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Stavros A. Kavouras, Lawrence E. Armstrong, Carl M. Maresh, Douglas J.

Casa,
Jorge A. Herrera-Soto, Timothy P. Scheett, James Stoppani, Gary W. Mack and
William J. Kraemer
J Appl Physiol 100:442-450, 2006. First published Oct 6, 2005; doi:10.1152/japplphysiol.00187.2005

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Hydration and Physical Performance
B. Murray
J. Am. Coll. Nutr., October 1, 2007; 26 (suppl_5): 542S-548S.
[Abstract] [Full Text] [PDF]

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Journal of Applied Physiology publishes original papers that deal with diverse areas of research in applied physiology, especially
those papers emphasizing adaptive and integrative mechanisms. It is published 12 times a year (monthly) by the American
Physiological Society, 9650 Rockville Pike, Bethesda MD 20814-3991. Copyright © 2005 by the American Physiological Society.
ISSN: 8750-7587, ESSN: 1522-1601. Visit our website at http://www.the-aps.org/.
J Appl Physiol 100: 442– 450, 2006.
First published October 6, 2005; doi:10.1152/japplphysiol.00187.2005.

Rehydration with glycerol: endocrine, cardiovascular, and thermoregulatory


responses during exercise in the heat
Stavros A. Kavouras,1,4 Lawrence E. Armstrong,1,2 Carl M. Maresh,1,2
Douglas J. Casa,1 Jorge A. Herrera-Soto,1 Timothy P. Scheett,1
James Stoppani,1 Gary W. Mack,3 and William J. Kraemer1,2
1
Human Performance Laboratory, Department of Kinesiology, and 2Departments of Physiology and Neurobiology and of
Nutritional Sciences, University of Connecticut, Storrs; 3The John B. Pierce Laboratory, Yale University School of Medicine,
New Haven, Connecticut; and 4Laboratory of Nutrition and Clinical Dietetics, Harokopio University, Athens, Greece
Submitted 15 February 2005; accepted in final form 29 September 2005

Kavouras, Stavros A., Lawrence E. Armstrong, Carl M. Maresh, and electrolyte restoration after exercise is a slow process that
Douglas J. Casa, Jorge A. Herrera-Soto, Timothy P. Scheett, James requires at least 3 h and ⬎100% of fluid losses, along with a
Stoppani, Gary W. Mack, and William J. Kraemer. Rehydration with significant amount of electrolytes (45).
glycerol: endocrine, cardiovascular, and thermoregulatory responses dur- However, ingesting a large volume of fluids, even in dehy-
ing exercise in the heat. J Appl Physiol 100: 442– 450, 2006. First drated subjects, rapidly decreases arginine vasopressin (AVP),
published October 6, 2005; doi:10.1152/japplphysiol.00187.2005.—The

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impact of rehydration with glycerol on cardiovascular and thermoregu-
even before plasma volume or osmolality have been restored,
latory responses during exercise in the heat was studied in eight highly leading to increased urinary output (12). Additionally, thirst
trained male cyclists. Each subject completed three dehydration-rehydra- sensation decreases in response to plasma volume and osmo-
tion experimental trials that differed only in the rehydration treatment, lality restoration, mouth wetness, and oral-pharyngeal stimu-
each separated by 7 days. Before each experimental day, subjects dehy- lation (33). For these reasons, rehydration is a major part of the
drated to ⫺4% of their body weight by exercise and water restriction. The recovery process after exercise, especially when individuals
experimental treatments were as follows: no fluid (NF), glycerol bolus (1 must undertake repeated bouts of exercise-heat stress.
g/kg body wt) followed by water (G), and water alone (W). Rehydration In 1987, glycerol, used with water, was first shown to induce
(3% body weight) was given over an 80-min period. After rehydration, hyperhydration more efficiently than water alone, and this
subjects cycled (74% peak O2 uptake) to exhaustion in a hot and wet effect persisted for 4 h (40). More recently, the effect of
(37°C and 48% relative humidity) environment. For G, plasma volume glycerol-induced hyperhydration on fluid-regulating hormones
was expanded (P ⬍ 0.05) during rehydration and remained higher than was investigated (14). The authors demonstrated a small,
W (P ⬍ 0.05) during exercise. Exercise time to exhaustion during G
(33 ⫾ 4 min) was longer (P ⬍ 0.05) compared with both W (27 ⫾ 3 min)
nonsignificant increase in plasma vasopressin and no change in
and NF (19 ⫾ 3 min). Cutaneous vascular conductance was significantly circulating aldosterone (Aldo) or atrial natriuretic factor. They
elevated (P ⬍ 0.05) during G, but G provided no other thermoregulatory reported that there may have been a direct glycerol effect on
or cardiovascular benefits compared with W and NF. Fluid-regulating the kidneys that increased fluid reabsorption and induced a
hormones (vasopressin, aldosterone, atriopeptin, and plasma renin activ- positive fluid balance.
ity) decreased during rehydration and increased during exercise (except Glycerol is a safe agent that does not approach toxic levels
atriopeptin), but there were no differences between G and W. These data when administered orally in doses of ⬍5 g/kg body wt (47).
indicated that glycerol had little or no major effect on fluid-regulating Glycerol also is an attractive compound for use in studies that
factors during rehydration or exercise, and the improved exercise capac- involve fluid balance and exercise, because it is not a major
ity in G was likely due to a greater plasma volume during exercise. energy source during intense exercise (15, 32, 34, 37). Fur-
fluid balance; plasma volume; osmoregulation; cycling; vasopressin thermore, glycerol-induced hyperhydration has been suggested
to 1) increase overall exercise performance (2, 9, 19) or time to
exhaustion (36) in both temperate (36) and hot (2, 9, 19)
IT IS WELL ESTABLISHED THAT intense physical exercise in the heat environments; 2) increase sweat rate (28); 3) decrease heart
increases the risk of heat illnesses and decreases exercise rate (HR) (36) and rectal temperature (Tre) during moderate-
performance, whereas significant dehydration can augment intensity cycling exercise (2); and 4) last for 32– 48 h after
these responses (4, 17, 38, 43). Additionally, since the 1940s, dosing (23). However, several other studies have shown no
scientists have observed that, when people exercise in the heat, performance (20, 29, 31, 48) or thermoregulatory benefit (9,
they become dehydrated, even if they have free access to water 19, 25, 26, 31, 36) following glycerol-induced hyperhydration.
(38). It is generally accepted that dehydration is difficult to The purpose of the present investigation was to examine the
prevent during heavy training or competition in the heat (7). In effect of partial oral rehydration with glycerol on a subsequent
prolonged events and in some individuals with high sweating exhaustive exercise test in the heat. Surprisingly, although
rate, total body water loss may be as much as 8% of initial there is a vast amount of information on the role of glycerol as
body weight (3). This phenomenon has been described as a hyperhydration agent, very limited information is available
involuntary dehydration (18). On the other hand, rapid fluid on its effect in rehydration (44). We hypothesized that glycerol

Address for reprint requests and other correspondence: S. A. Kavouras,


Laboratory of Nutrition and Clinical Dietetics, Dept. of Nutrition and Dietetics, The costs of publication of this article were defrayed in part by the payment
Harokopio Univ., 70 El. Venizelou Av., Athens 17671, Greece (e-mail: of page charges. The article must therefore be hereby marked “advertisement”
skav@hua.gr). in accordance with 18 U.S.C. Section 1734 solely to indicate this fact.

