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Scott K. Powers Edward T.

Howley

Theory and Application to Fitness and Performance


SEVENTH EDITION

Chapter

Exercise Metabolism

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Chapter 4

Objectives
1. Discuss the relationship between exercise intensity/duration and the bioenergetic pathways that are most responsible for the production of ATP during various types of exercise. 2. Define the term oxygen deficit. 3. Define the term lactate threshold. 4. Discuss several possible mechanisms for the sudden rise in blood-lactate concentration during incremental exercise.

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Chapter 4

Objectives
5. List the factors that regulate fuel selection during different types of exercise. 6. Explain why fat metabolism is dependent on carbohydrate metabolism. 7. Define the term oxygen debt. 8. Give the physiological explanation for the observation that the O2 debt is greater following intense exercise when compared to the O2 debt following light exercise.

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Chapter 4

Outline
Energy Requirements at Rest Rest-to-Exercise Transitions Recovery from Exercise: Metabolic Responses Metabolic Responses to Exercise: Influence of Duration and Intensity
Short-Term, Intense Exercise Prolonged Exercise Incremental Exercise

Factors Governing Fuel Selection


Exercise Intensity and Fuel Selection Exercise Duration and Fuel Selection Interaction of Fat/ Carbohydrate Metabolism Body Fuel Sources

Estimation of Fuel Utilization During Exercise

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Chapter 4

Energy Requirements at Rest

Energy Requirements at Rest


Almost 100% of ATP produced by aerobic metabolism Blood lactate levels are low (<1.0 mmol/L) Resting O2 consumption: 0.25 L/min 3.5 ml/kg/min

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Chapter

Rest-to-Exercise Transitions

Rest-to-Exercise Transitions
ATP production increases immediately Oxygen uptake increases rapidly Reaches steady state within 14 minutes After steady state is reached, ATP requirement is met through aerobic ATP production Initial ATP production through anaerobic pathways ATP-PC system Glycolysis Oxygen deficit Lag in oxygen uptake at the beginning of exercise
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Chapter 4

Rest-to-Exercise Transitions

The Oxygen Deficit

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Figure 4.1

Chapter 4

Rest-to-Exercise Transitions

Comparison of Trained and Untrained Subjects


Trained subjects have a lower oxygen deficit Better-developed aerobic bioenergetic capacity Due to cardiovascular or muscular adaptations Results in less production of lactic acid

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Chapter 4

Rest-to-Exercise Transitions

Differences in VO2 Between Trained and Untrained Subjects

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Figure 4.2

Chapter 4

Rest-to-Exercise Transitions

In Summary
In the transition from rest to light or moderate exercise, oxygen uptake increases rapidly, generally reaching a steady state within one to four minutes. The term oxygen deficit applies to the lag in oxygen uptake in the beginning of exercise. The failure of oxygen uptake to increase instantly at the beginning of exercise suggests that anaerobic pathways contribute to the overall production on ATP early in exercise. After a steady state is reached, the bodys ATP requirement is met via aerobic metabolism.

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Chapter 4

Recovery From Exercise: Metabolic Responses

Recovery From Exercise


Oxygen uptake remains elevated above rest into recovery Oxygen debt Term used by A.V. Hill
Repayment for O2 deficit at onset of exercise

Excess post-exercise oxygen consumption (EPOC) Terminology reflects that only ~20% elevated O2 consumption used to repay O2 deficit Many scientists use these terms interchangeably

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Chapter 4

Recovery From Exercise: Metabolic Responses

Oxygen Debt
Rapid portion of O2 debt Resynthesis of stored PC Replenishing muscle and blood O2 stores Slow portion of O2 debt Elevated heart rate and breathing = energy need Elevated body temperature = metabolic rate Elevated epinephrine and norepinephrine = metabolic rate Conversion of lactic acid to glucose (gluconeogenesis)
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Chapter 4

Recovery From Exercise: Metabolic Responses

EPOC is Greater Following Higher Intensity Exercise


Higher body temperature Greater depletion of PC Greater blood concentrations of lactic acid Higher levels of blood epinephrine and norepinephrine

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Chapter 4

Recovery From Exercise: Metabolic Responses

A Closer Look 4.1

Removal of Lactic Acid Following Exercise


Classical theory Majority of lactic acid converted to glucose in liver Recent evidence 70% of lactic acid is oxidized
Used as a substrate by heart and skeletal muscle

