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 Archives of Physiology and Biochemistry
2003, Vol. 111, No. 3, pp. 202–210
The Effects of Interval Training on Oxygen Pulse and
Performance in Supra-threshold Runs
Archives of Physiology and Biochemistry Downloaded from informahealthcare.com by University of Notre Dame Australia on 07/02/13
L.P. Laffite1, L. Mille-Hamard2, J.P. Koralsztein1 and V.L. Billat1,2
1
Centre de médecine du sport CCAS, Paris, France; 2Université d’Evry, Val d’Essonne, France
Abstract
The aim of this study was to examine (i) the effects of a
severe interval training
. period on oxygen pulse kinetics (O2-
p, the ratio between VO2 and heart rate), and (ii) to study the
consequences of these effects on the variation of perfor-
mance (time to exhaustion) during severe runs. Seven ath-
letes were tested before and after an eight-weeks period of
a specific intermittent training at vD50, i.e., the intermediate
velocity between the . lactate threshold
. (vLT) and the veloc-
For personal use only.
ity associated with VO2max (vVO2max). During the test ses-
sions, athletes performed an incremental test and an all-out
test at the pretraining vD50. After the training period they
also completed an additional all-out test at the posttraining
vD50 (vD50bis). Results showed that after training there was
i) an increase in the O2-p maximal value during the incre-
mental test (22.7 ± 1.5 mlO2.b-1 vs. 20.6 ± 1.5 mlO2.b-1; p <
0.04), ii) a decrease in the time to reach the O2-p steady state
(TRO2-p) at the same absolute vD50 (33 ± 7 s vs. 60 ± 27 s; p
< 0.04) and iii) an increase in the O2-p steady state duration
(TSSO2-p) at the same absolute vD50 (552 ± 201 s vs. 407 ±
106 s; p < 0.04). However, there was no relationship between
the improvement of these two O2-p kinetics parameters
(TRO2-p and TSSO2-p) and those of the performance. This study
found that after an individualised interval-training program
conducted at the same absolute velocity, the O2-p kinetics
reached a steady state quicker and for a longer duration than
before training. This is however not related with the improve-
ment of performance.
Keywords: Oxygen pulse, oxygen uptake, training, running.
Introduction
Several studies showed that improved performance after a
training period is not necessarily accompanied by a signifi-
Accepted: 29 May, 2003
Address correspondence to: Dr Laurent Laffite, Centre de Médecine du Sport, CCAS, 2 Avenue Richerand, 75010 Paris, France. Fax: +33
1 42 392083; E-mail: laurent.la@free.fr
1381-3455/03/11103-202$16.00
© Swets & Zeitlinger
.
cant modification of the maximal oxygen uptake (VO2max) or
the lactate threshold (vLT) (Flynn et al., 1994; Billat et al.,
1999).
. Thus, in addition to the monitoring of heart rate (HR)
or VO2, the analysis of a third cardio-vascular parameter may
deepen understanding of the relationship between modifica-
tion of these parameters and the improvement of the perfor-
mance. It is well .know that the stroke volume (SV) is the
major function of VO2max and then of the performance in exer-
cise lasting more than two minutes (Billat et al., 2001a for
marathon; di Prampero et al., 1993, for middle distance). Fur-
thermore, it has been reported. that in well endurance-trained
subjects, SV increase until vVO2max in an incremental exer-
cise (Billat et al., 2001b; Gledhill et al., 1994; Zhou et al.,
2001) and is increased by high intensity training (Billat et al.,
2001a; Wilmore et al., 2001). However, the measurement of
SV at high intensity cannot be done on the track and field,
where athletes are currently tested thanks to advent of the
portable
. gas exchange analyser. According to .the Fick equa-
tion, VO2 is the product of the cardiac output (Q) and the arte-
riovenous O2 difference ((a-v̄O
. 2)D). Thus, the oxygen pulse
(O2-p, the ratio between VO2 and HR) estimates the . product
of the stroke volume (SV) and (a-v̄O2)D. HR and VO2 are the
two most commonly cardiopulmonary parameters measured
easily and reliably on field and laboratory (Léger et
Thivierge, 1988). However, several authors have studied O2-
p and the theoretical relationship with SV and (a-v̄O2)D.
They suggested (Mahler et al., 1985; Sagiv, 1991; Whipp et
al., 1996) or showed (Bhambhani et al., 1994; Bhambhani,
1995) an important correlation between the evolution of SV
and O2-p, and other (Hossack et al., 1980) found a strong
relationship between (a-v̄O2)D and O2-p during maximal
treadmill exercise. The theoretical relationship between O2-
p, SV and (a-v̄O2)D was therefore confirmed and seems to
