
brachii EMG during ballistic isometric training. Figure 2B
shows traces of peak acceleration and biceps brachii EMG
during ballistic concentric training.
The subjects_ performance improved during training
(Figs. 2C, 2D). During ballistic isometric training, peak rate
of force development increased (F2, 18 = 7.717; P = 0.004),
whereas biceps brachii EMG was unchanged (F2, 18 = 0.417;
P = 0.67; Fig. 2C). During ballistic concentric training, peak
acceleration increased (F2, 18 = 66.047; P G 0.001), whereas
biceps brachii EMG was unchanged (F2, 18 = 0.154; P = 0.19;
APPLIED SCIENCES
Fig. 2D). Amplitude of biceps brachii EMG did not differ
(t9 = j0.153; P = 0.882) between ballistic isometric and
concentric training.
Corticospinal excitability. Figure 3A shows an indi-
vidual subject_s traces for biceps brachii MEPs, CMEPs, and
Mmax during ballistic isometric training. Figure 3B shows
traces of MEP, CMEP, and Mmax twitch forces.
Biceps brachii MEPs and MEP twitch forces tended to
increase after ballistic isometric training (Figs. 4A, 4B). For
biceps brachii MEPs, the main effect for condition was near
significant (F2, 18 = 3.431; P = 0.055), and a main effect was
found for time (F4.312, 38.808 = 3.707; P = 0.010; Fig. 4A).
For MEP twitch forces, a main effect was found for time
(F9, 63 = 3.431; P = 0.002), along with a condition–time
interaction (F9, 63 = 2.812; P = 0.008; Fig. 4B).
Biceps brachii CMEPs and CMEP twitch forces in-
creased after ballistic isometric training (Figs. 4C, 4D).
For biceps brachii CMEPs, main effects were found for
condition (F2, 16 = 5.540; P = 0.015; ballistic isometric
training 9 control; P = 0.026) and time (F9, 72 = 2.092;
P = 0.041), along with a condition–time interaction (F18, 144 =
1.721; P = 0.042). Biceps brachii CMEPs were greater after

FIGURE 3—Raw traces of evoked potentials and twitch forces in experiment 1. A, Biceps brachii MEPs, CMEPs, and Mmax in one subject after two
blocks of ballistic isometric training. The two training blocks are signified by gray boxes. Each waveform represents the average of five potentials.
Dashed lines indicate the amplitudes of the Pre values. For this subject, MEPs and CMEPs were facilitated. For MEPs, the greatest facilitation
occurred after the second block of training at Post-0 and Post-5. For CMEPs, the greatest facilitation occurred after the first block of training at Mid-5.
Mmax was depressed by the end of testing. B, MEP, CMEP, and Mmax twitch forces in the same subject. Each waveform represents the averages of five
twitches. For MEP twitch forces, the greatest facilitation occurred at Post-0. For CMEP twitch forces, facilitation occurred immediately after the first
block of training and remained throughout testing. Mmax potential twitch forces were potentiated at Mid-0 and Post-0.

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FIGURE 4—Group data (mean T SEM) of evoked potentials and twitch forces in experiment 1. The two training blocks are signified by gray boxes. A,
Biceps brachii MEP area. B, MEP twitch force amplitude. C, Biceps brachii CMEP area. D, CMEP twitch force amplitude. E, Biceps brachii Mmax area.
F, Mmax twitch force amplitude. For each subject, MEPs and CMEPs were normalized to Mmax and then expressed as a percentage of the Pre value.
MEP and CMEP twitch forces were not normalized to Mmax twitch forces before being expressed as a percentage of the Pre value. Twitch forces
were not available from the ballistic concentric training session. An asterisk (*) indicates statistically significant difference (all P e 0.004) between
ballistic isometric training and control at that time point; a cross (†) indicates statistically significant difference (all P e 0.002) between ballistic
isometric training and ballistic concentric training.