20% converted to glucose 10% converted to amino acids Lactic acid is removed more rapidly with light exercise in recovery Optimal intensity is ~3040% VO2 max
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Chapter 4

Recovery From Exercise: Metabolic Responses

Blood Lactate Removal Following Strenuous Exercise

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Figure 4.4

Chapter 4

Metabolic Responses to Exercise: Influence of Duration and Intensity

Metabolic Responses to Short-Term, Intense Exercise


First 15 seconds of exercise ATP through ATP-PC system Intense exercise longer than 5 seconds Shift to ATP production via glycolysis Events lasting longer than 45 seconds ATP production through ATP-PC, glycolysis, and aerobic systems 70% anaerobic/30% aerobic at 60 seconds 50% anaerobic/50% aerobic at 2 minutes

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Chapter 4

Metabolic Responses to Exercise: Influence of Duration and Intensity

In Summary
During high-intensity, short-term exercise (i.e., two to twenty seconds), the muscles ATP production is dominated by the ATP-PC system. Intense exercise lasting more than twenty seconds relies more on anaerobic glycolysis to produce much of the needed ATP. Finally, high-intensity events lasting longer than forty-five seconds use a combination of the ATP-PC system, glycolysis, and the aerobic system to produce the needed ATP for muscular contraction.

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Chapter 4

Metabolic Responses to Exercise: Influence of Duration and Intensity

Metabolic Responses to Prolonged Exercise


Prolonged exercise (>10 minutes) ATP production primarily from aerobic metabolism Steady-state oxygen uptake can generally be maintained during submaximal exercise Prolonged exercise in a hot/humid environment or at high intensity Upward drift in oxygen uptake over time Due to body temperature and rising epinephrine and norepinephrine

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Chapter 4

Metabolic Responses to Exercise: Influence of Duration and Intensity

Upward Drift in Oxygen Uptake During Prolonged Exercise

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Figure 4.6

Chapter 4

Metabolic Responses to Exercise: Influence of Duration and Intensity

Metabolic Responses to Incremental Exercise


Oxygen uptake increases linearly until maximal oxygen uptake (VO2 max) is reached No further increase in VO2 with increasing work rate VO2 max Physiological ceiling for delivery of O2 to muscle Affected by genetics and training Physiological factors influencing VO2 max Maximum ability of cardiorespiratory system to deliver oxygen to the muscle Ability of muscles to use oxygen and produce ATP aerobically
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Chapter 4

Metabolic Responses to Exercise: Influence of Duration and Intensity

Changes in Oxygen Uptake During Incremental Exercise

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Figure 4.7

Chapter 4

Metabolic Responses to Exercise: Influence of Duration and Intensity

Lactate Threshold
The point at which blood lactic acid rises systematically during incremental exercise Appears at ~5060% VO2 max in untrained subjects At higher work rates (6580% VO2 max) in trained subjects Also called: Anaerobic threshold Onset of blood lactate accumulation (OBLA)
Blood lactate levels reach 4 mmol/L

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Chapter 4

Metabolic Responses to Exercise: Influence of Duration and Intensity

Changes in Blood Lactate Concentration During Incremental Exercise

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Figure 4.8

Chapter 4

Metabolic Responses to Exercise: Influence of Duration and Intensity

Explanations for the Lactate Threshold


Low muscle oxygen (hypoxia) Accelerated glycolysis NADH produced faster than it is shuttled into mitochondria Excess NADH in cytoplasm converts pyruvic acid to lactic acid Recruitment of fast-twitch muscle fibers LDH isozyme in fast fibers promotes lactic acid formation Reduced rate of lactate removal from the blood
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Chapter 4

Metabolic Responses to Exercise: Influence of Duration and Intensity

Effect of Hydrogen Shuttle on Lactic Acid Formation

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Figure 4.9

Chapter 4

Metabolic Responses to Exercise: Influence of Duration and Intensity

Practical Uses of the Lactate Threshold


Prediction of performance Combined with VO2 max Planning training programs Marker of training intensity

