European Journal of
Eur J Appl Physiol (1985) 5 4 : 1 - 6
Plasma renin activity and serum aldosterone
during prolonged physical strain
The significance of sleep and energy deprivation
P. K. Opstad 1, O. Oktedalen 1, A. Aakvaag 2, F. Fonnum 1, and P. K. Lund 3
I Norwegian Defence Research Establishment Division for Toxicology, PO Box 25, N-2007 Kjeller
2 Hormone and Isotope Laboratory, Aker Hospital, Oslo 6
3 Hedmark Central Hospital, Division for Clinical Chemistry, N-2400 Elverum, Norway
Summary. Plasma renin activity (PRA), serum
aldosterone and the serum and urinary levels of
sodium and potassium have been investigated in 24
young men participating in a 5-day military training
course with heavy continuous physical exercise,
energy and sleep deprivation. The subjects were
divided into three groups. Group 1 did not get any
extra sleep or food, group 2 were compensated for
the energy deficiency, and group 3 slept 3 h each
night. The basic diet given to all the subjects was
about 5,000 kJ and 2 g NaC1 9 24 h -1 9 cadet -1. The
high calorie diet contained approximately
25,000-35,000 kJ and 20g of N a C 1 - 2 4 h -~. ca-
det -1.
The study showed that serum aldosterone and
P R A were extremely activated during such prolonged
physical strain combined with lack of food and salt,
whereas sleep deprivation did not seem to have any
large influence. Only small variations were found in
the serum levels of sodium and potassium and the
urinary level of potassium during the course, whereas
a decrease was seen in urinary sodium concentration.
The fairly good correlations between the decrease in
urinary sodium levels and the increase in PRA
(r = 0.7) and further between P R A and serum aldo-
sterone (r = 0.8) during the course indicate that there
is a causal connection between the decrease in urinary
sodium excretion and the increase in PRA and serum
aldosterone. An increased response was seen during a
short term exercise test in groups 1 and 3, whereas no
such increase was seen in the subjects of group 2,
probably because the higher salt intake abolishes the
renin-aldosterone response to exercise. In spite of
high catecholamine levels during the course, the
serum potassium response to physical exercise was
unchanged. The circulating catecholamines did not
seem to have any significance for renin secretion and
Offprint requests to: P. K. Opstad at the above address
Applied
Physiology
and Occupational Physiology
9 Springer-Veriag 1985
sodium reabsorption during the course. Further,
there was no sign of exhaustion in Na/K homeostasis
during the experiment.
Key words: Stress - Physical exercise - Sleep
deprivation - Fasting - Renin activity - Aldoste-
rone - Natrium - Kalium
Introduction
The renin-aldosterone system .is considered to be a
major regulatory system for the excretion of sodium
and potassium (Erlich 1979) and has been shown to
be activated during short-term exercise as well as
during long term exercise lasting for some hours
(Bozovid et al. 1967; Castenfors et al. 1967; Kotchen
et al. 1971; Sundsfjord et al. 1975; Adlercreutz et al.
1976; Costill et al. 1976). The plasma renin activity
(PRA) and aldosterone have also been shown to
increase during salt deprivation, dehydration, hypo-
volemia (erlich 1979), heat exposure (Kosunen et al.
19761 Bailey et al. 1972; Francis et al. 1978) and
psychosocial stress (Clamage et al. 1977).
There are no prior reports on alterations in Na/K
homeostasis, P R A and serum aldosterone during
prolonged multifactorial stress lasting for several days
and including heavy physical exercise, lack of sleep
and energy. To produce this experimentally, we have
used as a model the annual ranger training course of
The Norwegian Military Academy, which exposes
their cadets to continuous heavy physical activities for
5 days with minimal amounts of sleep and food.
During the course we have also studied if such
exhausting activities could lead to any sign of
exhaustion in Na/K balance by testing the PRA and
serum aldosterone responses to a standardized bicy-
cle exercise test.
2 P.K. Opstad et al.: Salt balance regulation during stress

