Eur J Appl Physiol (2002) 88: 214226

DOI 10.1007/s00421-002-0703-4

O R I GI N A L A R T IC L E

Jamie S.M. Pringle Andrew M. Jones

Maximal lactate steady state, critical power and EMG during cycling

Accepted: 19 July 2002 / Published online: 19 September 2002

Springer-Verlag 2002

Abstract We hypothesised that: (1) the maximal lactate

steady state (MLSS), critical power (CP) and electromyographic fatigue threshold (EMGFT) occur at the
same power output in cycling exercise, and (2) exercise
above the power output at MLSS (P-MLSS) results in
continued increases in oxygen uptake (V_ O2), blood lactate concentration ([La]) and integrated electromyogram
(iEMG) with time. Eight healthy subjects [mean (SD)
age 25 (3) years, body mass 72.1 (8.2) kg] performed a
series of laboratory tests for the determination of MLSS,
CP and EMGFT. The CP was determined from four
exhaustive trials of between 2 and 15 min duration. The
MLSS was determined as the highest power output at
which the increase in blood [La] was less than 1.0 mM
across the last 20 min of a series of 30-min trials. The
EMGFT was determined from four trials of 2 min duration at dierent power outputs. The surface electromyogram was recorded continuously from the vastus
lateralis muscle. The CP was signicantly higher than
the P-MLSS [242 (25) vs. 222 (23) W; P<0.05], although the two variables were strongly correlated
(r=0.95; P<0.01). The EMGFT could not be determined in 50% of the subjects. Blood [La], V_ O2 and
minute ventilation all increased signicantly with time
for exercise at power outputs above the P-MLSS. In
conclusion, the P-MLSS, and not the CP, represents the
upper limit of the heavy exercise domain in cycling.
During exercise above the P-MLSS, there is no association between changes in iEMG and increases in V_ O2
and blood [La] with time.
Keywords Fatigue threshold V_ O2 slow
component Endurance exercise
J.S.M. Pringle A.M. Jones (&)
Department of Exercise and Sport Science,
Manchester Metropolitan University, Hassall Road,
Alsager, ST7 2HL, UK
E-mail: a.m.jones@mmu.ac.uk
Tel.: +44-161-2475656
Fax: +44-161-2476375

Introduction
It is known that the tolerable duration (time limit, tlim)
of high-intensity exercise decreases hyperbolically as a
function of the power output both during exercise with
small muscle groups (Monod and Scherrer 1965) and
during whole-body exercise such as cycling (Poole et al.
1988). This relationship may be transformed into a linear relationship between the tlim and the total amount of
work performed during the task (work limit, Wlim) as
follows:
Wlim a b
tlim

where b is the critical power (CP; i.e. the power asymptote of the hyperbolic relationship) and a represents
the anaerobic work capacity (i.e. a nite quantity of
work that can be performed above the CP using energy
derived from anaerobic glycogenolyis, and phosphagen
and oxygen stores; Monod and Scherrer 1965; Moritani
et al. 1982; Poole et al. 1990). Moritani et al. (1982)
proposed that cycling at a power output below the CP
could be sustained for a long time without fatigue,
whereas cycling at a power output above CP would
result in the accumulation of blood lactate and depletion of the stored energy sources at a predictable rate
until exhaustion. The CP concept has also been applied
to other modes of exercise such as running (Hughson
et al. 1984; Smith and Jones 2001) and swimming
(Wakayoshi et al. 1993). However, it should be stressed
that there are a number of assumptions inherent in the
CP concept and it should not be applied uncritically, for
example to activities in which the relationship between
velocity and metabolic power is not constant (di
Prampero 1999).
The maximal lactate steady state (MLSS) has been
dened as the highest constant power output that can be
maintained without a progressive increase in blood lactate concentration ([La]) over time (Beneke and von
Duvillard 1996; Jones and Doust 1998). The physiological importance of the MLSS is that it denes the