442 8750-7587/06 $8.00 Copyright © 2006 the American Physiological Society http://www. jap.org
REHYDRATION WITH GLYCEROL 443
would stabilize cardiovascular function, reduce heat stress, and were presented in a random sequence to avoid order effect and were
enhance exercise capacity after partial rehydration. separated by a minimum of 1 wk to prevent a carryover effect.
Subjects were asked to maintain similar eating and training habits
MATERIALS AND METHODS during the 3 days before each trial, verified by 3-day dietary intake
and physical training records.
Subjects Euhydration baseline. Subjects reported to the laboratory between
Eight endurance-trained male cyclists agreed to serve as subjects in 1000 and 1200 in a euhydrated state, having ingested at least 30 ml/kg
this study. Their mean (⫾SE) characteristics were as follows: age, of their body weight of water (or other noncaffeinated fluids) during
24 ⫾ 1 (range 19 –29) yr; body mass, 70.1 ⫾ 1 (range 66.7–73.6) kg; the previous day. After each subject emptied his bladder, urine
height, 181 ⫾ 2 (range 174 –185) cm; fat-free mass, 56.6 ⫾ 0.4 (range specific gravity (USG) was determined by refractometry to verify
52.6 – 61.7) kg; and peak O2 uptake (V̇O2 peak), 61.4 ⫾ 0.8 (range good hydration status (⬍1.020), and baseline euhydrated body weight
58.9 – 65.9) ml 䡠 kg⫺1 䡠 min⫺1. Subjects were selected after their phys- was recorded to ⫾50 g (SRI, Instruments Precision Scales, Tonawanda,
ical activity and medical history questionnaires were reviewed, with- NY). Hydration status also was verified by plasma (OsmP ⫽ 285–295
out regard to race or ethnic origin. These men were heat acclimatized, mosmol/kgH2O) and urine osmolality (Osmu ⬍ 900 mosmol/kgH2O).
trained regularly, competed in road cycling or mountain bike races, The subjects entered the environmental chamber (model 2000, Minus
were nonsmokers, and reported no previous history of endocrine, Eleven, Malden, MA; ambient temperature, 36.7 ⫾ 0.2°C; relative
cardiovascular, renal, or thermoregulatory disorders. All athletes had humidity, 48.0 ⫾ 1.2%) and sat for 20 min before a blood sample was
completed their competitive season at least 2 mo before the study and taken without stasis using a 20-gauge butterfly needle.
were on their training season. The study was approved by the Dehydration. Following euhydration baseline, and for the remain-
Institutional Review Board for Studies Involving the Use of Human der of the day, subjects underwent fluid restriction and consumed food
Subjects at the University of Connecticut, and all volunteers gave their

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low in fluid content for lunch and dinner. Additionally, during the
written, informed consent after attending an informational meeting afternoon and evening of this day, subjects performed 2 h of low-
that addressed the study purpose, methods, and attendant risks and intensity cycling exercise to induce a body fluid loss equivalent to 4%
benefits. of body weight.
Dehydration baseline. The subject arrived at the laboratory 12 h
Preliminary Testing
postprandial, emptied his bladder, and had body weight measured to
Body composition. The hydrostatic weighing technique was used to verify the degree of dehydration. If dehydration was ⬍3.5 or ⬎4.5%,
determine body density. Each subject performed the measurement at the experiment was canceled for that day (see Table 1). USG also was
least 20 times, on 2 separate days, before the actual data were recorded to verify hypohydration. A 20-gauge, 32-mm indwelling
collected. Before this measurement, residual volume was calculated Teflon catheter was inserted in an antecubital vein, and an extension
based on the vital capacity measured with a hand-held spirometer. tube with a stopcock was attached to the catheter port for acquisition
Calculation of the percentage of body fat from body density was based of serial blood samples. The catheter was kept patent by flushing
on the Brozek equation (6). isotonic saline solution containing 20 heparin units/ml of solution.
V̇O2 peak test. V̇O2 peak was determined in a thermocomfortable From that point on, the subject remained seated in a wheelchair.
environment (27°C) using an incremental resistance exercise test on a Subjects were taken into the environmental chamber (36.7 ⫾ 0.2°C),
mechanically braked cycle ergometer (Monark ergomedic 818E, and, after 20 min of equilibration, a blood sample (25 ml) was taken.
Stockholm, Sweden). The exercise test consisted of continuous cy- Rehydration. The subject, remaining in the wheelchair, was taken
cling at a constant cadence (90 –100 rpm), while resistance increased from the environmental chamber and remained in a comfortable
by 0.5 kp every 2 min until volitional exhaustion. Breath-by-breath environment (27.3 ⫾ 0.3°C) for 20 min to allow body fluids to
analysis of the expired gases during the test was performed with an equilibrate. A blood sample was then taken, and one of the three
open-circuit respiratory apparatus (model CPXD, MedGraphics Car- randomly assigned, double-blind experimental trials began [i.e., NF
diopulmonary Exercise System, St. Paul, MN). Two of the following (control), G, and W]. One-third of the rehydrating fluid was admin-
three criteria were used to verify the attainment of V̇O2 peak: 1) no istered during the first 15 min. The remaining two-thirds of the fluid
increase in oxygen uptake (V̇O2; ⬍150 ml/min) with an increase in were administered in equal doses every 10 min, from minute 20 to 80
ergometer resistance, 2) HR ⬎90% of predicted maximal value (i.e., of the rehydration period. The first dose for G consisted of a 20%
220 ⫺ age), and 3) respiratory exchange ratio ⬎1.1. The average glycerol and water solution (Penta Manufacturing, Livingston, NJ)
duration of the V̇O2 peak test was 10.01 ⫾ 0.96 min. that was administered as 1 g glycerol/kg body wt. For W, the first dose
consisted of aspartame-flavored water to mimic the sweetness of the
Experimental Protocol
glycerol solution. All other doses in G and W were identical and
Each subject completed three experimental trials, which differed consisted of water. A noncaloric, nonsodium, flavored powder (Kool-
only with regard to rehydration. The rehydration treatments were as Aid) was added to both G and W to provide consistent sweetness,
follows: no fluid (NF), which served as a control; glycerol followed by flavor, and color. All fluids were served chilled (10°C). The total fluid
water (G), and water alone (W). To ensure double-blind design, a ingested was equal to 3% of the euhydrated body weight for each
noncaloric, nonsodium, flavored powder (Kool-Aid, White Plains, subject and was designed to partially rehydrate the subjects. Ten
NY) was added to both G and W drinks. The experimental treatments minutes following rehydration, subjects were taken back into the