be reliable. Hence, the interest to study the O2-p kinetics is
 The Effects of Interval Training on Oxygen Pulse
to appraise the evolution of the product
. of SV and (a-v̄O2)D.
Consequently, using simply the VO2 and HR measurement,
O2-p allows to get a third information
. on the cardiopul-
monary response in addition to VO2 and HR. On the other . given submaximal running velocity, Cavanagh & Williams,
hand, different studies showed that the parameters of VO2
kinetics may be modified after a training period (Hagberg et
al., 1980; Casaburi et al., 1987; Yoshida et al., 1992), and that
some of these modifications are correlated to the perfor-
mance. We hypothesize also that the O2-p kinetics during a
Archives of Physiology and Biochemistry Downloaded from informahealthcare.com by University of Notre Dame Australia on 07/02/13
severe intensity exercise is modified after a training session,
and that these modifications are correlated with the
performance.
Therefore, the aim of this study was (i) to examine the
effect of an eight-weeks severe endurance interval training
program on the O2-p kinetics during running; and (ii) to
examine the consequences of these modifications on the per-
formance during runs of severe intensity.
Materials and methods
Subjects
Seven endurance-trained male (25.3 ± 4.5 years, 175.0 ±
1.0 cm, 69.0 ± 4.0 kg and 60.6 ± 4.4 ml O2.kg-1.min-1; mean
For personal use only.
± SE) volunteered to participate in this study. They spe-
cialised in middle and long-distance running. All gave
written voluntary informed consent in accordance with the
guidelines of the University of Paris.
Experimental design
The test sessions were completed on a 400-m covered track
before and after an eight-weeks period of a specific
endurance interval-training. Before training, each test
session consisted of an incremental test and an all-out test at
vD50, the velocity midway between the velocity associated
with the
. lactate. threshold (vLT) and the velocity associated
with VO2max (vVO2max). After training, subjects completed the
same incremental and all-out tests, plus an additional all-out
test at vD50bis, the velocity calculated from the post-
Pre-tests with :
∑ An incremental test
∑ An all-out test at vD50
1 2 3 4
Eight weeks of
interval-training at vD50
Fig. 1. Experimental design.
203
.
training vLT and. the post-training
. vVO2max. The maximal
oxygen uptake (VO2max), vVO2max, vLT, vD50 and the running
economy (RE, defined as the rate of oxygen uptake for a
1982) were determined during the incremental test. The all-
out tests were carried out to determine the time to exhaus-
tion (Tex). Velocity was regulated with an audio-visual system
and strictly controlled by photoelectric cells (Brower Timing
Systems, USA, Utah, Salt Lake City). Each subject per-
formed tests at the same time of day, in a climate-controlled
environment.
Procedures
During each test, the respiratory and pulmonary gas
exchange variables were measured using a breath-by-breath
portable gas analyser (Cosmed K4b2, Roma, Italy), which
was calibrated before each test according to the manufac-
turer’s instructions (Hausswirth et al., 1996). Breath-by-
breath data were later reduced to 5-s stationary averages
(Data Management Software, Cosmed, Rome, Italy). Finger-
tip capillary blood samples were collected into a capillary
tube, and were analysed for lactate concentration using a
Doctor Lange analyser (GmbH, Berlin, Germany), which
was calibrated before the tests. HR was monitored by the
breath-by-breath gas analyser (Cosmed K4b2, Roma, Italy)
from the signal transmitted by a cardiofrequencemeter trans-
mitter (Polar, PE 4000, Finland) adjusted on the chest of the
athlete and was breath-by-breath measured. Thus, the O2-p
was calculated from these values.
The incremental test (3-min stages) was set to induce
exhaustion within 20 min for each subject. Thus, velocity of
the initial stage was set at the average velocity maintained
over
. a 3000-m race (which has been described close to the
vVO2max, Billat, 1996a) minus 6 km.h-1. The velocity incre-
ment was set at 1 km.h-1. All stages were followed by a 30-s
rest period. A fingertip capillary blood sample was collected
to measure the lactate concentration during each rest period,
and in addition,
. before the test, immediately and 3 min after
the test. VO2max and HRmax were defined respectively as being
Post-tests with :
∑ An incremental test
∑ An all-out test at vD50
∑ An all-out test at vD50bis
weeks
5 6 7 8 9 10
 204 L.P. Laffite et al.

