Sodium ion concentration vs.

sweat rate relationship in

humans
Michael J. Buono, Kimberly D. Ball and Fred W. Kolkhorst
J Appl Physiol 103:990-994, 2007. First published 28 June 2007;
doi: 10.1152/japplphysiol.00015.2007

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J Appl Physiol 103: 990994, 2007.
First published June 28, 2007; doi:10.1152/japplphysiol.00015.2007.
Sodium ion concentration vs. sweat rate relationship in humans
Michael J. Buono, Kimberly D. Ball, and Fred W. Kolkhorst
School of Exercise and Nutritional Sciences, San Diego State University, San Diego, California
Submitted 3 January 2007; accepted in final form 26 June 2007
Buono MJ, Ball KD, Kolkhorst FW. Sodium ion concentration vs.
Sweat rate relationship in humans. J Appl Physiol 103: 990994, 2007.
First published June 28, 2007; doi:10.1152/japplphysiol.00015.2007.
The purpose of this study was to determine the effect of active heat
acclimation on the sweat osmolality and sweat sodium ion concen-
tration vs. sweat rate relationship in humans. Eight healthy male
volunteers completed 10 days of exercise in the heat. The mean
exercising heart rate and core temperature were significantly de-
creased (P 0.05) by 18 beats/min and 0.6C, respectively, following
heat acclimation. Furthermore, sweat osmolality and the sweat sodium
ion concentration vs. sweat rate relationships were shifted to the right.
Specifically, the slopes of the relationships were not affected by heat
acclimation. Rather, heat acclimation significantly reduced the y-
intercepts of the sweat osmolality and sweat sodium relationships with
sweat rate by 28 mosmol/kgH2O and 15 mmol/l, respectively. Thus
there was a significantly lower sweat sodium ion concentration for a
given sweat rate following heat acclimation. These results suggest that
heat acclimation increases the sodium ion reabsorption capacity of the
human eccrine sweat gland.
heat acclimation; sweat rate; sweat osmolality; sweat sodium ion
concentration
NUMEROUS STUDIES (1, 9, 12, 17, 30) have reported that heat
acclimation results in a significant reduction in the sodium ion
concentration ([Na]) of human sweat. For example, the clas-
sic study by Kirby and Convertino (12) found that 10 days of
exercise in the heat (40C, 45% relative humidity) caused a
significant reduction in sweat [Na] from 88 to 42 mmol/l.
However, whole body sweat rate significantly increased by
14% during the same time period. More recently, Nielsen et al.
(17) found that humid heat acclimation caused sweat [Na] to
fall from 107 to 70 mM, whereas sweat rate significantly
increased by 26%. However, all of the above studies (1, 9, 12,
17, 30) did not attempt to control for the affect that changes in
sweat rate can have on sweat [Na]. This poses a concern
because it is well known that the [Na] in sweat is influenced
by sweat rate (21, 29, 33). Specifically, the amount of sodium
ions that escape reabsorption increases linearly with increases
in sweat rate (1, 6, 26, 28). The exact mechanism responsible
for this is unknown; however, it has been suggested that above
a certain sweat rate threshold there is insufficient time for
complete sodium ion reabsorption from the duct (28). It is
believed that this threshold reflects the capacity for ion reab-
sorption of the sweat duct (11, 28). In light of the above, it is
clear that changes in sweat [Na] following heat acclimation
can be misleading if they are not presented in relation to sweat
rate. The preferred method for doing this is to examine the
sweat [Na] vs. sweat rate relationship across a range of sweat
rates (11, 14).
Address for reprint requests and other correspondence: M. J. Buono,
C-7251, Diego State Univ., Diego, CA 92182 (e-mail: mbuono@mail.
sdsu.edu).
990 8750-7587/07 $8.00 Copyright 2007 the American Physiological Society
Unfortunately, however, only one study in the literature has
examined the effect of heat acclimation on the sweat [Na] vs.
sweat rate relationship in humans (1). They reported that the
slope of the sweat [Na] vs. sweat rate relationship was
reduced following heat acclimation. However, a number of
methodological concerns were associated with the study,
which limits the usefulness of the data. Specifically, the study
only used three subjects, and no explanation was given as to
how the data were statistically analyzed. Furthermore, the
Downloaded from http://jap.physiology.org/ by guest on June 26, 2013
study used an unorthodox, 3-wk passive heat acclimation
protocol in which the subjects were immersed up to their neck
in a hot-water (40C) bath for 1 h per day. Although the
subjects did have an increase in whole body sweat rate follow-
ing heat acclimation, no core temperature or heart rate data
