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    00009

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    s.si1e7/105 15082 <0 MEDION AND SCENCE IN SPORTS AND EXERGISE Connie Y85 by Ie moan Cotes ol Sports Mode Amino Acid and Protein Metabolism in Exercise CHAIRMAN: GeorGE A. Brooks Amino acid and protein exercise and recovery GEORGE A. BROOKS Exercise Physiology Laboratory, Department of Physical Education, University of California, Berkeley, CA 94720 ABSTRACT BROOKS,G, A, Amino acid and protein metabolism during exercise and recovery, Med. Sci. Sports Exerc., Vol 19, No. 5 (Supplement), pp. S150-$156, 1987. The integrated use of several energy sources allows high muscular power outputs to be sustained. Muscle glycogen provides the major fuel source for muscular exercise, but other fies an provide alternative energy sources which allow for muscle gly- coget-sparing and an increased potential for prolonged high meta- bolie rates Blood-borre glucose, derived fiom liver glycogenolysis and plyconeogenesis, as well as intra-muscular lipids and plasma tree fatty acids derived from adipose tissue provide the main energy alternatives fo muscle glycogen. Several amino acids, including the fesenlial amino acid leucine, arc also used directly as oxidizable fuels uring exercise, Depending on the duration and intensity of exercise ‘and other factors such as plycogen stores and energy intake, amino acids can provide from 2 few to approximately 10% of the total ‘energy for Sustained exercise. Additionally, many amino acids can be ‘converted to alutamate (via glutamate dehydrogenase) and then 10 ‘alanine (via glutamatepyruvate transaminase). Alanine, along with lactate and pyruvate, are recognized as the major gluconeogenic precursors, Via this mechanism, several amino acids play crucial roles in providing the carbon sources for maintaining blood glucose ho: ineostasis during exerese and glycogen restitution during recovery. ‘And finally, during exercise and recovery, amino acids likely play {important anaplerotic functions sustaining the whole metabolic ap- paratus EXERTION, FUEL, LACTATE SHUTTLE, GLUCOSE PARADOX, GLUCONEOGENESIS, ANAPLEROSIS, GLUCOSE HOMEOSTASIS, GLYCOGEN-SPARING, GLYCOGEN. SYNTHESIS In the last century, von Leibig (28) advanced the idea that amino acids derived from body proteins provided the major fuel source for exercise. In more recent times, the overwhelming evidence that carbohydrate and lipid Gerivatives constitute the dominant energy sources for ‘Smal or pobltonFebuan, 90, ‘Aezepeor palcslon Ape, 187 S150 metabolism during exercise led to the belief that amino acids and proteins are not used to sustain the energy requirements of exercise (3). In support of that hypothesis, exercise does not appear to result in a large increase in urinary nitrogen excretion (3). However, if careful determina- tion is made of urinary as well as sweat urea losses (22), the calculated equivalent contribution of amino acids tothe fuel supply becomes significant, In fact, estimates of 5 to 10% of the fuel for prolonged exercise being, supplied from amino acids have been provided by investigators using several independent experimental strategies (22, 25, 29). In addition, the pathways of nitrogenous exchanges between protein and amino acid pools can change dramatically without affecting meas- urements of urea excretion across tissues or at the whole-body level (see paper by Dr. Wolfe in this sym- posium issue). The following quotation is attributed to von Leibig (28) over 140 yr ago: ‘There can be no greater contradiction, with regard to the nutritive process, than to suppose that the nitrogen ofthe food can pass into Thea ue, without having previously become pr of an organized issue The contemporary view is like that of 140 yr ago; proteins and their amino acid derivatives are used along with other substrates to support the metabolic appara- tus which sustain prolonged muscular exercise (8). TECHNICAL CONSIDERATIONS For several reasons, it is difficult to estimate in a quantitative sense the roles played by amino acids and AMINO ACID METABOLISM proteins in sustaining exercise. As part of the continu- ous turnover process, the synthesis and degradation of proteins occur simultaneously. Further, free amino acids are released from proteins of different sizes and with different turnover rates. Moreover, particular amino acids such as leucine can be re-directed from synthetic to catabolic pathways, such as oxidation, In this way, the turnover rate ofan amino acid can remain constant, but its role as. fuel changes dramatically (21, 29), Therefore, from the study of one particular amino acid or protein, it is difficult to obtain a global picture of the effect of some stressor, such as exercise, on overall metabolic processes. For these reasons of complexity, several approaches have been developed to supplement the use of urinary urea collection in the study of amino acid and protein ‘metabolism in vivo. These include use of isotopes of carbon ("?C and '*C), hydrogen (7H and 7H), and nitro- gen ("#N) to study the effects of exercise on the metab- olism of different amino acids individually or simulta- neously, or to study the rates and pathways of carbon and nitrogen portions of the same amino acid. The results of such experiments will be elaborated on in the paper by Dr. Wolfe (in this symposium issue). Isotope labeling and techniques have also been ex- tremely valuable in allowing assessment of the effects of exercise on muscle protein synthesis and degrada- ion. In particular, the use of 3-methylhistidine as a marker for protein catabolism has become very useful, The paper by Dr. Dohm (in this symposium issue) summarizes