PREDICTION OF AEROBIC AND ANAEROBIC CAPACITIES
OF ELITE CYCLISTS FROM CHANGES IN LACTATE
DURING ISOCAPNIC BUFFERING PHASE
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MOHSEN HASANLI,1 ROHOLLAH NIKOOIE,1 MALIHE AVESEH,1
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1
Department of Exercise Physiology, Faculty of Physical Education and Sport Science, Shahid Bahonar University of Kerman,
Kerman, Iran; and 2Department of Exercise Physiology, Faculty of Physical Education and Sport Science, Tarbiat Modares
University, Tehran, Iran
ABSTRACT
Hasanli, M, Nikooie, R, Aveseh, M, and Mohammad, F. Pre-
diction of aerobic and anaerobic capacities of elite cyclists
from changes in lactate during isocapnic buffering phase.
J Strength Cond Res 29(2): 321–329, 2015—This study pre-
dicted aerobic and anaerobic capacities using relative changes
of arterial blood lactate during the isocapnic buffering phase
(relative [La]ISBP). Fourteen male professional cyclists (sprint-
trained [n = 6] and endurance [n = 8]) performed 2 exercise
sessions to exhaustion on a cycle ergometer; 1 incremental
standard test to determine the isocapnic buffering phase, buff-
ering capacities, and relative [La]ISBP and 1 supramaximal exer-
cise test to determine maximal accumulated oxygen deficit
(MAOD). The time between Lactate threshold (LT) and respi-
ratory compensatory threshold (RCT) was considered to be the
isocapnic buffering phase. Total buffering capacity was calcu-
lated as D[La]$DpH21. Bicarbonate buffering was calculated
as D[HCO32]$DpH21, and the difference between 2D[La]$D
pH21 and D[HCO32]$DpH21 was considered as nonbicarbon-
ate buffering. The lactate concentration for LT (p # 0.05) and
RCT (p # 0.05), and relative [La]ISBP (p , 0.01) were signif-
icantly lower for endurance cyclists than for sprint-trained
cyclists. A significant difference was found for bicarbonate
buffering capacity between groups (p , 0.01). A significant
correlation was found between relative [La]ISBP with V_ O2max
(r = 20.71, p # 0.05) and MAOD (r = 0.73, p , 0.01).
Relative [La]ISBP was useful for predicting aerobic power
(R2 = 51%) and anaerobic capacity (R2 = 53%). These results
demonstrated that relative [La]ISBP is an important variable in
intermediary metabolism and in addition to V_ O2max and LT is
recommended for better evaluation of performance of athletes
Address correspondence to Rohollah Nikooie, r_nikooie@uk.ac.ir.
29(2)/321–329
Journal of Strength and Conditioning Research
Ó 2015 National Strength and Conditioning Association
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AND FASHI MOHAMMAD2
who show nearly equal contributions from the aerobic and
anaerobic energy systems during exercise.
KEY WORDS buffering capacities, lactate kinetics, intermediary
metabolism
INTRODUCTION
E
xercise physiologists must have access to funda-
mental information on the qualities that contribute
to successful athletic performance to construct and
implement specific training programs. Several aero-
bic and anaerobic tests and parameters exist that allow coaches
and scientists to prescribe and monitor training programs. The
lactate threshold (LT) and maximal oxygen consumption
(V_ O2max) are important variables used to estimate endurance
performance (10). The Wingate anaerobic test (2) and maximal
accumulated oxygen deficit (MAOD) parameters (18) are also
used extensively to predict anaerobic performance.
Unlike endurance events in which most energy is pro-
vided by the aerobic system, many track events such as 1-,
3-, and 4-km cycling, 800- and 1,500-m runs require the
athletes to exhaust both aerobic and anaerobic metabolic
pathways (7). In such events, both the aerobic and anaerobic
systems are involved in energy production; their relative
contributions in adenosine triphosphate (ATP) production
are critical for a high level of performance. For example, it
has been shown that 4-km cycling performance is dependent
on V_ O2max, power output at LT, and MAOD variables (7);
Duffield et al. (8) have shown that the contribution of the
aerobic/anaerobic energy system to an 800-m run is 60/40%
in men and 70/30% in women. In this event, anaerobic ATP
formation is necessary to supplement oxidative metabolism
in the first few minutes of exercise until oxygen delivery
increases to meet demand and where the demand of power
output has a rate of energy utilization greater than that
which can be met by oxidative metabolism alone (11).
Therefore, the capacity for aerobic and anaerobic energy
production must be studied to predict performance in ath-
letes who have nearly equal contributions of aerobic and
anaerobic energy systems during exercise.
VOLUME 29 | NUMBER 2 | FEBRUARY 2015 | 321
 Lactate Kinetics and Performance
During a standard incremental exercise test, the time
between onset of LT and the respiratory compensatory
threshold (RCT) is called the isocapnic buffering phase
(14,22). During the isocapnic buffering phase, lactic acid,
which disassociates almost completely to lactate and hydro-
gen ions (H+) within the body physiological pH (power of
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hydrogen) range, is buffered by both bicarbonate and non-
bicarbonate buffers (14,22). Once both buffering systems
have been saturated, the accumulation of H+ leads to
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a decrease in pH and stimulates hyperventilation (14). The
duration of the isocapnic buffering phase is related to the
