Brief Review

International Journal of Sports Physiology and Performance, 2011, 6, 8-24

© 2011 Human Kinetics, Inc.

Blood Lactate Diagnostics in Exercise

Testing and Training
Ralph Beneke, Renate M. Leithäuser, and Oliver Ochentel

A link between lactate and muscular exercise was seen already more than 200
years ago. The blood lactate concentration (BLC) is sensitive to changes in exercise
intensity and duration. Multiple BLC threshold concepts define different points on
the BLC power curve during various tests with increasing power (INCP). The INCP
test results are affected by the increase in power over time. The maximal lactate
steady state (MLSS) is measured during a series of prolonged constant power
(CP) tests. It detects the highest aerobic power without metabolic energy from
continuing net lactate production, which is usually sustainable for 30 to 60 min.
BLC threshold and MLSS power are highly correlated with the maximum aerobic
power and athletic endurance performance. The idea that training at threshold
intensity is particularly effective has no evidence. Three BLC-orientated intensity
domains have been established: (1) training up to an intensity at which the BLC
clearly exceeds resting BLC, light- and moderate-intensity training focusing on
active regeneration or high-volume endurance training (Intensity < Threshold);
(2) heavy endurance training at work rates up to MLSS intensity (Threshold ≤
Intensity ≤ MLSS); and (3) severe exercise intensity training between MLSS and
maximum oxygen uptake intensity mostly organized as interval and tempo work
(Intensity > MLSS). High-performance endurance athletes combining very high
training volume with high aerobic power dedicate 70 to 90% of their training to
intensity domain 1 (Intensity < Threshold) in order to keep glycogen homeostasis
within sustainable limits.

Keywords: lactate threshold, maximal lactate steady state, training, intensity,

volume

The chemical basics of lactate diagnostics go back for more than 200 years.
The start was possibly in Stralsund, a Hanse town on the German Baltic sea. As
early as 1780, the Swedish pharmacist Carl Wilhelm Scheele discovered lactate in
sour milk and named it mjölksyra (milk acid). Twenty eight years later, Jöns Jakob
Berzelius described lactate in extracts from muscles of hunted stags and linked
it with muscular exercise.1 During the subsequent 50 years, Berzelius’s observa-
tions were confirmed and extended by Justus von Liebig, who found that lactate

Ralph Beneke is with the Centre for Sport and Exercise Science, Department of Biological Sciences,
University of Essex, Essex, England. Renate M. Leithäuser is with Biomedical Science, Department
of Biological Sciences, University of Essex, Essex, England. Oliver Ochentel is with the Centre for
Sport and Exercise Science, Department of Biological Sciences, University of Essex, Essex, England.

8
 Blood Lactate Diagnostics   9

is always present in dead muscle tissue,2 and by Emil Heinrich du Bois-Reymond,

who observed the effect of lactate on muscle contraction.3
In 1891, the concept of tissue hypoxia as a cause of lactate formation was first
put forward by Arkai and Zillessen.4,5 Tissue ischemia still serves as a highly effec-
tive way to activate glycolysis and achieve high lactate concentrations.6 However,
elevated lactate levels do not indicate ischemia per se. Any steady state or decrease
of the blood lactate concentration (BLC) as frequently observed during prolonged
constant power,7–11 irrespective of how much the BLC is elevated above resting levels,
is consistent with continuous lactate production under fully aerobic conditions.12–14
As early as at the beginning of the last century, a systematic stream of well-
coordinated in vitro and in vivo experiments by Archibald Vivian Hill and Otto
Fritz Meyerhof led to the first consistent and still widely accepted theory describing
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muscular metabolism as an open thermodynamic system15 with lactate production