ballistic isometric training versus control at Mid-0, Mid-5, and
Post-5 (all P e 0.004; Fig. 4C). For CMEP twitch forces, a
main effect was found for condition (F1, 7 = 22.235; P = 0.002;
ballistic isometric 9 control) and time (F9, 63 = 2.960; P =
0.005), along with a condition–time interaction (F9, 63 = 3.818;
P = 0.001). Biceps brachii CMEP twitch forces were greater
after ballistic isometric training versus control at Mid-5, Post-5,
Post-10, Post-15, Post-20, Post-25, and Post-30 (all P e 0.004;
Fig. 4D).
Biceps brachii M max and M max twitch forces decreased
after ballistic isometric training (Figs. 4E, 4F). For bi-
ceps brachii M max , main effects were found for condition
(F 2, 18 = 26.064; P G 0.001) and time (F9, 81 = 19.795;
APPLIED SCIENCES
P G 0.001), along with a condition–time interaction
(F 18, 162 = 7.717; P G 0.001). Biceps brachii M max was
smaller after ballistic isometric training versus both
control and ballistic concentric training at Mid-10, Post-
5, Post-10, Post-15, Post-20, Post-25, and Post-30 (all
P e 0.002; Fig. 4E). For M max twitch forces, a main effect
was found for time (F 9, 63 = 21.156; P G 0.001), along
with a condition–time interaction (F 9, 63 = 21.229; P G
0.001; Fig. 4F). Changes in brachioradialis MEPs,
CMEPs, and M max (see Figure, Supplemental Digital
Content 1, which illustrates brachioradialis evoked

STRENGTH TRAINING AND CORTICOSPINAL PATH Medicine & Science in Sports & Exercised 145

Copyright © 2015 by the American College of Sports Medicine. Unauthorized reproduction of this article is prohibited.
 FIGURE 5—Raw traces and group data (mean T SEM) of performance measures in experiment 2. A, Performance and biceps brachii EMG traces
(average trace for the eight contractions for that training set) for one subject during ballistic isometric training. A 72% improvement in peak RFD
from training sets 1 to 12 is indicated by the dashed circle. B, Performance and biceps brachii EMG traces for the same subject during slow-ramp
training. A 30% improvement in the ability to track the target force line from training sets 1 to 12 is indicated by the dashed circle. C, Group data for
peak RFD during ballistic isometric training. Each filled square represents the average of the eight contractions for that training set. D, Group data for
tracking performance during slow-ramp isometric training.

responses in experiment 1, http://links.lww.com/MSS/A559) Corticospinal excitability. Biceps brachii MEPs and

were generally similar to those found in biceps brachii.
Experiment 2: Ballistic Isometric Training,
Slow-Ramp Isometric Training
Training performance. Figures 5A and 5B show
an individual subject_s traces for force and biceps brachii
EMG during ballistic isometric and slow-ramp isometric
training, respectively. For the group, peak rate of force
development did not significantly change during ballistic
isometric training (t 12 = j1.509; P = 0.157; Fig. 5C).
Additionally, tracking performance did not significantly
change during slow-ramp isometric training (t 12 = 1.627;
P = 0.130; Fig. 5D). As intended, peak rate of force de-
velopment was greater (t 12 = 6.119; P G 0.001) in bal-
listic isometric (469.0 T 261.0 NImIs j1) compared to
slow-ramp isometric training (98.9 T 22.7 NImIs j1 ).
Additionally, the training protocols were similar in peak
APPLIED SCIENCES
MEP twitch forces increased after ballistic and slow-ramp
isometric training (Figs. 6A, 6B). For biceps brachii MEPs,
a main effect was found for time (F2.124, 25.485 = 5.809; P =
0.007). Planned contrasts showed that biceps brachii MEPs
were greater at Mid-0, Post-0, Post-5, Post-10, Post-20, and
Post-30 versus Pre (all P e 0.026; Fig. 6A). For MEP twitch
forces, a main effect was found for time (F2.712, 32.548 = 8.25;
P G 0.001). Motor-evoked potential twitch forces were
greater at Post-0, Post-5, Post-10, and Post-15 versus Pre (all
P e 0.050; Fig. 6B).
Biceps brachii CMEPs and CMEP twitch forces in-
creased after ballistic and slow-ramp isometric training
(Figs. 6C, 6D). For biceps brachii CMEPs, a main effect
was found for time (F3.199, 38.389 = 9.676; P G 0.001). Bi-
ceps brachii CMEPs were smaller at Mid-0 and Mid-5, and
greater at Post-5, Post-10, Post-15, and Post-25 versus Pre
(all P e 0.028; Fig. 6C). For CMEP twitch forces, a main
effect was found for time (F2.974, 35.685 = 11.088; P G 0.001).

