European Journal of

Eur J Appl Physiol (1982) 49:45-57

Applied
Physiology
and Occupational Physiofogy
9 Springer-Verlag 1982

Changes in Onset of Blood Lactate AccUmulation (OBLA)

and Muscle Enzymes After Training at OBLA*
Bertil Sj6din, Ira Jacobs, and Jan Svedenhag
National Defense Research Institute (Human Studies Department); Department of Clinical
Physiology, Karolinska Hospital; Department of Physiology III, Karolinska Institute,
S-t04 01 Stockholm, Sweden

Summary. Eight well-trained middle and long distance male runners added
to their regular training program a weekly 20-min treadmill run at a velocity
calculated to elicit a blood lactate concentration of 4 mmol x 1-1. i/o2 max,
the running velocity eliciting 4 mmol • 1-1 blood lactate (VoBLA), and the
activities of citrate synthase (CS), phosphofructokinase (PFK), lactate
dehydrogenase (LDH) and L D H isozymes in the M. vastus lateralis were
determined before and after 14 weeks of this training. Significant increases
were observed in VOBLA and the relative fraction of heart-specific LDH,
while the activity of PFK and the ratio of PFK/CS decreased after training.
The change in VOBLAwas negatively correlated to the mean rate of blood
lactate accumulation during the last 15 rain of the treadmill training runs,
and positively correlated to the percentage of slow twitch fibers in the M.
vastus lateralis. The data support the hypothesis that a steady state training
intensity which approximates VOBLAwill increase VOBLA, and will result in
measureable local metabolic adaptations in the active skeletal muscles of
well-trained runners without a significant change in maximal aerobic power.
Muscle fiber type composition may be an indicator of the "trainability" of the
musculature.
Key words: Physical training - Blood lactate - Muscle enzymes -
Anaerobic threshold - Muscle fiber composition

Introduction

It has been proposed that metabolic parameters measured during submaximal

exercise may be better indicators of endurance exercise capacity than the

* Supported in part by grants from the Research Council of the Swedish Sports Federation and the
Swedish Armed Forces Enrolment Board
Offprint requests to: Ira Jacobs, Dr. Med. Sc., Dept. of Clinical Physiology, Karolinska Hospital
(address see above)

0301-5548/82/0049/0045/$ 02.60
46 B. Sj6din et al.

determination of maximal aerobic power [8, 20, 21]. In this regard, the steady
state exercise intensity corresponding to a defined blood lactate concentration
has been investigated [17, 21, 23]. Endurance exercise performance is strongly
correlated to this exercise intensity [11, 29]. Many groups express this variable as
the exercise intensity corresponding to a blood lactate concentration of 4 mmol
x 1-1 [21, 23], and this intensity has been referred to as the onset of blood lactate
accumulation (OBLA) [29, 30]. Muscle metabolic characteristics related to
OBLA include muscle fiber type composition, capillary density, and the balance
between the glycolytic and oxidative enzyme activities [17], while the "lactate
threshold" has been demonstrated to be related to muscle respiratory capacity
[161.
It has been suggested that training intensity which approximates OBLA
would be optimal in eliciting the local changes in muscle metabolism
characteristic of adaptation to training [15, 21, 22]. The purpose of this study was
to examine in trained runners if the addition to their normal training program of
one session a week at OBLA would affect muscle enzyme activities, maximal
aerobic power, and the exercise intensity corresponding to OBLA.

Methods

Eight well-trained middle and long distance male runners participated in the present study. Mean
(range) age, height, weight and percent slow twitsch (% ST) muscle fibers in the M. vastus lateralis
were 19.8 (18-25) years, 182 (175-197) cm, 66.4 (58-75) kg, and 62.3 (45-74) % ST respectively.
Two runners were 800 m specialists while the others competed at distances from 1,500 m to 5,000 m.
All subjects were informed of the experimental procedures and associated risks, and that they were
free to withdraw from the study at any time.
The subjects' training programs were carefully recorded for 18 weeks prior to the OBLA training
period. The weekly training hours at exercise intensities above, below and corresponding to OBLA
calculated at the end of this 18 week period, were 6.1, 0, and 0.4 h respectively for the longer
distance runners, and 2.8, 0.1, 0.2 h respectively for the two 800 m runners. Subjects were asked to
maintain an identical training program during the entire study. This was strictly controlled by one of
the investigators, who also coached the subjects, and was confirmed by written training reports
submitted by the subjects.

