Scott K. Powers Edward T.

Howley

Theory and Application to Fitness and Performance

SEVENTH EDITION

Chapter

Bioenergetics

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Chapter 3

Objectives
1. Discuss the functions of the cell membrane, nucleus, and mitochondria. 2. Define the following terms: (1) endergonic reactions, (2) exergonic reactions, (3) coupled reactions, and (4) bioenergetics. 3. Describe the role of enzymes as catalysts in cellular chemical reactions. 4. List and discuss the nutrients that are used as fuels during exercise. 5. Identify the high-energy phosphates.
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Chapter 3

Objectives
6. Discuss the biochemical pathways involved in anaerobic ATP production. 7. Discuss the aerobic production of ATP. 8. Describe the general scheme used to regulate metabolic pathways involved in bioenergetics. 9. Discuss the interaction between aerobic and anaerobic ATP production during exercise. 10. Identify the enzymes that are considered rate limiting in glycolysis and the Krebs cycle.

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Chapter 3

Outline
Cell Structure Biological Energy Transformation
Cellular Chemical Reactions Oxidation-Reduction Reactions Enzymes

High-Energy Phosphates Bioenergetics

Anaerobic ATP Production Aerobic ATP production

Control of Bioenergetics
Control of ATP-PC System Control of Glycolysis Control of Krebs Cycle and Electron Transport Chain

Fuels for Exercise

Carbohydrates Fats Proteins

Aerobic ATP Tally Efficiency of Oxidative Phosphorylation Interaction Between Aerobic/Anaerobic ATP Production

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Chapter 3

Introduction
Metabolism Sum of all chemical reactions that occur in the body Anabolic reactions
Synthesis of molecules

Catabolic reactions
Breakdown of molecules

Bioenergetics Converting foodstuffs (fats, proteins, carbohydrates) into energy

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Chapter 3

Cell Structure

Cell Structure
Cell membrane Semipermeable membrane that separates the cell from the extracellular environment Nucleus Contains genes that regulate protein synthesis
Molecular biology

Cytoplasm Fluid portion of cell Contains organelles

Mitochondria

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Chapter 3

Cell Structure

A Typical Cell and Its Major Organelles

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Figure 3.1

Chapter 3

Cell Structure

In Summary
Metabolism is defined as the total of all cellular reactions that occur in the body; this includes both the synthesis of molecules and the breakdown of molecules. Cell structure includes the following three major parts: (1) cell membrane, (2) nucleus, and (3) cytoplasm (called sarcoplasm in muscle). The cell membrane provides a protective barrier between the interior of the cell and the extracellular fluid. Genes (located within the nucleus) regulate protein synthesis within the cell. The cytoplasm is the fluid portion of the cell and contains numerous organelles
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Chapter 3

Cell Structure

A Closer Look 3.1

Molecular Biology and Exercise Science

Study of molecular structures and events underlying biological processes Relationship between genes and cellular characteristics they control Genes code for specific cellular proteins Process of protein synthesis Exercise training results in modifications in protein synthesis Strength training results in increased synthesis of muscle contractile protein Molecular biology provides tools for understanding the cellular response to exercise
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Chapter 3

Biological Energy Transformation

Steps Leading to Protein Synthesis

1. DNA contains information to produce proteins. 2. Transcription produces mRNA. 3. mRNA leaves nucleus and binds to ribosome. 4. Amino acids are carried to the ribosome by tRNA. 5. In translation, mRNA is used to determine the arrangement of amino acids in the polypeptide chain.

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Figure 3.2

Chapter 3

Biological Energy Transformation

Cellular Chemical Reactions

Endergonic reactions Require energy to be added Endothermic Exergonic reactions Release energy Exothermic Coupled reactions Liberation of energy in an exergonic reaction drives an endergonic reaction

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Chapter 3

Biological Energy Transformation

The Breakdown of Glucose: An Exergonic Reaction

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Figure 3.3

Chapter 3

Biological Energy Transformation

Coupled Reactions
The energy given off by the exergonic reaction powers the endergonic reaction

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Figure 3.4

Chapter 3

Biological Energy Transformation

Oxidation-Reduction Reactions
Oxidation Removing an electron Reduction Addition of an electron Oxidation and reduction are always coupled reactions Often involves the transfer of hydrogen atoms rather than free electrons Hydrogen atom contains one electron A molecule that loses a hydrogen also loses an electron and therefore is oxidized Importance of NAD and FAD
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Chapter 3

Biological Energy Transformation

Oxidation-Reduction Reaction Involving NAD and NADH

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Figure 3.5

Chapter 3

Biological Energy Transformation

Enzymes
Catalysts that regulate the speed of reactions Lower the energy of activation Factors that regulate enzyme activity Temperature pH Interact with specific substrates Lock and key model