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Chapter 4

Metabolic Responses to Exercise: Influence of Duration and Intensity

In Summary
Oxygen uptake increases in a linear fashion during incremental exercise until VO2 max is reached. The point at which blood lactic acid rises systematically during graded exercise is termed the lactate threshold or anaerobic threshold. Controversy exists over the mechanism to explain the sudden rise in blood lactic acid concentrations during incremental exercise. It is possible that any one or a combination of the following factors might provide an explanation for the lactate threshold: (1) low muscle oxygen, (2) accelerated glycolysis, (3) recruitment of fast fibers, and (4) a reduced rate of lactate removal. The lactate threshold has practical uses such as in performance prediction and as a marker of training intensity.
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Chapter 4

Estimation of Fuel Utilization During Exercise

Estimation of Fuel Utilization During Exercise


Respiratory exchange ratio (RER or R)
R= VCO2 VO
2

R for fat (palmitic acid)


C16H32O2 + 23 O2 16 CO2 + 16 H2O R= VCO2 VO
2

16 CO2 23 O2

= 0.70

R for carbohydrate (glucose)


C6H12O6 + 6 O2 6 CO2 + 6 H2O R= VCO2 VO
2

6 CO2 6 O2

= 1.00

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Chapter 4

Estimation of Fuel Utilization During Exercise

In Summary
The respiratory exchange ratio (R) is the ratio of carbon dioxide produced to the oxygen consumed (VCO2/VO2). In order for R to be used as an estimate of substrate utilization during exercise, the subject must have reached steady state. This is important because only during steady-state exercise are the VCO2 and VO2 reflective of metabolic exchange of gases in tissues.

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Chapter 4

Factors Governing Fuel Selection

Exercise Intensity and Fuel Selection


Low-intensity exercise (<30% VO2 max) Fats are primary fuel High-intensity exercise (>70% VO2 max) Carbohydrates are primary fuel Crossover concept Describes the shift from fat to CHO metabolism as exercise intensity increases Due to:
Recruitment of fast muscle fibers Increasing blood levels of epinephrine

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Chapter 4

Factors Governing Fuel Selection

Illustration of the Crossover Concept

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Figure 4.11

Chapter 4

Factors Governing Fuel Selection

A Closer Look 4.2

The Regulation of Glycogen Breakdown During Exercise


Dependent on the enzyme phosphorylase Activation of phosphorylase Calmodulin activated by calcium released from sarcoplasmic reticulum
Active calmodulin activates phosphorylase

Epinephrine binding to receptor results in formation of cyclic AMP


Cyclic AMP activates phosphorylase

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Chapter 4

Factors Governing Fuel Selection

Clinical Applications 4.1

McArdles Syndrome: A Genetic Error in Muscle Glycogen Metabolism


Cannot synthesize the enzyme phosphorylase Due to a gene mutation Inability to break down muscle glycogen Also prevents lactate production Blood lactate levels do not rise during high-intensity exercise Patients complain of exercise intolerance and muscle pain

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Chapter 4

Factors Governing Fuel Selection

A Closer Look 4.3

Is Low-Intensity Exercise Best for Burning Fat?


At low exercise intensities (~20% VO2 max) High percentage of energy expenditure (~60%) derived from fat However, total energy expended is low
Total fat oxidation is also low

At higher exercise intensities (~50% VO2 max) Lower percentage of energy (~40%) from fat Total energy expended is higher
Total fat oxidation is also higher

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Chapter 4

Factors Governing Fuel Selection

Rate of Fat Metabolism at Different Exercise Intensities

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Figure 4.14

Chapter 4

Factors Governing Fuel Selection

Exercise Duration and Fuel Selection


Prolonged, low-intensity exercise Shift from carbohydrate metabolism toward fat metabolism Due to an increased rate of lipolysis Breakdown of triglycerides glycerol + FFA
By enzymes called lipases

Stimulated by rising blood levels of epinephrine

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Chapter 4

Factors Governing Fuel Selection

Interaction of Fat and CHO Metabolism During Exercise


Fats burn in the flame of carbohydrates Glycogen is depleted during prolonged highintensity exercise Reduced rate of glycolysis and production of pyruvate Reduced Krebs cycle intermediates Reduced fat oxidation
Fats are metabolized by Krebs cycle

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Chapter 4

Factors Governing Fuel Selection

The Winning Edge 4.2

Carbohydrate Feeding via Sports Drinks Improves Endurance Performance


The depletion of muscle and blood carbohydrate stores contributes to fatigue Ingestion of carbohydrates can improve endurance performance During submaximal (<70% VO2 max), long-duration (>90 minutes) exercise 3060 g of carbohydrate per hour are required May also improve performance in shorter, higher intensity events