some of the work in this field, 'No discussion of the effects of exercise on protein metabolism can be complete without recognition of the stimulatory effect of exercise on protein synthesis, Be- yond muscle hypertrophy, exercise and dietary carbo- hydrate have the effect of sparing use of dietary amino acids and body proteins (10). Over the long term, the transient increases in amino acid catabolism brought about by exercise may be compensated for by a more efficient utilization of dietary protein. In the final paper in this series, Dr. Butterfield summarizes the effects of exercise on dietary protein requirements, AMINO ACIDS AS FUELS In reviewing this and other papers in the symposium, the reader will note a great emphasis on the metabolism ‘ofleucine; there are several reasons for such an empha- sis. Leucine is an essential amino acid, Thus, in the ost-absorptive state, the supply of blood leucine must come from the catabolism of body proteins. Leucine is, unique as a purely ketogenic amino acid; so relative to other amino acids, the oxidation of leucine is relatively easy to study. And finally, leucine has been recognized as playing a potentially important role in the regulation of tissue protein metabolism (2, 11, 14). SISI The first demonstration of increased amino acid ox- idation with exercise ina mammal én vivo was by White and Brooks (29) who compared the oxidations of in- jected ['*CJleucine (Figure 1B) and ['*Clalanine with that of ['*C]glucose (Figure 1A). These studies followed several reports by Manchester (23), Buse et al. (9), and Odessey and Goldbery (24), who observed high rates of leucine de-carboxylation by rat muscle preparations int vitro. White and Brooks used pulse injection of “C tracers to study substrate oxidation in rats during rest and two levels of exercise. As with glucose, the relative oxidation of leucine was related to the metabolic rate with more label excreted as CO, during easy exercise than rest, and more label excreted during hard exercise than during easy exercise, Moreover, it was observed (29) that the excretion of "CO; occurred more rapidly and to a greater extent following [!*C]alanine injection than following labeled glucose injection, suggesting that alanine can be disposed of in ways independent of the glucose-alanine cycle. Increased leucine oxidation with exercise has also been demonstrated in humans (20, 31, 32). Figure 2 (from Hagg et al.) portrays results obtained on six women given a continuous infusion of [“CJleucine during rest and sub-maximal exercise, This figure re- veals much of what has been confirmed by other groups. During exercise, blood leucine specific activity (bottom panel) is the same as during rest. Therefore, blood leucine turnover rate is litle affected by exercise. Dur- ing exercise, specific activity of CO, in expired air (top panel) is about half of that during rest, but this result is interpretable only if it is known that the overall rate of COp production (VCO2) during exercise is 4 10 5 times greater than at rest. In the studies of Hage et al., the rate of leucine oxidation was calculated as the specific activity of CO; times the VCO>. In the subjects studied, during exercise this rate doubled, increasing from 474,000 to 967,000 dpm-h™'. However, despite the increase in leucine oxidation rate which occurred dur- ing exercise, the decrease in CO; specific activity indi- cates a relatively greater acceleration in the combustion of other substrates. And finally, with an unchanged leucine turnover rate, but an increased leucine oxida- tion rate, the rate of leucine incorporation into tissue proteins must decrease. The principles of interpreting kinetic tracer data are described more in the paper by Dr. Wolle. SPARING OF AMINO ACIDS BY CARBOHYDRATE AND QUANTITATIVE SIGNIFICANCE OF AMINO ACID METABOLISM AS suggested at the outset, amino acids act as alter- natives to the combustion of carbohydrate derivatives, so it is not surprising that increased availability of glucose reduces the oxidation of amino acids such as $152 COMULATWE % RECOVERY ‘Tm (aia) Figure 1—Excretion of label as "COs followis 1600 1200 800 épm/mmo! 400 wer c=: xeneise LEUCINE 30 60 90 120 150 TIME (MIND Figure 2—Specific activities of expired COs (top) and blood leucine (bottom) during contiauous infusion of {1-"Clcucine into women during rest (solid lines) and exercise (broken lines). [Data from agg, etal. (20), 180 MEDICINE AND SCIENCE IN SPORTS AND EXERCISE injetion of (A) [U-"C}glucose and (B) [U-MCHleucine. (Adapted from White and Brooks (27)1 leucine. Figure 3 is from the report of Davies et al. (12), who used continuous infusion of ['%C}leucine to study oxidation of that amino acid during rest and sub- maximal exercise. Leucine oxidation rate during exer- cise is approximately 3 times that during rest, but glucose ingestion rapidly reduces the oxidation of leucine. Although there are difficulties inherent in quantitat- ing urea losses in sweat during exercise, few reports exist on the subject. Lemon and Mullin (22) have demonstrated that, during prolonged exercise by hu- man subjects, amino acids provide approximately 5 to 10% of the fuet utilized. Moreover, in glycogen-depleted subjects, amino acid and protein utilization increase and account for a greater percentage of the total com- busted fuel (Figure 4). These data are consistent with those of Davies et al. (12) and support the concept of amino acid-sparing by carbohydrates. EFFECTS OF TRAINING ON AMINO ACID OXIDATION At present no reports exist on the effect of training on amino acid oxidation in humans during exercise; AMINO ACID METABOLISM 7 oH i ' rap ei

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