Materials and methods was performed indoors some months after the course with a room
temperature of about 21 ~ C. During the course, the exercise test
was performed outdoors in the training area at about 500 m
The ranger training course. The activities started on Monday at
altitude. The weather was fairly good with a temperature of about
08 : 00 in a forest area at about 500 m altitude. The cadets were
20~ C. Blood samples were drawn just before the test with the
exposed to continuous heavy physical exercise until the following
subjects sitting on their bicycles, and at the end of the exercise
Friday at about 18 : 00. The weather was fairly good during the
while still bicycling, and again after 15 and 25 min of recovery with
course with a maximum temperature of about 25 ~ C during the
the subjects recumbent. During a previous course (Opstad et al.
daytime and 5-10~ C during the nights. The continuous activities
1980; Rognum et al. 198l) we performed a 30-rain exercise test of
represented about 35% Vo2 max as a mean, and allowed for only
50% of Vo2 max"Blood samples were obtained at the following time
short periods of sleep estimated at a total of 1 - 3 h in the 5
schedule, 0, 15, 30, 35, 40, 70 rain. Serum was obtained for the
days.
measurement of sodium and potassium.
Continuous heart rate recordings have shown that each cadet
expended 35,000-45,000 kJ - 24 h -1 as a mean (Aakvaag et al.
Chemical analysis. Serum and urinary concentrations of sodium
1978).
and potassium were measured by flame photometry. Within assay
variation for sodium was 0.3% and for potassium 0.9%. Between
The subjects. Twenty-four cadets of the Norwegian Military
assay variation for sodium was 0.6% and for potassium 0.9%. PRA
Academy participated in the ranger training course as part of their
and aldosterone were estimated by radioimmunoassay (Sundsfjord
school program. The cadets had already studied at the Academy
1975).
for 1 year and were in excellent mental and physical condition.
They were divided into three groups at random: Group 1 consisted Statistics. The results are represented as mean + SEM. The results
of nine subjects with a mean age of 22.7 years (range 21 -25). They within the same group were treated according to the Wilcoxon
were exposed to continuous and heavy physical exercise, sleep signed rank test, and differences between the two groups were
deprivation and excessive calorie deficiency as described. treated to Wilcoxon rank-sum. Correlation coefficients were
Group 2 (n = 7) with a mean age of 24.8 years (range 22-27) calculated by the method of least squares. P-values < 0.01 were
were exposed to the same stress factors as group 1, but in addition considered statistically significant.
they received a special diet containing about 25,000-35,000
k J . 24 h -1 for each cadet. Group 3 (n = 8) with a mean age of 23.6
years (range 22-25) were similar to group 1 but were in addition
given 3 h of sleep each night between 00 : 00 and 06 : 00. When this Results
group was allowed to sleep, the subjects of the other two groups
had only very light activities. Plasma renin activity ( P R A )
(nmol Angiotensin I (AI) 9 1-1 9 h -z)
The diet. The basic food intake for each subject contained about
50 g of protein, 30 g of fat and 120 g of carbohydrate, representing
about 4,000-5,000 kJ 924 h -a. Group 1 and group 3 thus had a A 15-30 fold increase was found in the plasma levels
caloric deficiency of about 30,000-40,000 kJ. 24 h -1. The salt o f P R A d u r i n g t h e c o u r s e ( F i g . 1). F o r g r o u p 1 t h e
intake of the two groups was estimated to be approximately 2 g of PRA levels increased from 0.23 + 0.02 to a maximum
NaC1 and 3 g for KC1/day. In contrast, the special compound diet
o f 7 . 9 + 1.1 o n d a y 3, a n d t h e r e w a s a g r a d u a l
given to group 2 contained about 100 g proteins, 120 g fat, 1,200 g
carbohydrates, 2 0 - 3 0 g of NaC1 and 2 0 - 3 0 g of KCl, representing decrease in plasma level towards the end of the
about 25-30,000 kJ - 24 h -I for each cadet. A normal daily food c o u r s e . F o r g r o u p 3 t h e l e v e l i n c r e a s e d d u r i n g t h e 1st
consumption of the Norwegian population contains about 10-15 g d a y o f a c t i v i t i e s f r o m 0 . 2 8 + 0 . 0 8 t o 6.3 + 1.4 a n d
of NaCI and 3 - 4 g of KC1, which is considered to be a relatively from then on a constant level was kept throughout the
high degree of salt intake. This diet was given in the form of soup,
rest of the course. In group 2 the concentration
orange juice, cocoa, and milk shakes. The high carbohydrate
content was achieved by giving a substrate easily soluble in water, i n c r e a s e d f r o m 0 . 2 + 0 . 0 2 t o 2 . 9 +_ 1.0 d u r i n g t h e f i r s t
nearly tasteless and colourless (maltodextrin). Thus the members 24 h o f a c t i v i t i e s , a n d t h i s l e v e l w a s m a i n t a i n e d u n t i l
of group 2 were nearly in calorie balance, with no significant loss of d a y 5 w h e n a s i g n i f i c a n t d e c r e a s e t o 1.0 + 0.5
body weight in contrast to groups 1 and 3 (the low calorie groups)
was found. Group 1 and group 3 had significantly
where each cadet lost approximately 4 kg. The liquid intake of each
cadet during the course was estimated to be about 6 1/day. higher levels throughout the course when compared
t o g r o u p 2.
Blood sampling. Blood samples were taken each morning between
0 6 : 0 0 and 0 9 : 0 0 h . The blood samples were taken through
Veneflon| cannulae in vacuum tubes. Each sample of blood was The serum levels o f aIdosterone ( p m o l . l -~)
about 40 ml. Blood for plasma was collected in heparinized
ice-chilled tubes which were centrifuged immediately in a refri-
The serum levels of aldosterone increased dramati-
gerated centrifuge. Blood for serum was allowed to clot and then
centrifuged. The samples were frozen on dry ice immediately after cally during the course and followed a pattern very
centrifugation, and kept frozen at - 8 0 ~ until analyzed. s i m i l a r t o t h a t o f P R A ( F i g . 1) ( r = 0 . 7 ) .
A sixfold increase was observed in group 1 on day
Bicycle exercise test. The maximum oxygen uptake (Vo2ma~) was 3 with a subsequent gradual decrease. Group 3
estimated for each cadet. This was done before the course by reached a maximum on day 2 and maintained a
measuring the heart rate of each cadet during a bicycle ergometer
constant level throughout the course. Group 2
test at three maximal work loads. During the course on day 3 the
cadets were tested for 15 rain on the bicycle with a workload of showed a significantly lower increase during the
60% of Vo2~a (workload 100-130 W). The control experiment course when compared with the other groups.
P. K. Opstad et al.: Salt balance regulation during stress