force of each contraction (ballistic isometric: 74.8% T
9.6% MVC; slow-ramp isometric, 71.8% T 2.1% MVC;
t 12 = 1.908; P = 0.081) and area under the rectified bi-
ceps brachii EMG (ballistic isometric, 33.4% T 12.9%
MVC; slow-ramp isometric, 29.5% T 6.2% MVC; t 12 =
0.999; P = 0.338) but differed in force impulse (ballistic
isometric, 24.6 T 15.1 NImIs; slow-ramp isometric, 29.6 T
12.3 NImIs; t12 = j2.817; P = 0.016) and the RMS am-
plitude of biceps brachii EMG (ballistic isometric, 47.1% T
12.6% MVC; slow-ramp isometric, 36.3% T 8.4% MVC;
t12 = 2.583; P = 0.024).
Cervicomedullary motor–evoked potential twitch forces were
smaller at Mid-0 and Mid-5, and greater at Post-5, Post-10,
Post-15, and Post-25 versus Pre (all P e 0.014; Fig. 6D).
Biceps brachii Mmax and Mmax twitch forces decreased after
ballistic and slow-ramp isometric training (Figs. 6E, 6F).
For biceps brachii Mmax, a main effect was found for time
(F2.489, 29.866 = 18.129; P G 0.001). Biceps brachii Mmax was
smaller at Post-0, Post-5, Post-10, Pos-t15, Post-20, Post-25,
and Post-30 versus Pre (all P e 0.031; Fig. 6E). For Mmax
twitch forces, a main effect was found for time (F1.305, 15.662 =
20.542; P G 0.001). Mmax twitch forces were greater at

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FIGURE 6—Group data (mean T SEM) of evoked potentials and twitch forces in experiment 2. The control period is signified by the first gray box; the
training block is signified by the second gray box. A, Biceps brachii MEP area. B, MEP twitch force amplitude. C, Biceps brachii CMEP area. D,
CMEP twitch force amplitude. E, Biceps brachii Mmax area. F, Mmax twitch force amplitude. In the figure, for each subject, MEPs and CMEPs were
normalized to Mmax and then expressed as a percentage of the Pre value. MEP and CMEP twitch forces were not normalized to Mmax twitch force
before being expressed as a percentage of the Pre value. A number sign (#) indicates statistically significant difference (all P e 0.05) between the mean of
ballistic isometric and slow-ramp isometric training at that time point versus the mean of ballistic isometric and slow-ramp isometric training at Pre.
Note that significance is based on statistical analyses of data before normalization to Pre.

Post-0 versus Pre (P = 0.007) and were smaller at Post-10,
Post-15, Post-20, Post-25, and Post-30 versus Pre (all P e
0.014; Fig. 6F). Changes in brachioradialis MEPs, CMEPs,
and Mmax (see Figure, Supplemental Digital Content 2,
which illustrates brachioradialis evoked responses in ex-
periment 2, http://links.lww.com/MSS/A560) were gener-
ally similar to those found in biceps brachii.
DISCUSSION
The novelty of this study lies with the muscles targeted
by the training and the outcomes measured. Previous stud-
ies have assessed changes in corticospinal excitability after
one session of ballistic training, but they have involved
training of the hand muscles and/or assessments of MEPs
APPLIED SCIENCES
(5,10,13,18,20,24,25,27,28). In this study, we have demon-
strated, for the first time, that various types of high-force
and/or high rate of force development training of the elbow
flexors—with an intensity and volume of exercise consistent
with a real-world strength training session—lead to: 1) in-
creased CMEPs in two synergistic elbow flexor muscles
(biceps brachii and brachioradialis) and 2) increased CMEP
twitch forces that are specific to the training direction. These
findings suggest that acute strength training leads to in-
creased efficacy of corticospinal-motoneuronal synapses or
increased motoneuron excitability. Furthermore, we have

STRENGTH TRAINING AND CORTICOSPINAL PATH Medicine & Science in Sports & Exercised 147