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Chapter 3

Biological Energy Transformation

Enzymes Catalyze Reactions

Enzymes lower the energy of activation

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Figure 3.6

Chapter 3

Biological Energy Transformation

The Lock-and-Key Model of Enzyme Action

a) Substrate (sucrose) approaches the active site on the enzyme. Substrate fits into the active site, forming enzymesubstrate complex. The enzyme releases the products (glucose and fructose).

b)

c)

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Figure 3.7

Chapter 3

Biological Energy Transformation

Clinical Applications 3.1

Diagnostic Value of Measuring Enzyme Activity in the Blood

Damaged cells release enzymes into the blood Enzyme levels in blood indicate disease or tissue damage Diagnostic application Elevated lactate dehydogenase or creatine kinase in the blood may indicate a myocardial infarction

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Chapter 3

Biological Energy Transformation

Classification of Enzymes
Oxidoreductases Catalyze oxidation-reduction reactions Transferases Transfer elements of one molecule to another Hydrolases Cleave bonds by adding water Lyases Groups of elements are removed to form a double bond or added to a double bond Isomerases Rearrangement of the structure of molecules Ligases Catalyze bond formation between substrate molecules
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Chapter 3

Biological Energy Transformation

Example of the Major Classes of Enzymes

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Chapter 3

Biological Energy Transformation

Factors That Alter Enzyme Activity

Temperature Small rise in body temperature increases enzyme activity Exercise results in increased body temperature pH Changes in pH reduces enzyme activity Lactic acid produced during exercise

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Chapter 3

Fuels for Exercise

Carbohydrates
Glucose Blood sugar Glycogen Storage form of glucose in liver and muscle
Synthesized by enzyme glycogen synthase

Glycogenolysis
Breakdown of glycogen to glucose

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Chapter 3

Fuels for Exercise

Fats
Fatty acids Primary type of fat used by the muscle Triglycerides
Storage form of fat in muscle and adipose tissue Breaks down into glycerol and fatty acids

Phospholipids Not used as an energy source Steroids Derived from cholesterol Needed to synthesize sex hormones
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Chapter 3

Fuels for Exercise

Protein
Composed of amino acids Some can be converted to glucose in the liver Gluconeogenesis Others can be converted to metabolic intermediates Contribute as a fuel in muscle Overall, protein is not a primary energy source during exercise

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Chapter 3

Fuels for Exercise

In Summary
The body uses carbohydrate, fat, and protein nutrients consumed daily to provide the necessary energy to maintain cellular activities both at rest and during exercise. During exercise, the primary nutrients used for energy are fats and carbohydrates, with protein contributing a relatively small amount of the total energy used. Glucose is stored in animal cells as a polysaccharide called glycogen. Fatty acids are the primary form of fat used as an energy source in cells. Fatty acids are stored as triglycerides in muscle and fat cells.
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Chapter 3

High-Energy Phosphates

High-Energy Phosphates
Adenosine triphosphate (ATP) Consists of adenine, ribose, and three linked phosphates Synthesis

ADP + Pi ATP
Breakdown

ATP

ATPase

ADP + Pi + Energy

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Chapter 3

High-Energy Phosphates

Structure of ATP

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Figure 3.10

Chapter 3

High-Energy Phosphates

Model of ATP as the Universal Energy Donor

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Figure 3.11

Chapter 3

Bioenergetics

Bioenergetics
Formation of ATP Phosphocreatine (PC) breakdown Degradation of glucose and glycogen
Glycolysis

Oxidative formation of ATP Anaerobic pathways Do not involve O2 PC breakdown and glycolysis Aerobic pathways Require O2 Oxidative phosphorylation
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Chapter 3

Bioenergetics

Anaerobic ATP Production

ATP-PC system Immediate source of ATP

PC + ADP

Creatine kinase

ATP + C

Glycolysis Glucose 2 pyruvic acid or 2 lactic acid Energy investment phase

Requires 2 ATP

Energy generation phase

Produces 4 ATP, 2 NADH, and 2 pyruvate or 2 lactate

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Chapter 3

Bioenergetics

The Winning Edge 3.1

Does Creatine Supplementation Improve Exercise Performance?