sensitivity of the carotid bodies (26); however, factors such
as buffering capacity and lactate kinetics are also involved.
Buffering capacity and lactate kinetics are adaptable vari-
ables, and their response to exercise training is dependent
on exercise orientation (9,19). Previous studies have shown
that sprint-trained athletes who regularly perform high-
intensity interval training have a greater muscle buffering
capacity (nonbicarbonate buffer) than endurance athletes
or untrained subjects (9,28). Endurance athletes show
lower blood lactate kinetics than untrained subjects during
exercise (19).
The aerobic and anaerobic systems produce energy
simultaneously during the isocapnic buffering phase. The
study of physiological variables during isocapnic buffering
can provide good information about adaptation of the
aerobic and anaerobic systems in response to exercise. The
change in blood lactate kinetics during isocapnic buffering
among athletes depends on their training orientations (14).
Arterial blood lactate concentration during exercise is
a result of its production and accumulation in active tissues,
such as fast glycolytic muscles, and uptake and oxidation
by oxidative tissues, such as the heart and oxidative
muscles (6). Practically, this means that the increase in
blood and muscle lactate concentrations reflects anaerobic
capacity and its removal and oxidation is the symbol of
aerobic capacity. It is reasonable to assume that changes
in arterial blood lactate can be used to predict aerobic and
anaerobic capacity because the balance between lactate
production and removal determines the final arterial blood
lactate concentration.
Studies of the relationship between exercise capacity and
isocapnic buffering exist (14,23), but no study has been made
of relative [La]ISBP in relation to exercise capacity. Hirakoba
and Yanoki (14) examined this relationship using venous
blood samples and found that the properties of the blood
samples did not allow formulation of a precise conclusion.
This study compares buffering capacity and changes in arte-
rial blood lactate during the isocapnic buffering phase for
sprint-trained and endurance cyclists. The increase in blood
lactate concentration was derived using lactate kinetics and
buffering. The kinetics of lactate, pH, and bicarbonate during
the isocapnic buffering phase were measured simultaneously.
The study also predicted aerobic and anaerobic capacities
from changes in the relative [La]ISBP of elite cyclists.
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322 Journal of Strength and Conditioning Research
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METHODS
Experimental Approach to the Problem
This study predicted aerobic and anaerobic capacity using
relative [La]ISBP in professional cyclists. Elite endurance and
sprint-trained cyclists performed 2 exercise sessions to
exhaustion on a cycle ergonometer; 1 incremental test to
determine the isocapnic buffering phase and lactate, pH,
and bicarbonate changes during the isocapnic buffering
phase and 1 supramaximal test to determine their MAOD.
At first, the relative [La]ISBP was compared between endur-
ance and sprint-trained cyclists. This comparison allowed us
to conclude that the relative [La]ISBP is different at different
levels of aerobic and anaerobic capacities. For attribute the
relative [La]ISBP to anaerobic metabolism, relative contribu-
tion of buffering capacities against lactic acid during the
isocapnic buffering phase were also investigated. Finally,
the correlation between V_ O2max as the aerobic power and
MAOD as the anaerobic capacity with the relative [La]ISBP
was calculated by Pearson correlation moment, and simple
linear regression analysis was used to determine the success
of prediction.
Subjects
The subjects were 14 male professional cyclists. Experi-
enced athletes who were medalists in Asian championship
events were chosen. Only 14 cyclists satisfied the selection
criteria (sprint-trained [n = 6] and endurance [n = 8]).
Their mean (SD) age, height, and body mass were
21.8 6 3.7 years, 178.5 6 4.7 cm, and 73.4 6 4.7 kg, respec-
tively. All subjects were over 18 years old. Each participant
performed 2 exercise sessions to exhaustion on a cycle ergon-
ometer; 1 standard incremental test to determine the iso-
capnic buffering phase, buffering capacities, and arterial
blood lactate changes and 1 supramaximal exercise test to
determine MAOD. All participants had 1 week of rest
between the 2 tests and were requested to avoid alcohol,
drugs, caffeine, and smoking for a period of 12 hours before
testing. Each participant gave his written informed consent
before participating in the study, and the study protocol was
approved by the ethics committee of the Shahid Bahonar
University of Kerman (approval No. K/91/25).
Procedures
Incremental Test. After instructing participant about the
nature of experiment, a 3-Fr gauge catheter was inserted
in brachial artery by a physician, and first arterial blood
sample was collected for resting blood lactate, bicarbonate,
and pH measurement. Then, after cycling for 5 minutes at
50 W as warming up period, participants performed an
incremental test on a workload constant power bicycle
ergometer starting at 100 W, and work load was increased
25 W every 3 minutes until exhaustion. During the last
30 seconds of each 3-minute stage, arterial blood samples
were drawn, and some was immediately analyzed for
arterial bicarbonate concentration and pH by arterial blood

