906
Koopman et al.
97. Booth FW, Chakravarthy MV, Spangenburg EE. Exercise and
114. Garlick PJ. The role of leucine in the regulation of protein
gene expression: physiological regulation of the human gen-
metabolism. J Nutr 2005; 135: 1553S-6S
ome through physical activity. J Physiol 2002; 543: 399-411
115. Buse MG, Reid SS. Leucine: a possible regulator of protein
98. Dreyer HC, Fujita S, Cadenas JG, et al. Resistance exercise
turnover in muscle. J Clin Invest 1975; 56: 1250-61
increases AMPK activity and reduces 4E-BP1 phosphorylation
116. Li JB, Jefferson LS. Influence of amino acid availability on
and protein synthesis in human skeletal muscle. J Physiol
protein turnover in perfused skeletal muscle. Biochim Biophys
2006; 576: 613-24
Acta 1978; 544: 351-9
99. Koopman R, Zorenc AH, Gransier RJ, et al. The increase in
S6K1 phosphorylation in human skeletal muscle following
117. Morgan HE, Earl DC, Broadus A, et al. Regulation of protein
resistance exercise occurs mainly in type II muscle fibers. Am
synthesis in heart muscle I: effect of amino acid levels on
J Physiol Endocrinol Metab 2006; 290: E1245-52
protein synthesis. J Biol Chem 1971; 246: 2152-62
100. Evans WJ. Effects of exercise on body composition and func-
118. Anthony JC, Anthony TG, Kimball SR, et al. Orally adminis-
tional capacity of the elderly. J Gerontol A Biol Sci Med Sci
tered leucine stimulates protein synthesis in skeletal muscle of
1995; 50: 147-50
postabsorptive rats in association with increased eIF4F forma-
101. Fiatarone MA, Marks EC, Ryan ND, et al. High-intensity
tion. J Nutr 2000; 130: 139-45
strength training in nonagenarians: effects on skeletal muscle.
119. Proud CG. mTOR-mediated regulation of translation factors by
JAMA 1990; 263: 3029-34
amino acids. Biochem Biophys Res Commun 2004; 313:
102. Frontera WR, Meredith CN, O'Reilly KP, et al. Strength condi-
429-36
tioning in older men: skeletal muscle hypertrophy and im-
proved function. J Appl Physiol 1988; 64: 1038-44
120. Nair KS, Schwartz RG, Welle S. Leucine as a regulator of whole
body and skeletal muscle protein metabolism in humans. Am J
103. Ades PA, Ballor DL, Ashikaga T, et al. Weight training im-
proves walking endurance in healthy elderly persons. Ann
Physiol 1992; 263: E928-34
Intern Med 1996; 124: 568-72
121. Karlsson HK, Nilsson PA, Nilsson J, et al. Branched-chain
104. Bamman MM, Hill VJ, Adams GR, et al. Gender differences in
amino acids increase p70s6 kinase phosphorylation in human
resistance-training-induced myofiber hypertrophy among old-
skeletal muscle after resistance exercise. Am J Physiol En-
er adults. J Gerontol A Biol Sci Med Sci 2003; 58: 108-16
docrinol Metab 2004; 287: 1-7
105. Fiatarone MA, O'Neill EF, Ryan ND, et al. Exercise training
122. Liu Z, Jahn LA, Long W, et al. Branched chain amino acids
and nutritional supplementation for physical frailty in very
activate messenger ribonucleic acid translation regulatory pro-
elderly people. N Engl J Med 1994; 330: 1769-75
teins in human skeletal muscle, and glucocorticoids blunt this
106. Frontera WR, Hughes VA, Krivickas LS, et al. Strength training
action. J Clin Endocrinol Metab 2001; 86: 2136-43
in older women: early and late changes in whole muscle and
single cells. Muscle Nerve 2003; 28: 601-8
123. Matthews DE. Observations of branched-chain amino acid ad-
ministration in humans. J Nutr 2005; 135: 1580S-4S
107. Frontera WR, Meredith CN, O'Reilly KP, et al. Strength train-
ing and determinants of
VO
2max
in older men. J Appl Physiol
124. Dardevet D, Sornet C, Balage M, et al. Stimulation of in vitro rat
1990; 68: 329-33
muscle protein synthesis by leucine decreases with age. J Nutr
108. Lexell J, Downham DY, Larsson Y, et al. Heavy-resistance
2000; 130: 2630-5
training in older Scandinavian men and women: short- and
125. Dardevet D, Sornet C, Bayle G, et al. Postprandial stimulation
long-term effects on arm and leg muscles. Scand J Med Sci
of muscle protein synthesis in old rats can be restored by a
Sports 1995; 5: 329-41
leucine-supplemented meal. J Nutr 2002; 132: 95-100
109. Vincent KR, Braith RW, Feldman RA, et al. Resistance exercise
126. Combaret L, Dardevet D, Rieu I, et al. A leucine-supplemented
and physical performance in adults aged 60 to 83. J Am Geriatr
Soc 2002; 50: 1100-7
diet restores the defective postprandial inhibition of
proteasome-dependent proteolysis in aged rat skeletal muscle.
110. Meredith CN, Frontera WR, O'Reilly KP, et al. Body composi-
J Physiol 2005; 569: 489-99
tion in elderly men: effect of dietary modification during
strength training. J Am Geriatr Soc 1992; 40: 155-62
127. Koopman R, Verdijk LB, Manders RJF, et al. Co-ingestion of
111. Freyssenet D, Berthon P, Denis C, et al. Effect of a 6-week
protein and leucine stimulates muscle protein synthesis rates to
endurance training programme and branched-chain amino acid
the same extent in young and elderly lean men. Am J Clin Nutr
supplementation on histomorphometric characteristics of aged
2006; 84: 623-32
human muscle. Arch Physiol Biochem 1996; 104: 157-62
112. Campbell WW, Crim MC, Young VR, et al. Effects of resis-
tance training and dietary protein intake on protein metabolism
Correspondence: Dr RenŽe Koopman, Department of Move-
in older adults. Am J Physiol 1995; 268: E1143-53
ment Sciences, Maastricht University, PO Box 616, 6200 MD
113. Welle S, Thornton CA. High-protein meals do not enhance
Maastricht, The Netherlands.
myofibrillar synthesis after resistance exercise in 62- to 75-yr-
old men and women. Am J Physiol 1998; 274: E677-83
E-mail: R.Koopman@HB.unimaas.nl
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