Protocol. Immediately prior to and after the specific training period, the treadmill running velocity
eliciting a blood lactate concentration of 4 mmol x 1-1 (VOBLA) was determined as described
previously [29]. While running at the four different velocities to enable the calculation of VOBLA,
expired gases were collected in Douglas bags using open circuit spirometry simultaneously with
blood sampling during the last 60-90 s at each velocity. Expiratory minute volume was determined
with a wet spirometer, and 02 and COz concentrations were analysed with a mass spectrometer
(Centronic MGA200). Oxygen .consumption (Vo2) was calculated, as were linear regression
equations for the relationship of 1/o2 to treadmill running velocity for each subject. Based on these
equations, the Vo2 while running at 15 km x h -1 (Voz 15) and at VOBLA (Vo; OBLA) were
calculated. After a 20-rain rest, maximal oxygen consumption (1)o2 max) was determined using a
continuous protocol on the treadmill. Speed remained constant while the inclination was elevated by
0.5 ~ every 30s. Subjects were driven to exhaustion. Vo2 15, V o 2 0 B L A , and Vo2max were
determined prior to and after the 14 week training period.

Training. During the 14 weeks of the study, subjects reported to the laboratory once a week and ran
for 20 min on the treadmill at VOBLa. This training occurred during the winter months in order to
avoid the competitive season. During this run, blood samples were obtained after 5 and 20 rain for
OBLA and Muscle Enzymes After Training 47

lactate concentration determinations [33]. Based on the 5-rain sample, the treadmill velocity was
adjusted in the subsequent training session in order to approximate as closely as possible the running
velocity which would elicit a lactate concentration of 4 mmol x 1-1.

Test-Retest Reliability. Since no control group was employed in the present study, a "test-retest"
procedure was designed so that the subjects could act as their own controls. VOBLAwas determined
twice prior to the beginning of the VOBLAtraining. The first time was 18 weeks prior to, and the
second time one week prior to the beginning of the VOBLAtraining. After the 14 weeks of VOBLA
training the subjects were again tested twice: the week following the last VOBLAtraining session and
again 18 weeks later. Subjects maintained their routine training programmes as described above,
from the time of the first test 18 weeks prior to VOBLAtraining, until the last test 18 weeks after the
last VOBLAtraining session. Of the seven subjects involved in this test-retest portion of the study, five
are subjects who participated fully with all data reported in this paper, while an additional two
subjects participated fully in the training program but did not agree to have biopsies
performed.

Histochemical and Biochemical Procedures. Muscle tissue samples were taken from M. vastus
lateralis before and after the VOBLAtraining period by needly biopsy [2]. Muscle fiber type frequency
was determined histochemically by myofibrillar ATPase staining [25] after preincubation at a pH of
10.3 [3]. The activities of lactate dehydrogenase (LDH, E.C. 1.1.1.27), phosphofructokinase (PFK,
E.C. 2.7.1.11), and citrate synthase (CS, E.C. 4.1.3.7) were determined with fluorometric
techniques (for references see ref. [17]) on freeze dried muscle fibers which were dissected free of
connective tissue and blood clots. The samples were homogenized by sonication in an ice-cooled
medium of 0.1 M phosphate buffer (pH 7.3). In addition, absolute and relative LDH isozyme
activities were determined with an el~ctrophoretic technique [28] on this homogenate. Enzyme
activities are expressed per wet muscle weight with muscle water assumed to be 77% of total weight
[18].
Student's t-test for paired observations was used to determine the significance of differences
between mean values.