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Figure 1. Examples showing methods of determining the lactate threshold (LT) and respiratory compensatory threshold (RCT) (see Methods for detail). A) RCT
is the point where abrupt increase occurs in VE/V_ CO2-power curve. B) LT is the point where abrupt increase occurs in blood lactate-power curve.
gas analyzer (ABL800 FLEX analyzer). The rest of the
blood was collected by heparinized tube, and plasma
separation was performed by centrifuging at 3,000g for
10 minutes at 48 C for plasma lactate measurement. Also,
breath-by-breath measurement of respiratory gases was per-
formed during the test using a gas analyzer system (Cosmed
K4b2, Rome, Italy). The system was calibrated immediately
with calibration gases before each experiment. The power
attained in final completed stage during the test was con-
sidered as the participant’s pmax and used for determination
of MAOD.
Lactate Threshold Determination. Lactate threshold was
defined as the break point where blood lactate concen-
Figure 2. Example showing method of determining the isocapnic
buffering phase. The time between lactate threshold (LT) and respiratory
compensatory threshold (RCT) was defined as the isocapnic buffering
phase.
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tration abruptly increased (20). In other words, the
threshold was considered to be the highest V_ O2 that could
be attained before an abrupt increase in the blood lactate
level (Figure 1A). Equivalent lactate and power to this
V_ O2 was considered to be the lactate and power corre-
sponding to LT.
Respiratory Compensatory Threshold Determination. The VE /
V_ CO2 vs. time was graphed. Then, RCT was defined as the
time when VE /V_ CO2 value abruptly increased (Figure 1B).
From the beginning of the exercise until the end of the test,
a fifth-degree polynomial regression curve was fitted on data.
The polynomial coefficients were obtained by the least
squares method, and a second-order derivative was calcu-
lated analytically. An inflexion point was obtained through-
out the zero crossing from the second-order derivative of
VE /V_ CO2 vs. time representing the imbalance of the VE /
V_ CO2 response facing the time. Correspondingly, the inflex-
ion point of VE /V_ CO2-time represented the RCT (29). All
calculation was performed by MatLab 9 software (Math-
Works Inc., CA, USA). Because in all subjects the RCT
occurrence was not exactly concomitant with the end of
the stages (where blood samples were collected), the values
of La, bicarbonate, and pH corresponding to the RCT were
calculated from the interpolation of the relationship between
the time of exercise and each variable value at work rates
above the LT.
Isocapnic Buffering Phase Determination. The time between LT
and RCT was considered to be the isocapnic buffering phase
(22) (Figure 2). The absolute value of the change in lactate
during isocapnic buffering was calculated as RCTlac 2 LTlac.
Total lactate production depends on body size, and so the
values of lactate corresponding to RCT and LT were adjusted
for body mass. Relative [La]ISBP was used for all statistical
calculations because the RCT and LT for the 2 groups
VOLUME 29 | NUMBER 2 | FEBRUARY 2015 | 323
 Lactate Kinetics and Performance