Depletion of PC may limit short-term, high-intensity exercise Creatine monohydrate supplementation Increased muscle PC stores Some studies show improved performance in shortterm, high-intensity exercise
Inconsistent results may be due to water retention and weight gain

Increased strength and fat-free mass with resistance training Creatine supplementation does not appear to pose health risks
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Chapter 3

Bioenergetics

The Two Phases of Glycolysis

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Figure 3.13

Chapter 3

Bioenergetics

Glycolysis: Energy Investment Phase

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Figure 3.15

Figure 3.15

Chapter 3

Bioenergetics

Hydrogen and Electron Carrier Molecules

Transport hydrogens and associated electrons To mitochondria for ATP generation (aerobic) To convert pyruvic acid to lactic acid (anaerobic) Nicotinamide adenine dinucleotide (NAD)

NAD + 2H+ NADH + H+

Flavin adenine dinucleotide (FAD)

FAD + 2H+ FADH2

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Chapter 3

Bioenergetics

A Closer Look 3.3

NADH is Shuttled into Mitochondria

NADH produced in glycolysis must be converted back to NAD By converting pyruvic acid to lactic acid By shuttling H+ into the mitochondria A specific transport system shuttles H+ across the mitochondrial membrane Located in the mitochondrial membrane

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Chapter 3

Bioenergetics

In Summary
The immediate source of energy for muscular contraction is the high-energy phosphate ATP. ATP is degraded via the enzyme ATPase as follows:

ATP

ATPase

ADP + Pi + Energy

Formation of ATP without the use of O2 is termed anaerobic metabolism. In contrast, the production of ATP using O2 as the final electron acceptor is referred to as aerobic metabolism.

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Chapter 3

Bioenergetics

In Summary
Exercising skeletal muscles produce lactic acid. However, once produced in the body, lactic acid is rapidly converted to its conjugate base, lactate. Muscle cells can produce ATP by any one or a combination of three metabolic pathways: (1) ATP-PC system, (2) glycolysis, (3) oxidative ATP production. The ATP-PC system and glycolysis are two anaerobic metabolic pathways that are capable of producing ATP without O2.

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Chapter 3

Bioenergetics

Aerobic ATP Production

Krebs cycle (citric acid cycle) Pyruvic acid (3 C) is converted to acetyl-CoA (2 C)
CO2 is given off

Acetyl-CoA combines with oxaloacetate (4 C) to form citrate (6 C) Citrate is metabolized to oxaloacetate

Two CO2 molecules given off

Produces three molecules of NADH and one FADH Also forms one molecule of GTP
Produces one ATP

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Chapter 3

Bioenergetics

The Three Stages of Oxidative Phosphorylation

Figure 3.17

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Chapter 3

Bioenergetics

The Krebs Cycle

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Figure 3.18

Chapter 3

Bioenergetics

Fats and Proteins in Aerobic Metabolism

Fats Triglycerides glycerol and fatty acids Fatty acids acetyl-CoA
Beta-oxidation

Glycerol is not an important muscle fuel during exercise Protein Broken down into amino acids Converted to glucose, pyruvic acid, acetyl-CoA, and Krebs cycle intermediates

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Chapter 3

Bioenergetics

Aerobic ATP Production

Electron transport chain Oxidative phosphorylation occurs in the mitochondria Electrons removed from NADH and FADH are passed along a series of carriers (cytochromes) to produce ATP
Each NADH produces 2.5 ATP Each FADH produces 1.5 ATP

Called the chemiosmotic hypothesis H+ from NADH and FADH are accepted by O2 to form water

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Chapter 3

Bioenergetics

The Chemiosmotic Hypothesis of ATP Formation

Electron transport chain results in pumping of H+ ions across inner mitochondrial membrane Results in H+ gradient across membrane Energy released to form ATP as H+ ions diffuse back across the membrane

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Chapter 3

Bioenergetics

A Closer Look 3.4

Beta Oxidation is the Process of Converting Fatty Acids to Acetyl-CoA

Breakdown of triglycerides releases fatty acids Fatty acids must be converted to acetyl-CoA to be used as a fuel Activated fatty acid (fatty acyl-CoA) into mitochondrion Fatty acid chopped into 2 carbon fragments forming acetyl-CoA Acetyl-CoA enters Krebs cycle and is used for energy

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Chapter 3

Bioenergetics

Beta Oxidation

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Figure 3.21

Chapter 3

Bioenergetics

In Summary
Oxidative phosphorylation or aerobic ATP production occurs in the mitochondria as a result of a complex interaction between the Krebs cycle and the electron transport chain. The primary role of the Krebs cycle is to complete the oxidation of substrates and form NADH and FADH to enter the electron transport chain. The end result of the electron transport chain is the formation of ATP and water. Water is formed by oxygen-accepting electrons; hence, the reason we breathe oxygen is to use it as the final acceptor of electrons in aerobic metabolism.