Table 1. Indexes of fluid-electrolyte status during dehydration and rehydration


Euhyd Dhy-Base Rhy

Trial Weight, kg USG Osmp, mosmol/kgH2O Weight, kg Dehydration, % USG Osmp, mosmol/kgH2O Fluid intake, ml

No fluid 71.3⫾1.3 1.005⫾0.001 279.4⫾2.4 68.5⫾1.3 ⫺3.91⫾0.26 1.028⫾0.02 289.4⫾2.4 0


Glycerol 70.8⫾1.3 1.005⫾0.001 280.1⫾1.6 68.0⫾1.2 ⫺3.86⫾0.17 1.029⫾0.01 289.3⫾1.6 2,148⫾30
Water 71.8⫾1.1 1.005⫾0.002 284.0⫾1.0 68.9⫾1.1 ⫺4.00⫾0.35 1.029⫾0.01 291.6⫾1.0 2,142⫾40
Values are means ⫾ SE. Euhyd, euhydrated baseline; USG, urine specific gravity; Dhy-Base, dehydrated baseline; Osmp, plasma osmolality; Rhy, rehydration.

J Appl Physiol • VOL 100 • FEBRUARY 2006 • www.jap.org


444 REHYDRATION WITH GLYCEROL

environmental chamber (36.7 ⫾ 0.2°C) and sat quietly in the wheel- These samples were centrifuged immediately at 1,800 g and 4°C for
chair for 20 min before another 25-ml blood sample was obtained. 12 min. Plasma samples for hormonal assays were refrigerated at
Exercise test. Following the blood sample, subjects sat on a ⫺80°C for later analysis.
mechanically braked cycle ergometer (Monark ergomedic 818E). Seat Blood and urine analyses. USG and TPP were measured by
height was the same for all trials, and toe clips were used. The refractometer (model A300CL, Spartan). Hct was determined in
exercise test (time to exhaustion expressed in minutes) consisted of triplicate from whole blood by the microcapillary technique, follow-
steady-state cycling (74.0 ⫾ 1.1% V̇O2 peak) at 80 –100 rpm. Subjects ing centrifugation for 4 min at 9,500 g. No corrections were made for
continued cycling until one of the following criteria for termination trapped plasma or for peripheral venous sampling. Hb was measured
were met: inability to maintain cadence (decrease by ⬎20 rpm), in triplicate from whole blood with the cyanmethemoglobin technique
increase in Tre to 39.5°C for ⬎5 min, or signs or symptoms of heat (Sigma Diagnostics). Percent changes in plasma volume were calcu-
exhaustion. The following ambient conditions were maintained in the lated with the following formula (11):
climatic chamber during exercise: 36.8 ⫾ 0.1°C, 48.1 ⫾ 1.6% relative
humidity, and wind speed by fan of 2.54 m/s. All clocks and stop- ⌬PV ⫽ 100共HbB/HbA兲兵关1 ⫺ 共HctA/100兲兴/关共1 ⫺ HctB/100兲兴其 ⫺ 100
watches were removed from subject view.
where ⌬PV is percent change of plasma volume, subscript B is before
Tre and skin temperatures (Tsk) were recorded every 4 min,
(control), and subscript A is after (experimental). OsmP and OsmU
whereas HR was obtained at 2-min intervals during exercise. Arterial
were determined in duplicate by freezing point depression the same
blood pressure, forearm skin blood flow (SkBF), V̇O2, carbon dioxide
day (model 3DII, Advanced Digimatic Osmometer, Norwood, MA).
production (V̇CO2), respiratory quotient (R), respiratory rate (RR),
[Na⫹]P and plasma potassium concentration were determined by
pulmonary ventilation (V̇E), and cardiac output (Q̇) were measured at
ion-sensitive electrodes on fresh plasma samples (model 984-S, AVL
0, 5, 15, 30, 45, and 60 min of exercise. Blood samples (5 ml) were
Scientific, Roswell, GA). Plasma glucose and lactate concentrations
taken at 0, 5, 15, 30, 45, and 60 min of exercise, and a final blood

