Beruflich Dokumente
Kultur Dokumente
6 Ove Sollie1, Per B. Jeppesen2, Daniel S. Tangen1, Fredrik Jernerén3, Birgitte Nellemann1,
7 Ditta Valsdottir1,4, Klavs Madsen5, Cheryl Turner3, Helga Refsum3,6, Bjørn S. Skålhegg6,
8 John L. Ivy7, & Jørgen Jensen1,*
9 1 Department of Physical Performance, Norwegian School of Sports Sciences, Oslo, Norway
10 2 Department of Clinical Medicine, Aarhus University, Aarhus, Denmark
11 3 Department of Pharmacology, University of Oxford, UK
12 4 Department of Medical Sciences, Atlantis Medical University College, Oslo, Norway
13 5 Department of Public Health - Sport Science, Aarhus University, 8000 Aarhus C
14 6 Department of Nutrition, Section for Molecular Nutrition, University of Oslo, Norway
15 7 Department of Kinesiology and Health Education, University of Texas at Austin, Austin,
16 TX; USA
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324 Results
325 Exhaustive exercise on day one (EXH): The workload used during EXH was 298±12 W and
326 corresponded to 73.0±3.0% of VO2max. Voluntary exhaustion at W73% occurred after 112±9
327 and 108±6 min prior to CHO and CHO+PROT, respectively. The athletes thereafter
328 performed 1 min sprints at 372±14 W (13±3 and 9±1 prior to CHO and CHO+PROT,
329 respectively). There were no significant differences in EXH time or number of sprints before
330 interventions (p=0.66 and p=0.37, respectively). Total carbohydrate oxidation during EXH
331 was 426±34 and 405±32 g before CHO and CHO+PROT, respectively, and with no
332 significant difference in carbohydrate oxidation between interventions (p=0.36) (Table 2).
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608
610 Ratnatunga C, Tousoulis D, Stefanadis C, Refsum H and Channon KM. MTHFR 677 C>T
612 homocysteine, in regulation of vascular redox state and endothelial function in human
614 2. Berardi JM, Noreen EE and Lemon PW. Recovery from a cycling time trial is enhanced with
617 3. Berardi JM, Price TB, Noreen EE and Lemon PW. Postexercise muscle glycogen recovery
618 enhanced with a carbohydrate-protein supplement. Med Sci Sports Exerc 38: 1106-1113,
619 2006.
620 4. Betts J, Williams C, Duffy K and Gunner F. The influence of carbohydrate and protein
621 ingestion during recovery from prolonged exercise on subsequent endurance performance. J
623 5. Betts JA and Williams C. Short-term recovery from prolonged exercise: exploring the
624 potential for protein ingestion to accentuate the benefits of carbohydrate supplements.
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627 metabolism during exercise and recovery in human subjects. J Physiol 514 ( Pt 1): 293-302,
628 1999.
629 7. Blomstrand E and Saltin B. BCAA intake affects protein metabolism in muscle after but not
630 during exercise in humans. Am J Physiol Endocrinol Metab 281: E365-E374, 2001.
631 8. Bohe J, Low A, Wolfe RR and Rennie MJ. Human muscle protein synthesis is modulated by
632 extracellular, not intramuscular amino acid availability: a dose-response study. J Physiol 552:
634 9. Borsheim E, Cree MG, Tipton KD, Elliott TA, Aarsland A and Wolfe RR. Effect of
635 carbohydrate intake on net muscle protein synthesis during recovery from resistance
637 10. Breen L, Philp A, Witard OC, Jackman SR, Selby A, Smith K, Baar K and Tipton KD. The
640 11. Breen L, Tipton KD and Jeukendrup AE. No effect of carbohydrate-protein on cycling
641 performance and indices of recovery. Med Sci Sports Exerc 42: 1140-1148, 2010.
642 12. Brosnan JT and Brosnan ME. Branched-chain amino acids: enzyme and substrate regulation.
644 13. Cormie P, McGuigan MR and Newton RU. Developing maximal neuromuscular power: Part
645 1--biological basis of maximal power production. Sports Med 41: 17-38, 2011.