were presented to confirm the degree of acclimation that
occurred. The above concerns limit the practical applicability
of the data particularly because passive heat acclimation via
hot-water immersion and active heat acclimation results in
different physiological responses and adaptations (3). In light
of the above, it was the purpose of this study to determine the
effect a 10-day active heat acclimation protocol had on the
sweat osmolality and sweat [Na] vs. sweat rate relationships
in humans. Unlike most past studies, the present study em-
ployed increases in temperature, relative humidity, and exer-
cise intensity during three successive exercise bouts to pur-
posely ensure that three different sweat rates would be pro-
duced, thus allowing for the construction of sweat osmolality
and sweat [Na] vs. sweat rate graphs. It was hypothesized that
heat acclimation would shift both the sweat osmolality and
sweat [Na] vs. sweat rate relationships to the right, thus
supporting a greater sodium ion reabsorptive capacity in the
eccrine sweat gland following heat acclimation.
METHODS
The subjects for the study were eight healthy male volunteers. They
had a mean ( SD) age, height, and weight of 26 7 yr, 174 5 cm,
and 69.8 11.2 kg, respectively. The San Diego State University
institutional review board approved the study, and written informed
consent was obtained from each subject before participation. The
subjects were asked to refrain from initiating new exercise programs
during the duration of the study. No restrictions were placed on their
diet because it has been reported that electrolyte supplementation is
not necessary during heat acclimation (24). Furthermore, unless the
subjects are on very low-sodium-intake diets, manipulations of dietary
sodium do not appear to greatly effect sweat sodium ion secretion
during heat acclimation (2, 3).
Each subject completed a 10-day heat acclimation protocol. On a
separate day, before heat acclimation, each subject performed a
graded treadmill test to volitional exhaustion to determine maximal
oxygen uptake (VO2max). During the treadmill test oxygen uptake was
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 HEAT ACCLIMATION REDUCES SWEAT SODIUM CONCENTRATION
measured continuously using a calibrated metabolic cart (TrueMax
2400, Parvomedics, Sandy, UT). All subjects achieved a respiratory
exchange ratio of 1.1 during the test. The mean ( SD) VO2max for
the group was 55.6 12.7 ml kg1 min1.
Within 7 days of completing the treadmill test to exhaustion, each
subject began 10 consecutive days of exercise in the heat. Each day
consisted of three 30-min exercise bouts of walking on a motorized
treadmill in an environmental chamber. The chamber temperature and
relative humidity for the three exercise bouts were 36, 40, and 42C
and 40, 40, and 60%, respectively. Ten minutes of seated rest outside
the chamber (23C and 50% relative humidity) occurred between the
three exercise bouts. During this time, water was given to completely
replace the fluids lost via sweating, thus preventing dehydration
during each exercise session. Furthermore, before the first day of the
heat acclimation period, the subjects were asked to consume 1 liter of
water before going to bed and on arising in the morning. Preexercise
nude body weight was measured to the nearest 0.1 kg on the first day
before the start of the first exercise bout. This was considered the
baseline, euhydrated nude body weight, and all successive days were
991
the torso, which prevented leakage and sample contamination. The
Macroduct collectors had a surface area of 5.2 cm2 and were fitted
with an additional airtight reservoir to increase their sweat collection
capacity to 1.5 ml. A new Macroduct was applied to the same chest
site immediately before each of the 3 exercise bouts and sweat was
collected for the complete 30 min of each exercise bout. Whole body
sweat rate was calculated on days 1 and 10 using pre- and postexercise
dry nude body weights, which were corrected for the amount of fluids
consumed.
The skin was cleaned with deionized water and dried immediately
before securing each collector. Chest sweat rate was calculated by the
change in volume within the sample collection tubing and expressed
in milligrams per square centimeter per minute. The chest sweat
samples were analyzed in duplicate for osmolality (mosmol/kgH2O)
using a vapor pressure osmometer (Wescor) and for [Na] using a
flame photometer (Cole-Palmer, Chicago, IL).
Regression equations were developed for each subject, and the
mean slope and y-intercept for the group were calculated. Pre- and
postacclimation data were analyzed using paired t-tests. The sweat