215

exercise intensity above which anaerobic metabolism

makes an increasingly important contribution to the
energy demand of exercise (Antonutto and di Prampero
1995). At power outputs up to and including the MLSS,
there is a balance between the rate of lactate production
and the rate of lactate removal, whereas at power outputs above the MLSS, the rate of lactate production
exceeds the rate of lactate clearance. It can be calculated
that the net anaerobic energy yield is negligible when
there is a balance between lactate production and removal, even if blood [La] is elevated (Antonutto and di
Prampero 1995). The MLSS is determined by measuring
the blood [La] response to a series of constant-load exercise bouts of up to 30 min duration performed on
dierent days, and is identied as the highest power
output at which blood [La] increases by <1.0 mM after
between 10 and 30 min of exercise (Beneke and von
Duvillard 1996; Jones and Doust 1998). The power
output at MLSS (P-MLSS) demarcates the boundary
between the heavy exercise domain [in which oxygen
uptake (V_ O2), blood [La] and hydrogen ion concentration ([H+]) can be maintained at an elevated but steady
level] and the severe exercise domain [in which both
blood [La] and V_ O2 increase continuously, and maximum oxygen uptake (V_ O2max) may be reached unless
volitional exhaustion ensues earlier; Poole et al. 1988].
The CP has also been used to demarcate the boundary
between the heavy and severe exercise domains (Poole et
al. 1988; Vandewalle et al. 1997). However, surprisingly
few studies have tested the supposition that the P-MLSS
and the CP occur at the same power output. Indeed,
several studies have shown that exercise at the CP can
only be maintained for some 1540 min before exhaustion occurs or that power output has to be reduced for
exercise to continue (Housh et al. 1989; Jenkins and
Quigley 1990). This latter observation is clearly at odds
with the concept that the CP and the P-MLSS are
equivalent.
The highest power output that can be maintained
without an increase in the integrated electromyogram
signal (iEMG) over time has been termed the electromyogram (EMG) fatigue threshold (EMGFT; Moritani et al. 1993). This concept is based on the
observation of a linear relationship between external
power output and the rate of increase in iEMG with

Table 1 Subject characteristics.

(V_ O2max Maximal oxygen
uptake, Thla lactate threshold,
F female, M male)

time (Housh et al. 1991), and is determined by measuring the iEMG response to four short bouts of highpower exercise (Moritani et al. 1993; de Vries et al.
1982). Simultaneous increases in iEMG and pulmonary
V_ O2 during high-intensity exercise have been taken as
evidence that the V_ O2 slow component is related to
the serial recruitment of additional (type II) motor units
(Saunders et al. 2000; Shinohara and Moritani 1992).
No previous study has determined the relationship between the EMGFT, the CP and the MLSS.
The purpose of this study was to test the hypotheses
that: (1) the MLSS, CP and EMGFT occur at the same
power output in cycling exercise and (2) exercise above
the P-MLSS results in continued increases in V_ O2, blood
[La] and iEMG with time.

Methods
Subjects
Eight healthy subjects (one female) were briefed as to the benets
and risks of participation and gave their written informed consent
to participate in the study, which was approved by the Manchester
Metropolitan University Ethics Committee. The subjects were all
involved in regular exercise training and were familiar with laboratory exercise testing procedures. The subjects physical and
physiological characteristics are shown in Table 1. Subjects were
instructed to avoid strenuous exercise in the 48 h preceding a test
session and to arrive at the laboratory in a rested and fully hydrated state, and at least 3 h postprandial. For each subject, tests
took place at the same time of day (2 h).
Design of the study
Subjects rst completed an incremental exercise test to exhaustion
to determine the lactate threshold (Thla) and V_ O2max. Over the
subsequent 2 weeks, each subject visited the laboratory a further
nine times to determine the CP (four trials), P-MLSS (four trials)
and the power output at EMGFT (P-EMGFT, one trial). These
trials were presented in random order.
All cycle tests were conducted on an electrically braked cycle
ergometer (Ergoline, Jaeger, Germany), with seat and handlebar
height and angle kept constant for individual subjects. A 5-min
warm-up of pedalling at 50 W was allowed before all tests. At the
start of a test, subjects increased their pedal rate to 90 revmin1
(this pedal rate was used for all tests) and the necessary loading was
applied, at which point timing commenced. Throughout all tests,
heart rate was recorded every 5 s using a telemetric heart rate
monitor (Polar Electro Oy, Kemple, Finland).

Subject
no.

Gender

Age
(years)

Mass
(kg)

Height
(cm)

V_ O2max
(mlmin1)

% V_ O2max
at Thla

Training
background

1
2
3
4
5
6
7
8
Mean
SD

F
M
M
M
M
M
M
M

26
29
25
27
20
25
21
24
25
3

66.2
87.0
65.6
73.2
61.2
76.7
70.4
76.8
72.1
8.2

172
177
174
178
167
180
179
182
176
5

2450
3190
3141
3917
3492
3920
4872
4860
3730
844

47
67
53
57
61
56
57
45
55
7

Recreational exercise
General tness training
Recreational cyclist
Competitive duathlete
Competitive cyclist
Competitive duathlete
Competitive cyclist
Competitive runner