the highest 30-s oxygen uptake. and beat values reached
during the incremental test.. vVO2max was defined as the
minimal velocity at which VO2max was achieved (Billat &
Koralsztein,
. 1996b). In the case of the subject did not reached
VO2max during the previously
. stage and if he stopped to run
close to the half stage, vVO2max was considered as being the
median velocity between the last two stages (Kuipers et al.,
1985). vLT was defined as the velocity associated with an
increase in blood lactate concentration corresponding to
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.
50% of the pretraining vVO2max for the active recovery. To set
the number of repetitions of the interval-training sessions,
subjects ran until exhaustion during the first and the eighth
session in order to measure the maximal number of repeti-
tion (n). Thus, to vary and to avoid an overload of the inter-
val-training sessions the number of repetitions of the other
interval-training sessions was set at (n) minus 2 repetitions
(n - 2) or 1 repetition (n - 1) during respectively the first or
second interval-training session of each week. Intensity was

1 mmol.l-1 which occurred at approximately 3.5 to 5 mmol.
l-1 (Aunola & Rusko, 1984). vD50 .was defined as the veloc-
ity midway between vLT and vVO2max (Gaesser &. Pool,
1996). Running economy (RE), defined as the rate of VO2 for
a given submaximal work rate (Cavanagh
. & Williams, 1982),
was calculated using rate of VO2 averaged between the
second and the third minute of the stage run at 13 km.h-1.
The aim of the all-out constant test at vD50 was to deter-
mine the time until exhaustion at vD50 and vD50bis, i.e., the
time until the subject was unable to maintain the fixed veloc-
ity. Before each all-out test, .subjects completed a 15-min
warm-up period at 60% of vVO2max then observed a 5-min
rest period. Each subject was verbally encouraged. A finger-
tip capillary blood sample was collected before the test,
immediately and 3 min after the test.
For personal use only.
unchanged during the training period, nevertheless, the train-
ing volume was fitted according to the improvements of each
subject with an adjustment of (n) during the eighth interval-
training session. The other sessions (recovery) were an . easy
run during one hour at 60–70% of the pre-training vVO2max.
A professional trainer controlled each training session in
order to be sure that these instructions were respected.
Oxygen pulse kinetics determination
The oxygen pulse (O2-p) kinetics was analysed with the soft-
ware Sigmat Plot (SPSS, Chicago, IL, USA). Data were fitted
to the mono-exponential equation based on the model of
Linnarson (1974). According to Barstow et Mole (1991), the
equation is as follows:

Training program
Subjects were already well trained in endurance before their
participation. They completed a eight-weeks endurance train-
ing program, which was composed of two interval-training
sessions and three easy continuous running sessions per
week. The exercise duration of the interval-training sessions
was set at 50% of the time to exhaustion (Tex) of the all-out
pre-test at vD50 and the active recovery duration at 25%.
Thus, the duration ratio was of 2/1. The intensity of the inter-
val-training sessions was set at vD50 for exercise runs and
y = y 0 + A ¥ (1 - e - t t
) (1)
with y0, the y value at time = 0; A, the amplitude of the mono
exponential i.e., the y maximal value above y0; t, the time
constant of O2-p response defined as the time required to
attain 63% of the O2-p steady state; t, the time (s).
Calculation of the time to reach the O2-p steady
state (TRO2-p)
If a 5% error is admitted in the determination of the
O2-p value (the same error as for the determination of

Table 1. Weekly training program.

