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Abstract
The aim of this study was to test the hypothesis that prior heavy exercise results in a higher oxygen cost during a subsequent bout of
moderate exercise due to changes in muscle activity. Eight male subjects (25 2 yr, SE) performed moderatemoderate and moderate
heavymoderate transitions in work rate (cycling intensity, moderate = 90% LT, heavy = 80% VO2 peak). The second bout of moderate
exercise was performed after 6 min (C) or 30 s (D) of recovery. Pulmonary gas exchange was measured breath-by-breath and surface
electromyography was obtained from the vastus lateralis and medialis muscles. Root mean square (RMS) and median power frequency
_ 2 =DWR (C: +2.0 0.8 ml min1 W1, D: +3.4 0.8 ml min1 W1;
(MDPF) were computed. Prior heavy exercise increased DVO
P < 0.05) and decreased exercise eciency (C: 13.3 5.6%, D: 22.2 4.9%; P < 0.05) during the second bout of moderate exercise
in the absence of changes in RMS. MDPF was slightly elevated (2%) during the second bout of moderate exercise, but MDPF was not
_ 2 (r = 0.17). These ndings suggest that the increased oxygen cost during moderate exercise following heavy exercise is
correlated with VO
not due to increased muscle activity as assessed by surface electromyography.
2006 Elsevier Ltd. All rights reserved.
Keywords: Electromyography; Prior heavy exercise; Constant work rate exercise; Oxygen cost
1. Introduction
During the adjustment to an abrupt increase in exercise
_ 2 increases, after
intensity, pulmonary oxygen uptake (VO
a short delay, towards a new steady-state if the exercise is
of moderate intensity (i.e. below the lactate threshold, LT).
Results from a number of studies (Barstow and Mole,
1991; Barstow et al., 1993) suggest that the characteristics
_ 2 work rate relationship during moderate exercise
of the VO
can be described as a linear dynamic system, that is with an
invariant time constant and a proportional change in ampli-
*
Corresponding author. Present address: Cardiopulmonary and Metabolism Research Laboratory, Department of Kinesiology, MS 119, Health
and Human Services Building, The University of Toledo, Toledo, OH
43606-3390, USA. Tel.: +1 419 530 2741; fax: +1 419 530 4759.
E-mail address: barry.scheuermann@utoledo.edu (B.W. Scheuermann).
1050-6411/$ - see front matter 2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jelekin.2006.09.002
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J.U. Gonzales, B.W. Scheuermann / Journal of Electromyography and Kinesiology 18 (2008) 99107
2. Methods
2.1. Subjects
J.U. Gonzales, B.W. Scheuermann / Journal of Electromyography and Kinesiology 18 (2008) 99107
volume (3.0 l). Prior to each exercise session, the O2 and CO2
analyzers were calibrated using gases of known concentrations.
Corrections for ambient temperature and water vapor were made
for conditions measured near the mouth.
2.4. Measurement of surface electromyography (sEMG)
During each of the protocols, surface electromyography
(sEMG) was obtained from the vastus lateralis and vastus medialis muscle groups using a commercially available data acquisition
system (PowerLab 8SP, ADInstruments, Grand Junction, CO).
The analog sEMG signal was sampled at a rate of 2000 Hz,
amplied (common mode rejection ratio: 96 dB, input impedance:
1 MX, gain: 5000; Model 408 Dual Bio Amplier-Stimulator,
ADInstruments, Grand Junction, CO), passed through a frequency window of 3-3000 Hz, digitized by a 12-bit A/D converter,
and stored on a computer for later analysis. The raw sEMG signal
was sampled using bipolar (2 9 mm discs, 15 mm diameter
sample area) AgAgCL surface electrodes (DDN-30 Norotrode,
Myotronics-Noromed, Inc., Tukwila, WA) with a xed interelectrode spacing of 30 mm placed on the right leg. The sEMG
electrodes were positioned over the distal half of the muscle belly
aligned longitudinally to the muscle bers. A reference electrode
was placed over the tibial tuberosity or over the head of the bula.
Electrode sites were shaved and cleaned with alcohol prior to
electrode placement in order to reduce inter-electrode resistance
(<10 kX). All wiring attached to the electrodes was securely fastened to prevent motion artifact. The sEMG signal was checked
for motion artifact by moving and tapping the area surrounding
the electrode. The site was cleaned again and a new electrode
applied if motion artifact was detected in the signal.
2.5. Measurement of plasma lactate
Prior to testing, subjects rested in a supine position while a
percutaneous Teon catheter (22 gauge, Insyte I.V. Catheters,
Becton Dickinson, Inc.) was placed into a dorsal hand vein. The
blood sample was arterialized by heating the forearm and hand
throughout the exercise protocol by use of a heating lamp.
Samples were obtained at rest and at 2 min intervals during
exercise and recovery in each protocol. Samples were placed in an
ice-water slurry and analyzed for plasma lactate concentration
([Lac]) within 510 min (Stat Prole M Blood Gas and Electrolyte
Analyzer, Nova Biomedical, Inc., Waltham, MA).