216
Incremental exercise test
The starting power output for the incremental test was 50 W, and
this was increased by 25 W at the end of each minute until subjects
reached volitional exhaustion and/or the pedal rate could no longer
be maintained at 90 revmin1. Strong verbal encouragement was
provided during the latter stages of the test. For each minute of
exercise, pulmonary gas exchange and minute ventilation (V_ E) were
measured, and at the end of each stage a ngertip blood sample was
taken to determine whole blood [La] (see later).
The V_ O2 at Thla was determined from plots of blood [La]
against V_ O2 as the rst clear and sustained increase in blood [La]
above the near-resting baseline concentrations. The highest V_ O2
measured in the test was accepted as the V_ O2max. The power outputs corresponding to various percentages of V_ O2max were estimated by extrapolation of the linear regression of V_ O2 versus
power output for the sub-Thla portion of the incremental test. The
power outputs corresponding to Thla and V_ O2max were determined
with account taken of the lag in V_ O2 that occurs during incremental exercise (Davis et al. 1982).
Determination of the MLSS
To determine the MLSS, subjects completed four 30-min constantload transitions at power outputs calculated to require between
100% of the V_ O2 at Thla and 50% of the dierence between the
V_ O2 at Thla and V_ O2max (50% D). The dierence in power output
between the trials used to determine the MLSS was 19 (5) W. Expired air was collected for a timed period every 5 min and ngertip
capillary blood samples were collected at the start and end of exercise and every 5 min throughout exercise. The P-MLSS was determined as the highest power output at which the increase in blood
[La] was less than 1.0 mM across the last 20 min of the 30-min trial
(Beneke and von Duvillard 1996; Jones and Doust 1998). A surface
EMG signal was recorded from the vastus lateralis muscle over the
last 48 s of each 1-min period during each trial (see later). The
electrode site was marked with reference to anatomical landmarks
and distinguishing skin markers, and in subsequent sessions the
EMG electrodes were replaced in the same position. Data collected
during the determination of MLSS were normalised relative to the
average iEMG in the 1st min for that particular test, allowing a
relative comparison between tests.

y-axis of this graph. The P-EMGFT could not be calculated for all
subjects (see results).
Measurement of pulmonary gas exchange and V_ E
Subjects wore a nose clip and breathed through a Salford lowresistance respiratory valve/mouthpiece assembly (both tted at
least 30 s before expired air collection began). The mouthpiece was
attached to a 1-m-long piece of 3.75-cm-bore Falconia tubing.
Expired air was collected into 150 l Douglas bags (Hans Rudolph,
Kansas City, Mo., USA) for a whole number of breaths over a
hand-timed period, and closing and opening of the Douglas bag
was synchronised with inspiration. Expired air was collected for at
least 45 s of each 1-min period during the incremental test and for
@60 s preceding the withdrawal of each blood sample (i.e. 45 min,
910 min, etc.) during the MLSS trials.
Expired air was analysed for percentage of oxygen and carbon
dioxide by sampling through a paramagnetic transducer and an
infrared analyser, respectively (Servomex, Crowborough, UK, series 1400). Both gas analysers were calibrated with BOC-certied
precision gases immediately before each experimental session. Gas
volume was determined by a dry gas volume meter (Harvard Apparatus, Edenbridge, UK) and was frequently calibrated and
checked for linearity with a high-precision 7-l graduated gas syringe
(Hans Rudolph). Pulmonary gas exchange variables [V_ O2, carbon
dioxide output, V_ E and the respiratory exchange ratio] were determined using standard formulae.
Measurement of blood [La]
Finger-prick blood samples were collected every minute during the
incremental test and every 5 min during the MLSS determination.
The puncture site was cleaned with an alcohol swab, dried with
tissue, and a small skin puncture approximately 2 mm in depth was
made using a disposable safety lancet. The rst drops of blood were
wiped away, and approximately 2025 ll of arterialised blood was
collected into capillary tubes containing an anticoagulant agent
(Hawksley and Sons, Lancing, UK). Whole-blood [La] was determined using an automated analyser (YSI 1500, Yellow Springs
Instruments, Ohio, USA), which was calibrated prior to the test
sessions using a 5 mM lactate standard provided by the manufacturer. The coecient of variation for blood [La] measurement was
1.3% (0.06 mM) for 20 samples in the physiological range (5 mM).