First week Second week Third week Fourth week

Monday Easy continuous running Easy continuous running Easy continuous running Easy continuous running
Tuesday IT session: n intervals IT session: n - 1 intervals IT session: n - 2 intervals IT session: n - 1 intervals
Wednesday Easy continuous running Easy continuous running Easy continuous running Easy continuous running
Thursday IT session: n - 2 intervals IT session: n - 2 intervals IT session: n - 1 intervals IT session: n - 2 intervals
Friday Easy continuous running Easy continuous running Easy continuous running Easy continuous running
Saturday Rest Rest Rest Rest
Sunday Rest Rest Rest Rest

Note These four weeks of training has been repeated two . times, with a reappraisal of the number n at the eighth interval training session (IT).
Example of the IT session with a subject having a vVO2max of 18.5 km.h-1, a vLT of 16 km.h-1, a vD50 of 17.3 km.h-1 and a tlim at vD50 of
425 s.
During these sessions, the subject would run at a hard intensity during 213 s, whereas the length of the light runs was 107 s.
The interval-training was run at 17.3 km.h-1 for the hard runs and at 9.3 km.h-1 for the light runs.
Subjects completed n - 1 or n - 2 intervals, where n is the maximal repetition number that the subject is able to achieve; n was recorded
during the first and the eighth interval-training sessions. The easy continuous running involved running during one hour under 13 km.h-1.
 The Effects of Interval Training on Oxygen Pulse 205

.
VO2), it may be accepted that a steady state in O2-p is as means and standard deviation (SD). The level of signifi-
attained above 95% of the asymptotic value reached during cance was established at 0.05.
the test.
Thus, to calculate TRO2-p, Equation (1) was changed to the
following: Results
O2 - p(t) = O2 - p(base) + A ¥ (1 - e -t t
) (2) Training effect on velocities and physiological maximal
with O2-p(t) is the oxygen pulse as a function of time; O2- values in incremental and vD50 test
p(base), the oxygen pulse at rest; A, the amplitude of the
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Training improved significantly vD50 in mean by 3% (17.6

mono exponential which corresponded to the maximal O2-p ± 1 km.h-1 vs. 17.2 ± 1 km.h-1; p = 0.03) and running
value above rest; t, the slope; t, the time. economy in mean by 4,7% (0.194 ± 0.01 ml.m.-1.kg-1 vs.
Given that the plateau value is equal to the maximal O2- 0.205 ± 0.02 ml.m-1.kg-1; p = 0.04), whereas vVO2max and
p value reached during the test (O2-pmax), TRO2-p is equal to vLT remained unchanged (respectively 18.5 ± 0.9 km.h-1 and
95% O2-pmax. Moreover, 16.1 ± 1.1 km.h-1). The other aerobic fitness values were not
95%O2 - p max = 0.95 [ A + O2 - p(base)] significantly improved.
Training improved significantly O2-pmax during the
Therefore, by expressing t as a function of the other vari- incremental test (22.7 ± 1.5 mlO2.b-1 vs. 20.6 ± 1.5 mlO2.b-1:
ables, Equation (2) was changed to the following: p = 0.048) and the amplitude of O2-p during the all-out test
TA95%O2 p max = TR O2 - p at v50 (13.4 ± 1.9 mlO2.b-1 vs. 11.9 ± 2.7 mlO2.b-1; p = 0.01).
This last observation is the consequence of the decrease of
È 0.95 ¥ A - 0.05 ¥ O2 p(base) ˘ the maximal HR value at vD50 (187. ± 6 bpm vs. 193 ± 4
= - t ¥ ln Í1 - ˙˚
Î A bpm; p = 0.02) whereas the maximal VO2 and [Lmax] values
(3) were not significantly different.
For personal use only.

Time spent at the O2-p steady state (TSSO2-p)
This time corresponds to the duration of the plateau of the
mono exponential. Thus, TSSO2-p is calculated by the differ-
ence between the total time and TRO2-p.
. the O2-p steady state was reached earlier and was maintained
Heart rate (HR) and VO2 kinetics
. in mean during 88% ± 4% of the time until exhaustion (Tex)
HR and VO2 kinetics parameters were analysed according to
the identical
. model of the O2-p kinetics replacing O2-p by HR
and VO2 respectively.
Statistical analysis
To compare data before and after training, data were com-
pared using a Student’s t-test for paired values and an analy-
sis of variance with repeated measures. Results are presented
.
Training effect on the O2-p, V O2 and HR kinetics
parameters at vD50 test
Time to reach the O2-p steady state (TR O2-p) and t were sig-
nificantly lower after training (respectively 60 ± 27 s vs. 33
± 7 s; p = 0.04; and 24 ± 10 s vs. 13 ± 3 s; p = 0.03). Thus,
(between 6 ± 2% and 94 ± 3% of Tex) vs. 80% ± 5% (between
13 ± 6% and 94 ± 3% of Tex) before training. Time spent at
the O2-p steady state (TSSO2-p) increased significantly after
training (552 ± 201 s vs. 407 ± 107 s; p = 0.04). There was
no significant difference in O2-p kinetic parameters to
compare results of the vD50 test before training and the
vD50bis test. .
The HR and VO2 kinetics parameters during each all-out
test (before and after training) are presented in Table 5.