101
3. Results
3.1. Subjects
On average, subjects weighed 72.1 4.2 kg, were 178.1
2.4 cm tall, and had a body mass index of 20.2 1.2
_ 2;peak meakg m2. The group mean aerobic capacity (VO
sured during the preliminary ramp exercise test was 45.1
3.2 ml kg1 min1 and the estimated LT was 52.9 2.4%
_ 2;peak , corresponding to a VO
_ 2 of 1700 14 ml
of the VO
min1. The mean work rate for moderate and heavy exercise
was 88 10 W and 214 20 W, respectively.
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J.U. Gonzales, B.W. Scheuermann / Journal of Electromyography and Kinesiology 18 (2008) 99107
Table 1
_ 2 , DE, and sEMG between the rst and second bouts of moderate exercise within each protocol
Comparison of steady-state VO
Protocol A
_ 2 (ml min1)
VO
_ 2 (ml min1)
DVO
_ 2 =DWR (ml min1 W1)
DVO
DE (%)
RMS (lV s1)
DRMS/DWR (%W)
_ 2 (lV l1 min1)
DRMS=DVO
MDPF (Hz)
Protocol B
Protocol C
Protocol D
First
Second
First
Second
First
Second
First
Second
1411 126
588 95
9.7 0.4
29.8 1.3
0.11 0.02
1.7 1.9
0.09 0.01
69.1 2.9
1438 135
616 102
10.1 0.4
28.7 1.1
0.11 0.02
1.8 0.3
0.09 0.02
71.2 3.3*
1413 133
635 107
10.3 0.4
28.0 1.0
0.10 0.02
2.0 0.3
0.09 0.01
68.2 1.8
1422 139
645 109
10.4 0.7
28.9 2.9
0.11 0.01
2.0 0.4
0.10 0.02
70.0 2.0*
1420 132
633 112
10.3 0.4
28.4 1.3
0.10 0.01
1.8 0.2
0.09 0.02
70.3 2.5
1500 115*
714 89*
12.2 0.7*
24.4 1.5*
0.11 0.01
2.1 0.3
0.09 0.02
71.9 2.3*
1418 127
651 115
10.7 0.4
27.2 1.1
0.11 0.02
2.0 0.2
0.10 0.02
72.2 3.5
1584 124*,#
818 103*,#
14.1 0.8*,#
21.0 1.4*
0.11 0.01
2.2 0.3
0.08 0.02
72.0 3.3
_ 2
exercise (P < 0.05). Interestingly, the higher absolute VO
during the second bout of moderate exercise in the moderateheavymoderate transitions was negatively correlated
with aerobic capacity (i.e. tness level) such that individu_ 2;peak exhibited the largest increase in
als with the lower VO
_ 2 following heavy exercise (r = 0.54, F = 5.87,
VO
P < 0.05).
3.3. Vastus muscle sEMG activity
The average RMS and MDPF response from the vastus
lateralis and vastus medialis muscles during moderate constant work rate cycling exercise are presented in Table 1.
No dierence in RMS or the percent change in RMS from
20 W cycling was observed between bouts of moderate
exercise within any of the four constant work rate exercise
protocols despite changes in the O2 cost of exercise (Fig. 1).
In contrast, MDPF was increased by 23% during the second bout of moderate exercise in the moderatemoderate
transitions (Protocols A and B), and also during the moderateheavymoderate transition with the 6 min recovery
(Protocol C). However, MDPF remained unchanged
between the rst and second bouts of moderate exercise
when the second bout of moderate exercise was preceded
by 30 s of recovery from heavy exercise (Protocol D; see
Table 1). Interestingly, MDPF expressed as the percent
change from 20 W cycling was found to be 46.8 8.7%
lower than the average response during the rst minute
of the second bout of moderate exercise when preceded
by 30 s of recovery from heavy exercise (Protocol D,
Fig. 1). The change in MDPF was not correlated with the
_ 2 (r = 0.17, F = 0.80, P = 0.38).
increase in VO
The ratio of RMS to cycling work rate (DRMS/DWR)
showed a constant level during all moderate exercise bouts
indicating a coupling between motor unit recruitment and
cycling work rate (Table 1). The relationship between
_ 2 was examined by the DRMS=DVO
_ 2 ratio
RMS and VO
which was increased during the rst minute of moderate
exercise when preceded by 6 min of recovery from either
moderate or heavy exercise, but reduced when preceded
_ 2 (i.e. metabolic rate) was eleby 30 s of recovery when VO
Fig. 1. Changes in vastus muscle activity during the rst and last minute
of the second bout of moderate exercise. Upper panel: The increase in
RMS from 20 W cycling (DRMS) reached a stable level of motor unit
recruitment at exercise onset that did not vary or change with prior heavy
exercise. Lower panel: The increase in MDPF from 20 W cycling (DMDPF)
was similar between the rst and last minute of moderate exercise during
the moderatemoderate transitions, but varied from the average response
during the moderateheavymoderate transitions. Dotted lines represent
the average steady-state value reached during the rst bout of moderate
exercise in the four protocols combined. (*) Signicant dierence between
rst and last minute of moderate exercise within each protocol (P < 0.05).