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were determined in triplicate with an enzymatic technique (model
sample (25 ml) was drawn immediately after the end of exercise.
2003, Yellow Springs Instruments). [Glyc]P was measured in dupli-
Thermoregulatory measurements. Tre was measured by using a
cate with a colorimetric technique (Sigma Diagnostics). [AVP]P was
rectal thermistor (model 401, Yellow Springs Instruments, Yellow
determined by a commercially available radioimmunoassay kit (Ni-
Springs, OH) inserted 10 cm past the external anal sphincter. Tsk were
chols Institute, San Juan Capistrano, CA). The within- and between-
recorded via skin thermistors (model 409, Yellow Springs Instru-
assay coefficients of variation for [AVP]P at midrange (13 pg/ml)
ments) at four sites: chest, triceps, thigh, and calf. Mean-weighted Tsk
were 1.8 and 5.8%, respectively. The sensitivity of the assay and the
was calculated according to the equation of Ramanathan [Tsk ⫽ 0.2
average extraction recovery were 1.3 pg/ml and 63.8%, respectively.
(Tchest ⫹ Ttriceps) ⫹ 0.3 (Tthigh ⫹ Tcalf), where T is temperature] (39).
PRA was evaluated in duplicate by radioimmunochemical determina-
Forearm SkBF was measured for 2 min with a laser Doppler flow-
tion of plasma angiotensin I, generated during 1 h of incubation at pH
meter for microvascular perfusion (model Periflux PF2B, Perimed,
6.0 (Incstar, Stillwater, MN). The within- and between-assay coeffi-
Stockholm, Sweden). During the measurement, subjects rested their
cients of variation of this assay at midrange (1.2 ng 䡠 l⫺1 䡠 s⫺1) were 4.6
forearm in a neutral position with 90° flexion of the elbow. Data were
and 4.8%, respectively. The sensitivity of the assay was 0.05
expressed as cutaneous vascular conductance (CVC) by dividing
ng 䡠 l⫺1 䡠 s⫺1. [AP]P was determined by a radioimmunoassay technique
SkBF, in volts, over mean arterial pressure (MAP), in 100 mmHg.
(Peninsula Laboratories, Belmont, CA), after extraction on octadecyl-
Whole body sweat rate during exercise was calculated from body
silane C18 cartridges (Sep-Pak C18, Waters Associates, Milford, MA).
weight changes, corrected for estimated respiratory water losses (35),
The coefficient of variation of the assay was ⬍5%. [Aldo]P was
blood sample volume, and urine output.
determined with a radioimmunoassay technique (Coat-A-Count, Di-
Cardiovascular variables. HR was measured by a lead I configu-
agnostic Products, Los Angeles, CA) that had a within-assay coeffi-
ration, using a telemetric cardiotachometer (model Vantage XL, Polar cient of variation for the mid- (395 pmol/l) and the high-range (1,020
Electro). Breath-by-breath analysis of the expired gases was per- pmol/l) of 3.3 and 1.9%, respectively. The sensitivity of the assay was
formed with an open-circuit respiratory apparatus (model CPXD, 44 pmol/l. To reduce interassay variations, each subject’s plasma
MedGraphics Cardiopulmonary Exercise System) to determine V̇O2, samples were analyzed within the same assay run. All hormonal
V̇CO2, R, RR, and V̇E. Q̇ was measured via CO2 rebreathing, employ- analyses were performed in duplicate.
ing the exponential method (1, 10). V̇O2 and V̇CO2 were measured, as
described above, for 3 min before each rebreathing procedure. To
Statistical Analysis
calculate stroke volume (SV), HR was recorded just before the onset
of the rebreathing technique. Arterial systolic (SBP) and diastolic Statistical evaluation of the data was accomplished with a
blood pressures (DBP) also were measured with an aneroid sphyg- two-way analysis of variance with repeated measures (treatment ⫻
momanometer and stethoscope. MAP was calculated as shown in the time). Significant differences between the means were determined by
following formula: MAP ⫽ (SBP ⫺ DBP)/3 ⫹ DBP. Newman-Keuls post hoc test. Statistical differences were determined
Blood collections. All blood samples were drawn without stasis at at the P ⬍ 0.05 level of confidence. All values were reported as
the points described above. From the 5-ml blood samples, a 4-ml means ⫾ SE.
aliquot was transferred to a test tube containing lithium heparin to
determine OsmP, plasma sodium concentration ([Na⫹]P), plasma RESULTS
potassium concentration, plasma glucose concentration, plasma lac-
tate concentration, total plasma proteins (TPP), and plasma Aldo Hydration indexes, hydration status, and amount of fluid
concentration ([Aldo]P). The remaining aliquot (1 ml) was analyzed ingested by the subjects are presented in Table 1. The euhy-
immediately for hematocrit (Hct) and hemoglobin (Hb). From the drated body weight and fluid status of subjects before each
25-ml blood sample, a 5-ml aliquot was treated as just described. trial, the degree of dehydration achieved, pretest USG, and
From the remaining sample, a 7-ml aliquot was transferred into an
EDTA-treated test tube for plasma AVP ([AVP]P) and glycerol
OsmP after dehydration did not differ significantly (P ⬎ 0.05).
([Glyc]P) analysis. A separate 5-ml aliquot was transferred into a The amount of fluid ingested during G and W was almost
chilled EDTA-treated test tube that contained 2,000 kallikrein-inhib- identical. During rehydration, urine output for the NF, G, and
itor units of aprotinin (Sigma Diagnostics, St. Louis, MO) for plasma W were 0, 138 ⫾ 63, and 523 ⫾ 93 ml, respectively. It should
atriopeptin ([AP]P) analysis. An 8-ml aliquot was transferred into a be noted that, during G, only two subjects were able to urinate.
chilled EDTA-treated tube for plasma renin activity (PRA) analysis. As a result of the urine output, although the amount of fluid
J Appl Physiol • VOL 100 • FEBRUARY 2006 • www.jap.org
REHYDRATION WITH GLYCEROL 445
intake was the same, hydration levels before the beginning of mmol/l) above NF (from 0.06 ⫾ 0.01 to 0.08 ⫾ 0.02 mmol/l)
exercise were ⫺4.06, ⫺0.89, and ⫺1.37% for NF, G, and W, and W (from 0.06 ⫾ 0.01 to 0.11 ⫾ 0.04 mmol/l). Glycerol
respectively. No statistically significant differences were found rehydration induced a higher level of OsmP than W at 0, 10,
in urine output or level of hydration between G and W before and 15 min of exercise and immediately postexercise (Fig. 1).
the beginning of exercise. The dehydration levels at the end of However, [Na⫹]P was lower (P ⬍ 0.05) during G than W.
the exercise were greater in NF (⫺5.5 ⫾ 0.3%) compared with [Na⫹]P during exercise was significantly higher for NF (P ⬍
G (⫺3.3 ⫾ 0.4%) and W (⫺3.4 ⫾ 0.4%), whereas no differ- 0.05) than both G and W (Fig. 1). TPP also was higher before
ences were found between G and W. Because volunteers and during exercise during NF (vs. G and W). Rehydration
exercised more during G, it seems that the rate of dehydration with glycerol significantly increased plasma volume (vs. W
is likely reduced during the G trial. Urine output after exercise and NF; P ⬍ 0.05), and those differences were maintained
for NF (118 ⫾ 20 ml) was significantly lower (P ⬍ 0.05) than throughout the exercise test (Fig. 1). Plasma glucose did not
for G (205 ⫾ 32 ml) and W (336 ⫾ 112 ml), but was not change significantly during any of the trials (Table 2), but was
different between G and W. higher than the 15-min value during all trials. However, at 15
None of our subjects reported nausea, headache, or gastro- min of exercise and immediately postexercise, plasma glucose
intestinal problems during, or the 24 h following, G. One was significantly higher during NF compared with G and W.
subject was stopped by the investigators during the G exercise During NF, plasma lactate was higher than G and W at 15 min
test (time: 51 min and 45 s), as his core temperature reached of exercise, whereas immediately postexercise this was true
39.5° for ⬎5 min (see criteria for termination in Experimental only compared with G (Table 2).
Protocol section).