GROUP I
GROUP 2
PRA and serum aldosterone response
GROUP 3 o I ~ to the bicycle exercise test
T
P R A increased significantly during the exercise test
7-

O-
(Fig. 2). This response (nmol AI 1-1. h -1) was
PLASMA BENIN
ACTIVITY 5-
increased greatly during the course from 0.69 to 4.3 in
.tool A l -F 1 h "I ~-
group i and from 1.5 to 3.4 in group 3, whereas a
3-
2-
reduced response from 1.6 to 0.7 was seen in group 2.
The percentage increase was reduced in all
groups.
2500.
In the control experiment all levels were normal-
ized within 20 min of recovery, in contrast to those
2OOO- levels taken during the course which were signifi-
cantly higher after 25 min of recovery than before the
ALDOSTERONE 1500-

pmol.I "l test.

I000-
i ~ ~ ~ ~ z4 GAYS
Fig. 1. Changes in ~.hemorning levels of plasma renin activity and
serum aldosterone during a 5-days ranger training course in young
male cadets. The subjects were divided in three groups. Group 1
was both sleep- and calorie deprived. Group 2 was compensated to
the calorie deficiency and group 3 was given 3 h of organized sleep
each night. The results are given as mean (_+ SEM). Day to day
variations significant at P < 0.01 are indicated by a thich line while
no significant alterations as indicated by a dotted line
GROUP 1 GROUP 2
PLASMA BENIN
ACTIVITY
nrnol A z .I "l .h "t
In the control experiment, aldosterone showed a
2-fold increase during the 15-min bicycle test. During
the course, the aldosterone response to the same
exercise test was increased 3 - 4 - f o l d in group 1 and 3
and in group 2 a decrease was seen.
The plasma levels of catecholamines
There was a gradual increase in plasma levels of
nor-adrenaline during the course from 1.1 _+ 0.1
nmol 9 1-1 on day I to 11.4 + 1.0 nmol 9 1-1 on day 5.
Small but significantly lower levels were found on
days 3 and 4 in group 2.
GROUP 3 Adrenaline also showed a gradual increase during
the course from 0.38 + 0.02 nmol 9 1-1 on day i to
1.27 + 0.25 n m o l . 1-1 on day 5. Significantly lower
levels were found for group 2 on days 3 and 4.