and respiration as integral factors of metabolic energy production.16 In 1922, Hill
and Meyerhof were awarded the still only Nobel Prize crediting exercise physi-
ological/biochemical research. Their work laid the foundation also for subsequent
research on lactate as substrate of aerobic metabolism based on intra- and intercell
and intertissue shuttle processes, including lactate as substitute for glucose as fuel
for oxidation in the brain during exercise.8,12,13,17–23
Recent studies discussed lactate as an indicator of the activation of the use of
carbohydrate as fuel for aerobic metabolism,24 and suggested oxygen radical gen-
eration by metabolic stress-induced increases in lactate and mitochondrial oxygen
uptake as factor of structural impairment of cellular integrity25 and as transcriptional
activators and signals of adaptive cell response.14
Experimental basics of most current testing protocols have already been seeded
during the third decade of the last century by investigators like Ole Bang, who care-
fully described and analyzed BLC during and after incremental (INCP), short-term
high-intensity (STHIP), and prolonged constant power (CP) tests.7 Clear experi-
mental evidence that lactate production is possible even with no increase in lactate,
and that aerobic skeletal muscle metabolism may be regarded as the main factor
of lactate removal during exercise was provided by Hermansen and Stensvold.8
Subsequently, the development of an enzymatic-photometric micromethod to
analyze lactate reduced the required volume of specimen to 20 µL of capillary blood
usually taken from the fingertip or the ear lobe.26 This new method was feasible
for BLC measurements under laboratory and also field conditions. This method,
however, is now being almost replaced by less manual work–intensive enzymatic-
amperometric and microphotometric techniques.27 The fact that the BLC shows
specific responses to INCP, STHIP, and CP tests7,8 led to a still lasting boom in
lactate measurements in exercise physiology and sports medicine. Consequently,
nowadays blood lactate diagnostics serves as standard element of exercise testing
and prescription in top-class endurance athletes but also of numerous commercial
leisure exercise programs.

Testing
The BLC discriminates the metabolic response to exercise intensity and duration
clearly during INCP and CP tests. The possibly most common endurance testing
procedure is the INCP combined with the analysis of changes in BLC associated
 10   Beneke, Leithäuser, and Ochentel

with the increase in power. Usually INCP tests start at a power reflecting moder-
ate exercise intensity. The exercise intensity is subsequently increased by given
increments at given times. Over a range of low power increments, the BLC remains
more or less at resting level before it increases slowly but progressively during
subsequent steps in power (Figure 1).
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Figure 1 — Example of BLC during INCP; the BLC does not increase during increasing
power up to an intensity of 48%; at 4 mmol⋅L–1 exercise intensity is 66%, which is lower
than the MLSS intensity of 83% of maximum power at test termination.

Lactate thresholds are generally accepted performance indicators. The first

BLC-based threshold concept termed the aerobic-anaerobic threshold28 detected
the power at a BLC of 4.0 mmol⋅L–1 during an INCP test with relatively long
lasting stages of 5 min at minimum. This threshold, also termed the 4 mmol⋅L–1
threshold, was based upon empirical observations that long lasting constant
exercise was sustainable if the BLC stabilized after an initial increase indicating
equilibrium between lactate appearance and elimination (Figure 2). A constant
BLC indicates that the rate of glycolytic lactate generation is similar to its use
as fuel for the aerobic metabolism. A decrease in BLC appears if more pyruvate
and lactate are aerobically consumed than generated via anaerobic glycolysis.
If the rate of glycolytic lactate generation exceeds the corresponding rate of
aerobic lactate utilization, the resulting net lactate accumulation and increase in
the BLC indicate a demand of anaerobic metabolic energy. During prolonged
running exercise, a constant BLC was frequently seen above BLC baseline but
rarely above 4.0 mmol⋅L–1.28
Shortly after Mader et al28 had published their threshold idea, an excessive
number of other threshold concepts were put forward and this process is still ongo-
ing. Due to this “threshold inflation,” it is nowadays almost impossible to verify the
correct number of different threshold concepts published internationally. However,
 Blood Lactate Diagnostics   11
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Figure 2 — BLC during CP according to INCP (Figure 1).

most threshold concepts reflect just slight variations of procedures developed in