Determination of the CP
CP was determined from four exhaustive transitions at power
outputs calculated to require between 50% D and 110% V_ O2max
and always included a trial at 100% V_ O2max. These trials were
performed on separate days. Subjects were instructed to maintain
the power output for as long as possible and exercise was terminated when the pedal rate dropped below 85 revmin1 for more
than 5 s. In all cases, this drop-o was precipitous. The time to
exhaustion was recorded to the nearest second and the CP was
calculated according to the linear model of power output versus 1/
time (Fig. 1). Those conditions eliciting volitional exhaustion
within 215 min were included in the CP determination (typically
this included four trials, but in two of the subjects only three trials
were within this time criterion).
Determination of the EMGFT
The EMGFT was determined from four square-wave transitions,
each 2 min in duration and separated by at least 25 min of rest.
These transitions required power outputs between 75% D and
115% V_ O2max (equivalent to 230460 W for males and 150275 W
for the female). The surface EMG was recorded continuously from
the vastus lateralis muscle. All EMG data collection was completed
within one laboratory visit (see below). The power output and the
rate of increase of the vastus lateralis iEMG for each 2-min trial
were plotted and the P-EMGFT was dened as the intercept on the

Electromyography
Surface electrodes were applied to the skin of the right leg over the
vastus lateralis muscle. The muscle belly was palpated during a
functional isometric contraction and the site selected at the visual
midpoint of the muscle belly. The skin was shaved, lightly roughened with abrasive electroencephalogram gel and cleaned with
cotton wool dipped in mild detergent and water. Bipolar silver/
silver chloride surface electrode stickers (30 mm20 mm, BIOTAB) were placed on the selected site, with a centre-to-centre interelectrode distance of 30 mm, along a line approximately parallel
to the direction of the underlying muscle bres. Two 50 mm leads
were used to connect the electrodes to a subminiature preamplier,
which was connected directly on top of an electrocardiogram-type
press-stud electrode located on the skin as far as possible from the
EMG electrodes. The electrode wires and preamplier were further
secured to the skin using surgical tape where necessary.
The preamplier was connected to a lightweight transmitter on
a waistband worn by the subjects. This eight-channel FM MT8
radio telemetry system (MIE Medical Research, Leeds, UK)
transmitted the signal to the nearby receiver, which was connected
to an IBM PC via a 12-bit analog-to-digital converter, and the
signal was sampled at 625 Hz (Myo-dat software, MIE). The raw
EMG data were band-pass ltered, full-wave rectied and scrutinised visually for remaining movement artefacts. One subjects data
for exercise above the P-MLSS was discarded due to technical

217
Fig. 1 A Hyperbolic relationship between power output and
time to exhaustion for a single
subject (no. 2). B Critical power
(CP) was determined by linear
transformation of the data
where CP is the y-axis intercept
of the linear trend line of power
versus 1/time to exhaustion

problems. The rectied EMG was integrated with respect to time

(iEMG), with an iEMG value computed every 2 s (EMGFT test) or
10 s (MLSS trials).

Results
MLSS, CP, and EMGFT

Statistical analysis
Analysis of variance with two-tailed, paired Student t-tests where
appropriate were used to test the signicance of dierences between
the P-MLSS, CP and P-EMGFT. In the trials used for determination of P-MLSS, paired Student t-tests were used to compare the
physiological variables at 10 min with those at the end of exercise.
Pearson product moment correlation coecients were used to assess the signicance of relationships between selected variables.
Bland and Altman plots (Bland and Altman 1986) were used to
determine the bias and limits of agreement where appropriate.
Statistical signicance was accepted at 5%. Results are presented as
mean (SEM) unless stated otherwise.

Table 2 shows the power output at the MLSS, CP and

EMGFT for each subject. The CP and the P-MLSS were
signicantly dierent (P<0.05), although they were very
strongly correlated (r=0.95, P<0.01; Fig. 2; Table 2).
In addition, CP occurred at a higher percentage of the
power output at V_ O2max (P-V_ O2max) than did P-MLSS
[CP 71 (3)% vs P-MLSS 65 (3)%; P<0.05]. The EMGFT
could not be determined in half of the subjects. On average, these four individuals had a signicantly higher

218
Table 2 Power output at the
Thla, maximal lactate steady
state (MLSS), critical power
(CP), the electromyogram
fatigue threshold (EMGFT) and
at V_ O2max (P-V_ O2max) in the
subjects in whom the EMGFT
could be determined (EMGFT
group, and in those in whom it
could not (Non-EMGFT group)

a
Signicantly dierent to MLSS
(P <0.05);
*signicantly dierent to
EMGFT group (P<0.05);
**signicantly dierent to
EMGFT group (P<0.01)

Subject no.
EMGFT group

Thla (W)

1
95
2
157
3
105
4
200
Mean
139
SEM
24
Non-EMGFT group
5
161
6
169
7
198
8
150
Mean
170
SEM
10
All mean
154
All SEM
14

P-MLSS, CP and P-V_ O2max than their counterparts in

whom an EMGFT was determined (Table 2). The anaerobic work capacity [@19 (1) kJ] and the average time
to exhaustion at 100% P-V_ O2max [@215 (15) s] were not
signicantly dierent between the two subgroups. Reasons for the inability to determine the EMGFT were that:
(1) iEMG occasionally decreased over time at some
power outputs and (2) the rate of iEMG increase over
time was sometimes less at higher compared to lower
power outputs, and thus the slope of the linear regression line was negative. The method for determination of
the EMGFT is illustrated in Fig. 3.