The serum levels of sodium and potassium retool. 1-1

(Table 1)

The serum levels of sodium and potassium showed

3500. only small variations during the course. However,
CONTROL
EXPERIMENT
UUBINGTHE
group 2 tended to have slightly higher sodium levels
3000- COURSE
and lower potassium levels when compared with
SERUM 2500- groups 1 and 3.
ALDOSTERONE
pmol '1"1 2000-

Urinary excretion of sodium and potassium mmol 91-1

iOOO
500 ~
EX~R~E RECOVERY EXERCISE RECOVERY EXERCISE RECOVERY
MINUTES
Fig. 2. Alterations in the response of plasma renin activity and
serum aldosterone to a standardized bicycle exercise test with 60%
of Vo2 max during the course (for details, see Fig. 1). Blood samples
were taken before and at the end of 15 rain of exercise and during
the recovery period after 25 and 40 rain
(Table 2)
A dramatic decrease was found in the urinary levels
of sodium from 100 + 15 to 10 + 3 on day 4 in group 1
and from 144 + 19 to 32 + 5 in group 3, whereas the
decrease in group 2 was from 139 + 19 to 52 + 23.
The urine volume of the subjects decreased by about
5 0 - 7 0 % during the course, the decrease in urinary
sodium excretion therefore appeared to be even more
pronounced. There was no significant alteration in
4 P.K. Opstad et al.: Salt balance regulation during stress

Table 1. Shows the changes in the sodium and potassium levels in serum and urine during the course (see Fig. 1). Because of technical
reasons the samples were not obtained on all days during the course

Day 1 Day 2 Day 3 Day 4 Day 5 Day 24

1. Serum
a) Sodium mmol. 1-1
GR 1 (stress) 139.7+ 0.9 136.1 0.4 138.0-- 0.8 139.4+ 0.5 137.3+ 1.1
GR 2 (isocalorie) 139.7 + 0.6 142.1 1.2 140.9 _+ 1.7 139.3 _+ 0.6 140.0 + 0.7
G R 3 (sleep) 139.0 1.1 137.0 0.9 136.3 + 1.4 135.9 + 0.8 137.9 0.6
b) Potassium mmol. 1-1
GR 1 4 . 3 + 0.1 4.1_+ 0.1 4.8+ 0.1 4 . 5 + 0.1 4 . 2 + 0.1
GR2 4 . 1 + 0.1 3.9_+ 0.1 4.5 0.2 4 . 0 + 0.1 4 . 0 + 0.1
GR 3 4.4_+ 0.1 4.1 0.1 4.4 0.1 4.3 _+ 0.1 4.2_+ 0.1

2. Urine
a) Sodium mmol 91-1
GR 1 100.5 + 14.5 9.8 -+ 3.4 51.0 _+ 6.7 130.3 26.4
GR 2 139.2 + 19.0 51.9 _+ 22.5 69.3 + 22.5 121.0 +_ 14.0
GR 3 143.9 + 18.9 32.0 _+ 7.5 43.5 3.5 115.3 _+ 12.4
a) Potassium mmol. 1-a
GR 1 36.8 + 6.6 56.4 5.5 63.8 + 5.7 52.6 + 4.8
GR 2 50.7 11.6 54.3 _+ 11.1 58.1 12.6 70.0 17.4
GR 3 49.7 +_ 8.7 54.3 _+ 4.2 60.1 +_ 3.8 59.3 _+ 7.8

Table 2. The serum levels of sodium and potassium during 30 min of bicycle exercise corresponding to 50% of Vo2 max before and on day 3
and day 5 during the course. Blood samples were obtained before and after 15 and 30 min of exercise with the subjects sitting on the bicycle,
and further after 5, 10, 20, and 40 min of recumbent recovery