as D[HCO32]$DpH21. The dif-

ference between D[La]$DpH21
TABLE 1. Selected physiological variables of participants during the incremental
ergometer test.*† and D[HCO32]$DpH21 was
the nonbicarbonate buffering
Variable Endurance Sprint-trained capacity (4,5).
V_ O2max (ml$kg21$min21) 6 6
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63.51 5.24 57.94 4.51

pmax (W) 334 6 24.5 327 6 Maximal Accumulated Oxygen
22.7
Isocapnic buffering timez (s) 220.5 6 31.74 165.67 6 Deficit Test. Participants per-
19.07
O2 at LT (ml$kg21$min21)§ 44.24 6 5.11 38.92 6 4.05
formed a warming up period
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O2 at RCT (ml$kg21$min21)§ 51.21 6 4.12 46.58 6 4.18

Maximum lactate accumulation (mmol$L21)z
*LT = lactate threshold; RCT = respiratory compensatory threshold.
†Values are given as mean 6 SD (sprint-trained cyclists: n = 6 and endurance cyclists:
n = 8).
zSignificant difference between groups (p # 0.05).
§Significant difference between groups (p , 0.01).
occurred at different work intensities. Relative [La]ISBP was
calculated as previously described by Rocker et al. (27)
RCTlac 2LTlac
Relative  ½LaISBP ¼ 3100;
LTlac
where relative [La]ISBP is the relative lactate changes during
isocapnic buffering phase, RCTlac and LTlac are the lactate
concentration corresponding to RCT and LT, respectively.
Buffering Capacity and Its Component Calculation. Buffering
capacity and its components were calculated for each
participant. The blood lactate concentration and pH corre-
sponding to the LT and RCT were used to determine the
relative contribution of the buffering components during the
isocapnic buffering phase. Total buffering capacity was calcu-
lated as D[La]$DpH21. Bicarbonate buffering was calculated
for 10 minutes at 100 W fol-
8.9 6 1.2 11.01 6 1.62
lowed by 5 minutes of rest.
Then, subjects exercised on
the cycle ergometer at inten-
sity corresponding to %120 of
their pmax until exhaustion.
Breath-by-breath O2 consump-
tion was collected during the
test, and subjects were encour-
aged verbally to perform their maximal effort. Submaximal
oxygen uptake corresponding to intensities ranging from 100
W to individual LT powers in incremental test was used to
construct the power-O2 regression for each subject. The
individual linear relationship between power and O2 uptake
was extrapolated to the intensity at the supramaximal test,
and the corresponding O2 demand was predicted for each
subject. The MAOD was defined as the sum of the difference
between the predicted oxygen demand and actual oxygen
uptake for each 10-second period, for the duration of the
supramaximal test (18).
Biochemical Measurements. The arterial bicarbonate concen-
tration and pH were measured using an arterial blood gas
analyzer, and the plasma lactate concentration was deter-
mined using a lactate assay kit (K607-100; Biovision)
according to the manufacturer’s instructions.