Physiological responses to exercise relative to the MLSS

Four subjects (nos. 1, 2, 4 and 5) could not complete the
full 30 min for the trial above the P-MLSS. Thus, the
average exercise time was 21.9 (3.3) min at this exercise
power output. Over the nal 20 min of exercise below
the P-MLSS, blood [La] decreased by @0.4 mM (nonsignicant; Fig. 4). At the P-MLSS, blood [La] reached a
plateau at 3.8 (0.5) mM for the nal 20 min. Above the
P-MLSS, blood [La] increased signicantly by 1.2
(0.8) mM from 10 min to the end of exercise (P<0.05).
Figure 4 shows that below and at the P-MLSS, a
steady state in V_ O2 and V_ E was achieved. On average,
V_ O2 attained 80 (3)% V_ O2max across the nal 20 min for
exercise at the P-MLSS. This was equivalent to 56 (4)%
D and exceeded the V_ O2 values predicted from the linear
extrapolation of the power output/V_ O2 relationship
obtained during the incremental test by 225 (60) mlmin1.
During the trial above the P-MLSS, V_ O2 increased signicantly from 3126 (270) mlmin1 [85 (2)% V_ O2max] at
10 min to 3301 (274) mlmin1 [89 (2)% V_ O2max] at the
end of exercise. At the respective time points, these
values were 245 (67) and 371 (62) mlmin1 higher, respectively, than the predicted value.
Above the P-MLSS, V_ E increased signicantly from
80 (8) to 92 (7) lmin1 from 10 min to the end of exercise

MLSS (W)

CP (W)

EMGFT (W)

P-V_ O2max (W)

115
180
165
255
179
29

118
205
210
237
192
26

128
114
228
296
192
43

223
283
280
365
288
29

220
240
310
290
265*
21
222
23

219
289
341
314
291**
26
242a
25

192
43

308
355
423
433
380*
30
334
27

(P<0.05), in comparison to the non-signicant increase

from 68 (7) to 73 (7) lmin1 at P-MLSS.

The iEMG response

The large interindividual variability in iEMG responses
meant that there were no signicant changes over time at
any of the power outputs studied. Figure 5 shows that
the normalised iEMG increased by 6 (3) and 12 (7)%
from the 1st to the 30th min for exercise just below and
at the P-MLSS, respectively. Normalised iEMG increased by 3 (1)% (<P-MLSS) and 4 (5)% (at P-MLSS)
between 10 and 30 min. Above the P-MLSS, the iEMG
response was extremely variable across subjects (Fig. 5).
On average, the peak increase [11 (8)%] occurred at
10 min. After this, iEMG decreased over time, reaching
a value 3 (9)% above the initial value by the end of
exercise. This was due to the intersubject variability in
both the time to end exercise and the iEMG response
(Fig. 6). When the data were normalised by expressing
time as a percentage of the overall exercise time, the
average iEMG was remarkably constant and the increasing variability in the response can be seen in the
divergence of the iEMG scatter towards the end of exercise (Fig. 6).

Discussion
There were three main ndings to this study. Firstly,
the CP was signicantly higher than the P-MLSS, although the two variables were strongly correlated.
Secondly, the EMGFT could not be determined in four
subjects, and it was not related to CP or P-MLSS.
Thirdly, for subjects cycling at @20 W above the PMLSS, close to their CP, blood [La], V_ O2 and V_ E all
increased signicantly over time. This was not the case
for iEMG due to the large intersubject variability in the
EMG response.

219
Fig. 2 A The relationship between the power output at the
maximal lactate steady state
(P-MLSS) and the CP (n=8).
B The bias and the limits of
agreement between the two
variables using a Bland and
Altman analysis (Bland and
Altman 1986)