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Exercise time to exhaustion during G (32.5 ⫾ 3.8 min) was Cardiovascular Responses
19% longer than during W (27.1 ⫾ 3.3 min) and 72% longer Q̇ was maintained to a similar level from 5 min (NF: 21.0 ⫾
than during NF (18.9 ⫾ 2.7 min). Whereas seven out of the 0.8, W: 20.6 ⫾ 0.9, and G: 21.4 ⫾ 0.8 l/min) to 15 min of
eight subjects exercised longer in G than in W, all subjects exercise (NF: 20.0 ⫾ 0.7, W: 21.0 ⫾ 0.9, and G: 20.5 ⫾ 0.8
exercised longer in G and W than in NF. However, no differ- l/min) for all trials. HR responses during exercise (Fig. 2) were
ences were observed among the three trials for V̇O2, R, VE, or not different between the G and W trials (P ⬎ 0.05). However,
RR, both at rest and during exercise (Table 2). during the first 15 min of NF, HR was significantly higher than
Plasma Responses both the G and W values. HR at the end of the exercise test did
not differ among trials. The SV at 15 min of exercise (Fig. 2)
Glycerol ingestion significantly increased (P ⬍ 0.05) for NF was significantly (P ⬍ 0.05) lower than the G and W.
[Glyc]P by ⬃100 times (from 0.06 ⫾ 0.01 to 9.94 ⫾ 0.23 During the G, there was a slight trend to maintain a higher SV

Table 2. Selected measurements at Dhy-Base, preexercise, and 5, 15, and 30 min of exercise
End (Post-Ex for
Variable Title Dhy-Base Preexercise 5 min 15 min Glucose, Lactate, and Glycerol)

V̇O2, l/min
No fluid 0.45⫾0.03 3.17⫾0.12 3.13⫾0.12 3.17⫾0.12
Glycerol 0.53⫾0.06 3.19⫾0.11 3.25⫾0.10 3.20⫾0.11
Water 0.47⫾0.03 3.21⫾0.12 3.23⫾0.11 3.20⫾0.12
R
No fluid 0.81⫾0.04 0.97⫾0.02 0.99⫾0.02 0.98⫾0.02
Glycerol 0.80⫾0.05 0.97⫾0.02 0.97⫾0.02 0.97⫾0.02
Water 0.81⫾0.04 0.99⫾0.02 0.96⫾0.02 0.98⫾0.02
V̇E, l/min
No fluid 14.0⫾1.2 91.4⫾5.3 108.0⫾6.5 99.2⫾5.1
Glycerol 15.7⫾2.5 86.7⫾4.5 98.4⫾4.7 95.3⫾4.1
Water 14.8⫾1.6 93.0⫾5.5 103.2⫾6.5 99.1⫾5.8
MAP, mmHg
No fluid 84.0⫾1.7 89.6⫾3.1 90.7⫾3.5 90.5⫾3.1
Glycerol 82.4⫾2.7 90.0⫾2.1 88.0⫾2.8 85.8⫾2.9
Water 85.6⫾2.0 92.2⫾3.7 89.1⫾3.8 87.6⫾3.8
RR, breaths/min
No fluid 17⫾2 42⫾4 52⫾4 46⫾4
Glycerol 17⫾2 37⫾3 45⫾4 44⫾4
Water 17⫾2 40⫾4 48⫾4 45⫾4
Plasma glucose, mmol/l
No fluid 4.9⫾0.2 4.8⫾0.1 4.9⫾0.2 6.4⫾0.4 8.0⫾0.5†
Glycerol 4.8⫾0.1 5.0⫾0.1 4.9⫾0.2 5.4⫾0.4* 6.8⫾0.8*†
Water 5.0⫾0.1 4.7⫾0.1 4.9⫾0.1 5.5⫾0.4* 7.0⫾0.5†
Plasma lactate, mmol/l
No fluid 1.0⫾0.1 1.0⫾0.0 5.1⫾1.0 8.8⫾2.0 7.9⫾1.7
Glycerol 1.0⫾0.1 1.3⫾0.1 4.5⫾0.9 6.8⫾1.3* 6.8⫾1.5*
Water 1.0⫾0.1 1.1⫾0.1 5.2⫾1.0 7.3⫾1.9* 7.5⫾1.8
Values are means ⫾ SE. V̇O2, oxygen uptake; R, respiratory quotient; V̇E, pulmonary ventilation; MAP, mean arterial pressure; RR, respiratory rate; End, end
of exercise; Post-Ex, postexercise. Significantly different from *no fluid and †preexercise value: P ⬍ 0.05.

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446 REHYDRATION WITH GLYCEROL

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Fig. 2. Heart rate (HR) and stroke volume (SV) responses during exercise.
Values are means ⫾ SE. *Significantly different from no fluid: P ⬍ 0.05.

than for W, but the difference was not statistically significant.


We were unable to collect enough Q̇ data after 15 min, since
several subjects stopped exercising before the 30-min mark.
Thermoregulatory Responses
Tre (Fig. 3) was significantly elevated (P ⬍ 0.05) during NF
compared with G and W at 0, 4, 8, and 12 min of exercise. Tsk
(Fig. 3) was significantly higher before the beginning of
exercise in NF, but not different during exercise. SkBF and
CVC were similar among the three experimental trials imme-
diately before the exercise test (Fig. 4). During exercise,
however, both SkBF and CVC were significantly higher (P ⬍
0.05) during G vs. W and NF. Sweating was higher (P ⬍ 0.05)
during G and W (G: 1,426 ⫾ 152 ml and W: 1,395 ⫾ 128 ml)
than NF (798 ⫾ 72 ml).
Hormonal Responses
[AVP]P was significantly decreased after rehydration during
Fig. 1. Plasma osmolality (OsmP), plasma sodium concentration ([Na⫹]P),
total plasma proteins (TPP), and percent change of plasma volume (PV) at
G and W, but increased (P ⬍ 0.05) in response to exercise for
dehydrated baseline (Dhy-Base), during, and immediately postexercise (Post- all of the trials. The postexercise level of vasopressin for NF
Ex). Rhy, rehydration. Values are means ⫾ SE. Significantly different from was significantly higher (P ⬍ 0.05) than for G and W (Fig. 5).
*no fluid and †water: P ⬍ 0.05. [AP]P level was similar before rehydration and did not change
during rehydration and exercise (Fig. 5).
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REHYDRATION WITH GLYCEROL 447
Similar improvements in exercise time to exhaustion have
been observed in the only other study in which glycerol was
used as a rehydration agent (44), as well as in a hyperhydration
study (36). Increase in exercise performance has been observed
in some studies that utilized glycerol-induced overhydration
before exercise (2, 9, 19), whereas others reported no perfor-
mance improvement (20, 29, 31, 48). Although previous glyc-
erol-induced overhydration studies have reported gastrointes-
tinal discomfort, headaches, or even blurred vision (9, 25, 26),
in our study, none of the subjects experienced any of those
symptoms during G or the 24 h following the study. The
absence of side effects has been also shown by others (27, 36,
40), and it has been speculated that high-concentration glycerol
solutions (i.e., 50%) are likely to induce these symptoms.
Although rehydration during G did not induce a statistically
significant better fluid balance compared with W, the small
difference in hydration (due to smaller urinary output) may
have provided physiologically greater body water “availabil-
ity” during the glycerol trial. Furthermore, since volunteers

