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NUTRITION IN SPORT - Index of

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tion. This decline occurs principally via deamination<br />

<strong>of</strong> AMP to inosine monophosphate (IMP)<br />

but also by the dephosphorylation <strong>of</strong> AMP to<br />

adenosine. The loss <strong>of</strong> AMP may initially appear<br />

counterproductive because <strong>of</strong> the reduction in<br />

the total adenylate pool. However, it should be<br />

noted that the deamination <strong>of</strong> AMP to IMP only<br />

occurs under low ATP/ADP ratio conditions<br />

and, by preventing excessive accumulation <strong>of</strong><br />

ADP and AMP, enables the adenylate kinase<br />

reactions to continue, resulting in an increase in<br />

the ATP/ADP ratio and continuing muscle force<br />

generation. Furthermore, it has been proposed<br />

that the free energy <strong>of</strong> ATP hydrolysis will<br />

decrease when ADP and P i accumulate, which<br />

could further impair muscle force generation.<br />

For these reasons, adenine nucleotide loss has<br />

been suggested to be <strong>of</strong> importance to muscle<br />

function during conditions <strong>of</strong> metabolic crisis;<br />

for example, during maximal exercise and in the<br />

later stages <strong>of</strong> prolonged submaximal exercise<br />

when glycogen stores become depleted (Sahlin &<br />

Broberg 1990).<br />

Glycolysis<br />

Under normal conditions, muscle clearly does<br />

not fatigue after only a few seconds <strong>of</strong> effort, so a<br />

source <strong>of</strong> energy other than ATP and PCr must be<br />

available. This is derived from glycolysis, which<br />

is the name given to the pathway involving the<br />

breakdown <strong>of</strong> glucose (or glycogen), the end<br />

product <strong>of</strong> this series <strong>of</strong> chemical reactions being<br />

biochemistry <strong>of</strong> exercise 23<br />

Table 2.2b Maximal rates <strong>of</strong> adenosine triphosphate (ATP) resynthesis from anaerobic and aerobic metabolism and<br />

approximate delay time before maximal rates are attained following onset <strong>of</strong> exercise.<br />

PCr, phosphocreatine.<br />

Max rate <strong>of</strong> ATP resynthesis<br />

(mmol ATP · kg -1 dm · s -1) Delay time<br />

Fat oxidation 1.0 > 2h<br />

Glucose (from blood) 1.0 Approx. 90min<br />

oxidation<br />

Glycogen oxidation 2.8 Several minutes<br />

Glycolysis 4.5 5–10s<br />

PCr breakdown 9.0 Instantaneous<br />

pyruvate. This process does not require oxygen,<br />

but does result in energy in the form <strong>of</strong> ATP being<br />

available to the muscle from reactions involving<br />

substrate-level phosphorylation. In order for the<br />

reactions to proceed, however, the pyruvate<br />

must be removed; in low-intensity exercise,<br />

when adequate oxygen is available to the muscle,<br />

pyruvate is converted to carbon dioxide and<br />

water by oxidative metabolism in the mitochondria.<br />

In some situations the majority <strong>of</strong> the pyruvate<br />

is removed by conversion to lactate, a<br />

reaction that does not involve oxygen.<br />

A specific transporter protein (GLUT-4) is<br />

involved in the passage <strong>of</strong> glucose molecules<br />

across the cell membrane. Once the glucose molecule<br />

is inside the cell, the first step <strong>of</strong> glycolysis<br />

is an irreversible phosphorylation catalysed by<br />

hexokinase to prevent loss <strong>of</strong> this valuable nutrient<br />

from the cell: glucose is converted to G6P.<br />

This step is effectively irreversible, at least as far<br />

as muscle is concerned. Liver has a phosphatase<br />

enzyme which catalyses the reverse reaction,<br />

allowing free glucose to leave the cell and enter<br />

the circulation, but this enzyme is absent from<br />

muscle. The hexokinase reaction is an energyconsuming<br />

reaction, requiring the investment <strong>of</strong><br />

one molecule <strong>of</strong> ATP per molecule <strong>of</strong> glucose.<br />

This also ensures a concentration gradient for<br />

glucose across the cell membrane down which<br />

transport can occur. Hexokinase is inhibited by<br />

an accumulation <strong>of</strong> the reaction product G6P, and<br />

during high-intensity exercise, the increasing<br />

concentration <strong>of</strong> G6P limits the contribution that

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