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compared with it. Thus, if the subject reported to the laboratory and
preexercise body weight was less than the baseline measure, water
was administered until rehydration occurred and baseline weight was
reestablished. Ensuring euhydration, both before and throughout the
heat acclimation period was critical, because dehydration has been
previously shown (16) to significantly effect sweat [Na]. This was
achieved as evidence by the fact that preexercise nude body weight for
each of the 10 heat acclimation days fluctuated by 1% (Table 1).
The intensity for the three exercise bouts was calculated to approx-
imate 30, 40, and 50% of the previously determined VO2max, respec-
tively. This was achieved by altering the treadmill speed and grade for
each subject. On the first day of heat acclimation, expired air samples
were collected from the 8th to the 10th min of each exercise bout, and
oxygen uptake was determined to ensure that the proper intensity was
obtained. The treadmill speed and grade for each of the three exercise
bouts remained constant for each individual subject throughout the
10-day heat acclimation protocol. The oxygen uptake data showed
that as a group the subjects worked at 28 3, 38 2, and 47 3%
of their preacclimation VO2max during the three exercise bouts. The
increases in temperature, relative humidity, and exercise intensity for
the three exercise bouts were purposely used to ensure that three
different sweat rates would be produced.
Heart rate and core temperature were continuously measured each
day during the three exercise bouts. Heart rate was measured using a
Polar monitor while core temperature was measured using a rectal
thermistor (YSI series 400) inserted by the subject to 10 cm past the
anal sphincter.
Chest sweat samples were also collected during each of the three
30-min exercise bouts on day 1 and day 10 of the heat acclimation
protocol using separate Macroduct sweat collectors (Wescor, Logan,
UT). The collectors were held in place by elastic straps placed around
osmolality vs. sweat rate and sweat [Na] vs. sweat rate relationships
obtained before and after heat acclimation were compared using a
two three repeated-measures ANOVA. Significance was set at the
P 0.05 level.
RESULTS
The results of the present study showed that 10 days of heat
acclimation significantly reduced the heart rate and rectal
temperature during exercise in the heat. The mean ending heart
rate and core temperature for each of the three different
exercise bouts on each of the 10 heat acclimation days are
presented in Table 1. As can be seen, there were significant
reductions in core temperature and heart rate as a result of heat
acclimation. For example, on day 1, the group had a mean
( SD) heart rate following the third exercise bout of 158 18
beats/min. On day 10, the heart rate at the same time point was
significantly reduced to 140 17 beats/min. Similarly, the
mean ending rectal temperature was significantly reduced from
39.0 0.4C on day 1 to 38.4 0.4C on day 10. The
magnitude of the decreases in rectal temperature and heart rate
are consistent with the results of previous studies (5, 10, 18).
Such reductions strongly suggest that the 10-day protocol used
in the present study was successful in conferring heat acclima-
tion in the subjects. There were no significant changes in chest
sweat rate over the course of the 10-day heat acclimation
period. Specifically, the mean chest sweat rate for the three
exercise bouts on day 1 were 0.26 0.06, 0.59 0.17 and
0.95 0.26 mg cm2 min1, respectively. On day 10 the

Table 1. Physiological responses during 10 days of heat acclimation

Day

1 2 3 4 5 6 7 8 9 10

Ending exercise core temperature, C

Bout 1 37.6 (0.3) 37.4 (0.4) 37.5 (0.4) 37.5 (0.4) 37.5 (0.3) 37.4 (0.4) 37.4 (0.4) 37.4 (0.4) 37.4 (0.3) 37.3 (0.4)
Bout 2 38.2 (0.3) 38.0 (0.4) 38.0 (0.2) 38.0 (0.3) 38.0 (0.2) 37.8 (0.3) 37.9 (0.3) 37.9 (0.3) 37.8 (0.3) 37.9*(0.3)
Bout 3 39.0 (0.4) 38.8 (0.4) 38.7 (0.5) 38.6 (0.5) 38.6 (0.5) 38.6 (0.3) 38.5 (0.4) 38.4 (0.4) 38.4 (0.4) 38.4*(0.4)
Ending exercise heart rate, beats/min
Bout 1 108 (12) 108 (9) 106 (11) 104 (11) 105 (9) 104 (11) 100 (8) 103 (11) 101 (10) 100*(9)
Bout 2 129 (15) 127 (14) 122 (12) 122 (13) 120 (10) 115 (11) 114 (12) 116 (14) 115 (11) 115*(11)
Bout 3 158 (18) 154 (17) 149 (16) 146 (17) 142 (17) 143 (15) 138 (17) 141 (17) 140 (17) 140*(17)
Preexercise body weight, kg
69.8 (11.2) 69.5 (11.3) 69.7 (11.5) 69.4 (11.3) 69.6 (11.6) 69.8 (11.7) 69.7 (11.8) 69.4 (11.8) 69.5 (10.8) 69.4 (11.1)
Values are means (SD) for 8 subjects. *Day 10 value was significantly less than day 1, P 0.05.