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exercised more during G but sweated similarly with W, we
found that the rate of dehydration was reduced. Glycerol, on
the other hand, is a solute that increases the tonicity of blood,
and, although 2 liters of water were consumed, the 100-fold

Fig. 3. Rectal temperature (Tre) and mean weighted skin temperature (Tsk)
during exercise. Values are means ⫾ SE. Significantly different from *no fluid
trial and ‡0-min value: P ⬍ 0.05.

PRA (Fig. 6) decreased during rehydration for G and W and


increased significantly (P ⬍ 0.05) at the end of exercise (vs.
preexercise) in all experimental conditions. No differences
were found among the different trials. [Aldo]P was greater
(P ⬍ 0.05) after exercise (vs. preexercise) for all trials. At the
end of exercise, [Aldo] was lower (P ⬍ 0.05) in NF than W and
G (Fig. 6).
DISCUSSION

This study examined the endocrine, thermoregulatory, and


cardiovascular responses to exercise in the heat after oral
rehydration with glycerol. We hypothesized that rehydration
with glycerol (G trial) would prolong exercise time to exhaus-
tion, elicit greater cardiovascular stability, and lower thermo-
regulatory strain compared with NF or W trials. Rehydration
with glycerol did prolong exercise time to exhaustion in the
heat. However, this improved exercise capacity in the heat with
glycerol rehydration was not associated with any clear thermo-
regulatory or cardiovascular advantage compared with rehy-
dration with water only. Although glycerol rehydration re-
sulted in a higher SkBF during exercise in the heat, the
environmental conditions prevented any heat loss by radiation Fig. 4. Skin blood flow (SkBF) and cutaneous vascular conductance (CVC) in
or convection. As such, the improved SkBF had no thermo- the forearm. Values are means ⫾ SE. Significantly different from *no fluid,
regulatory impact. †water, and ‡0-min value: P ⬍ 0.05.

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448 REHYDRATION WITH GLYCEROL

exercise-heat stress offered no cardiovascular advantage com-


pared with water-induced hyperhydration.
It was hypothesized that glycerol-related ergogenicity could
enhance thermoregulatory responses, despite the fact that most
studies have not reported thermoregulatory benefits (9, 19, 36,
44). In the present study, because the environmental tempera-
ture was always higher than the Tsk, sweat evaporation was the
only means of heat dissipation. In contrast to the work of Lyon
et al. (28), which reported that glycerol-induced hyperhydra-
tion increased sweating and decreased exercise core tempera-
ture, no difference was found in the sweat rates of W and G.
Glycerol is a gluconeogenic substance that can be metabo-
lized in the liver and provides energy. Studies in experimental
animals have shown that rats fed with high doses of glycerol
performed better: they were protected from hypoglycemia and
exhibited both liver and muscle glycogen sparing (46). This
finding was not reported for humans, during 90 min of contin-
uous running (34), cycling to exhaustion for ⬃90 min (15), or
even cycling to exhaustion (80 –92 min) following a 36-h fast

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(32). These data suggest that the human liver may not be able
to metabolize glycerol fast enough to provide sufficient energy
during intense exercise. Similarly, we found no differences
between G and W in plasma glucose, lactate, V̇O2, and R,
indicating that glycerol did not contribute substantially as an
energy substrate.

Fig. 5. Plasma arginine vasopressin concentration ([AVP]P) and atriopeptin


concentration ([AP]P) responses to rehydration and exercise. Values are
means ⫾ SE. Significantly different from *no fluid, ‡Dhy-Base value, and
§preexercise value: P ⬍ 0.05.

increase in plasma glycerol (Table 2) led to maintenance of the


dehydration-induced elevation of Osmp. This osmotic effect,
along with a slight improvement in hydration level, could
induce the increased plasma volume observed during G com-
pared with W. A similar response of plasma volume has been
reported in other glycerol studies when plasma volume changes
were calculated from Hct and Hb values, like in the present
study (21, 29, 37, 44), or with Evans blue dilution technique (21).
Both Montner et al. (36) and Anderson et al. (2) indicated
that glycerol-induced hyperhydration maintained HR at a lower
level than a placebo. In the present study, although plasma
volume for G was always greater than that for W, HR, V̇O2,
blood pressure, and SV were not significantly different (G vs.
W), although SV showed a nonsignificant trend to be higher
during G (Fig. 2). It should be noted that we were unable to
measure SV for all subjects after 15 min of exercise, since few
of the volunteers managed to reach the 30-min time point at
which the CO2 rebreathing maneuver was performed for the
SV estimation (NF: 1, G: 5, and W: 3 subjects). We speculate
that SV would have been higher in the G compared with W and
NF at a later exercise stage, when cardiovascular strain was
likely greater. Based on the first 15 min of exercise, our data Fig. 6. Plasma renin activity (PRA) and aldosterone concentration ([Aldo]P)
are in agreement with a previous study by Latzka et al. (26), in responses to rehydration and exercise. Values are means ⫾ SE. Significantly
which glycerol-induced hyperhydration before uncompensable different from *no fluid, ‡Dhy-Base value, and §preexercise value: P ⬍ 0.05.