Table 2. Training effects on the aerobic fitness parameters.

.
vVO2max vLT .% vD50 .% RE at 13 km.h-1 Tex at vD50 Tex at
(km.h-1) (km.h-1) vVO2max (km.h-1) vVO2max (ml.kg-1.m-1) (s) vD50bis (s)

Before training 18.5 ± 0.9 16.1 ± 1.1 87 ± 2 17.2 ± 1 93 ± 1 0.205 ± 0.02 509 ± 131 –
After training 18.6 ± 0.9 16.2 ± 1.0 87 ± 3 17.6 ± 1† 94 ± 3 0.194 ± 0.01† 617 ± 201 470 ± 78*
Test t NS NS NS p = 0.03 NS p = 0.04 NS p = 0.04
. . .
Note VO2max, maximal oxygen uptake; vV.O2max, velocity associated with VO2max; vLT, velocity associated with the lactate threshold. (LT);
vD50, median velocity between vLT and vVO2max; vD50bis, the velocity calculated from the post-training vLT and the post-training vVO2max;
RE, running economy; Tex, time to exhaustion. NS: not significantly different. † Significant difference between before and after training. * Sig-
nificant difference between the test at the new vD50 and the one at old vD50 after training.
 206 L.P. Laffite et al.

Table 3. Maximal responses during incremental and all-out tests at vD50.

.
VO2max O2-pmax HRmax [Lmax]
(ml.kg-1.min-1) (mlO2.b-1) (b.min-1) (mmol.l-1)

Incremental test 60.6 ± 4.4 20.6 ± 1.5 198 ± 7 11.4 ± 1.9

before training
Incremental test 63.0 ± 5.0 22.7 ± 1.5* 196 ± 7 11.2 ± 2.4
after training
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All-out test at vD50 62.3 ± 4.8 20.1 ± 1.6 193 ± 5 11.8 ± 2.5
before training
All-out test at vD50 61.7 ± 4.3 21.5 ± 1.1 187 ± 6† 11.9 ± 2.0
after training
All-out test at vD50bis 64.6 ± 4.8 22.3 ± 1.5† 190 ± 4 11.7 ± 2.9
after training
ANOVA with NS p = 0.04 p < 0.001 NS
repeated measures
.
Note VO2max, the maximal oxygen uptake reached during the test; O2-pmax, the maximal oxygen
pulse reached during the test; HRmax, the maximal heart rate reached during the test; [Lmax]; the
maximal blood lactate concentration reached during the test. NS: no significantly different.
* Significant difference between this test and the Incremental test before training (p < 0.04).
†
Significant difference between this test and the all-out test at the vD50 before training (p <
0.04).
For personal use only.

Table 4. Training effects on O2-p kinetics parameters.

O2-p base TRO2p t A TSSO2-p TSSO2-p (% of

(ml.b-1) (s) (s) (ml.b-1) (s) the total time)

Before training 8.0 ± 1.9 60 ± 27 24 ± 10.0 13.4 ± 1.9 407 ± 106 80 ± 4

After training at vD50 (17.2 km.h-1) 8.1 ± 1.7 33 ± 7† 13 ± 3.0† 11.9 ± 2.7 552 ± 201† 88 ± 4
After training at vD50bis (17.6 km.h-1) 6.9 ± 2.1 65 ± 21 24 ± 8.0 17.2 ± 2.6* 392 ± 63 84 ± 4
ANOVA with repeated measures NS p < 0.05 p = 0.01 p = 0.01 p = 0.03 NS

Note O2-p base, the oxygen pulse at rest; TRO2p, the time to reach 95% of O2-pmax of the all-out test; t, time to reach 63% of the O2-p steady
state value; A, the amplitude of O2-p kinetics curve; TSSO2-p, time spent at the O2-p steady state. NS: no significantly different. * Significant
difference between before training and after training at the new vD50 (p < 0.01). † Significant difference between before training and after
training at the old vD50 (p < 0.04).