the 1970s and 1980s.29–43
The idea of the aerobic-anaerobic threshold28 was that it may detect an exercise
intensity at or close to the highest prolonged CP at which, after an initial increase, a
constant BLC is observed. The intention to verify this idea led to the concept of the
maximal lactate steady state (MLSS).10,44 The MLSS is measured during a series
of prolonged CP tests and defines an exercise intensity that is usually sustainable
for 30 to 60 min.45,46 It detects the highest aerobic power without metabolic energy
from continuing net lactate production. This MLSS power is highly correlated
with the maximum aerobic power. The between-subject variability of the MLSS
is rather high and the MLSS is rarely found at a BLC of 4 mmol⋅L–1. MLSS values
between 1.9 and 7.5 mmol⋅L–1 have been reported, which corresponded to exercise
intensities between 54 and 83% of the maximum power measured during INCP
testing. Surprisingly, MLSS, and MLSS intensity are independent of maximum
power.47 There is evidence that the MLSS differs between different sports even
if performed by the same athlete.48,49 Consequently, depending on the individual
but also on testing conditions, the MLSS may reflect a BLC close to that typically
found at rest or at a higher BLC, which may be significantly beyond 4 mmol⋅L–1
(Figure 2). Under selected conditions, the average MLSS power may be close or
similar to the power corresponding to a specific BLC threshold measured during a
selected INCP testing program. However, even then the preciseness of individual
MLSS power predictions via INCP testing remains moderate. Therefore, threshold
determinations are likely to under- or overestimate individual MLSS power.10,44,48,50
Obviously differently defined thresholds determine different points on the
BLC power curve. These points reflect either (a) selected BLC values, (b) defined
increases in BLC above baseline or (c) specific changes in the steepness of the BLC
power curve or (d) combinations of BLC measurements during INCP testing, at
INCP test termination and during recovery.30,31,51 Thresholds detecting the power
 12   Beneke, Leithäuser, and Ochentel

at which the BLC increases above resting BLC and the MLSS indicate a range of
exercise intensities that require a significant increase in the glycolytic rate to fuel
aerobic pyruvate utilization. However, this increase in BLC during the initial phase
of steady-state exercise does not indicate anaerobic metabolism. The corresponding
dynamic steady-state BLC response is functionally identical to the pharmacokinetic
principle that explains an increase of a drug in the blood when applied via a constant
flow infuser and metabolized at the rate of infusion.51 Consequently, the analysis
of the BLC response during INCP and CP discriminates between three intensity
domains reflecting three qualitatively different metabolic situations: (1) intensities
that do not require an increase in the glycolytic rate that increases the BLC above
resting BLC (Intensity < Threshold), (2) intensities causing an increase in the
glycolytic rate that leads to a BLC steady state higher than resting BLC, which is
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matched by the rate of aerobic pyruvate utilization and therefore an aerobic meta-
bolic condition (Threshold ≤ Intensity ≤ MLSS), and (3) intensities that require a
glycolytic rate that cannot be matched by aerobic pyruvate combustion indicating
the demand of anaerobic glycolytic energy (Intensity > MLSS) (Figure 2).
From a practical point of view, the direct determination of the MLSS is no real
option for regular routine performance testing. Direct determination of the MLSS
requires three to seven different CP tests of sufficient duration all conducted at dif-
ferent days with sufficient recovery periods in between.11 The duration of MLSS tests
should be 30 min or longer because the time required to establish a BLC steady state
increases if the steady state of the BLC is increased. A MLSS of about 7 mmol⋅L–1
may require approximately 15 to 20 min until a steady state is established.11,52 This
is hardly compatible with normal training routines and is incompatible with eco-
nomical workflow of a performance testing service unit. Although discriminating
well between steady-state and non-steady-state conditions, attempts with two tests
only, an INCP test plus a second INCP test with very few but long-lasting stages
(Figure 3 and 4) were not fully convincing in detecting MLSS power.53–55 Therefore,
threshold testing will keep its value for endurance testing just by clear definition of
how a specifically defined point on the BLC power curve can be found.
Any point on the BLC power curve as specified by current threshold concepts is
an objective test interpretation which does not require maximum effort of the tested
individual toward the end of the INCP. It is therefore motivation independent and
also less stressful than testing other endurance indicators requiring maximum effort.
However, each specific threshold concept detects its own point on the BLC power
curve.29–31 Depending on the specific threshold concept, these may reflect exercise
intensities close to baseline BLC or higher. There is no convincing evidence that
any concept is better than another. Most thresholds are well correlated with other
threshold concepts, and more importantly with performances in real and simulated
competition in various endurance events.29,56–66 However, those threshold concepts
corresponding to higher BLC values tend to provide slightly higher coefficients of
correlation with nonthreshold measures of endurance.29 The latter is explained by
the more or less exponential shape of the BLC to power relationship during INCP.
Threshold concepts detecting the power which causes an increase in BLC above
baseline analyze the flat part of the BLC power curve. Here small variations in
BLC are associated with substantial changes in power. The variability of modern
lactate analyzers is low with ∼2% at BLC below 2 mmol⋅L–1, ∼1% at BLC 2 to 6
mmol⋅L–1 and ∼1.5% if repetitive analyses of the same specimen cover a range of
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Figure 3 — BLC during INCP. A: not specifically trained subject (maximum oxygen
uptake mL⋅kg–1⋅min–1). B: internationally successful endurance athlete (maximum oxygen
uptake 77 mL⋅kg–1⋅min–1).