J Appl Physiol VOL 103 SEPTEMBER 2007 www.jap.org

992 HEAT ACCLIMATION REDUCES SWEAT SODIUM CONCENTRATION
mean values were 0.31 0.14, 0.66 0.22, and 1.03 0.34
mgcm2 min1, respectively. In addition, the mean ( SD)
whole body sweat rate was 0.92 0.46 l/h on day 1 and
1.08 0.30 l/h on day 10. The 17% increase is consistent in
magnitude with past heat acclimation studies (12, 17); how-
ever, it was not significant.
The relationship between sweat osmolality and chest sweat
rate obtained on day 1 and day 10 of heat acclimation is
presented in Fig. 1. As can be seen, the relationship was shifted
to the right following heat acclimation, thus sweat osmolality
was reduced for a given sweat rate as a result of heat acclima-
tion. The interaction, however, was not significant, which
suggests that the slope of the relationship was not affected by
heat acclimation. Rather, heat acclimation significantly re-
duced the mean y-intercept from 83 31 to 55 34 mosmol/
kgH2O.
The relationship between sweat [Na] and chest sweat rate
obtained on days 1 and 10 of heat acclimation is presented in
Fig. 2. Similarly, the relationship was shifted to the right
following heat acclimation; thus sweat [Na] was reduced for
a given sweat rate. The slope of the relationship was not
changed by heat acclimation, as suggested by a nonsignificant
interaction. Heat acclimation, however, significantly reduced
the mean y-intercept from 22 11 to 7 16 mmol/l. It was
not surprising that the two figures were similar because it has
previously been shown that sweat osmolality and [Na] are
highly correlated, with sodium usually accounting for approx-
imately half of the sweat osmolality (7).
DISCUSSION
The most important finding of the present study was that 10
days of heat acclimation shifted the sweat osmolality and sweat
[Na] vs. sweat rate relationship to the right; thus both
variables were reduced for a given sweat rate, up to 1
mgcm2 min1. More specifically, the slopes of the sweat
osmolality and sweat [Na] vs. sweat rate relationships were
not significantly changed by heat acclimation. However, there
were significant reductions in the mean y-intercept of both
relationships following heat acclimation.
Sweating has the potential to cause significant amounts of
sodium and chloride ions to be lost; therefore, they must be
conserved. Structurally, the human eccrine sweat gland con-
sists of two distinct regions. These are the secretory coil, which
produces an isosmotic precursor sweat by the electrogenic
Fig. 1. Sweat osmolality vs. chest sweat rate relationship on days 1 and 10 of
heat acclimation. Values are means SD for 8 subjects. Day 10 mean sweat
osmolality was significantly different from day 1, P 0.05.
J Appl Physiol VOL
Fig. 2. Sweat sodium ion concentration ([Na]) vs. chest sweat rate
relationship on days 1 and 10 of heat acclimation. Values are means SD
for 8 subjects. Day 10 mean sweat [Na] was significantly different from
day 1, P 0.05.
Downloaded from http://jap.physiology.org/ by guest on June 26, 2013
movement of sodium and chloride ions and the coupled move-
ment of water via osmosis, and the reabsorptive duct, which
actively reabsorbs salt from the precursor sweat (19, 29). The
reabsorption depends on Na being actively transported into
the interstitial fluid via an ouabain-sensitive Na-K ATPase
localized on the basolateral membrane of the ductal cells.
Sodium ions enter the duct cell via an amiloride-sensitive
channel expressed in the apical membrane. Chloride ions
follow down their electrochemical gradient during reabsorption
using the cystic fibrosis transmembrane conductance regulator
located in the luminal membrane (19, 29).
During low levels of sweat production significant amounts
of sodium and chloride ions can be reabsorbed from sweat
during its passage along the duct, resulting in a very hypos-
motic sweat appearing at the skin surface. However, the rate of
sodium and chloride ion reabsorption has a finite capacity;
thus, at high sweat rates, there is insufficient time for complete
ion reabsorption. This is evidenced by the fact that the con-
centration of sodium ions in the sweat increases linearly with
increases in sweat rate (1, 6, 11). The results of the present
study agree with the above findings. As can be seen in Fig. 2,
sweat [Na] increased linearly with increases in sweat rate
both before and after heat acclimation. Heat acclimation, how-
ever, shifted the relationship to the right, significantly reducing
the y-intercept, without changing the slope. These findings
suggest that heat acclimation increases the reabsorptive ability
of the eccrine sweat duct. Shamsuddin et al. (28) have previ-