Dierence between the P-MLSS and the CP

The relationship between power output and the inverse
of the time to exhaustion was highly linear (r=0.986),
indicating that the calculated CP was representative of
true maximal eorts in each trial. The CP theoretically
represents the highest power output that can be maintained without fatigue (Monod and Scherrer 1965;
Moritani et al. 1981) and has been considered to be
coincident with the P-MLSS (Poole et al. 1988, 1990).
Experimental data suggesting that the CP or critical
velocity (CV) and MLSS power/velocity are coincident
has been presented for cycling (Poole et al. 1988, 1990;
Vandewalle et al. 1997), swimming (Wakayoshi et al.
1993) and local knee-extension exercise (Le Chevalier et
al. 2000), although it should be noted that the MLSS
was not determined directly in any of these studies. In
contrast, a number of studies indicate that the CP or CV

cannot be sustained beyond approximately 1030 min,

presumably due to the accumulation of muscle and
blood [La] and [H+] (Housh et al. 1989, 1991; Jenkins
and Quigley 1990; Pepper et al. 1992). In running, the
CV and velocity at MLSS (V-MLSS) were shown to be
similar by Sid-Ali et al. (1991) and Smith and Jones
(2001). However, Sid-Ali et al. (1991) did not determine
the MLSS directly, and Smith and Jones (2001) stressed
that whilst the V-MLSS [13.8 (1.1) kmh1] and CV [14.4
(1.1) kmh1] were, on average, not signicantly dierent
in a group of trained runners, large interindividual differences in the variables indicated that the terms were
not necessarily interchangeable. It is unclear why the
present data (signicant dierence between CP and PMLSS) dier from those of Smith and Jones (no signicant dierence between CV and V-MLSS), when the
experimental methods employed were similar. It is possible that dierences in exercise mode or precision in the

220

Fig. 3 Determination of the electromyogram fatigue threshold

(EMGFT) in a subject (no. 2) in whom the increase in integrated
electromyogram (iEMG) over time was relatively proportional to
the power output. The EMGFT was determined as the y-intercept
of the linear regression relating power output to the slope of the
iEMG increase with time at that power output

determination of MLSS inuenced the relationship between MLSS and CV/CP in the two studies.
The present study revealed CP to be highly correlated
with, but signicantly higher than P-MLSS. Although,
on average, the CP was only 20 W higher than the PMLSS, the physiological signicance of this dierence is
evident in the subjects responses to exercise just above

221
Fig. 4 Blood lactate concentration ([La]), oxygen uptake
(V_ O2) and minute ventilation
(V_ E) responses over time for
cycling exercise below [201
(23) W; closed circles], at [222
(23) W; open squares] and
above [241 (24) W; closed triangles] the P-MLSS. Values are
mean (SEM) (n=8). Above PMLSS, V_ O2 and V_ E increased
signicantly from 10 min to the
end of exercise

222
Fig. 5 Normalised iEMG responses over time for cycling
exercise below (closed circles),
at (open squares) and above
(closed triangles) the P-MLSS.
Values are mean (SEM). The
iEMG was normalised relative
to the average iEMG in the
1st min of exercise for each
condition. The 612% increases
in iEMG below and at P-MLSS
were not signicant. After an
initial increase, iEMG decreased towards the end of
exercise for exercise at intensities above P-MLSS

their P-MLSS. That is, when subjects exercised at a

power output >P-MLSS [241 (24) W] and close to their
CP [242 (25) W], blood [lactate], V_ E and V_ O2 increased
signicantly with time (Fig. 4) and some subjects fatigued before 30 min had elapsed. This suggests strongly
that exercise at the CP cannot be sustained without an
increasing lactic acidosis. In summary, our rst hypothesis that the highest power output at which a steady
state in blood [La] can be maintained coincides with the
CP was not supported.
It should be pointed out here that the CP concept
contains a number of implicit assumptions that may be
questionable (di Prampero 1999; Smith and Jones 2001).
For example, it is assumed that: (1) power is innite as
time to exhaustion approaches zero; (2) the anaerobic
energy stores are depleted at exhaustion; (3) the eciency of exercise is constant; (4) V_ O2 reaches the required rate instantaneously at exercise onset (see di
Prampero 1999 for discussion). Furthermore, the CP
concept should only be applied to modes of exercise
where the energy cost per unit of distance covered is
independent of velocity.
One reason for the dissociation between the P-MLSS
and the CP in the present study may be the training
status of our subjects. Compared to sedentary subjects,
trained subjects may have higher tolerance to exhaustive
exercise (and the ensuing intramuscular and systemic
acidosis and high rates of ventilation) and a greater experience of exercising to absolute (i.e. reaching mechanical failure) rather than voluntary exhaustion.
Theoretically, the point of absolute exhaustion must
occur at or later than the actual or voluntary point of
exhaustion. Therefore, the true or theoretical absolute CP can only be higher than the CP determined
from trials to voluntary exhaustion. It should be pointed
out that the mean dierence between CP and P-MLSS in
this study (20 W) was similar to the precision with which
P-MLSS was determined. Therefore, there might be no
dierence between CP and P-MLSS if (but only if) the PMLSS was systematically underestimated in our study.