.
Results of the training effects were similar to VO2 and HR
kinetics parameters. Thus, during the vD50 test, time to reach
the steady state (TR) was significantly. lower in HR (104.2 ±
41.6 s vs. 59.6 ± 19.4 s; p < 0.05) and VO2 (109.5 ± 11.2 s vs.
71.8 ± 25.7 s) after training that before training. The same
evolution was observed . in t of HR (42.9 ± 18.5 s vs. 25.4 ±
7.7 s; p < 0.05) and VO2 kinetics (40.4 ± 5.5 s vs. 26.5 ±
9.9 s; p < 0.05). Identical results were observed when com-
pared results of vD50 test before training and result of
vD50bis test (Table 5).
Relationship between the training effect on cardio
respiratory parameters and the performance (Tex at vD50)
There was no relationship between the modifications in O2-
p kinetics and the improvement in Tex at vD50. There was no
correlation between the improvement in RE and the improve-
ment in Tex at vD50 (r = -0.459; p = 0.321) and between RE
and the other variables measured in this study. There was also
no
. correlation between the O2-p kinetics parameters and the
VO2 kinetics parameters as well as between the O2-p kinet-
ics parameters and the HR kinetics parameters.
Discussion
The main results of this study showed that: i) endurance
training accelerated the oxygen pulse (O2-p) kinetics; ii) and
there was no relationship between the modification in the O2-
p kinetics and the increase in endurance.
The O2-p kinetics
O2-p may represent
. a third cardiopulmonary information in
addition to VO2 and HR (i.e., the product of the stroke
 The Effects of Interval Training on Oxygen Pulse 207

.
Table 5. Training effects on heart rate and VO2 kinetics parameters.

Before training at After training at After training at ANOVA with

vD50 (17.2 km.h-1) vD50 (17.2 km.h-1) vD50bis (17.6 km.h-1) repeated measures

Heart rate kinetics parameters

HR base (b.min-1) 76.6 ± 13.9 84.1 ± 15.6 79.3 ± 12.6 NS
TRHR (s) 104.2 ± 41.6 59.6 ± 19.4* 64.7 ± 20.8* p < 0.05
t (s) 42.9 ± 18.5 25.4 ± 7.7* 26.7 ± 8.2* p < 0.05
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A (b.min-1) 109.1 ± 10.2 91.4 ± 10.7* 100.7 ± 14.5 p < 0.05

.
VO.2 kinetics parameters
VO2 base (ml.kg-1.min-1) 12.3 ± 6.6 12.8 ± 3.4 9.6 ± 4.7 NS
TRVO2 (s) 109.5 ± 11.2 71.8 ± 25.7* 84.1 ± 24.7* p < 0.01
t (s) 40.4 ± 5.5 26.5 ± 9.9* 29.8 ± 8.5* p < 0.05
A (ml.kg-1.min-1) 41.6 ± 7.3 42.0 ± 4.7 ** 49.3 ± 5.8* p < 0.05

Note * Significant difference between before training and after training.

**. Significant difference after training between vD50 and vD50bis (the velocity calculated from the post-training vLT and the post-training
vVO2max).
NS: no significantly different. .
HR base, the heart. rate at rest; VO2, the oxygen consumption at rest; TRHR, the time to reach 95% of HRmax of the all-out test; TRVO2, the time
to reach 95% of VO2max of the all-out test; t, time to reach 63% of the steady state value; A, the amplitude of the kinetics curve.

volume (SV) and the arteriovenous O2 difference (a-v̄O2)D) The present study showed that after a eight-weeks period
For personal use only.