Figure 3. Temporal course of changes in arterial blood HCO3, pH, and lactate during the incremental test in the groups of study. Baseline values: endurance
cyclists: [La2] 2.1 6 0.2 mmol$L21, [HCO32] 24.3 6 0.7 mmol$L21, pH 7.38 6 0.004 and sprint-trained cyclists: [La2] 2.5 6 0.2 mmol$L21, [HCO32] 23.4 6
0.8 mmol$L21, pH 7.37 6 0.004. *Significant difference between groups (p # 0.05).

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324 Journal of Strength and Conditioning Research

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TABLE 2. Lactate, bicarbonate concentration, and pH corresponding to LT and RCT and their relative changes during
the isocapnic buffering phase.*†

Variable Group LT RCT Relative changesz

Lactate concentration§k¶ (mmol$L21) 6 6 0.39 0.41 6 0.03

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Endurance 3.28 0.36 4.72

Sprint-trained 3.78 6 0.27 5.69 60.27 0.63 6 0.04
Bicarbonate (mmol$L21)¶ Endurance 20 6 2.2 17.7 6 1.6 211.7 6 1.3
Sprint-trained 19.6 6 1.2 17.9 6 1.4 28.97 6 0.95
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pH Endurance 7.28 6 0.05 7.23 6 0.04 20.66

Sprint-trained 7.27 6 0.04 7.22 6 0.04 20.68

*LT = lactate threshold; RCT = respiratory compensatory threshold.

†Values are given as mean 6 SD (sprint-trained cyclists: n = 6 and endurance cyclists: n = 8).
zRelative changes = ([value at RCT 2 value at LT]/[value at LT]) 3 100, for lactate relative changes, the attained values were also
adjusted for body mass.
§Significant difference between groups for the values corresponding to LT.
kSignificant difference between groups for the values corresponding to RCT.
¶Significant difference between groups for the relative change of each variable.

Statistical Analyses
The data are presented as mean 6 SD. The statistical anal-
ysis was initially performed using the Shapiro-Wilk nor-
mality test and the homoscedasticity test (Levene’s test).
All variables presented a normal distribution and homo-
scedasticity. Comparisons of variables between study
groups were performed using 2 independent sample t-tests.
The Pearson correlation moment was calculated to deter-
mine the common variances shared between predictors
(relative [La]ISBP) and the dependent variables (MAOD
and V_ O2max). Simple linear regression analysis was used
to determine the success of prediction. The least square
method was used to find the line of best fit. The residuals
were examined for a normal distribution using SPSS histo-
grams, normal probability plots, and a Shapiro-Wilks test.
The strength of each model fit to the data (R of Model)
was compared with use of the means by analysis of vari-
ance and considered satisfactory where p # 0.05. A
Durbin-Watson value of around 2.0 was used to confirm
the assumption that errors in regression were independent.
Effect size (ES) was calculated on variables exhibiting sig-
nificant results by dividing the difference between means of
the study groups by the pooled SD. This was performed to
determine the strength of the effects observed in these
results. Cohen’s d ESs are classified as small (0.20), mod-
erate (0.50), and large (0.80).

RESULTS
Table 1 lists the results of the
physiological variables for the
incremental test. No significant
TABLE 3. Variable values during the isocapnic buffering phase and total defense differences were found for pmax
throughout the incremental test.*
and V_ O2max between the
Variable Group Values sprint-trained and endurance
cyclists. O2 for LT (p , 0.01,
Bicarbonate buffering (mmol$L21 per pH unit)† Endurance 46.88 6 2.58 ES = 1.13) and RCT (p ,
Sprint-trained 40.02 6 2.75
Nonbicarbonate buffering (mmol$L21 per pH unit) Endurance 19.13 6 2.74 0.01, ES = 1.11) was signifi-
Sprint-trained 22.33 6 3.07 cantly higher for the endur-
Total buffering Endurance 65.8 6 3.24 ance cyclists. Total oxygen
Sprint-trained 62.3 6 3.37 deficit that developed during
Respiratory buffering (mmol$L21 per pH unit) Endurance 21.4 6 2.74 the supramaximal test was
Sprint-trained 24.5 6 3.14
Total defense Endurance 87.2 6 5.84 26.7 6 3.04 and 33.2 6
Sprint-trained 86.8 6 7.24 1.62 (ml$kg21$min21) in the
endurance and sprint-trained
*Values are given as mean 6 SD (sprint-trained cyclists: n = 6 and endurance cyclists: cyclists, respectively, and the
n = 8).
†Significant difference between groups (p , 0.01). significant difference was found
between the 2 groups (p #
0.05, ES = 2.55).