21
647 strenuous exercise when fed carbohydrate. J Appl Physiol 61: 165-172, 1986.
648 15. Currell K and Jeukendrup AE. Validity, reliability and sensitivity of measures of sporting
650 16. Di Donato DM, West DW, Churchward-Venne TA, Breen L, Baker SK and Phillips SM.
651 Influence of aerobic exercise intensity on myofibrillar and mitochondrial protein synthesis in
652 young men during early and late postexercise recovery. Am J Physiol Endocrinol Metab 306:
654 17. Donges CE, Burd NA, Duffield R, Smith GC, West DW, Short MJ, Mackenzie R, Plank LD,
655 Shepherd PR, Phillips SM and Edge JA. Concurrent resistance and aerobic exercise stimulates
656 both myofibrillar and mitochondrial protein synthesis in sedentary middle-aged men. J Appl
658 18. Ferguson-Stegall L, McCleave EL, Ding Z, Doerner PG, III, Wang B, Liao YH, Kammer L, Liu Y,
660 improves subsequent exercise performance and intracellular signaling for protein synthesis. J
663 exercise during road race pro-cycling competition. Med Sci Sports Exerc 32: 1002-1006, 2000.
664 20. Goh Q, Boop CA, Luden ND, Smith AG, Womack CJ and Saunders MJ. Recovery from cycling
665 exercise: effects of carbohydrate and protein beverages. Nutrients 4: 568-584, 2012.
22
667 K. Protein intake during training sessions has no effect on performance and recovery during a
668 strenuous training camp for elite cyclists. J Int Soc Sports Nutr 13: 9, 2016.
669 22. Hermansen L, Hultman E and Saltin B. Muscle glycogen during prolonged severe exercise.
671 23. Hill KM, Stathis CG, Grinfeld E, Hayes A and McAinch AJ. Co-ingestion of carbohydrate and
672 whey protein isolates enhance PGC-1alpha mRNA expression: a randomised, single blind,
673 cross over study. J Int Soc Sports Nutr 10: 8, 2013.
674 24. Howarth KR, Moreau NA, Phillips SM and Gibala MJ. Coingestion of protein with
675 carbohydrate during recovery from endurance exercise stimulates skeletal muscle protein
677 25. Howarth KR, Phillips SM, MacDonald MJ, Richards D, Moreau NA and Gibala MJ. Effect of
678 glycogen availability on human skeletal muscle protein turnover during exercise and
680 26. Ivy JL. Muscle glycogen synthesis before and after exercise. Sports Med 11: 6-19, 1991.
681 27. Ivy JL. The regulation and synthesis of muscle glycogen by means of nutrient intervention. In:
682 The Encyclopaedia of Sports Medicine: An IOC Medical Commission Publication, Volume
683 19, edited by Maughan RJ. Oxford, UK: Wiley-Blackwell Science Ltd, 2013, p. 113-125.
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687 29. Jacobs RA, Rasmussen P, Siebenmann C, Diaz V, Gassmann M, Pesta D, Gnaiger E,
688 Nordsborg NB, Robach P and Lundby C. Determinants of time trial performance and
689 maximal incremental exercise in highly trained endurance athletes. J Appl Physiol (1985 )
691 30. Kato H, Suzuki K, Bannai M and Moore DR. Protein Requirements Are Elevated in Endurance
692 Athletes after Exercise as Determined by the Indicator Amino Acid Oxidation Method. PLoS
694 31. Kirk B, Mitchell J, Jackson M, Amirabdollahian F, Alizadehkhaiyat O and Clifford T. A2 Milk
695 Enhances Dynamic Muscle Function Following Repeated Sprint Exercise, a Possible Ergogenic
697 32. Kjeldahl J. Neue Methode zur Bestimmung des Stickstoffs in organischen Körper. Analytische
699 33. Lemon PWR and Mullin JP. Effect of initial muscle glycogen levels on protein catabolism
701 34. Levenhagen DK, Carr C, Carlson MG, Maron DJ, Borel MJ and Flakoll PJ. Postexercise protein
702 intake enhances whole-body and leg protein accretion in humans. Med Sci Sports Exerc 34:
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705 Jones' factors. Crit Rev Food Sci Nutr 48: 177-184, 2008.