ously shown that a sweat rate threshold exists above which
sweat [Na] increases rapidly. Furthermore, this threshold may
reflect the ion reabsorptive capacity of the sweat gland duct and
be the result of an imbalance between sodium accumulation in
the precursor sweat and sodium reabsorption from the proximal
duct (28). The reduction in the y-intercept of the sweat [Na]
vs. sweat rate relationship found in the present study is con-
sistent with an increase the sodium ion reabsorption capacity.
This would delay the imbalance between sodium accumulation
in the precursor sweat and sodium reabsorption to a greater
sweat rate following heat acclimation. The exact mechanism
responsible for the increased sodium reabsorption following
heat acclimation is currently unknown. Most likely, it involves
aldosterone-induced increases (8) in the Na-K-ATPase and
the sodium channels in the reabsorption duct (22, 23). This is
103 SEPTEMBER 2007 www.jap.org
 HEAT ACCLIMATION REDUCES SWEAT SODIUM CONCENTRATION
evidenced by the fact that administration of the aldosterone
antagonist, spironalactone, has been shown to increase sweat
[Na] during heat acclimation (13). This finding is in contrast
to the results of Allan and Wilson (1), who reported that heat
acclimation reduced the slope of the sweat sodium concentra-
tion vs. sweat rate relationship. However, as discussed earlier,
that study had several methodological concerns, including a
small sample size and an unorthodox heat acclimation
protocol.
The unchanged slope, coupled with increased electrolyte
reabsorption, found in the present study would be consistent
with the data of Shamsuddin et al. (27). They altered sodium
reabsorption of the eccrine sweat gland by changing skin
temperature. Increases in temperature are known to increase
sodium channel excitability (20). Thus they reported that sweat
ion reabsorption capacity was significantly increased during
exercise in a 25 vs. a 15C environment. However, the slope of
the sweat ion concentration and sweat rate relationship was
unchanged. Rather the increased sweat ion reabsorption was
the result of a lower y-intercept on the sweat ion concentration
vs. sweat rate relationship in the 25C condition.
Furthermore, a recent study (14) measured the sweat [Na]
vs. sweat rate relationship before and after 21 days of heat
acclimation in horses. The results clearly showed that the slope
of the relationship was unchanged with heat acclimation,
whereas the y-intercept was reduced 15 mmol/l for the six
animals studied. Such findings are in agreement with the
present results (Fig. 2).
The primary advantages of the reduced sweat [Na] seen
following heat acclimation appear to be threefold. First, by
virtue of having more dilute sweat, heat-acclimated individuals
have additional solutes (primarily sodium and chloride) re-
maining within the extracellular space to exert an osmotic
pressure to redistribute fluid from the intracellular space (24).
This is evidence by the fact that following heat acclimation,
subjects have a significantly smaller reduction in plasma vol-
ume (5 vs. 12%) for a given level of dehydration (25). Second,
an increased sweat [Na] has been one of the factors associated
with the development of hyponatremia during prolonged exer-
cise (15). For example, cystic fibrosis patients, who secrete
high levels of sodium and chloride in their sweat, are predis-
posed to the development of hyponatremia when exposed to
heat (24, 31). Thus heat acclimation has the potential to
attenuate the development of hyponatremia by reducing the
sodium lost via sweating (15). Lastly, it has recently been
shown (32) that subjects with lower sweat [Na] had attenu-
ated osmotic inhibition of various thermoregulatory responses
to heat stress. Specifically, the hyperosmotic-induced increases
in the core temperature threshold for sweating and cutaneous
vasodilatation were smaller in those subjects with lower sweat
[Na]. It was hypothesized that such results would be benefi-
cial in maintaining thermoregulatory sweating and skin blood
flow during prolonged heat stress, thus preventing hyperther-
mia (32).
In conclusion, the results of the present study are the first in
the literature to show that active heat acclimation significantly
reduces both the sweat osmolality and sweat [Na] for a given
sweat rate up to 1 mgcm2 min1 in humans. Specifically
heat acclimation reduces the y-intercept of both the sweat
osmolality and sweat [Na] vs. sweat rate relationships, with-
J Appl Physiol VOL
993
out changing their slopes. Such a finding suggests that heat
acclimation increases ion reabsorption in the sweat gland duct.
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