VOLUME 29 | NUMBER 2 | FEBRUARY 2015 | 325

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 Lactate Kinetics and Performance
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Figure 4. Correlation between arterial blood lactate changes during isocapnic buffering phase (relative [La]ISBP) with (A) V_ O2max (r = 20.7, p , 0.01) as the
aerobic power and (B) maximal accumulated oxygen deficit (MAOD) as the anaerobic capacity (r = 0.71, p , 0.01).
Figure 3 shows the course of change by time for arterial
blood pH, HCO32, and lactate concentration during the
incremental test for both groups. The decrease in [HCO32]
throughout the incremental test was less in the sprint-
trained group than for the endurance group (7.2 6 0.6 vs.
8.9 6 0.71 [mmol$L21]), and the difference between
groups was significant at 3 final stages (p # 0.05). The
increase in arterial blood lactate concentration throughout
the incremental test was significantly greater in the sprint-
trained group than for the endurance athletes (6.7 6 0.52
vs. 8.4 6 0.72 [mmol$L21]). There was no significant dif-
ference in pH change throughout the test between 2
groups.
Table 2 presents the value of lactate and bicarbonate con-
centration, and pH corresponding to LT and RCT and their
relative changes during the isocapnic buffering phase. The
lactate concentration for LT (p # 0.05, ES = 1.53) and RCT
(p # 0.05, ES = 2.81) was significantly lower for endurance
cyclists than for sprint-trained cyclists. Relative [La]ISBP was
significantly different between groups (p , 0.01). Bicarbon-
ate values for LT and RCT were not statistically different
between groups but showed a relative change that was sig-
nificantly higher for endurance athletes (p # 0.05, ES =
2.33). The duration of the isocapnic buffering phase was
significantly longer for endurance athletes in comparison
with sprint-trained cyclists (p # 0.05, ES = 2.02).
Table 3 lists the results of the buffering capacities during
the isocapnic buffering phase. A significant difference was
found for bicarbonate buffering capacity between groups
(p , 0.01). Nonbicarbonate buffering capacity during the
isocapnic buffering phase was not significantly different
between groups.
A significant negative correlation was found between relative
[La]ISBP and V_ O2max (r = 20.71, p , 0.01) (Figure 4A).
the
326 Journal of Strength and Conditioning Research
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The single-factor regression model using relative [La]ISBP
was useful for predicting aerobic power (R2 = 51%,
SEE = 3.18 ml$kg21$min21). The model had an Akaike’s
information criterion (AIC) value of 75.3, a Bayesian infor-
mation criterion (BIC) value of 77.3, and a Durbin-Watson
value of 2.13.
In addition, a significant positive correlation was found
between relative [La]ISBP and MAOD (r = 0.73, p , 0.01)
(Figure 4B). The single-factor regression model using rela-
tive [La]ISBP was useful for predicting anaerobic capacity
(R2 = 53%, SEE = 3.85 ml$kg21$min21). The model had
an AIC value of 61.3, a BIC value of 63.2, and Durbin-Watson
value of 1.83.
DISCUSSION
This study predicted aerobic and anaerobic capacities
using relative [La]ISBP in professional cyclists. The fol-
lowing results are novel findings of this study: (a) Relative
[La]ISBP was lower for endurance than for sprint-trained
cyclists; (b) the relative contribution of bicarbonate and
nonbicarbonate buffers against lactic acid during the iso-
capnic buffering phase was different for endurance and
sprint-trained cyclists; (c) relative [La] ISBP can be used
in some level of prediction of aerobic and anaerobic
capacity.
Endurance cyclists recorded significantly lower LT and
RCT values for arterial blood lactate concentrations in
response to their higher workloads. Similar results have
been reported in previous studies (12,19,27). The lower
blood lactate concentrations for LT and RCT in endurance
cyclists resulted from more effective lactate removal (1,19).
Phillips et al. (24) measured the percentage of fiber and
suggested that this may be a result of the increased presence
of slow twitch muscle fibers that more efficiently use lactate
TM