Results

After 14 weeks of training at VOBLAonce a week for 20 min, in addition to their

normal training, a significant increase was observed in VOBLA(Table 1). It is of
interest that the two 800 m runners demonstrated a decreased Vo2 max (from
68.9 to 66.5 ml x kg -1 x min -1) while the other runners demonstrated an
increase from 68.7 to 71.0 ml x kg -1 x min -1 (p < 0.01). ~702 15 was lower in
seven of the eight subjects following training and when expressed relative to Vo2
max all subjects ran 15 km x h -1 at a lower exercise intensity following training
(from 73.8 to 70.1% Vo2 max, p < 0.01, Table 1).
During the controlled VOBLAtraining the mean blood lactate concentration
after 5 min was 4.1 + SD 0.3 mmol x 1-a and increased to 5.9 _+ 1.0 mmol x 1-1
after 20 min of running9 Thus, the mean rate of blood lactate accumulation
during the last 15 min of running was 0.12 + 0.06 mmol x 1-1 x min -t. The
changes in VOBLAobserved after the OBLA training correlated negatively with
this rate of lactate accumulation during the training sessions (r = -0.81, p <
0.01) (Fig. 1) and positively to % ST (r = 0.83, p < 0.01) (Fig. 2).
The mean activity of PFK decreased significantly while the activities of total
LDH and CS were unchanged (Table 2). However, the longer distance runners
exhibited an increased CS activity (+ 22%) (p < 0.01), if their data are examined
separately from the two 800 m runners who demonstrated a decreased CS
activity (-13%) 9 In addition, the decrease in PFK activity was 50% of the
Table. 1. 902 max, oxygen cost of running 15 km x h-* (//o2 15) in absolute terms, and relative to ~ZQ2max (~/rQ215 x ]Zo2 max -1, %), the running velocity and g
Vo2 corresponding to 4 mmol x 1-1 blood lactate (VoBLAand Vo20BLA), V O 2 0 B L A relative to V02 max (V02 OBLA x Vo2 max -1, %) before and after 14
weeks of "VoBLA training" in six long-distance and two middle-distance runners

Subject Vo2 max Vo2 max V02 15 .V02 15 x VOBLA V02 OBLA V02 OBLA x
1/o2 max -1 Vo2 max <

pre post pre post pre post pre post pre post pre post pre post

1 x min -1 ml x kg -1 x min -1 mlxkg-lxmin 1 % m x s -1 m l x k g - l x m i n -1 %

1 4.65 4.95 72.5 75.4 48.9 48.7 67.5 64.6 4.88 5.24 61.7 65.3 85.1 86.6
2 4.24 4.58 68.8 72.2 45.2 44.7 65.4 61.9 5.03 5.29 59.4 63.3 86.4 87.7
3 4.48 4.55 66.6 68.9 49.7 50.3 74.7 73.0 4.67 4.66 57.3 58.6 86.0 85.0
4 4.85 4.91 64.5 64.9 51.3 50.3 79.5 77.5 4.55 4.84 56.9 61.1 88.2 94.2
5 3.77 4.16 67.7 71.5 53.5 52.4 79.0 73.3 4.78 4.88 61.5 62.5 90.9 87.4
6 4.54 4.58 71.8 73.1 49.3 46.7 68.8 63.9 4.83 4.95 59.7 59.4 83.1 81.4
7a 4.86 4.66 68.4 65.1 51.9 48.7 75.8 74.8 4.52 4.74 56.7 56.7 82.8 87.1
8~ 4.92 4.71 69.3 67.9 55.0 51.4 79.3 75.7 4.22 4.52 55.5 56.4 80.2 83.0

Mean 4.54 4.64 68.7 69.9 50.6 49.2 73.8 70.1 4.69 4.89 58.6 60.6 85.3 86.6
SD 0.39 0.29 2.6 3.8 3.0 2.5 5.7 2.6 0.25 0.27 2.3 3.2 3.4 3.8
p n.s. n.s. < 0.05 < 0.01 < 0.01 < 0.05 n.s.