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REHYDRATION WITH GLYCEROL 449
In the present study, glycerol ingestion (G) altered fluid- 4. Armstrong LE, Maresh CM, Gabaree CL, Hoffman JR, Kavouras
regulating hormone responses to exercise in the heat (AVP, SA, Kenefick RW, Castellani JA, and Ahlquist LE. Thermal and
circulatory responses during exercise: effects of hypohydration, dehydra-
AP, and Aldo) and PRA in ways that were not observed when tion, and water intake. J Appl Physiol 82: 2028 –2035, 1997.
water alone was consumed (W). Glycerol increased OsmP; 5. Brandenberger G, Candas V, Follenius M, and Kahn JM. The influ-
therefore, we expected a consequent increase in plasma AVP ence of the initial state of hydration on endocrine responses to exercise in
(42) (i.e., osmoreceptors are potent stimulators of AVP), de- the heat. Eur J Appl Physiol Occup Physiol 58: 674 – 679, 1989.
6. Brozek J, Grande F, Anderson J, and Keys A. Densiometric analysis of
spite a plasma volume increase that could otherwise hamper
body composition: revision of some quantitative assumptions. Ann NY
AVP secretion (41). We also considered the observation that Acad Sci 110: 113–140, 1963.
the supraoptic and paraventricular nuclei of the hypothalamus 7. Convertino VA, Armstrong LE, Coyle EF, Mack GW, Sawka MN,
are more sensitive to sodium than to substances like glycerol, Senay LC, and Sherman WM. Position stand on exercise and fluid
glucose, or urea (14). At the end of exercise during the NF trial, replacement. Med Sci Sports Exerc 28: i–vii, 1996.
8. Convertino VA, Keil LC, and Greenleaf JE. Plasma volume, renin, and
however, AVP was approximately three times greater than G vasopressin responses to graded exercise after training. J Appl Physiol 54:
and W, likely because of a dehydration-induced hyperosmotic- 508 –514, 1983.
hypernatremic hypovolemia (22, 30). 9. Coutts A, Reaburn P, Mummery K, and Holmes M. The effect of
AP tended to decrease with rehydration and increase in glycerol hyperhydration on Olympic distance triathlon performance in
response to exercise (P ⬎ 0.05). This may have occurred high ambient temperatures. Int J Sport Nutr Exerc Metab 12: 105–119,
2002.
because plasma AP increases in response to atrial stretch (16). 10. Defares JG. Determination of Pv៮ CO2 from the exponential CO2 rise during
Freund et al. (13) reported an increased [AP]P as a result of rebreathing. J Appl Physiol 13: 159 –164, 1958.
maximal exercise in both trained and untrained subjects, 11. Dill DB and Costill DL. Calculation of percentage changes in volumes of

Downloaded from jap.physiology.org on November 2, 2007


whereas Kraemer et al. (24) reported that submaximal exercise blood, plasma, and red cells in dehydration. J Appl Physiol 37: 247–248,
(71% maximum V̇O2) did not alter [AP]P. In the present study, 1974.
12. Figaro MK and Mack GW. Regulation of fluid intake in dehydrated
AP remained unchanged during exercise, suggesting that sub- humans: role of oropharyngeal stimulation. Am J Physiol Regul Integr
maximal exercise in the heat, with or without glycerol, is not a Comp Physiol 272: R1740 –R1746, 1997.
potent secretory stimulus. In agreement with this finding, 13. Freund BJ, Claybaugh JR, Dice MS, and Hashiro GM. Hormonal and
Freund et al. (14) showed that glycerol did not affect ANP and vascular fluid responses to maximal exercise in trained and untrained
males. J Appl Physiol 63: 669 – 675, 1987.
other fluid-regulating hormones. 14. Freund BJ, Montain SJ, Young AJ, Sawka MN, DeLuca JP, Pandolf
PRA decreased during both rehydration conditions (P ⬍ KB, and Valeri CR. Glycerol hyperhydration: hormonal, renal, and
0.05), whereas Aldo did not change significantly. In response vascular fluid responses. J Appl Physiol 79: 2069 –2077, 1995.
to exercise, both PRA and Aldo increased significantly as 15. Gleeson M, Maughan RJ, and Greenhaff PL. Comparison of the effects
previously reported (8), but no changes were found among the of pre-exercise feeding of glucose, glycerol and placebo on endurance and
fuel homeostasis in man. Eur J Appl Physiol Occup Physiol 55: 645– 653,
trials. Brandenberger et al. (5) also observed that the initial 1986.
state of hydration does not affect exercise-induced responses of 16. Goetz K. Physiology and pathophysiology of atrial peptides. Am J Physiol
Aldo and PRA. Endocrinol Metab 254: E1–E15, 1988.
In conclusion, this study supported the hypothesis that re- 17. Gonzalez-Alonzo J, Mora-Rodriguez R, Below PR, and Coyle EF.
hydration with glycerol increased exercise time to exhaustion Dehydration markedly impairs cardiovascular function in hyperthermic
endurance athletes during exercise. J Appl Physiol 82: 1229 –1236, 1997.
during intensive cycling in the heat, likely by maintaining 18. Greenleaf JE. Problem: thirst, drinking behavior, and involuntary dehy-
greater plasma volume. Glycerol did not provide significant dration. Med Sci Sports Exerc 24: 645– 656, 1992.
cardiovascular or thermoregulatory advantages, although CVC 19. Hitchins S, Martin DT, Burke L, Yates K, Fallon K, Hahn A, and
was significantly increased. Furthermore, glycerol did not have Dobson GP. Glycerol hyperhydration improves cycle time trial perfor-
mance in hot humid conditions. Eur J Appl Physiol Occup Physiol 80:
a significant effect on fluid-regulating hormones, before or 494 –501, 1999.
during exercise. 20. Inder WJ, Swanney MP, Donald RA, Prickett TC, and Hellemans J.
The effect of glycerol and desmopressin on exercise performance and
ACKNOWLEDGMENTS hydration in triathletes. Med Sci Sports Exerc 30: 1263–1269, 1998.
The authors gratefully acknowledge the technical assistance of Frank T. 21. Jimenez C, Melin B, Koulmann N, Allevard AM, Launay JC, and
Hacker Jr., Dr. Tabbatha Elliott, Dr. NiCole Keith, Dr. Melissa Roti, Costas Savourey G. Plasma volume changes during and after acute variations of
Anastasiou, Timothy Bilodeau, David Blair, Sheri Huckleberry, Jeff Mace, and body hydration level in humans. Eur J Appl Physiol Occup Physiol 80:
Jason Sutherland. We also thank all of our subjects for their cooperation. 1– 8, 1999.
22. Kenefick RW, Maresh CM, Armstrong LE, Castellani JW, Riebe D,
GRANTS Echegaray ME, and Kavorous SA. Plasma vasopressin and aldosterone
responses to oral and intravenous saline rehydration. J Appl Physiol 89:
The study was partially supported by General Nutrition, Pittsburgh, PA; the 2117–2122, 2000.
University of Connecticut Research Foundation, Storrs, CT; and National 23. Koenigsberg PS, Martin KK, Hlava HR, and Riedesel ML. Sustained
Heart, Lung, and Blood Institute Grant HL-39818.
hyperhydration with glycerol ingestion. Life Sci 57: 645– 653, 1995.
24. Kraemer WJ, Armstrong LE, Hubbard RW, Marchitelli LJ, Leva N,
REFERENCES
Rock PB, and Dziados JE. Responses of plasma human atrial natriuretic
1. Alves da Silva G, el-Manshawi A, Heigenhauser GJ, and Jones NL. factor to high intensity submaximal exercise in the heat. Eur J Appl
Measurement of mixed venous carbon dioxide pressure by rebreathing Physiol Occup Physiol 57: 399 – 403, 1988.
during exercise. Respir Physiol 59: 379 –392, 1985. 25. Latzka WA, Sawka MN, Montain SJ, Skrinar GS, Fielding RA,
2. Anderson MJ, Cotter JD, Garnham AP, Casley DJ, and Febbraio Matott RP, and Pandolf KB. Hyperhydration: thermoregulatory effects
MA. Effect of glycerol-induced hyperhydration on thermoregulation and during compensable exercise-heat stress. J Appl Physiol 83: 860 – 866,
metabolism during exercise in heat. Int J Sport Nutr Exerc Metab 11: 1997.
315–333, 2001. 26. Latzka WA, Sawka MN, Montain SJ, Skrinar GS, Fielding RA,
3. Armstrong LE, Hubbard RW, Jones BH, and Daniels JT. Preparing Matott RP, and Pandolf KB. Hyperhydration: tolerance and cardiovas-
Alberto Salazar for the heat of the 1984 Olympic marathon. Phys cular effects during uncompensable exercise-heat stress. J Appl Physiol
Sportsmed 14: 73– 81, 1986. 84: 1858 –1864, 1998.