as a substrate for oxidative metabolism. In addition, their
higher percentage of oxidative fibers provided more energy
from aerobic metabolism until the higher intensity decreased
lactate accumulation. Moreover, previous studies have
shown that lower blood [La] values during submaximal
exercise in endurance athletes are mediated by the decrease
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in lactate appearance and increased lactate metabolic clear-
ance (17,19).
The relative [La]ISBP values were lower for endurance
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cyclists than for sprint-trained cyclists. The initial lactate
concentration for LT was different between groups and
RCT occurred at different percentages of V_ O2max; thus,
relative [La]ISBP was used in all calculations instead of
absolute values. This allowed us to conclude that relative
[La]ISBP can be attributed to anaerobic energy production.
The difference in relative [La]ISBP can be explained by
different buffering capacities and lactate kinetics. The
arterial blood lactate kinetic is the result of balance
between its production and removal (6). It has been
shown that lactate production during intense submaximal
exercise is lower in endurance athletes (15). This may be
caused by the higher concentration of mitochondria in the
oxidative fibers that theoretically increases the oxidative
capacity of the muscle. This adaptation was also found in
the endurance cyclists in this study; they recorded higher
O2 consumption at workloads corresponding to RCT.
When O2 consumption for RCT was adjusted for work-
load (because RCT occurred at higher intensities in endur-
ance athletes), its values were higher for endurance
cyclists than for sprint-trained cyclists. This finding indi-
cates that endurance cyclists received more energy from
aerobic metabolism during the isocapnic buffering phase
and consequently experienced a lower increase in relative
[La]ISBP.
An alternative explanation may be related to the
amount of monocarboxylate transporter proteins (MCTs)
in the muscle fibers. MCT1 and MCT4 are predominant
isoforms of human skeletal muscle involved in lactate
exchange (6). MCT1 is highly expressed in oxidative tis-
sue and facilitates lactate uptake at the sarcolemmal and
mitochondrial membranes (6). MCT4 is found predomi-
nantly in glycolytic fibers, and its putative role is lactate
extrusion (6). Nikooie et al. (21) observed that endurance
training can increase MCT1 expression in the sarcolem-
mal and mitochondrial membrane in rat skeletal muscle
and facilitate lactate oxidation (21). By contrast, repeated
high-intensity exercise routinely performed by sprint-
trained cyclists increases sarcolemmal MCT4 expression
that facilitates the lactate export from skeletal muscle
(3,25). This adaptation to high-intensity interval training
can partially explain the higher values for relative [La]ISBP
in these athletes.
Buffering against lactic acid is another important factor
that affects blood lactate concentration during exercise.
Because this study attributes relative [La]ISBP to anaerobic
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metabolism, the relative contribution of buffering during
the isocapnic buffering phase was also investigated.
Endurance cyclists showed greater bicarbonate buffering
during the isocapnic buffering phase, and sprint-trained
cyclists showed that more H+ was buffered by the non-
bicarbonate buffer system. Previous studies have noted the
former and have shown that high-intensity interval train-
ing increases muscle buffering capacity (3,9,25). Initial
buffering of lactic acid by creatine formed from the split-
ting of creatine phosphate (13), an increase in intracellular
pH buffering as the result of increased muscle carnosine
levels (30), and an increased lactate/H+ cotransport due
to an increased expression of MCTs and Na+/H+ proteins
(3,16,25) are the responsible mechanisms. Although the
increase in relative [La]ISBP in sprint-trained cyclists was
higher than that for endurance athletes, the change in
blood [HCO32] during the isocapnic buffering phase
was significantly lower in sprint-trained cyclists. Taken
together, the initial increase in the blood lactate concen-
tration without a concomitant decrease in the blood
