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Glucose e Frutose Durante Exercicio fisico

Glucose e Frutose Durante Exercicio fisico

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96:1277-1284, 2004. First published Dec 2, 2003; doi:10.1152/japplphysiol.00974.2003
 J Appl Physiol
Asker E. JeukendrupRoy L. P. G. Jentjens, Luke Moseley, Rosemary H. Waring, Leslie K. Harding and
 
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40 articles, 23 of which you can access free at:This article citeshttp://jap.physiology.org/cgi/content/full/96/4/1277#BIBL6 other HighWire hosted articles, the first 5 are:This article has been cited by[PDF] [Full Text] [Abstract] , August 1,2005; 289(2): E206-E211.
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JeukendrupL. Moseley, R. L. P. G. Jentjens, R. H. Waring, R. M. Harris, L. K. Harding and A. E. 
[U-13C]glucose tracersMeasurement of exogenous carbohydrate oxidation: a comparison of [U-14C]glucose and
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 J Appl Physiol
S. E. Yeo, R. L. P. G. Jentjens, G. A. Wallis and A. E. Jeukendrup
Caffeine increases exogenous carbohydrate oxidation during exercise
R. L. P. G. Jentjens, K. Underwood, J. Achten, K. Currell, C. H. Mann and A. E. Jeukendrup
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A. E. Jeukendrup, L. Moseley, G. I. Mainwaring, S. Samuels, S. Perry and C. H. Mann
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J. Achten, R. L. Jentjens, F. Brouns and A. E. Jeukendrup
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including high-resolution figures, can be found at:Updated information and serviceshttp://jap.physiology.org/cgi/content/full/96/4/1277can be found at:
 Journal of Applied Physiology
aboutAdditional material and informationhttp://www.the-aps.org/publications/japplThis information is current as of November 2, 2007 .
http://www.the-aps.org/.ISSN: 8750-7587, ESSN: 1522-1601. Visit our website atPhysiological Society, 9650 Rockville Pike, Bethesda MD20814-3991. Copyright© 2005 by the American Physiological Society.those papers emphasizing adaptive and integrative mechanisms. It is published 12 times a year (monthly) by the Americanpublishes original papers that deal with diverse areas of research in applied physiology, especially
 Journal of Applied Physiology
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Oxidation of combined ingestion of glucose and fructose during exercise
Roy L. P. G. Jentjens,
1
Luke Moseley,
1
Rosemary H. Waring,
2
Leslie K. Harding,
3
andAsker E. Jeukendrup
1
1
 Human Performance Laboratory, School of Sport and Exercise Sciences, and 
2
Schoolof Biosciences, University of Birmingham, Birmingham B15 2TT; and 
3
Physics and  Nuclear Medicine Department, City Hospital, Birmingham B18 7QH, United Kingdom
Submitted 9 September 2003; accepted in final form 3 November 2003
Jentjens, Roy L. P. G., Luke Moseley, Rosemary H. Waring,Leslie K. Harding, and Asker E. Jeukendrup.
Oxidation of com-bined ingestion of glucose and fructose during exercise.
J ApplPhysiol
96: 1277–1284, 2004. First published December 2, 2003;10.1152/japplphysiol.00974.2003.—The purpose of the present studywas to examine whether combined ingestion of a large amount of fructose and glucose during cycling exercise would lead to exogenouscarbohydrate oxidation rates
1 g/min. Eight trained cyclists (maxi-mal O
2
consumption: 62
3 ml
kg
1
min
1
) performed four exer-cise trials in random order. Each trial consisted of 120 min of cyclingat 50% maximum power output (63
2% maximal O
2
consumption),while subjects received a solution providing either 1.2 g/min of glucose (Med-Glu), 1.8 g/min of glucose (High-Glu), 0.6 g/min of fructose
1.2 g/min of glucose (Fruc
Glu), or water. The ingestedfructose was labeled with [U-
13
C]fructose, and the ingested glucosewas labeled with [U-
14
C]glucose. Peak exogenous carbohydrate oxi-dation rates were
55% higher (
P
0.001) in Fruc
Glu (1.26
0.07 g/min) compared with Med-Glu and High-Glu (0.80
0.04 and0.83
0.05 g/min, respectively). Furthermore, the average exogenouscarbohydrate oxidation rates over the 60- to 120-min exercise periodwere higher (
P
0.001) in Fruc
Glu compared with Med-Glu andHigh-Glu (1.16
0.06, 0.75
0.04, and 0.75
0.04 g/min,respectively). There was a trend toward a lower endogenous carbo-hydrate oxidation in Fruc