Figure 4 — BLC during three-step INCP test with 12 min stage duration. A: not specifi-
cally trained subject. B: internationally successful endurance athlete.

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 14   Beneke, Leithäuser, and Ochentel

BLC between 0.6 and 11.7 mmol⋅L–1.27 However, this does not include the vari-
ability of repeated blood sampling. The combined variability of blood sampling
and analysis is in the magnitude of 3 to 7%. Analyzing the steeper part of the BLC
power curve reduces the effect of this variability on power predictions.
The identical threshold concept will provide different results if the INCP
test varies in terms of increase in power over time.29,50,52,67 A steeper increase in
power over time shifts the BLC power curve to the right, indicating an effectively
nonexistent improvement in aerobic fitness. Therefore, it may lead to confusion
and inappropriate conclusions about testing results if it is not clearly documented
which specific threshold concept is used in combination with which testing program.
However, as long as the identical threshold concept is used under identical testing
conditions, this is a minor issue for cross-sectional or longitudinal assessments of
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changes in the endurance capacity.

Identical testing conditions include also the identical setup of handlebars
and saddle of regularly calibrated cycle ergometers and identical treadmills used
at identical inclinations. These factors affect the biomechanical efficiency and
therefore the shape of the BLC power curve.29,52 In selected buildings even the
position of a treadmill in the laboratory may affect running economy due to modi-
fied resonance oscillations of floor and building. Environmental factors get even
more important under field conditions. The BLC power curve is highly sensitive
to running surfaces,52 temperature,68–70 altitude, and timing of testing at altitude.71
A further pitfall for comparing field and laboratory testing is that running tests
on treadmills include given increments in running velocity after constant periods
of running, which results in a linear increase in speed over time. Contrary to the
latter, field running tests are mostly designed as numerous constant distances and
running velocity is increased by shortening the time set for each subsequent identi-
cal distance. Compared with treadmill and ergometer testing in the laboratory, the
decreasing duration of, for example, running or cycling at subsequent increased
velocities over a given distance, shifts the BLC power curve to the right under field
conditions. Furthermore, high-speed events like cycling and speed skating require
test interruptions for blood sampling, which include relative long deceleration and
acceleration periods before and after BLC sampling, which affect the power profile
of a test and therefore the BLC–speed relationship.48,52,72

Training
Exercise testing should ideally transfer directly into training. Consequently, the idea
that a lactate threshold may identify a clearly defined metabolic state made it tempt-
ing to utilize a point on the BLC power curve as orientation for training advice.28
The original idea was that a running speed below the 4 mmol⋅L–1 threshold intensity
would be appropriate for endurance training covering distances between 5 and 8
km. Distances between 3 and 5 km should reflect threshold intensity, and highly
intensive running training requires running speeds above threshold intensity.28 How
this led to proposals that training at threshold intensity might be particularly effective
remains unclear. Very early anecdotal evidence suggested that top athletes did not
respond favorably to substantial training volumes at threshold intensity. This led to
speculations that threshold BLC may decrease with increasing endurance, which
was one reason for boosting the search for individual threshold concepts particularly
 Blood Lactate Diagnostics   15