On the other hand, however, the criterion for MLSS

determination we used (<1 mM change in blood [La]
from 10 to 30 min) is relatively liberal and may lead to
an overestimation of the true MLSS. The more
stringent criteria applied by some investigators (i.e. dening MLSS as the power output at which the slope of
the linear regression through the [La] values is equal to
zero) would have served to lower the P-MLSS in the
present study, thus further increasing the dierence between P-MLSS and CP. The duration of the exhaustive
trials used to determine CP can also inuence the calculated CP and, therefore, the relationship between CP
and P-MLSS. There is some evidence that longer trials
to exhaustion tend to lower the CP (Jenkins et al. 1998;
Vandewalle et al. 1997), and in our study this would
have reduced the dierence between CP and P-MLSS.
However, it has been argued that longer trials can be
aected by factors such as motivation, and we therefore
followed the recommendations of Hill (1993) that the
trials to exhaustion used in the determination of CP
should be in the range of @215 min.
Theoretically, the CP should be independent of the
mathematical model used to express the hyperbolic relationship between power output and the time limit for
which it can be sustained. In the present study, CP was
calculated using the linear model of power output versus
1/time. It has been suggested that two-component
mathematical models overestimate CP (Bull et al. 2000)
and that a three-component mathematical model would
be more appropriate (Morton 1996). Furthermore, the
power output versus 1/time model may yield a higher CP
than the linear model of work done (power time)
versus time (Gaesser et al. 1995). In the present study,
however, the CP calculated from the latter model was
almost identical to that calculated from the power output versus 1/time model [238 (26) vs 242 (25) W]. Furthermore, the bias and limits of agreement for
comparisons between the CP and P-MLSS using either
model of CP determination [work limit model: 16
(56) W; time limit model: 20 (46) W] suggest that the CP

223
Fig. 6 Normalised iEMG
responses over time for cycling
exercise below (lower panel), at
(middle panel) and above
(upper panel) the P-MLSS. The
iEMG was normalised relative
to the average iEMG in the
1st min of exercise for each
condition, and time was
normalised relative to the end
exercise time. The 612%
increases in iEMG below and at
P-MLSS were not signicant

224

and P-MLSS are dierent and should not be used interchangeably.

Physiological responses at and above the P-MLSS
For exercise below and at the P-MLSS, a V_ O2 steady
state was attained by 10 min of exercise, which was elevated by 154 to 225 mlmin1 above the value predicted
from the linear extrapolation of the V_ O2/power output
relationship from the incremental exercise test. Above
the P-MLSS, V_ O2 was elevated above the expected value
by 181 (67) mlmin1 at 5 min and 245 (67) mlmin1 at
10 min, and increased by another 126 (32) mlmin1 to
the end of exercise. Therefore, for cycling exercise, it
appears that the maximal steady states for blood [La]
and V_ O2 occur at approximately the same power output.
However, whilst the P-MLSS appears to represent the
upper limit of the heavy exercise domain, it should be
pointed out that V_ O2max was not attained in those
subjects reaching volitional exhaustion for exercise
above the P-MLSS.
Simultaneous measurement of pulmonary and leg
V_ O2 demonstrated that @86% of the slow rise in V_ O2
between 3 and 21 min of constant-power heavy cycling
exercise originates from within the exercising limb
(Poole et al. 1991). The oxygen cost of systemic support
processes such as ventilatory and cardiac work, gluconeogenesis of lactate, extra postural muscle activity
and elevations of plasma catecholamines and body
temperature, therefore makes a relatively small contribution to the development of the V_ O2 slow component
(Casaburi et al. 1987; Poole et al. 1991). In the present
study, it was calculated that the oxygen cost of V_ E could
account for @30% of the DV_ O2 from 5 min to the end of
exercise. In contrast to previous studies (e.g. Poole et al.
1988), DV_ O2 was negatively related to Dblood [La] in the
>P-MLSS trial. The lack of a signicant positive relationship between DV_ O2 and Dblood [La] has been reported previously (e.g. Carter et al. 2000). The similarity
in the time course of the two variables noted in earlier
studies may have been a feature of the use of shorter
exercise bouts.
EMG responses to exercise
The highest power output that can be maintained
without an increase in iEMG over time has been dened
as the EMGFT (P-EMGFT; Moritani et al. 1993). Le
Chevalier et al. (2000) reported that the CP in local knee
extension exercise was signicantly correlated (r=0.96
0.98) and not signicantly dierent (<3%) from the
power output associated with a steady state in V_ O2,
blood [La] and iEMG (predicted from the change in
these variables during constant-load, supra-CP trials).
However, de Vries et al. (1982) reported that the PEMGFT was @12% higher than the CP (191 vs 170 W),
although they were highly correlated (r=0.87). In the