Copyright © 2015 by the American College of Sports Medicine. Unauthorized reproduction of this article is prohibited.
 demonstrated that these facilitated responses occur despite a
decrease in spinal-level excitability that occurs with 20 min
of sitting. The acquisition of MEPs in the elbow flexors after
one session of strength training is also novel, and the ob-
served increase in these responses suggests that a concurrent
change in motor cortical excitability is likely.
Our main aim was to determine whether one session of
strength training of the elbow flexors changes the spinal
cord. We identified spinal level changes, as CMEPs in bi-
ceps brachii and brachioradialis increased after various
types of strength training, and these changes were accom-
panied by an increase in elbow flexion twitch forces. Cervi-
comedullary motor–evoked potentials have a large monosynaptic
component in biceps brachii (23), and corticospinal synapses
onto motoneurons lack conventional presynaptic inhibition
(15,21). Therefore, facilitation of CMEPs suggests either
improved efficacy of the corticospinal-motoneuronal syn-
apses or increased excitability of the motoneurons. For the
current findings, changes in excitability of spinal moto-
neurons cannot be ruled out. Previous studies, which have
reported increases, decreases, or no change in H-reflexes
after a single session of various types of training, have as-
cribed such findings to presynaptic mechanisms (i.e., al-
tered presynaptic inhibition or homosynaptic postactivation
depression of Ia afferents), rather than altered motoneuron
excitability (17,22,34). Such processes, which change the
level of facilitation from muscle spindle input to the mo-
toneurons, are unlikely to influence the CMEP measured in
a relaxed muscle, as in the current study.
Although definitive evidence is limited, studies using
paired repeated stimuli suggest that plastic changes can occur
at corticospinal-motoneuronal synapses (2,31,32). Facilitation
of CMEPs, thought to occur through spike-timing–dependent
plasticity, was associated with enhanced voluntary force and
EMG of the elbow flexors (32). Thus, in the current study,
improved efficacy of corticospinal-motoneuronal synapses
may explain the facilitation in CMEPs and improved perfor-
mance during training. Twitch force responses accompanying
CMEPs represent the summed twitches evoked from multiple
arm muscles and likely include contributions from elbow
extensor as well as elbow flexor muscles. Thus, the increase
in these CMEP twitches emphasizes that changes in the effi-
cacy of corticospinal input occurred preferentially for the
trained muscles either through synaptic or postsynaptic me-
chanisms in the motoneurons.
APPLIED SCIENCES
Conclusions can be made about the time course of change
of the CMEPs. Cervicomedullary motor evoked potentials
were generally unchanged immediately after training. This
is consistent with a previous study, which found that 5-s and
2-min MVCs of the elbow flexors initially depressed CMEPs
in resting biceps brachii for 1–2 min after contraction (12). In
experiments 1 and 2, CMEP and CMEP twitch forces were
then facilitated 5 min after isometric training and remained
elevated for an additional 20 min. However, CMEPs after
ballistic concentric training (experiment 1) did not demon-
strate the same magnitude or time course of facilitation. These
148 Official Journal of the American College of Sports Medicine
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differences are, in part, due to the normalization to Mmax.
Mmax decreased after isometric training but was either in-
creased or unchanged after concentric training. Nevertheless,
even without normalizing to the twitch force produced by
Mmax, the CMEP twitch force after isometric training was
significantly facilitated. This suggests that the prolonged
facilitation in CMEPs after isometric training is not simply
an artifact of normalization but an indication of a physiolog-
ical change in the spinal cord. Additionally, with ballistic
concentric training, the return of facilitated CMEPs back to
Pre levels 15 min after training is consistent with the time
course of CMEPs after ballistic concentric training with the
index finger (13). Taken together with evidence of less
change in biceps brachii EMG after 5 d of training in a con-
centric skill task compared to an isometric task (29,30), the
current study results, although not statistically significant,
suggest differences in the magnitude and time course of
changes in the motor pathway between ballistic isometric and
concentric training.
Another important observation on the time course of
change of biceps brachii CMEPs is their reduction after
control periods. In experiment 1, CMEPs on the control
day were consistently depressed from Pre to Mid (i.e., after
17.5 min of sitting), and they remained depressed over the
remaining 60 min of the experiment. To check this finding,
experiment 2 included a control period of 17.5 min of sitting