in the late 70s and early 80s.29 The subsequent inflation of the number of different
threshold concepts made the term threshold-orientated training even more confusing.
Selected studies on moderately trained, untrained or deconditioned subjects
described that threshold training works.32,73–76 However, it remains unclear whether
threshold intensity really matters in untrained subjects. In untrained subjects and
recreational athletes also various other training programs lasting between 2 and
12 weeks were comparably successful as long as the training generated additional
but also sustainable regular metabolic demand.77–79 The supposedly rather blunt
consideration that endurance training works as long as the mix in exercise intensities
accumulates substantial but sustainable metabolic demand irrespective of threshold
intensities may get credibility by the fact that successful endurance athletes do
not spend much effort on threshold training. Depending on seasonal variations,
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top athletes competing in middle and long distance events train 70 to 90% of their
endurance training at intensities corresponding to BLCs below 2 mmol⋅L–1. Also,
the remaining training includes more effort at intensities corresponding to BLCs
above 6 mmol⋅L–1, which is likely to be higher than usually measured thresholds.80–92
Based on these observations, a number of BLC-oriented endurance training
zones have been developed. They basically mirror the above-mentioned three
BLC-related intensity domains: (1) training up to an intensity at which the BLC
clearly exceeds resting BLC, equivalent to light and moderate training focusing on
active regeneration or high-volume endurance work mostly lasting between 1 and
6 h (Intensity < Threshold); (2) heavy endurance training at work rates suppos-
edly up to MLSS intensity lasting 30 to 90 min (Threshold ≤ Intensity ≤ MLSS);
and (3) severe exercise intensity training between MLSS and maximum oxygen
uptake intensity organized as interval and tempo work normally lasting up to 30
min (Intensity > MLSS). Interestingly, the volume of training spent on domain 2
is rather low even compared with that of domain 3.86,91
Studies that tried to intensify the training in athletes by increasing the training
volume at threshold intensity combined with a reduction of hours spent at intensities
near resting BLC did not report any favorable and some reported even detrimental
effects compared with more low intensity training.93,94 The fact that selected subjects
already hit MLSS intensity at BLC levels of around 2 mmol⋅L–1 may partly explain
why some do not tolerate substantial training volumes at intensities corresponding
to BLC levels significantly higher than 2 mmol⋅L–1. It is possibly also reasonable
to speculate that the high between-subject variability of the MLSS may explain
why in highly trained endurance athletes and also in less endurance trained but
physically active individuals the postexercise autonomic response discriminated
between training below and above threshold intensity independent of endurance
capacity and individual training volume, but not between training below or above
MLSS intensity.95 However, this does not explain why untrained subjects and rec-
reational athletes seem to tolerate these higher training intensities better than top
athletes. Nevertheless, the above-mentioned training studies suggest that identical
BLC-oriented intensity domains, such as (1) Intensity < Threshold, (2) Threshold
≤ Intensity ≤ MLSS, and (3) Intensity > MLSS, have different metabolic meanings
and adaptive consequences in untrained and highly endurance trained individuals.
The test results displayed in Figure 3 present BLC acute responses as frequently
observed during INCP tests with untrained and highly endurance trained individu-
als. In untrained subject A (maximum oxygen uptake: 48 mL⋅kg–1⋅min–1), the BLC
 16   Beneke, Leithäuser, and Ochentel

increased during the first and with each subsequent increment in power. Therefore,
intensity domain 1 (Intensity < Threshold) could not be clearly identified. In highly
trained athlete B (maximum oxygen uptake: 77 mL⋅kg–1⋅min–1), BLC increased
during the initial stage but decreased at the subsequent stage. Above 200 W, the
BLC clearly increased with every increase in power. In both individuals, the MLSS
power of 160 and 305 W, respectively, was near a power at 4 mmol⋅L–1 correspond-
ing to 63 and 69% of maximum oxygen uptake in subjects A and B. Subject A
terminated a CP test at 150 W after 55 min whereas athlete B was able to exercise
at 295 W for more than an hour. As expected, both A and B terminated a three-step
test with 12-min stages prematurely at a power higher than MLSS intensity (Figure
4). In spite of the fact that MLSS power and maximum power were 90% and 67%
higher in athlete B than in athlete A, the BLC to exercise intensity (% of maximum
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oxygen uptake) relationship was very similar and did not discriminate convincingly
between untrained and highly trained individuals. Both show a similar pattern of
intensity-related increases in BLC during INCP; both show similar BLC responses
at similar exercise intensities between resting BLC and MLSS; and both terminate
exercise above MLSS intensity rather quickly (Figure 3 and 4). Neither a specific
point on the BLC power curve nor the time to exhaustion at power slightly below
and above MLSS or the corresponding BLC provide a clue why untrained subjects
and recreational athletes seem to tolerate training at intensities between threshold
and MLSS better than high-performance athletes.
However, the highly endurance trained athlete B obviously performed at higher
power than the untrained individual A. Therefore, the test results of subjects A and
B reflect greatly different metabolic demands at similar intensities related to BLC-
oriented intensity domains. Estimation of the relative pyruvate combustion based on
indirect calorimetry during INCP24,96 reveals that subject A oxidizes approximately
30 mmol⋅min–1 of pyruvate at MLSS power of 160 W combined with a maximum
pyruvate combustion rate of approximately 53 mmol⋅min–1 at 270 W at test termi-
nation (Figure 5A). This is substantially less than athlete B. With approximately
29 mmol⋅min–1, athlete B already matches the carbohydrate combustion of subject
A at the intensity where the BLC started to increase above resting level. The cor-
responding power of 200 W was 25% higher than the MLSS power of subject A,
indicating that at the same carbohydrate use athlete B covered only 73% of the total
metabolic rate from carbohydrate whereas subject A relied to 95% on carbohydrate
oxidation (Figure 5B). At MLSS power of 305 W, athlete B matched the carbohy-
drate demand of 53 mmol⋅min–1 (91% carbohydrate oxidation) seen in subject A at
maximum power (Figure 5B). At MLSS intensity, the carbohydrate demand of B
is approximately 75% higher than that of A although both had a fairly similar BLC
response at MLSS intensity during INCP and CP tests (Figure 3 and 4).
Recreational athletes usually train three to five times weekly, equivalent to
3 to 10 h/wk. Ambitious programs for beginners comprise three to four training
sessions per week accumulating 1 to 4 h/wk.77 This is significantly less than high
performance endurance athletes may train. Most top endurance athletes train 15 to
20 h/wk but selected endurance events seem to require up to 30 h of training and
up to 12 training sessions per week.91 The combination of substantial differences
in metabolic demand at given exercise intensity and training volume between
untrained, recreational, and high-performance athletes seems to provide the clue for
different adaptive responses to training at given BLC-related exercise intensities.
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Figure 5 — Glycolytic rate, rate of aerobic pyruvate and fatty acid combustion as a func-
tion of cycling power. A: not specifically trained subject. B: internationally successful
endurance athlete.