J Appl Physiol • VOL 100 • FEBRUARY 2006 • www.jap.org


450 REHYDRATION WITH GLYCEROL

27. Lyons TP and Riedesel ML. Glycerol-induced hyperhydration: its effects 37. Murray R, Eddy DE, Paul GL, Seifert JG, and Halaby GA. Physio-
on fluid compartments in the rat. Life Sci 53: 1779 –1787, 1993. logical responses to glycerol ingestion during exercise. J Appl Physiol 71:
28. Lyons TP, Riedesel ML, Meuli LE, and Chick TW. Effects of glycerol- 144 –149, 1991.
induced hyperhydration prior to exercise in the heat on sweating and core 38. Pitts GC, Johnson RE, and Consolazio FC. Work in the heat as affected
temperature. Med Sci Sports Exerc 22: 477– 483, 1990. by intake of water, salt and glucose. Am J Physiol 142: 253–259, 1944.
29. Magal M, Webster MJ, Sistrunk LE, Whitehead MT, Evans RK, and 39. Ramanathan NL. A new weighting system for mean surface temperature
Boyd JC. Comparison of glycerol and water hydration regimens on of the human body. J Appl Physiol 19: 531–533, 1964.
tennis-related performance. Med Sci Sports Exerc 35: 150 –156, 2003. 40. Riedesel ML, Allen DY, Peake GT, and Al-Qattan K. Hyperhydration
30. Maresh CM, Gabaree-Boulant CL, Armstrong LE, Judelson DA, with glycerol solutions. J Appl Physiol 63: 2262–2268, 1987.
Hoffman JR, Castellani JW, Kenefick RW, Bergeron MF, and Casa 41. Robertson GL and Athar S. The interaction of blood osmolality and
DJ. Effect of hydration status on thirst, drinking, and related hormonal blood volume in regulating plasma vasopressin in man. J Clin Endocrinol
responses during low-intensity exercise in the heat. J Appl Physiol 97: Metab 42: 613– 620, 1976.
39 – 44, 2004. 42. Robertson GL, Shelton RL, and Athar S. The osmoregulation of
31. Marino FE, Kay D, and Cannon J. Glycerol hyperhydration fails to vasopressin. Kidney Int 10: 25–37, 1976.
improve endurance performance and thermoregulation in humans in a 43. Sawka MN and Coyle EF. Influence of body water and blood volume on
warm humid environment. Pflügers Arch 446: 455– 462, 2003. thermoregulation and exercise performance in the heat. Exerc Sport Sci
32. Maughan RJ and Gleeson M. Influence of a 36 h fast followed by Rev 27: 167–218, 1999.
refeeding with glucose, glycerol or placebo on metabolism and perfor- 44. Scheett TP, Webster MJ, and Wagoner KD. Effectiveness of glycerol
mance during prolonged exercise in man. Eur J Appl Physiol Occup as a rehydrating agent. Int J Sport Nutr Exerc Metab 11: 63–71, 2001.
Physiol 57: 570 –576, 1988. 45. Shirreffs SM, Taylor AJ, Leiper JB, and Maughan RJ. Post-exercise
33. McKinley MJ and Johnson AK. The physiological regulation of thirst rehydration in man: effects of volume consumed and drink sodium
and fluid intake. News Physiol Sci 19: 1– 6, 2004. content. Med Sci Sports Exerc 28: 1260 –1271, 1996.
34. Miller JM, Coyle EF, Sherman WM, Hagberg JM, Costill DL, Fink 46. Terblanche SE, Fell RD, Juhlin-Dannfelt AC, Craig BW, and Holloszy

Downloaded from jap.physiology.org on November 2, 2007


WJ, Terblanche SE, and Holloszy JO. Effect of glycerol feeding on JO. Effects of glycerol feeding before and after exhausting exercise in
endurance and metabolism during prolonged exercise in man. Med Sci rats. J Appl Physiol 50: 94 –101, 1981.
Sports Exerc 15: 237–242, 1983. 47. Tourtellotte WW, Reinglass JL, and Newkirk TA. Cerebral dehydra-
35. Mitchell JW, Nadel ER, and Stolwijk JA. Respiratory weight losses tion action of glycerol. Clin Pharmacol Ther 13: 159 –171, 1972.
during exercise. J Appl Physiol 32: 474 – 476, 1972. 48. Wingo JE, Casa DJ, Berger EM, Dellis WO, Knight JC, and McClung
36. Montner P, Stark DM, Riedesel ML, Murata G, Robergs R, Timms JM. Influence of a pre-exercise glycerol hydration beverage on perfor-
M, and Chick TW. Pre-exercise glycerol hydration improves cycling mance and physiologic function during mountain-bike races in the heat. J
endurance time. Int J Sports Med 17: 27–33, 1996. Athl Train 39: 169 –175, 2004.

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