Glu compared with the other two carbo-hydrate trials, but this failed to reach statistical significance (
P
0.075). The present results demonstrate that, when fructose andglucose are ingested simultaneously at high rates during cyclingexercise, exogenous carbohydrate oxidation rates can reach peak values of 
1.3 g/min.substrate utilization; carbohydrate absorption; isotopic tracers; metab-olism
IT HAS BEEN SHOWN THAT CARBOHYDRATE
(CHO) feedings duringprolonged moderate- to high-intensity exercise can postponefatigue and enhance exercise performance when the exerciseduration is
45 min or longer (2, 17, 41). The observedimprovements in performance with CHO ingestion have con-tributed to better maintenance of plasma glucose concentra-tions and high rates of CHO oxidation late in exercise (3, 5),when muscle and liver glycogen levels are low.Studies that have used either stable (16, 21, 29, 36, 40) orradioactive isotopes (3, 11) to quantify exogenous CHO oxi-dation during exercise have found peak oxidation rates of 
1g/min (for review, see Refs. 10, 18, 21). Even when CHO wasingested at high rates (up to 3.0 g/min), oxidation rates did notexceed 1 g/min (16, 21, 29, 36, 40). In most of the above-citedstudies, glucose or glucose polymer was the type of CHOingested. Although direct evidence is lacking, it has beensuggested that the absorption capacity of glucose in the intes-tine (31) is a limiting factor for the oxidation of ingestedglucose (18, 21).A number of studies have compared the oxidation rates of various types of ingested CHO with the oxidation of exogenousglucose during exercise (for review, see Ref. 18). From thesestudies, it appears that the rate of oxidation of ingested maltose(11), sucrose (27, 40), glucose polymer (26), and maltodextrin(32) is fairly similar to the oxidation rate of ingested glucose.However, significantly lower exogenous CHO oxidation rateshave been reported for fructose (
20–25% lower) (1, 15, 25,26) and galactose (
50% lower) (23) compared with glucose.There have been suggestions that the lower oxidation rate of fructose is due to a lower rate of absorption (1, 18). Further-more, both fructose and galactose have to be converted intoglucose in the liver before they can be oxidized, and this mayslow down the rate of oxidation.Intestinal transport of glucose occurs via a sodium-depen-dent glucose transporter (SGLT1), which is located in thebrush-border membrane (8). It is likely that SGLT1-transport-ers are saturated at a glucose ingestion rate of 
1 g/min, whichcould explain why higher glucose intake rates do not result inoxidation rates higher than 1.0–1.1 g/min (21, 40). In contrastto glucose, fructose is absorbed from the intestine by GLUT-5,a sodium-independent facilitative fructose transporter (4, 8). Itis not unlikely that, when a mixture of glucose and fructose isingested, there is less competition for absorption comparedwith the ingestion of an isoenergetic amount of glucose (orfructose), and this may increase the availability of CHO intothe bloodstream for oxidation.To our knowledge, only two studies have attempted toinvestigate the oxidation of combined ingestion of glucose andfructose (1, 33). The results of these studies are contradictory,which might be due to differences in study design or subjectselection. Riddell et al. (33) studied 12 children who ingested67.5 g of a mixture of glucose and fructose or an isoenergeticamount of glucose during 90 min of exercise at 55% maximalO
2
consumption (V˙
O
2 max
). No difference was found in exog-enous CHO oxidation between the two CHO trials, and peak oxidation rates did not exceed 0.36 g/min. In a study by Adopoet al. (1), six trained subjects cycled for 120 min at 61%V˙
O
2 max
while ingesting 100 g of glucose or fructose or amixture of 50 g glucose and fructose. The addition of fructoseto the glucose drink increased total exogenous CHO oxidationby 27% compared with the isoenergetic glucose drink (0.61 vs.
Address for reprint requests and other correspondence: A. E. Jeukendrup,School of Sport and Exercise Sciences, Univ. of Birmingham, Edgbaston, B152TT, Birmingham, UK (E-mail: A.E.Jeukendrup@bham.ac.uk).The costs of publication of this article were defrayed in part by the paymentof page charges. The article must therefore be hereby marked “
advertisement 
in accordance with 18 U.S.C. Section 1734 solely to indicate this fact.