   17
 18   Beneke, Leithäuser, and Ochentel

During 20 min exercise at 150 W (60% VO2max, 4.0 mmol⋅L–1), which is

slightly below MLSS intensity, subject A would reduce his muscle glycogen by
approximately 54 g based on a fuel mix of 94% carbohydrate (CHO) and 6% fatty
acid oxidation (FAO). This is equivalent to a decrease of muscle glycogen of 13%
in a well-nourished untrained 75 kg male. Increasing the training volume to 1 h on
4 d/wk would be a hard training regime for this subject but still without any risk
of accumulative glycogen depletion at a normal diet (Figure 6).
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Figure 6 — BLC and daily cycling distance as function of training hours per day; training
BLC levels higher than indicated by the dotted lines A and B or cycling distances higher
than solid lines A and B would cause accumulated unsustainable glycogen depletion. A:
not specifically trained subject—metabolic demand is too low to deplete glycogen stores
unsustainably, irrespectively of during 4 h training per week. B: internationally successful
endurance athlete— metabolic demand is so high that 10 to 15 h of training at BLC levels
between 2 and 4 mmol⋅L–1 is a risk of accumulated glycogen depletion.

Under the assumption that MLSS power is sustainable for up to approximately

1 h of exercise, 15 h of training at MLSS intensity is hardly possible. However,
neither recreational nor top athletes train continuously at the same intensity. For
recreational athletes, a threshold-oriented training regimen had been proposed com-
prising 46% training in intensity domain 1 (Intensity < Threshold), 35% in domain
2 (Threshold ≤ Intensity ≤ MLSS), and 19% in domain 3 (Intensity > MLSS).92
Reducing the training of top athlete B from 15 to 10 h/wk, a training volume
equivalent to that of a very ambitious recreational athlete, could result in 4.5 h at
150 W (37% VO2max, 1.8 mmol⋅L–1, 66% CHO, 34% FAO), 3.5 h at 250 W (58%
VO2max, 3.0 mmol⋅L–1, 84% CHO, 16% FAO) and 2 h at 400 W (90% VO2max, >
7.0 mmol⋅L–1, 98% CHO, 2% FAO) with an accumulated training induced glycogen
demand of 2.1 kg equivalent to 360 km of road cycling per week.97 Assuming a
 Blood Lactate Diagnostics   19

fully recovered muscle glycogen content of 100 mmol⋅kg–1 of wet muscle mass
and five training sessions per week, approximately 78% of muscle glycogen needs
to be refilled during and after every training day. The latter carbohydrate demand
is high but still manageable.98,99 Keeping the latter intensity profile and increasing
the training volume by 50% to 15 h, the resulting 540 km of road cycling per week
is still a relative low training volume for a top endurance athlete. However, this
would extend the training-related glycogen demand to approximately 630 g per
training day. This demand is only sustainable if the training is distributed over two
training sessions per day separated by about 4 to 6 h, combined with a carefully
designed nutritional program maximizing glycogen replenishment and/or substantial
carbohydrate intake during the training session.98,99 At the given intensity profile,
any further increase in training volume would be likely to result in unsustainable
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accumulated glycogen depletion.