Exercise Recovery

0 15 30 35 40 50 70

Sodium mmol 91 1
Control exp 135.1 + 1.1 138.0 -+ 1.0 136.2 + 1.1 136.0 + 0.8 138.6 + 0.9 138.2 + 0.9 138.0 + 1.1
Day 3 134.9 _+ 1.1 134.6 +__0.6 134.5 + 1.0 138.0 + 2.4 134.6 + 0.6 135.5 + 0.5 135.1 + 0.8
Day 5 135.7 + 0.7 135.0 ___1.8 134.9 + 1.1 135.1 + 1.9 133.0 + 2.9 137.7 + 2.2 132.9 2.4

Potassium mmol- 1-1

Control exp 4.4 + 0.1 4.8 ___0.1 4.8 _+ 0.1 4.2 + 0.1 4.2 + 0.1 4.2 + 0.1 4.2 _+ 0.1
Day 3 4.3 _+ 0.1 4.7 0.1 4,7 + 0.6 4.5 + 0.1 4.3 + 0.1 4.2 +_ 0.1 4.2 + 0.1
Day 5 4.1 _+ 0.1 4.5 + 0.1 4.5 + 0.1 4.1 +_ 0.1 4.1 + 0.2 4.0 + 0.1 3.7 + 0.1

the potassium levels in the urine. Taking into account Correlations

that the mean urinary volume decreased greatly
during the course, the total urinary excretion of
p o t a s s i u m is p r o b a b l y a l s o d e c r e a s e d .
The sodium and potassium response
to a 30-rnin exercise test
D u r i n g a p r e v i o u s c o u r s e ( O p s t a d e t al. 1 9 8 0 ) , w e
studied the changes in sodium and potassium during a
3 0 - r a i n e x e r c i s e t e s t w i t h 5 0 % o f 1/o2 max" O n l y s m a l l
variations were found in the serum levels of sodium
and potassium, and the potassium response to the
exercise was unchanged during the course.
There was a high degree of correlation (P < 0.01)
between the changes in PRA and serum aldosterone
during the course, r = 0.7-0.8. There was also a
fairly high correlation between the increase in PRA
or serum aldosterone and the decrease in urinary
sodium concentration, r = 0.6-0.7. On the other
hand there were no significant correlations between
the serum levels of sodium or potassium on one side
and serum aldosterone, PRA or the urinary concen-
tration of sodium and potassium on the other side.
Furthermore there was no significant correlation
b e t w e e n t h e s e r u m levels o f a l d o s t e r o n e o r P R A a n d
plasma catecholamines during the course. During the
P. K. Opstad et al.: Salt balance regulation during stress
exercise test there were no significant correlations
between the pre-exercise levels and the response to
exercise either in the control study or during the
course. Similarly, there were no significant correla-
tions between the aldosterone-renin response to
exercise and the plasma levels of sodium or potas-
sium.
Discussion
The present investigation shows that the regulation of
serum sodium during prolonged strain is so good that
even large alterations in salt intake or losses will be
compensated for by an almost complete reabsorption
of sodium in the tubules of the kidneys. A significant
decrease in the urinary sodium excretion was found
even in the subjects receiving a high calorie diet,
indicating that 20-30 g of salt each day was not
sufficient to cover the subjects losses of salt during
such extreme conditions, without activating salt
sparing mechanisms. One of the most important
regulating mechanisms for sodium and potassium
homeostasis is the renin-aldosterone system, which is
shown to be activated strongly during the present
course. This activation is due mainly to salt deficien-
cy, since the subjects receiving the high calorie and
high salt diet had significantly lower levels of serum
aldosterone and PRA during the course compared to
the others.
The increased response of PRA and serum
aldosterone to the exercise test in the subjects
receiving the low calorie and low salt diet compared
to the subjects receiving the high calorie and high salt
diet, and compared to the control experiment,
showed that the renin-aldosterone responses to
short-term physical exercise were dependent on the
salt balance of the subjects. This conclusion is in
accordance with other investigations showing that a
salt overload extinguishes renin-aldosterone activa-
tion during physical exercise (Aurell et al. 1971;
Engquist et al. 1978; Leenen et al. 1978). The
increases in PRA and serum aldosterone during the
course are therefore due to a combination of the
effects of salt deficiency and physical exercise,
whereas sleep does not seem to play any major part.
Hypovolemia and dehydration are known to stimu-
late the renin-aldosterone system (Dumoulin et al.
1982). Hypovolemia has probably only a minor role,