706 36. Moberg M, Apro W, Ekblom B, van HG, Holmberg HC and Blomstrand E. Activation of
707 mTORC1 by leucine is potentiated by branched-chain amino acids and even more so by
708 essential amino acids following resistance exercise. Am J Physiol Cell Physiol 310: C874-C884,
709 2016.
710 37. Moore DR, Robinson MJ, Fry JL, Tang JE, Glover EI, Wilkinson SB, Prior T, Tarnopolsky MA
711 and Phillips SM. Ingested protein dose response of muscle and albumin protein synthesis
712 after resistance exercise in young men. Am J Clin Nutr 89: 161-168, 2009.
713 38. Moss BM, Refsnes PE, Abildgaard A, Nicolaysen K and Jensen J. Effects of maximal effort
714 strength training with different loads on dynamic strength, cross-sectional area, load-power
715 and load-velocity relationships. Eur J Appl Physiol Occup Physiol 75: 193-199, 1997.
716 39. Nelson AR, Phillips SM, Stellingwerff T, Rezzi S, Bruce SJ, Breton I, Thorimbert A, Guy PA,
717 Clarke J, Broadbent S and Rowlands DS. A protein-leucine supplement increases branched-
718 chain amino acid and nitrogen turnover but not performance. Med Sci Sports Exerc 44: 57-68,
719 2012.
720 40. Overmyer KA, Evans CR, Qi NR, Minogue CE, Carson JJ, Chermside-Scabbo CJ, Koch LG,
721 Britton SL, Pagliarini DJ, Coon JJ and Burant CF. Maximal oxidative capacity during exercise is
722 associated with skeletal muscle fuel selection and dynamic changes in mitochondrial protein
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726 42. Phillips SM, Atkinson SA, Tarnopolsky MA and MacDougall JD. Gender differences in leucine
727 kinetics and nitrogen balance in endurance athletes. J Appl Physiol (1985 ) 75: 2134-2141,
728 1993.
729 43. Romano-Ely BC, Todd MK, Saunders MJ and Laurent TS. Effect of an isocaloric carbohydrate-
730 protein-antioxidant drink on cycling performance. Med Sci Sports Exerc 38: 1608-1616, 2006.
731 44. Romijn JA, Coyle EF, Sidossis LS, Gastaldelli A, Horowitz JF, Endert E and Wolfe RR.
732 Regulation of endogenous fat and carbohydrate metabolism in relation to exercise intensity
734 45. Rowlands DS, Nelson AR, Phillips SM, Faulkner JA, Clarke J, Burd NA, Moore D and
735 Stellingwerff T. Protein-leucine fed dose effects on muscle protein synthesis after endurance
737 46. Rowlands DS, Rossler K, Thorp RM, Graham DF, Timmons BW, Stannard SR and
738 Tarnopolsky MA. Effect of dietary protein content during recovery from high-intensity
739 cycling on subsequent performance and markers of stress, inflammation, and muscle damage
740 in well-trained men. Appl Physiol Nutr Metab 33: 39-51, 2008.
741 47. Rowlands DS, Thomson JS, Timmons BW, Raymond F, Fuerholz A, Mansourian R, Zwahlen
743 Transcriptome and translational signaling following endurance exercise in trained skeletal
744 muscle: impact of dietary protein. Physiol Genomics 43: 1004-1020, 2011.
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746 feeding on recovery after intense exercise. Int J Sport Nutr Exerc Metab 17: 521-543, 2007.
747 49. Rowlands DS and Wadsworth DP. Effect of high-protein feeding on performance and
748 nitrogen balance in female cyclists. Med Sci Sports Exerc 43: 44-53, 2011.