J.U. Gonzales, B.W. Scheuermann / Journal of Electromyography and Kinesiology 18 (2008) 99107
103
_ 2 (DRMS=DVO
_ 2 ) was increased at the
Fig. 2. The ratio of RMS to VO
onset of exercise when 6 min of recovery was present before the second
_ 2 showed a reduction under
bout of moderate exercise. DRMS=DVO
conditions of high metabolic rate at exercise onset. During the last minute
_ 2 was similar to the average steady-state value
of exercise, DRMS=DVO
calculated during the rst bout of moderate exercise shown as the dotted
line. (*) Signicant dierence between rst and last minute of moderate
exercise within each protocol (P < 0.05).
vated (Fig. 2). During the last minute of the second bout of
_ 2 ratio returned to the
moderate exercise, DRMS=DVO
average steady-state ratio measured during the rst bout
of moderate exercise in all the exercise protocols irrespective of the prior exercise conditions.
3.4. Gain and exercise eciency
_ 2 =DWR for moderate exercise paralThe gain or DVO
_ 2 response and is presented in Table 1. Moderleled the VO
atemoderate exercise transitions (Protocol A and B) did
_ 2 =DWR which remained
not lead to a dierence in DVO
at 10 ml min1 W1 in spite of the dierent recovery metabolic rates prior to the onset of the second bout of moderate exercise. In contrast, prior heavy exercise increased
_ 2 =DWR to 12.2 0.7 ml min1 W1 and 14.1 0.8
DVO
ml min1 W1 during the second bout of moderate exercise
as compared to the rst bout of moderate exercise when
preceded by 6 min and 30 s of recovery from heavy exer_ 2 =DWR
cise, respectively (P < 0.05). The increase in DVO
during the second bout of moderate exercise was greater
after 30 s of recovery (i.e. high metabolic rate) as compared
to 6 min of recovery from heavy exercise (Protocol
D > Protocol C, P < 0.05; Fig. 3).
Net eciency or DE calculated during steady-state
moderate exercise is presented in Table 1. During the moderatemoderate exercise transitions, DE was not dierent
between the rst and second bout of moderate exercise.
However, prior heavy exercise resulted in a decrease in DE
by 13.3 5.6% and 22.2 4.9% during the second bout
of moderate exercise as compared to the average steadystate response for Protocols C and D, respectively
(Fig. 3, P < 0.05).
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4. Discussion
Prior heavy exercise has consistently been shown to
_ 2 and the O2 cost (as both
increase both absolute VO
_ 2 =DWR and DE) during a subsequent bout of exercise
DVO
for both moderate (Sahlin et al., 2005) and heavy constant
work rate exercise (Burnley et al., 2001, 2002; Scheuermann
et al., 2001). The contribution of motor unit recruitment to
_ 2 remains uncertain and has received relathe elevated VO
tively little attention during the steady-state of moderate
intensity exercise. Consistent with the recent study by Sahlin et al. (2005), the present study found prior heavy exercise
to decrease exercise eciency during a subsequent bout of
moderate exercise. We further showed heavy exercise to
increase the gain during a subsequent bout of moderate
exercise. However, in contrast to our hypothesis, the
increased O2 cost during moderate exercise was not associated with either the recruitment of additional motor units,
since RMS remained unchanged, or the recruitment of less
ecient type II motor units, since the change in MDPF was
_ 2 . These ndings suggest that
not related to changes in VO
the mechanism(s) causing the increased O2 cost during prior
heavy exercise conditions is not solely related to alterations
in motor unit recruitment patterns as assessed by sEMG.
4.1. sEMG and moderate constant work rate exercise
The observation of a rapid increase of RMS to a constant
level during moderate constant work rate exercise is not a
new nding and has been demonstrated to occur in both
untrained and trained subjects during cycling exercise (Jammes et al., 1998). Petrofsky (1979) has further shown RMS of
the quadriceps to remain unchanged during prolonged
(80 min) moderate exercise at cycling intensities of 2040%
_ 2 max and for at least 20 min during moderate exercise at
VO
_ 2 max . It is at high intensities (>60% VO2max) that
60% VO
RMS has been shown to increase with time during moderate
J.U. Gonzales, B.W. Scheuermann / Journal of Electromyography and Kinesiology 18 (2008) 99107
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J.U. Gonzales, B.W. Scheuermann / Journal of Electromyography and Kinesiology 18 (2008) 99107
J.U. Gonzales, B.W. Scheuermann / Journal of Electromyography and Kinesiology 18 (2008) 99107
Joaquin U. Gonzales is a Ph.D. candidate in the
Department of Kinesiology at The University of
Toledo. He received his BS in Kinesiology
(2000) at the University of Texas of the Permian Basin in Odessa, TX and his MS in Exercise
Physiology at Texas Tech University in Lubbock, TX. His current research interests include
examining the eect of dierent blood ow
patterns during exercise on endothelial function
in humans.
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