[HCO32] in sprint-trained cyclists reflects the onset of
a nonbicarbonate buffering mechanism in the skeletal
muscle. The decrease in arterial blood pH during the iso-
capnic buffering phase was similar between groups, how-
ever, indicating that both groups experienced the same
total buffering during the isocapnic buffering phase. It is
reasonable to conclude that the higher values for relative
[La]ISBP in sprint-trained cyclists were the result of higher
anaerobic energy production.
Relative [La]ISBP has a negative correlation with
V_ O2max as the aerobic power index and a positive corre-
lation with MAOD as the anaerobic capacity index. The
linear relationship between power and O2 uptake for inten-
sities below LT was established to calculate MAOD, and
this relationship was used to extrapolate the oxygen
demand for supramaximal intensity. There is a nonlinear
V_ O2-power relationship with a decreasing slope at higher
exercise intensities; therefore, estimation of O2 demand
at supramaximal intensity based on power-O2 uptake
attained from intensities below LT can overestimate O2
uptake for supramaximal intensity. Nevertheless, MAOD
is a valid index of anaerobic capacity and has been used
extensively (8,18). The negative correlation between rela-
tive [La]ISBP and V_ O2max may be related to the greater
percentage of energy production from the aerobic system
during exercise. It has been shown that athletes with high-
er capacity of the aerobic system show decreased contri-
butions from the anaerobic pathway in energy production
during intense exercise (11). It can be concluded that rel-
ative [La]ISBP is an integrated variable that reflects
the contribution of both aerobic and anaerobic systems
in energy production during exercise. This finding is essen-
tial to the evaluation of performance in athletes who
have similar contributions from the aerobic and
anaerobic energy system during exercise. In such athletes,
VOLUME 29 | NUMBER 2 | FEBRUARY 2015 | 327
 Lactate Kinetics and Performance
simultaneous evaluation of V_ O2max, LT, and relative
[La]ISBP can provide better insight into their performance
and adaptation to training.
Athletes training for 800-m runs, or 2-, 3-, and 4-km
cycling perform mainly at intensities between LT and
RCT. For these athletes, optimal performance depends on
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improvement in both aerobic and anaerobic capacities. To
better evaluate their performance, coaches and athletes
can use relative [La]ISBP, which reflects the combination
CX1AWnYQp/IlQrHD3i3D0OdRyi7TvSFl4Cf3VC1y0abggQZXdtwnfKZBYtws= on 04/18/2023
of aerobic and anaerobic systems in energy production.
Monitoring relative [La]ISBP throughout the conditioning
phase allows coaches to evaluate the adaptation of their
athletes and modify their training programs according to
physiological demands. However, because only 14 cyclists
met the selection criteria for this study, the sample size
was small. This is a clear limitation of this study that may
have influenced the results, especially for regression anal-
ysis; however, the data provide a useful platform for future
research.
PRACTICAL APPLICATIONS
The results of this study showed that relative [La]ISBP is an
important variable in intermediary metabolism and can be
used to predict anaerobic and aerobic capacities. Athletes
involved in sports such as 3- and 4-km cycling usually train
at intensities between LT and RCT. Their optimal perfor-
mance depends on improving both aerobic and anaerobic
capacities. Coaches can use relative [La]ISBP, which reflects
the combination of aerobic and anaerobic systems in energy
production, to better evaluate the performance of these ath-
letes. Monitoring relative [La]ISBP throughout the condition-
ing phase allows coaches to evaluate their training and helps
adapt and modify training programs to improve either the
aerobic or anaerobic system. Relative [La]ISBP in addition to
V_ O2max and LT is recommended for better evaluation of
performance of athletes who show nearly equal contribu-
tions from the aerobic and anaerobic energy systems during
exercise.
ACKNOWLEDGMENTS
The authors thank all test subjects for their willing
cooperation and also thank all collaborators involved in
this study.
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