a Middle-distance runner

C3:
OBLA and Muscle Enzymes After Training 49

.40 r = -0.81
p< 0.01
y= - 1.70x §
.35

.30

25

t~ .20

<
m .15

<~

.10

.05

010s o'1o 0.'is 0120 012s

Rate of blood l a c t a t e a c c u m u l a t i o n mmolxl'lxmin "1
Fig. 1. The relationship of the change in VOBLAafter the 14 weeks of VOBLAtraining to the mean rate
of blood lactate accumulation during the weekly 20-rain treadmill run. The two 800 m runners are
represented by open circles

pre-training values in the 800 m runners while only a 21% decrease was observed
in the longer distance runners. The end result was a decrease in the PFK/CS ratio
in all runners.
A shift in the LDH isozyme pattern was also observed. The relative activity
of the heart specific LDH isozyme (H-LDH) had increased in all subjects, due to
either an increase in absolute H-LDH activity or a decrease in muscle specific
LDH (M-LDH) or both (Table 2). The changes in VOBLAdid not covary with the
changes in enzyme activity. However, the change in absolute VO2 ]-5 was
correlated with the increase in relative activity of H-LDH after VoBI~Atraining
(r = - 0 . 7 5 , p < 0.05, Fig. 3).
The test-retest correlation coefficients for VOBLApre- and post-training were
0.90 and 0.89 respectively (Fig. 4). A significant change in Vo~I~Awas observed
only after the VoBi~a training period (p < 0.01, Fig. 5).

Discussion
The present study describes how training at an exercise intensity corresponding
to 4 m m o l • 1-1 blood lactate once a week for 20 rain affects various muscle
enzyme activities and the exercise intensity corresponding to OBLA in
50 B. Sj6din et al.

/
0.3 ~ /
r : 0.83
p< 0.01
= , - .

0.2

w
x
E

0.~

~.r I
50 60 7 80
%ST

Fig. 2. The relationship of the change in VOBLA after the 14 weeks of VOBLA training to the
percentage of slow twitch muscle fibers (% ST) in the M. vastus lateralis. Closed circles represent the
long distance runners and open circles represent the 800 m runners

well-trained runners. The changes observed in the different enzyme markers

were smaller than those reported after training of less well-trained subjects [12,
31]. None the less, following the training period in the present study, the rate of
glycogenolysis during exercise would be relatively slower and the potential to
oxidize pyruvate and/or lactate increased. This is suggested by the decreased
PFK/CS ratio and the increased relative activity of the H-LDH isozyme. It has
been previously demonstrated that H-LDH bound to the inner membrane of the
mitochondrion will facilitate an oxidative translocation reaction of lactate in the
presence of NAD [32]. The net result should be lower muscle and/or blood
lactate accumulation at a given absolute and relative exercise intensity, as has
been previously demonstrated [10, 19], and consequently an increase in
VOBLA.
The lack of correlation between the changes in VOBLA and the changes in
enzyme activities may be a function of differences between the variables in the
 9

g~

e~

Table 2. Enzyme activities before and after 14 weeks of " V OBLA training" in six long-distance and two middle-distance runners

Subject CS PFK PFFUCS LDH M-LDH H-LDH H-LDH

pre post ;>

pre post pre post pre post pre post pre post pre post

tool x g 1 x rain -a x 10 -6 mol x g-1 x min -1 x 10 -4 %

P~
1 4,67 5.45 6.69 5.33 1.43 0.98 0.73 0.74 0.32 0.33 0.41 0,41 55.8 55.9
2 5.62 6.37 5.73 5.53 1,02 0.87 0.52 0.74 0.21 0.27 0.31 0.47 59.6 63.3
3 4.86 5.40 6.89 5,32 1.42 0.99 0.83 0,75 0.43 0.37 0.40 0.38 48.6 51.1
4 4.86 7.56 4.78 5.60 0.98 0.74 0.87 1.54 0.50 0.82 0.37 0,72 42.2 46.8
5 5.27 5.64 9.97 6.00 1.89 1.06 1.56 1,00 1.09 0.60 0.47 0.40 30,2 40.1
6 4.96 6.36 7.57 5.30 1.53 0.83 0.63 0.76 0.35 0.31 0.28 0.45 44.5 59.2
7a 7.24 6.26 9.94 5.42 1.37 0.87 1.05 0.81 0.62 0.44 0.43 0.37 41.1 46.0
8a 5.53 4.87 9.62 4.32 1.74 0,89 0.85 0.49 0.47 0.18 0.38 0.31 44.7 62.5