because plasma volume alteration has been shown to
be less than 10% during previous courses (Opstad et
al., unpublished). In contrast, dehydration may have
some significance because the subjects complainted
of thirst during the course in spite of unrestricted
water access. However, this might also be explained
by the increase in angiotensin II levels in plasma,
which is known to stimulate thirst centers in the
hypothalamus. There was a good correlation between
the serum levels of aldosterone and PRA before
(r=0.75) as welt as during the course (r=0.71),
suggesting that the main regulatory mechanism for
serum aldosterone was PRA and angiotensin II.
ACTH and serum potassium, which also is known to
regulate serum aldosterone (Erlich 1979; Espiner et
al. 1978), did not seem to play any major role because
there was no significant correlation between serum
aldosterone and cortisol or serum potassium (Opstad
et al. 1985). Furthermore, there was a good corre-
lation between serum aldosterone or PRA and the
urinary concentration of sodium during the course
(r=0.6-0.7), indicating a causal connection. There
were no significant correlations between the simul-
taneous determination of circulating nor-adrenaline,
adrenaline or dopamine and PRA or serum aldo-
sterone (Opstadt et al. 1985). This is well in accordance
with other investigators, who found that even large
alterations in plasma catecholamines gave only small
variations in PRA (Johnsen et al. 1982). The lack of
correlation between the serum levels of sodium or
potassium and PRA or serum aldosterone suggests
that alterations in the serum levels of sodium and
potassium do not have any significance for increased
activation of the renin-aldosterone system. If, how-
ever, aldosterone is the major regulatory mechanism
for sodium reabsorption during the course, it is
surprising to find such small variations in the serum
and urinary levels of potassium, because normally
aldosterone exchanges sodium for potassium in the
distal tubules. Sodium might, however, be exchanged
by other ions such as NH~- and H +, which are
probably produced in larger amounts in energy
deficient groups 1 and 3 than in the energy compen-
sated group 2 (Erlich 1972). Another explanation
might be that other regulatory mechanisms are
involved, reducing sodium excretion without
affecting potassium excretion. Such a mechanism
might be by the catecholamines, because it has been
shown that dopamine has a natriuretic effect, whereas
nor-adrenaline has an antinatriuretic effect by its
action through the renal nerves (Johnson et al. 1979,
1981; Johnson 1982; Schrier et al. 1979; Bello-Reuss
et al. 1976). However, we did not find any significant
correlation between the plasma levels of catechol-
amines and the urinary sodium levels, which conse-
quently excludes circulating catecholamines acting as
a major regulator of sodium excretion during the
course. Still, the possibility is open of an influence of
the renal sympathetic nerves on sodium reabsorbtion.
It has also been suggested that antiotensin II can
bring about sustained renal sodium sparing without
inducing potassium loss (B~rresen et al. 1982).
The potassium response to short term exercise
which has been shown to be affected by /31 and/32

blockers ( C a r l s o n et al. 1978; S t e r n h e i m et al. 1982)
was n o t c h a n g e d d u r i n g a p r e v i o u s course ( T a b l e 2) in
spite of i n c r e a s e d c a t e c h o l a m i n e levels ( O p s t a d et al.
1980), which m i g h t indicate a d o w n w a r d s r e g u l a t i o n
of these receptors b e c a u s e of s u s t a i n e d high levels of
the agonists.
I n c o n c l u s i o n , the study shows that r e n i n - a l d o -
s t e r o n e is strongly activated by p r o l o n g e d exercise
c o m b i n e d with severe e n e r g y a n d salt deficiency,
whereas sleep d e p r i v a t i o n does n o t seem to have a n y
m a j o r i n f l u e n c e . T h e r e is n o sign of e x h a u s t i o n of the
s o d i u m a n d p o t a s s i u m h o m e o s t a t i c m e c h a n i s m dur-
ing these e x t r e m e c o n d i t i o n s , in spite of limited
i n t a k e a n d great losses of salt.
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Accepted November 22, 1984