749 50. Rustad PI, Sailer M, Cumming KT, Jeppesen PB, Kolnes KJ, Sollie O, Franch J, Ivy JL, Daniel H
750 and Jensen J. Intake of Protein Plus Carbohydrate during the First Two Hours after Exhaustive
751 Cycling Improves Performance the following Day. PLoS One 11: e0153229, 2016.
752 51. Samuelsson H, Moberg M, Apro W, Ekblom B and Blomstrand E. Intake of branched-chain
753 or essential amino acids attenuates the elevation in muscle levels of PGC-1alpha4 mRNA
754 caused by resistance exercise. Am J Physiol Endocrinol Metab 311: E246-E251, 2016.
755 52. Sanchez-Munoz C, Zabala M and Muros JJ. Nutritional intake and anthropometric changes of
756 professional road cyclists during a 4-day competition. Scand J Med Sci Sports 26: 802-808,
757 2016.
758 53. Saunders MJ, Kane MD and Todd MK. Effects of a carbohydrate-protein beverage on cycling
759 endurance and muscle damage. Med Sci Sports Exerc 36: 1233-1238, 2004.
760 54. She P, Zhou Y, Zhang Z, Griffin K, Gowda K and Lynch CJ. Disruption of BCAA metabolism in
761 mice impairs exercise metabolism and endurance. J Appl Physiol (1985 ) 108: 941-949, 2010.
762 55. Stisen AB, Stougaard O, Langfort J, Helge JW, Sahlin K and Madsen K. Maximal fat oxidation
763 rates in endurance trained and untrained women. Eur J Appl Physiol 98: 497-506, 2006.
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765 M. The importance of isometric maximum strength and peak rate-of-force development in
767 57. Tarnopolsky MA, MacDougall JD and Atkinson SA. Influence of protein intake and training
768 status on nitrogen balance and lean body mass. J Appl Physiol (1985 ) 64: 187-193, 1988.
769 58. Thomson JS, Ali A and Rowlands DS. Leucine-protein supplemented recovery feeding
770 enhances subsequent cycling performance in well-trained men. Appl Physiol Nutr Metab 36:
772 59. van Loon LJ, Greenhaff PL, Constantin-Teodosiu D, Saris WH and Wagenmakers AJ. The
773 effects of increasing exercise intensity on muscle fuel utilisation in humans. J Physiol 536:
775 60. Wilkinson SB, Phillips SM, Atherton PJ, Patel R, Yarasheski KE, Tarnopolsky MA and Rennie
776 MJ. Differential effects of resistance and endurance exercise in the fed state on signalling
777 molecule phosphorylation and protein synthesis in human muscle. J Physiol 586: 3701-3717,
778 2008.
779 61. Williams M, Raven PB, Fogt DL and Ivy JL. Effects of recovery beverages on glycogen
780 restoration and endurance exercise performance. J Strength Cond Res 17: 12-19, 2003.
781 62. Wolfe RR. Does exercise stimulate protein breakdown in humans?: isotopic approaches to
782 the problem. Med Sci Sports Exerc 19: 172-178, 1987.
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784 and urea metabolism in exercising humans. J Appl Physiol Respir Environ Exerc Physiol 52:
786 64. Zawadzki KM, Yaspelkis BB, III and Ivy JL. Carbohydrate-protein complex increases the rate
787 of muscle glycogen storage after exercise. J Appl Physiol 72: 1854-1859, 1992.
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796 Table 2: Blood glucose, blood lactate, VO2, RER, carbohydrate and fat oxidation, HR and
797 RPE during EXH. During the first interval (30 min), measurements were taken at 3-4 min and
798 the last two min of the interval. On the rest of the intervals (20 min), measurements were
799 taken during the two last min of each interval. Measurements were also taken the last two min
800 before voluntary exhaustion (W73%End). Furthermore, blood lactate and blood glucose were
801 measured before and after the one min bouts at 90% of VO2max.