during tests on the track and field. This is true only if changes
in O. 2-p kinetics after training do not reflect the modifications
in VO2 or HR kinetics. Thus, three cases may occur: the first
would be that only the HR kinetics is. modified with training,
the second would be that only the VO2 kinetics. is modified
with training and the third would be that both VO2 and HR
kinetics are modified with training. The two first cases imply
that the modification of the O2-p kinetics . correlates and
reflects only the modification of HR or VO2 kinetics. Thus,
in this case the interest to study the O2-p kinetics is poor. .On
the other hand, the third case supposes both variables (VO2
and HR) are modified with training. Thus, the O2-p kinetics
modifications do not correlated with any kinetics of the sen-
sible variables. Only in that case. O2-p brings new informa-
tion compared with the HR or VO2 kinetics. In this study,
none
. of O2-p kinetics parameters are correlated with HR or
VO2 kinetics parameters, which confirm the interest to study
the O2-p kinetics. We can therefore conclude that. the O2-p
kinetics is no redundant with the study of HR or VO2 kinet-
ics, and that O2-p kinetics is the reflect of the product of SV
and (a-v̄O2)D. However, a next study would be carried out to
determine the role of these two variables in the O2-p kinet-
ics while measuring them.
To our knowledge, only one publication (Mille-Hamard et
al., 2000) has studied the O2-p kinetics during a constant and
severe intensity exercise but without training period. Mille-
Hamard et al. (2000) have . studied the O2-p kinetics during
an all-out test at 95% vVO2max and they showed that the time
spent at the O2-p steady state (TSSO2-p) was included between
10 and 90% of the time to exhaustion (Tex). Indeed, they did
not study the effects of an endurance-training period on
TSSO2-p and on the modification of the kinetics parameters.
of interval-training and during a test at vD50), TSSO2-p was
increased while starting earlier. Thus, we find that after train-
ing subjects maintained TSSO2-p from 6% (± 2) to 94% (± 3)
of Tex (vs. 13 ± 6% and 94 ± 3% of Tex before training). The
time to reach the O2-p steady state (TRO2-p) and t decreased
significantly during the vD50 test after training, but this two
kinetics parameters were very similar and no significantly
different during exercise at vD50bis. Thus, in accordance
with Sietsema et al. (1989), results showed that the endurance
training influenced significantly de O2-p kinetics and that the
O2-p kinetics in response to exercise is dependent on work
rate and also influenced
. by the fitness of the subject (char-
acterized by peak VO2 or anaerobic threshold values).
Finally, in accordance to different longitudinal studies
(Kasch et al., 1973; Mahler et al., 1985), we could have
observed an increase in O2-pmax after eight weeks of
endurance training during the incremental test.
Relationship between Tex and O2-p kinetics
In the present study, O2-pmax was increased after training.
Thus, the specific interval training allowed to improve
specifically vD50 and O2-p kinetics. Other studies may be
carried out to evaluate the role of SV and (a-v̄O2)D in the
modification of the O2-p kinetics. Tex at vD50 was improved
after training (617 ± 201 vs. 509 ± 131 s before training), but
not significantly, due to a large SE. This increase of Tex was
109 s in average, which represents an average increase of
23%. However, the modifications of the O2-p kinetics para-
meters were no correlated with the improvement of perfor-
mance after training at vD50. Billat et al. (1994) showed that
the measurement of Tex at the maximal aerobic speed (MAS)
 208 L.P. Laffite et al.

24 A study, athletes were able to improve their performances

22 (however not significantly) after training during the vD50
20 tests
. and during incremental test, whereas the values of the
18 VO2max and the maximal blood lactate concentration were not
16 significantly modified. HR decreased significantly after train-
O2-p (mL.b )
-1

14 ing during the vD50 test and O2-pmax increased significantly

12 during the incremental test, which showed an adaptation
10 to training (Wolfe & Cunningham, 1982; Mahler et al., 1985;
Sietsema et al., 1989; Krzeminski et al., 1991; Sagiv et al.,
Archives of Physiology and Biochemistry Downloaded from informahealthcare.com by University of Notre Dame Australia on 07/02/13

6 1991). Thus, it cannot be excluded that Tex at vD50 could be

4 related to the improvement of the O2-p kinetics or to other
2 bioenergetics characteristics in a wider sample of runners.
0 200 400 600 800 1000
The improvement in Tex could be explained also by the
70 B improvement in RE. However, regarding the effects of
60
training on RE, some studies reported no difference
(Overend et al., 1992; Ramsbottom et al., 1989), whereas
VO2 (ml.kg .min )

50
-1

others found an improvement in RE (Wilcox & Bulbulian,

40 1984; Kirwan et al., 1988) as result of training. Recently, a
-1

30
study showed an improvement in RE after six weeks of ply-
ometric training (Turner et al., 2003), whereas other studies
20
showed also an improvement in RE but after an endurance
10 training (Billat et al., 1999). Thus, responsible mechanism
0
of the improvement in RE must still be determined but are
0 200 400 600 800 1000 out of the scope of this study. Further researches are still
For personal use only.