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Chapter 4

Factors Governing Fuel Selection

Sources of Carbohydrate During Exercise


Muscle glycogen Primary source of carbohydrate during high-intensity exercise Supplies much of the carbohydrate in the first hour of exercise Blood glucose From liver glycogenolysis Primary source of carbohydrate during low-intensity exercise Important during long-duration exercise
As muscle glycogen levels decline

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Chapter 4

Factors Governing Fuel Selection

Sources of Fat During Exercise


Intramuscular triglycerides Primary source of fat during higher intensity exercise Plasma FFA From adipose tissue lipolysis
Triglycerides glycerol + FFA

FFA converted to acetyl-CoA and enters Krebs cycle Primary source of fat during low-intensity exercise Becomes more important as muscle triglyceride levels decline in long-duration exercise

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Chapter 4

Factors Governing Fuel Selection

Influence of Exercise Intensity on Muscle Fuel Source

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Figure 4.15

Chapter 4

Factors Governing Fuel Selection

Effect of Exercise Duration on Muscle Fuel Source

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Figure 4.16

Chapter 4

Factors Governing Fuel Selection

Sources of Protein During Exercise


Proteins broken down into amino acids Muscle can directly metabolize branch chain amino acids and alanine Liver can convert alanine to glucose Only a small contribution (~2%) to total energy production during exercise May increase to 510% late in prolonged-duration exercise

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Chapter 4

Factors Governing Fuel Selection

Lactate as a Fuel Source During Exercise


Can be used as a fuel source by skeletal muscle and the heart Converted to acetyl-CoA and enters Krebs cycle Can be converted to glucose in the liver Cori cycle Lactate shuttle Lactate produced in one tissue and transported to another

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Chapter 4

Factors Governing Fuel Selection

A Closer Look 4.4

The Cori Cycle: Lactate as a Fuel Source


Lactic acid produced by skeletal muscle is transported to the liver Liver converts lactate to glucose Gluconeogenesis Glucose is transported back to muscle and used as a fuel

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Chapter 4

Factors Governing Fuel Selection

The Cori Cycle: Lactate As a Fuel Source

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Figure 4.17

Chapter 4

Factors Governing Fuel Selection

In Summary
The regulation of fuel selection during exercise is under complex control and is dependent upon several factors, including diet and the intensity and duration of exercise. In general, carbohydrates are used as the major fuel source during high-intensity exercise. During prolonged exercise, there is a gradual shift from carbohydrate metabolism toward fat metabolism. Proteins contribute less than 2% of the fuel used during exercise of less than one hours duration. During prolonged exercise (i.e., three to five hours duration), the total contribution of protein to the fuel supply may reach 5% to 10% during the final minutes of prolonged work.
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Chapter 4

Study Questions
1. Identify the predominant energy systems used to produce ATP during the following types of exercise:
a. Short-term, intense exercise (i.e., less than ten seconds duration) b. 400-meter dash c. 20-kilometer race (12.4 miles)

2.

Graph the change in oxygen uptake during the transition from rest to steady-state, submaximal exercise. Label the oxygen deficit. Where does the ATP come from during the transition period from rest to steady state? Graph the change in oxygen uptake and blood lactate concentration during incremental exercise. Label the point on the graph that might be considered the lactate threshold or lactate inflection point. Discuss several possible reasons why blood lactate begins to rise rapidly during incremental exercise.
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3.

4.

Chapter 4

Study Questions
5. Briefly, explain how the respiratory exchange ratio is used to estimate which substrate is being utilized during exercise. What is meant by the term nonprotein R? List two factors that play a role in the regulation of carbohydrate metabolism during exercise. List those variables that regulate fat metabolism during exercise. Define the following terms: (a) triglyceride, (b) lipolysis, and (c) lipases. Graph the change in oxygen uptake during recovery from exercise. Label the oxygen debt.

6. 7. 8. 9.

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Chapter 4

Study Questions
10. How does the modern theory of EPOC differ from the classical oxygen debt theory proposed by A.V. Hill? 11. Discuss the influence of exercise intensity on muscle fuel selection. 12. How does the duration of exercise influence muscle fuel selection?

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