802
803
804 Figures
805 Figure 1: Design of the interventions. The protocol was completed twice in a double-blinded,
806 randomized, balanced, crossover experimental design. The athletes reported to the laboratory
807 11:30 AM on day 1 and started an exhaustive training protocol (EXH) at 12:00 PM. EXH
808 consisted of exhaustive cycling at 73% and 90% of VO2max, and two 10 s sprints (at the start
809 and at the end of EXH). After EXH, an 18 h standardized isoenergetic recovery period started,
810 where the subjects where fed either carbohydrate alone (CHO group) or carbohydrate and
811 protein (CHO+PROT group) the first 2 h. The diet during the rest of the recovery period was
812 similar for both interventions. On day 2, the athletes ate a standardized breakfast at 07:00 AM
813 and reported to the laboratory 08:30 AM. The performance tests started 09:00 AM, and
814 consisted of a 10 s sprint, 30 min at W73%, a time-trial performance test, and a second 10 s
815 sprint 5 min after the time-trial.
816
817 Figure 2: Plasma metabolites before and after exhaustive exercise, and during the dietary
818 interventions. Carbohydrate or carbohydrate and protein were provided during the immediate
819 recovery (0-120 min) as described. Plasma levels of glucose (A), insulin (B), FFA (C),
820 glycerol (D), uric acid (E), leucine (F), isoleucine (G), valine (H) were measured before
821 exercise (-110 min) and during the 2 h of recovery from exhaustive exercise when participants
822 received energy drinks. Values are mean ± SEM. E: p<0.05 compared to basal value before
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826 Figure 3. Performance from the time-trial and 10 s sprint tests the day after the diet
827 interventions. Average time spent on the time trial (A), power curve during TT (B), mean
828 power during the sprints (C). Values are mean ± SEM. * In Fig. 3A and 3C p<0.05 compared
829 to CHO. In Fig. 3B, * p<0.05 comparing CHO and CHO+PROT for each 10% interval.
830
831 Figure 4: Measurements from the performance test on day 2. Time 0 is at arrival before warm
832 up. Time 15 and 30 is after 15 and 30 min cycling at of W73%. Time 80 is at the end of TT,
833 and 95 is 15 min after completion of TT. RPE (A) and heart rate (B) were taken at 4, 18, 30
834 min of W73%, and at 10%, 20%, 60%, 80% and at 15 min and at the end of the time trial. VO2
835 (C), RER (D), carbohydrate oxidation (E), fat oxidation (F), blood lactate (G), plasma
836 glucose (H), were taken at 4, 18 and 30 min of W73%, and at 15 min and at the end of TT.
837 Glucose was also taken at time 0. Leucine (I), valine (J), isoleucine (K), FFA (L), glycerol
838 (M), uric acid (N) and insulin (O) were taken before, at 15 min, and at the end of W73%, and
839 at15 min and at the end of TT and 15 min after TT. Values are mean ± SEM. T= Time effect;
840 P=treatment effect between groups. * Time points with differences between CHO and
841 CHO+PROT (p<0.05).
842
843 Figure 5: Nitrogen balance and urine excretion of nitrogen, urea and BCAA during the 18 h
844 recovery period. Nitrogen balance was calculated from protein intake and urine nitrogen.
845 Nitrogen balances for CHO and CHO+PROT during the recovery period with individual
846 responses (A) and responses for the different time periods for urine collection (B), total
847 nitrogen excretion (C), total urea excretion (D), * Significant differences between groups.
848 Values are mean ± SEM. p<0.05.
849
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852 Table 1
Time Time after Carbohydrate Protein Fat Energy content
EXH (g·kg-1) (g·kg-1) (g·kg-1) (kJ·kg-1)
854 Table 2
0 min 3-4 min 28-30 min 48-50 min 68-70 min W73% End Exhaustion
VO2 (ml·min-1)
4354±172b 4352±173b α 4509±213b,c,d,e
CHO 4163±162 4336±203b,
CHO+PRO 4149±177 4280±164b 4360±177b 4422±192b 4516±248 b,c,d,e
HR (beats·min-1)
169±3b 170±4b α 178±3 b,c,d,e
CHO 156±4 167±2b,
CHO+PRO 155±4 166±3b 166±4b 168±4b 175±3 b,c,d,e
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859
860 Figures
861
862 Figure 1
863
864
865
866
867
868
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870
871
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873
874
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876
877
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880
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