220 C necessary to specify the mechanisms of running economy

improvements. Thus, given results of the present study, it is
200
not possible to hypothesize objectively a peripheral improve-
180 ment as result of training.
HR (b.min-1)

160

140 Role of SV and (a-v̄O2)D in the O2-p kinetics

120
100
80
0 200 400 600
Test at vD50 after training
Test at vD50 before training
. phenomena: the first would be that both SV and (a-v̄O2)D
Fig. 2. (A) O2-p, (B) VO2 and (C) HR kinetics before and after
training pf a typical subject.
in a laboratory setting was reproducible for a group of
runners at 1-week interval and that the average value could
be used for studying the effects of training, tapering, and/or
different warm-up procedures in a group of runners. . Fur-
thermore, they showed that the time spent at the VO2 steady
state was reproducible at 1-week interval. Thus, it is not very
probable that the reproducibility of the Tex may influence the
hypothetical relationship between the improvement in O2-p
kinetics and the improvement of the performance. Flynn et
al. (1994) showed that athletes and swimmers were able to
improve their Tex at the same velocity during the sport season,
whereas some physiological parameters (the serum concen-
tration of creatine kinase, cortisol . . .) did not vary in rela-
tion to the beginning of the season. It would seem that in this
According to the Fick equation, the O2-p is the product
between stroke volume (SV) and arterio-venous oxygen dif-
ference (a-v̄O2)D. Our results showed that O2-pmax deter-
800 1000 mined during the incremental . test increased significantly
Time (s) after training while HR and VO2max remained unchanged. The
increase in O2-pmax could be explained by three different
increased; the second would be that only SV increased; and
the last would be that only (a-v̄O2)D increased. It is difficult
to estimate how SV and (a-v̄O2)D evolved during training
because one limitation of the present study was that SV and
(a-v̄O2)D are not measured. Bhambhani et al. (1994) and
Hossack et al. (1980) have shown that during cycle or tread-
mill exercises in trained as well as in untrained subjects, O2-
p was significantly correlated with SV, but not necessarily
with (a-v̄O2)D. Moreover, Wolfe et al. (1992) have showed
that training during 11 weeks increased SV significantly
whereas (a-v̄O2)D was not increased. Therefore, many
authors (Sagiv et al., 1991; Wolfe et al., 1992; Bhambhani et
al., 1994; Bhambhani, 1995; Whipp, 1996) suggested that
O2-p may be a good means of appraising indirectly SV during
a submaximal exercise in trained and untrained subjects since
O2-p seems poorly influenced by the variability of (a-v̄O2)D:
according to Whipp et al. .(1996), SV may be estimated as 5
times the slope of the VO2-HR relationship (where 5 is
 The Effects of Interval Training on Oxygen Pulse
approximately 1/CaO2). In this context, we hypothesised that
the improvement in O2-p is mainly due to the increase in SV
in the present study, but other studies measuring SV and (a-
v̄O2)D during severe exercises, should be carried out to
confirm the validity of O2-p in order to predict SV.
In conclusion, the present study showed that after
endurance training, O2-p kinetics was faster and the steady
state was sustained for a longer period
. during an all-out test
at vD50, and also both for HR and VO2. Secondly, there was
Archives of Physiology and Biochemistry Downloaded from informahealthcare.com by University of Notre Dame Australia on 07/02/13
no relationship between modification of the O2-p kinetics and
improvement of the performance during exercise at vD50.
The hypothesis according to which the improvement of the
O2-p kinetics reflects mainly the improvement of SV may be
validated with SV measured during severe exercise before
and after a training session. These measures are today possi-
ble by the non invasive kinetics (Charloux et al., 2000; Chris-
tensen et al., 2000).
Acknowledgements
This study was supported by grants from Caisse Centrale des
Activités Sociales d’Electricité et Gaz de France.
For personal use only.
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