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Chapter 3

Aerobic ATP Tally

A Closer Look 3.5

A New Look at the ATP Balance Sheet

Historically, 1 glucose produced 38 ATP Recent research indicates that 1 glucose produces 32 ATP Energy provided by NADH and FADH also used to transport ATP out of mitochondria. 3 H+ must pass through H+ channels to produce 1 ATP Another H+ needed to move the ATP across the mitochondrial membrane

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Chapter 3

Efficiency of Oxidative Phosphorylation

Efficiency of Oxidative Phosphorylation

One mole of ATP has energy yield of 7.3 kcal 32 moles of ATP are formed from one mole of glucose Potential energy released from one mole of glucose is 686 kcal/mole
32 moles ATP/mole glucose x 7.3 kcal/mole ATP 686 kcal/mole glucose x 100 = 34%

Overall efficiency of aerobic respiration is 34% 66% of energy released as heat

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Chapter 3

Efficiency of Oxidative Phosphorylation

In Summary
The aerobic metabolism of one molecule of glucose results in the production of 32 ATP molecules, whereas the aerobic yield for glycogen breakdown is 33 ATP. The overall efficiency of aerobic of aerobic respiration is approximately 34%, with the remaining 66% of energy being released as heat.

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Chapter 3

Control of Bioenergetics

Control of Bioenergetics
Rate-limiting enzymes An enzyme that regulates the rate of a metabolic pathway Modulators of rate-limiting enzymes Levels of ATP and ADP+Pi
High levels of ATP inhibit ATP production Low levels of ATP and high levels of ADP+Pi stimulate ATP production

Calcium may stimulate aerobic ATP production

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Chapter 3

Control of Bioenergetics

Example of a Rate-Limiting Enzyme

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Figure 3.22

Chapter 3

Control of Bioenergetics

In Summary
Metabolism is regulated by enzymatic activity. An enzyme that regulates a metabolic pathway is termed a rate-limiting enzyme. The rate-limiting enzyme for glycolysis is phosphofructokinase, while the rate-limiting enzymes for the Krebs cycle and electron transport chain are isocitrate dehydrogenase and cytochrome oxidase, respectively. In general, cellular levels of ATP and ADP+Pi regulate the rate of metabolic pathways involved in the production of ATP. High levels of ATP inhibit further ATP production, while low levels of ATP and high levels of ADP+Pi stimulate ATP production. Evidence also exists that calcium may stimulate aerobic energy metabolism.
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Chapter 3

Interaction Between Aerobic/Anaerobic ATP Production

Interaction Between Aerobic/Anaerobic ATP Production

Energy to perform exercise comes from an interaction between aerobic and anaerobic pathways Effect of duration and intensity Short-term, high-intensity activities
Greater contribution of anaerobic energy systems

Long-term, low to moderate-intensity exercise

Majority of ATP produced from aerobic sources

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Chapter 3

Interaction Between Aerobic/Anaerobic ATP Production

In Summary
Energy to perform exercise comes from an interaction of anaerobic and aerobic pathways. In general, the shorter the activity (high intensity), the greater the contribution of anaerobic energy production. In contrast, long-term activities (low to moderate intensity) utilize ATP produced from aerobic sources.

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Chapter 3

Study Questions
1. List and briefly discuss the functions of the three major components of cell structure. 2. Briefly explain the concept of coupled reactions. 3. Define the following terms: (1) bioenergetics, (2) endergonic reactions, and (3) exergonic reactions. 4. Discuss the role of enzymes as catalysts. What is meant by the expression energy of activation? 5. Where do glycolysis, the Krebs cycle, and oxidative phosphorylation take place in the cell? 6. Define the terms glycogen, glycogenolysis, and glycolysis.
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Chapter 3

Study Questions
7. What are the high-energy phosphates? Explain the statement that ATP is the universal energy donor. 8. Define the terms aerobic and anaerobic. 9. Briefly discuss the function of glycolysis in bioenergetics. What role does NAD play in glycolysis? 10. Discuss the operation of the Krebs cycle and the electron transport chain in the aerobic production of ATP. What is the function of NAD and FAD in these pathways?
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Chapter 3

Study Questions
11. What is the efficiency of the aerobic degradation of glucose? 12. What is the role of oxygen in aerobic metabolism? 13. What are the rate-limiting enzymes for the following metabolic pathways: ATP-PC system, glycolysis, Krebs cycle, and electron transport chain? 14. Briefly discuss the interaction of anaerobic versus aerobic ATP production during exercise. 15. Discuss the chemiosmotic theory of ATP production.
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Chapter 3

Study Questions
16. List and define the six classes of enzymes identified by the International Union of Biochemistry. 17. Briefly discuss the impact of changes in both temperature and pH on enzyme function. 18. Discuss the relationship between lactic acid and lactate.

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