 J Appl Physiol
96: 1277–1284, 2004.First published December 2, 2003; 10.1152/japplphysiol.00974.2003.8750-7587/04 $5.00 Copyright
©
2004 the American Physiological Societyhttp://www.jap.org 1277
  on o em b  e , 0  0  j   a p. p y  s i   ol   o g y . o g ownl   o a d  e d f   om 
 
0.48 g/min). It should be noted that the amounts of CHOingested in the study of Adopo et al. were relatively small(
0.8 g/min), and, therefore, glucose (and fructose) transport-ers were probably not fully saturated. At present, there are nostudies available in the literature that have investigated whethercombined ingestion of large amounts of glucose and fructosecan result in exogenous CHO oxidation rates that exceed 1g/min. Furthermore, in the study of Adopo et al., it was notpossible to determine the oxidation rate of both glucose andfructose in one trial, because only one of the ingested CHO wasisotopically labeled. To measure the total oxidation rate of theingested glucose plus fructose mixture, two separate trials wereperformed, and this may have confounded the results.We hypothesized that combined ingestion of a large amountof fructose (72 g) and glucose (144 g) during 2 h of exercisewould lead to exogenous CHO oxidation rates higher than 1g/min. The ingested fructose was labeled with [U-
13
C]fructose,and the ingested glucose was labeled with [U-
14
C]glucose,which enabled us to measure the oxidation rates of glucose andfructose when ingested simultaneously.
METHODS
Subjects.
Eight trained male cyclists or triathletes, aged 29.0
2.7yr and with a body mass of 75.1
1.8 kg, took part in this study.Before participation, each of the subjects was fully informed of thepurpose and risks associated with the procedures, and a written,informed consent was obtained. All subjects were healthy, as assessedby a general health questionnaire. The study was approved by theSouth Birmingham Local Research Ethics Committee and the UKadministration of Radioactive Substance Advisory Committee.
Preliminary testing.
At least 1 wk before the start of the experi-mental trials, an incremental cycle exercise test to volitional exhaus-tion was performed to determine the individual maximum poweroutput (W
˙
max) and V
˙
O
2 max
. This test was performed on an electro-magnetically braked cycle ergometer (Lode Excalibur Sport, Gro-ningen, The Netherlands), modi
ed to the con
guration of a racingbicycle with adjustable saddle height and handlebar position. After thesubjects reported to the laboratory, body mass (Seca Alpha, Hamburg,Germany) and height were recorded. Subjects then started cycling at95 W for 3 min, followed by incremental steps of 35 W every 3 minuntil exhaustion. Heart rate (HR) was recorded continuously by aradiotelemetry HR monitor (Polar Vantage, Kempele, Finland).W
˙
max was calculated from the last completed work rate, plus thefraction of time spent in the
nal, noncompleted work rate, multipliedby the work rate increment. The results were used to determine thework rate corresponding to 50% W
˙
max, which was later employed inthe experimental exercise trials. Breath-by-breath measurements wereperformed throughout exercise by using an online automated gasanalysis system (Oxycon Alpha, Jaeger, Wuerzberg, Germany). Thevolume sensor was calibrated by using a 3-liter calibration syringe,and the gas analyzers were calibrated by using a 5.03% CO
2
-94.97%N
2
gas mixture. Oxygen consumption (V
˙
O
2
) was considered to bemaximal (V
˙
O
2 max
) when at least two of the three following criteriawere met:
1
) a leveling off of V
˙
O
2
with increasing workload (increaseof no more than 2 ml
kg
1
min
1
),
2
) a HR within 10 beats/min of predicted maximum (HR 220 minus age), and
3
) a respiratory ex-change ratio (RER)
1.05. V
˙
O
2 max
was calculated as the averageoxygen uptake over the last 60 s of the test. The V
˙
O
2 max
and W
˙
maxachieved during the incremental exercise test were 62
3ml
kg
1
min
1
and 374
12 W, respectively.
 Experimental design.
Each subject performed four exercise trials,which consisted of 120 min of cycling at 50% W
˙
max, while ingestingan 8.7% glucose drink (Med-Glu), a 13.1% glucose drink (High-Glu),an isoenergetic fructose plus glucose drink (Fruc
Glu) (the ingestedfructose-to-glucose ratio was 1:2), or plain water (Wat). To quantifyexogenous CHO oxidation, the ingested fructose was enriched with[U-
13
C]fructose, and the ingested glucose was labeled with[U-
14
C]glucose. The order of the experimental drinks was counter-balanced in a crossover design. Experimental trials were separated byat least 7 days.