Mean 5.38 5.99 7.65 5.35 1.42 0.90 0.88 0.85 0.50 0.41 0.38 0.44 45,8 53.1
SD 0.83 0.83 2.00 0.48 0.31 0.10 0.32 0.31 0.27 0.21 0.06 0.12 9.1 8.5
p n.s. < 0.05 < 0.001 - n.s. n.s. n.s. < 0.025

Middle-distance runner
 52 B. Sj6din et al.

A % H-LDH

-t r 9
9~ " p < 0.02
y =-0.18x-0.13
.E
E
x
-2
"7

x
w
E -3

o"
.>
-4
<~

-5

Fig. 3. The relationship of the change in r)o2 while running at 15 km x h -1 after the 14 weeks of
VOBLA training to the change in the relative proportion of total LDH activity consisting of the
H-LDH isozyme

5.5
PRE- TRAINING
5.5 POST TRAINING
-

9/

~
50 9

~--/,
4.5
I I
5.0
J
5.5
4s~'~4's so s15
VOBLA, m x s -~ Test I VOBLA , m x s -1 Test I

Fig. 4. Test-retest values for pre- and post-VoBLA training

OBLA and Muscle Enzymes After Training 53

5.5

5.0

x
E

>~m
4.5 '~Vo BL A -
training"

4.0
/
0 18 32 50
Weeks

l~go 5. VOBLA(mean, SD) measured twice before and twice after 14 weeks of "VOBLAtraining" in
seven well-trained runners. The "xx" indicates a significant difference (p < 0.01) from the previously
measured value

time course of their respective changes. Such differences have been reported
earlier with regard to the training-induced changes in muscle enzymes vs. Vo2
max [12, 14, 31]. In an effort to minimize the number of muscle biopsies, the
post-training biopsy was taken after 14 weeks of OBLA training. Five of the
subjects performed the protocol to determine VOBLA after only 7 weeks of
training and had achieved 66% of the change observed after 14 weeks. Whether
or not the changes in VOBLAobserved at that time were correlated to changes in
enzyme activities is speculative.
Changes in capillary density and/or enzymatic adaptation in other muscles
used in running may also have contributed to the lack of covariation observed
between VOBLA and enzyme activities. The M. vastus lateralis was the muscle
sampled in the present study. Significant recruitment of this muscle during level
running has been previously demonstrated [4, 7] although the gastrocnemius and
soleus muscles are probably stressed to a greater extent [6]. Thus the changes in
enzyme activities observed in the vastus lateralis tissue may have been even
more marked in the gastrocnemius and soleus.
An increase in VOBLA of 0.20 m x s -1 could also be partially due to an
improved running economy after training. This is suggested by the decreased
 54 B. Sj6din et al.

76

,3 9 '=~ 8.00,
,_x ,+/
! . /7
,

!ZOO /

i>o67 /
f" o 9 . / o

I 9

s.~176
If-~// 1 I I I
5.00 6.00 7.00 8.00
Pre-Vo2max, ml,~kg "~• min "1 Pre- CS, mmolxg'l x rain-1• 10-6
Fig. 6. Pre- and post-training values for maximal oxygen consumption ( V o 2 max) during treadmill
running and the activity of citrate synthase (CS). The two 800 m runners are represented by open
circles and the longer distance runners by closed circles