Although borderline in terms of sustainability, the latter training volume is
still low for a top endurance athlete training 10 to 12 times per week.92 However,
the volume of training can only increase if the demand of glycogen is kept within
sustainable limits. This is supported by convincing empiric evidence that top ath-
letes dedicate approximately 70 to 90% of their training volume to the intensity
domain 1 (Intensity < Threshold), although the distribution of the remaining training
volume between intensity domain 2 (Threshold ≤ Intensity ≤ MLSS) and domain
3 (Intensity > MLSS) remains less clear. If the weekly training volume of 15 h is
redesigned, for example, to an intensity profile of 80% in domain 1 (Intensity <
Threshold), 5% in domain 2 (Threshold ≤ Intensity ≤ MLSS), and 15% in domain
3 (Intensity > MLSS), then the corresponding weekly glycogen demand decreases
to 2.4 kg and is well sustainable even with seven to nine training sessions per week
but combined with a reduction in the weekly cycling distance to 500 km. However,
if the training volume is pushed to the sustainable limit, the latter intensity profile
provides a reserve to increase training volume of athlete B to roughly 20 h and 650
km/wk. An increase of the training volume to approximately 30 h as seen in some
training weeks of professional cyclists would require that athlete A limits cycling
intensity to a BLC not higher than 1.5 mmol⋅L–1 throughout the entire week. Under
these relative unusual conditions, A could cycle almost 900 km per week (Figure 6).

Conclusions
The BLC is sensitive to changes in exercise intensity and duration. Multiple BLC
threshold concepts define objectively testable but different points on the BLC power
curve of various INCP test programs. The identical threshold concept will provide
different results if the INCP test varies in terms of increase in power over time.
Many BLC thresholds detect a point on the BLC power curve that is between the
lowest power at which the BLC starts to increase above resting BLC and MLSS
power. The MLSS is measured during a series of prolonged CP tests and detects
the highest aerobic power without metabolic energy from continuing net lactate
production. MLSS power is usually sustainable for 30 to 60 min. MLSS power is
highly correlated with the maximum aerobic power, athletic endurance performance,
and BLC thresholds. However, MLSS and MLSS intensity are highly variable
and no threshold concept can predict MLSS power precisely. BLC thresholds are
 20   Beneke, Leithäuser, and Ochentel

useful for cross-sectional or longitudinal endurance assessments as long as the

same threshold concept is used under identical testing conditions.
The idea that training at threshold intensity may be particularly effective is
a myth, and without evidence. Three BLC-oriented intensity domains have been
established: (1) training up to an intensity at which the BLC clearly exceeds rest-
ing BLC, equivalent to light and moderate training focusing on active regenera-
tion or high-volume endurance work mostly lasting between 1 and 6 h (Intensity
< Threshold); (2) heavy endurance training at work rates supposedly up to MLSS
intensity lasting 30 to 90 min (Threshold ≤ Intensity ≤ MLSS); and (3) severe
exercise intensity training between MLSS and maximum oxygen uptake intensity
normally lasting up to 30 min (Intensity > MLSS). During short-term training
interventions in untrained subjects and recreational subjects, a focus on intensity
domain 2 (Threshold ≤ Intensity ≤ MLSS) was comparably as successful as various
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combinations of all three intensity domains. In high-performance endurance ath-

letes, it is essential to link training intensity and volume in order to keep glycogen
homeostasis within sustainable limits. The combination of very high training volume
and high aerobic potential of top-performance endurance athletes reasons that 70
to 90% of their training is dedicated to intensity domain 1 (Intensity < Threshold).
The remaining training is variably spent on domains 2 (Threshold ≤ Intensity ≤
MLSS) and 3 (Intensity > MLSS).

References
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