 Diet and activity before testing.
Subjects were asked to record theirfood intake and activity pattern 2 days before the
rst exercise trialand were then instructed to follow the same diet and exercise activitiesbefore the other three trials. In addition, 5
7 days before eachexperimental testing day, they were asked to perform an intensetraining session (
glycogen-depleting
exercise bout) in an attempt toempty any
13
C-enriched glycogen stores. Subjects were further in-structed not to consume any food products with a high naturalabundance of 
13
C (CHO derived from C4 plants: corn, sugar cane) atleast 1 wk before and during the entire experimental period to reducethe background shift (change in
13
CO
2
) from endogenous substratestores.
Protocol.
Subjects reported to the Human Performance Laboratoryin the morning (between 7:00 and 9:00 AM) after an overnight fast(10
12 h) and having refrained from any strenuous activity or drink-ing any alcohol in the previous 24 h. For a given subject, all trialswere conducted at the same time of the day to avoid any in
uence of circadian variance. On arrival in the laboratory, a
exible 21-gaugeTe
on catheter (Quickcath, Baxter BV, Norfolk, UK) was inserted inan antecubital vein of an arm and attached to a three-way stopcock (Sims Portex, Kingsmead, UK) to allow for repeated blood samplingduring exercise. The catheter was kept patent by
ushing with 1.0
1.5ml of isotonic saline (0.9% Baxter) after each blood sample collection.After voiding, subjects were weighed in cycling shorts to the nearest0.1 kg by using a platform scale (Seca Alpha, Hamburg, Germany).The subjects then mounted a cycle ergometer, and a resting breathsample was collected in 10-ml Exetainer tubes (Labco Brow Works,High Wycombe, UK), which were
lled directly from a mixingchamber in duplicate to determine the
13
C-to-
12
C ratio (
13
C/ 
12
C) inthe expired air. A second resting breath sample was collected for laterdetermination of 
14
CO
2
-speci
c activity. The expired air of eachsecond breath sample was collected in a 6-liter anesthetic gas bag byusing a two-way Hans Rudolph valve and subsequently passedthrough a CO
2
trapping solution, containing 1 ml of hyamine hydrox-ide in 1 M methanol (Zinsser Analytic, Berkshire, UK), 2 ml of 96%ethanol (BDH Laboratory Supplies, Poole, UK), and one to two dropsof phenolphthalein (Riedel-de Hae
¨
n, Seeize, Germany). The expiredair was bubbled for 2
3 min through the pink-colored CO
2
trappingmixture until the solution became clear, at which point exactly 1 mmolof CO
2
was trapped (37). Seventeen milliliters of liquid scintillationcocktail (Ready Gel, Beckman Coulter, High Wycombe, UK) wasthen added to the solution, and
14
CO
2
radioactivity in disintegrations/ min (dpm, later converted to dpm/mmol) was subsequently counted ina liquid scintillation counter (Beckman, LS 1800).Next, a resting blood sample (10 ml) was taken and stored on iceuntil centrifugation. Subjects then started a 120-min exercise bout ata work rate equivalent to 50% W
˙
max (63
2% V
˙
O
2 max
). Additionalblood samples were drawn at 15-min intervals during exercise. Ex-pired breath samples were collected every 15 min until the end of exercise. V
˙
O
2
, carbon dioxide production (V
˙
CO
2
), and RER weremeasured every 15 min for periods of 4 min by using an onlineautomated gas analysis system, as previously described.During the
rst 3 min of exercise, subjects drank an initial bolus(600 ml) of one of the four experimental drinks: Med-Glu, High-Glu,Fruc
Glu, or Wat. Thereafter, every 15 min, a beverage volume of 150 ml was provided. The total
uid provided during the 120-minexercise bout was 1.65 liters. The average rate of glucose intake in theMed-Glu and High-Glu trial was 1.2 and 1.8 g/min, respectively. Inthe Fruc
Glu trial, subjects ingested, on average, 0.6 and 1.2 g/minof fructose and glucose, respectively.
1278
OXIDATION OF COMBINED INGESTION OF GLUCOSE AND FRUCTOSE
 J Appl Physiol
VOL 96
APRIL 2004
www.jap.org
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