present study, the EMGFT could only be determined in

the less-t half of the subject group. For these four
subjects, the similar power output at CP and EMGFT
was entirely coincidental and there was no consistent
trend to suggest that the EMGFT was related to the
other physiological variables investigated. The power
outputs used in the four, 2-min trials were adjusted to
the tness of the subject, as suggested by de Vries et al.
(1982) and Moritani et al. (1993), and in all cases were
between the CP and the power output at 115% V_ O2max.
However, in the well-trained subjects exercising at high
power outputs, the linear trend line tted to the iEMG
versus time response over the 2-min bouts was either
negative or did not describe the response well. Furthermore, the proportional linear relationship between
external power output and the rate of increase in iEMG
reported previously (Moritani et al. 1982; de Vries et al.
1982) could not be reproduced.
It has been proposed that the EMGFT is more closely
associated with the steady state of lactate metabolism in
the active muscle (i.e. the MLSS) than with the lactate or
ventilatory threshold, given that it occurred at a V_ O2
just above the ventilatory threshold (Shinohara and
Moritani 1992; de Vries et al. 1982). However, the present study provides little support for this hypothesis.
Whilst V_ O2 and blood [La] did not increase during exercise below and at the P-MLSS, iEMG showed a nonsignicant increase with time. Also, during exercise
above the P-MLSS, where V_ O2, blood [La] and V_ E increased signicantly over time, there was generally little
change in iEMG when the response was normalised to
time to exhaustion.
It is thought that the increase in iEMG represents the
recruitment of previously inactive motor units and/or
increased ring rate (rate coding) of the activated motor
units to compensate for a decrease in contractility of
impaired or fatigued motor units (Edwards and Lippold
1956). If additional motor units are recruited, the size
principle dictates that they will be larger motor units of a
higher threshold (Beelen et al. 1993; Vollestad and Blom
1985). A concurrent slow rise in pulmonary V_ O2 and
increases in iEMG from the exercising muscles during
heavy and severe exercise has been taken as evidence
that the serial recruitment of the less-ecient type II
motor units is related to the V_ O2 slow component
(Barstow et al. 1996; Saunders et al. 2000; Shinohara
and Moritani 1992). However, Scheuermann et al.
(2001) were unable to detect signicant changes in EMG
during heavy cycle exercise that elicited a V_ O2 slow
component, and Takaishi et al. (1996) reported an increase in iEMG without an increase in V_ O2 in some
subjects.
The nding that iEMG occasionally decreased over
time in both the 2- and 30-min exercise bouts, especially
in the tter subjects, suggests that a degree of mechanical
failure occurred in the most powerful bres of the vastus
lateralis muscle, masking any increases in recruitment
or rate coding or synchronisation of bres lower in the
recruitment hierarchy.

225

It should be acknowledged that our interpretation of

the neuromuscular response to heavy and severe exercise
is based solely on our measurement of EMG activity in
the vastus lateralis. Whilst the vastus lateralis probably
produces a large proportion of the propulsive forces
generated in cycling (Broker and Gregor 1994), Housh
et al. (1995) reported that for some subjects the PEMGFT was lower in the rectus femoris muscle compared to the vastus lateralis muscle. Thus, it is possible
that other muscles, including the gluteus maximus,
contribute proportionally more to the slow rises in V_ O2
and blood [La] during exercise above the P-MLSS. The
calculation of EMGFT in the vastus lateralis did not
accurately predict the cardiorespiratory or iEMG responses to longer-term exercise. The usefulness of EMG
variables as indices of fatigue is disputed and the present
study indicates that EMG activity and cardiorespiratory
responses are not related in any simple way. This may be
particularly true in cycling in which the same power
output can be generated using dierent combinations of
the muscle groups around the hip, knee and ankle, and
in which the contributions from the left and right legs
can uctuate. This might also explain the dierence
between our results and those of Le Chevalier et al.
(2000), who studied EMG responses during single kneeextension exercise.
In conclusion, the CP was signicantly greater than the
P-MLSS. The P-MLSS appears to represent the upper
limit of the heavy exercise domain in cycling. At a power
output @6% above the P-MLSS, blood [La] and V_ O2 increased signicantly with time. The EMGFT could only be
determined in four of the eight subjects. The large intersubject variability in the iEMG responses recorded during
the determination of both P-EMGFT and P-MLSS suggests that surface EMG cannot be used to distinguish
between increased recruitment, rate coding and mechanical failure occurring in the active muscles.

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