before training. Again, biceps brachii CMEPs decreased
from Pre to Mid. This finding is unlikely to result from the
subjects_ familiarity or unfamiliarity with the stimulation or
from the Pre value itself. First, sessions were completed in
random order. Second, Pre responses were obtained after
approximately 45 min of setup procedures, which included
sitting and numerous stimuli. Third, the Pre value itself is
the average of three sets of responses 5 min apart. In these
three sets, CMEPs were stable, indicating that the decrease
only occurred after 17.5 min of sitting. The mechanisms
underlying the depression in the CMEP are unknown, and it is
unclear if minimal low intensity activity (e.g., flexing the el-
bow during walking) would abolish the decrease.
In contrast to our hypothesis, that training with contrac-
tions with a high rate of force development would produce
the greatest increase in responses to stimulation of corti-
cospinal axons, we found that ballistic and slow-ramp con-
tractions produced similar results. Our hypothesis was based
on studies showing long-term synaptic potentiation after
high-frequency trains of electrical stimuli in vitro (7), and
facilitated CMEPs after ballistic, but not visuomotor, training
in humans (13). In experiment 1, we confirmed that CMEPs
are facilitated by ballistic training. However, in experiment 2,
we discovered that when ballistic and slow-ramp training are
comparable in peak force and amount of biceps brachii EMG,
they lead to a similar level and time course of facilitation in
CMEPs. Similarly, changes in the magnitude and direction of
MEP twitch forces are comparable after single sessions of
ballistic isometric and slow-ramp isometric strength training
of the forearm muscles (28). Therefore, the voluntary activity
http://www.acsm-msse.org
conveyed through corticospinal-motoneuronal synapses is
apparently not different enough during ballistic and slow-
ramp isometric contractions to alter the magnitude of change
in CMEPs of the exercised arm muscles.
Biceps brachii MEPs and MEP twitch forces increased
after isometric training. Motor-evoked potentials have also
been shown to increase in hand muscles after various bal-
listic training tasks (5,10,18,20,24,25,27). Facilitation of
MEPs was greatest immediately after training and then de-
clined toward Pre levels. However, in experiment 2, facili-
tation remained 30 min after training. This time course is
consistent with previous reports in the hand muscles (10,25).
Increased MEPs produced by cortical stimulation are often
thought to represent increased motor cortical excitability.
However, because the MEP must be influenced by spinal
factors (11,14,33), a change in the MEP cannot be inter-
preted simply as a cortical change. Here, we observed
changes in CMEPs, but it is not clear how much of the
facilitation in MEPs reflects a spinal level change. Un-
fortunately, we cannot directly compare the MEPs and
CMEPs because their peak-to-peak amplitudes were different
(MEPs, ~1%–2% of Mmax; CMEPs, ~15% of Mmax). How-
ever, the time courses of change of the MEPs and CMEPs
differed after training. For example, in both experiments,
MEPs were facilitated immediately after training, whereas
CMEPs were not. Furthermore, after ballistic concentric
training in experiment 1, facilitation of MEPs continued to
the end of testing, whereas CMEPs returned to the Pre level.
Thus, an increase in motor cortical excitability, together
with changes at a spinal level, likely contributed to facilitation
of MEPs.
Subjects improved or maintained performance throughout
the various training protocols. Presumably, this indicates
that the nature of the inputs to the motoneurons remained
consistent for a given training type. However, changes in
training performance were not directly associated with
changes in CMEPs or MEPs. The meaning of this lack of
correlation between size of evoked potentials and motor
performance is not certain. With a bout of strength training,
muscle fatigue and an acute decrease in performance is
usual, yet such training leads to longer-term improvements.
Here, our protocol attempted to minimize fatigue, but is
unlikely to have eliminated it. The twitch force elicited by
the Mmax stimulus was greater immediately after the protocol
but was then reduced. This suggests that both potentiation
and fatigue-related processes are likely to have occurred
in the muscle during training. Additionally, increases and
decreases in the size of Mmax suggest changes in the muscle
fiber action potentials. However, there is little evidence on
whether this affects neural adaptations (8). We propose that
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Research Council of Australia. James L. Nuzzo is supported by a
University of New South Wales International Postgraduate Research
Scholarship and a Neuroscience Research Australia Supplementary
Scholarship.
All authors affirm there are no competing interests, financial or
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