oxygen consumption while running at 15 km x h -1. We cannot ascertain,

however, what proportion of this decrease is due to. an improved running style.
The significant relationship between changes in 11o2 15 and changes in the
relative activity of H-LDH may indicate that the improved running economy was
at least partly due to an improved intracellular oxidative capacity, which would
increase the rate of lactate oxidation.
The two 800 m runners deviated from the longer distance runners in their
response to VOBLAtraining. This is exemplified in Fig. 6, where it is shown that
these two runners are the only ones to demonstrate a decrease in both 17o2 max
and CS activity following the training period. In spite of this deviation, the extent
of the changes in VOBLA w e r e of the same magnitude as those observed in the
other subjects.
Hollman et al. [15] have recently reported that training at OBLA for 30 rain,
five times weekly will s!gnificantly increase VOBLA, while training at a higher
exercise intensity (95% Vo2 max) with a similar duration and frequency results in
no changes after 6 weeks of training. Mader [22] conceives of the exercise
intensity corresponding to 4 mmol • 1-1 lactate as representing a maximal
threshold for the maintenance of a steady state and therefore as an optimal
training intensity. This concept is supported by Kindermann et al. [21], who
described the effects on blood lactate of prolonged running at a constant speed
which initially elicits the 4 mmol • t -1 level. Indeed, in the present study, the
greatest increases in VOBLA w e r e observed in the subjects with the lowest rate of
OBLA and Muscle EnzymesAfter Training 55

blood lactate accumulation during the training session. This suggests that within
the limitations of the present study the training effect was greatest when a
"steady state" blood lactate concentration approximating 4 mmol • 1-1 could be
maintained during the training sessions. The variation among subjects in the rate
of blood lactate accumulation suggests that individual "steady state" lactate
levels may deviate from the 4 mmol x 1-1 level in some cases.
Successful athletes in endurance running possess a higher frequency of slow
twitch muscle fibers in the leg musculature than do athletes participating in
non-endurance sports or untrained subjects [1, 5, 13]. The present results
suggest that % ST may not be a characteristic which determines success in
endurance sports as much as it is an indicator of the potential "trainability" of the
musculature. This is demonstrated by the positive relationship observed
between % ST and the increase in exercise intensity required to elicit a blood
lactate accumulation of 4 mmol x 1-1 after the training period. Orlander et al.
[24] have reported a similar relationship between % ST and the increase in
Vo2 max observed in sedentary men after 7 weeks of low intensity training.
A limitation of the present study is the lack of a control group of subjects
tested at the same intervals as the experimental group. However, the
experimental subjects have essentially acted as their own controls with the
test-retest design both preceding and following the training part of the present
study. The high test-retest correlation coefficients and the insignificant
differences between test-retest values are evidence of the high reproducibility of
VOBLAin well-trained subjects. Similar strong test-retest correlation coefficients
were calculated by Davis et al. [9] for another indicator of blood lactate
accumulation during steady state exercise - the anaerobic threshold.
The subjects in the present study were well-trained competitive runners, in
contrast to other training studies which have examined changes in enzyme
activities and oxygen transport capacity in less well-trained or sedentary subjects
[12, 14, 31]. Daniels et al. [8] suggest that factors representing local muscular
metabolic characteristics should be considered when attempting to evaluate the
adaptation to training in trained subjects. This is in the light of their finding that,
after training well-trained runners at an intensity higher than their normal
training program, performance, improvements were noted without any signif-
icant alteration of the runners' Vo2 max. This may be due in part to the inverse
relationship existing between initial physical training status and the increase
observed in maximal aerobic power following a training program [26, 27]. The
present group of subjects increased their Vo2 max by 3% at the most, while
changes in VOBI.A ranged from 0 to 7% of pre-training values. Based on the
observed changes in muscle enzyme activities and VOBLA, and the established
strong correlation between VOBLAand endurance exercise performance, we feel
that for the purpose of laboratory experiments VOBLAmay be a more sensitive
parameter for the monitoring of adaptations to training than is the determination
of V02 max.
56 B. Sjrdin et al.

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Accepted February 1, 1982