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Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer
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Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

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Page 1: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Fundamentals of Human Energy Transfer

Fundamentals of Human Energy Transfer

Chapter 5Chapter 5

Section 3: Energy Transfer

Section 3: Energy Transfer

Page 2: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Fundamental DefinitionsFundamental Definitions• Bioenergetics studies diverse

means for energy transfer for biologic work within living organisms.

• Aerobic and anaerobic breakdown of food nutrients provides energy for synthesizing the chemical fuel for all biologic work.

Page 3: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Energy: The Capacity for Work

Energy: The Capacity for Work

Page 4: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

First Law of Thermodynamics

First Law of Thermodynamics

• Conservation of energy

• Dictates that the body does not produce, consume, or use up energy;rather, energy transforms from one form into another as physiologic systems undergo continual change.

Page 5: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Definition of EnergyDefinition of Energy• Potential Energy: stored, inactive; ability to

do work even if it is not doing work at the time.

• Kinetic Energy: energy at work, active.

Page 6: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Potential Energy as– Energy of position

(gravitational)– Mechanical potential

energy (elastic deformation)

– Bound energy within internal structure

Releasing potential energy transforms into kinetic energy of motion.

Page 7: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Energy-Releasing and Energy-Conserving

Processes

Energy-Releasing and Energy-Conserving

Processes• Exergonic reactions

– Chemical processes that release energy to its surroundings

– Downhill processes

• Endergonic reactions– Chemical processes that store or

absorb energy – Uphill processes

Page 8: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Coupled ReactionsCoupled Reactions

Page 9: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Second Law of Second Law of ThermodynamicsThermodynamics

• Tendency to degrade potential energy to kinetic energy with lower capacity for work (i.e., increase entropy).

• Food and other chemicals represent excellent sources of potential energy, yet energy decreases as compounds decompose via normal oxidation.

• Example: flashlight battery.

Page 10: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Forms of EnergyForms of Energy

• During energy conversions, a loss of potential energy from one source often produces increase in potential energy of another source.

Page 11: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Biologic Work Biologic Work in Humansin Humans

1. Performance of mechanical work

2. Chemical work in biosynthesis of macromolecules

3. Active transport of molecules and ions concentrating various substances in & out of cells.

Page 12: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

• The limits of exercise intensity ultimately depend on the rate that cells, extract, conserve, and transfer chemical energy in the food nutrients to the contractile filaments of skeletal muscle

Key PointKey Point

Page 13: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Factors Affecting Bioenergetics

Factors Affecting Bioenergetics

• Enzymes – Reaction rates: operation rate of

enzymes– Enzyme mode of action: how an

enzyme reacts with its specific substrate

• Coenzymes

Page 14: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

EnzymesEnzymes• Are highly specific protein catalysts• Accelerate the forward and reverse

reactions• Are neither consumed nor changed in

the reaction• pH and temperature dramatically

affect enzyme activity• Named for functions they perform “-

ase”

Page 15: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Reaction RatesReaction Rates• The rate of exergonic and endergonic

reactions depends on:

– Substrate availability

– Enzyme availability

– Metabolic state of the cell

– Cellular conditions (temperature, pH)

• Energy charge is maintained.

Page 16: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Page 17: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Coenzymes Coenzymes • Complex nonprotein organic substances

facilitate enzyme action by binding the substrate with its specific enzyme

• Coenzymes are smaller molecules than enzymes

• Many vitamins serve as coenzymes, e.g. riboflavin (flavin adenine dinucleotide) and niacin (nicotinamide adenine dinucleotide).

Page 18: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Phosphate-Bond EnergyPhosphate-Bond Energy

Page 19: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

ATP: The Energy CurrencyATP: The Energy Currency

• Adenosine triphosphate provides the required

energy for all cellular functions

• Cell’s “energy currency”

• ATP + H2O ↔ ADP + Pi + ∆ G (free energy) +

heat

• ATP hydrolysis yields 7.3 kcal of free energy.

Page 20: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Structure of Adenosine Structure of Adenosine Triphosphate (ATP)Triphosphate (ATP)

Page 21: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

ATP ADP Cycle is the fundamental mode of energy release in biological systems.

Anabolic: use extracted chemical energy from ATP to synthesize new compounds.

Catabolic: release energy for biologic work.

Page 22: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Catabolism-Anabolism Catabolism-Anabolism InteractionsInteractions

Page 23: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Synthesized

end products

Carbohydrates, fats,

proteins, and O2

ADP+Pi

ATPBuilding

blockprecursors

H2O + CO2

AnabolismAnabolism(endergoni(endergoni

c)c)

CatabolisCatabolismm

(exergonic)

Page 24: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

How much ATP can the body store?Why do cells store small

quantity?

Page 25: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

PhosphocreatinePhosphocreatine• Energy rich phosphate compound closely related to ATP.• Contains an energy rich phosphoanhydride bond.

Insert Figure 3.7

Page 26: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

• Released energy is coupled with energy requirement for re-synthesis of ATP.– For every mole of PCr broken down, 1 mole of ATP

synthesized.– The coupled reaction is:

Where isPCr stored?

Page 27: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Intramuscular High Energy Intramuscular High Energy PhosphatesPhosphates

1. How long can high energy phosphates sustain all-out activity?

2. Mobilization of ATP and PCr important in determining anaerobic power.

Page 28: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Cellular OxidationCellular OxidationPhosphorylationPhosphorylation: the energy transfer through the phosphate bonds of ATP to other compounds to raise them to a higher activation level

OxidationOxidation: biologic burning of macronutrients in the body for the energy needed for phosphorylation

Occurs on inner lining of mitochondrial membranesInvolves transferring electrons from NADH and FADH2 to molecular oxygen, which release and transfer chemical energy to combine ATP from ADP plus a phosphate ion.During aerobic ATP resynthesis, oxygen combines with hydrogen to form water.More than 90% of ATP synthesis takes place in the respiratory chain by oxidative reactions coupled with phosphorylation.

Page 29: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Cellular Oxidation

• Human energy dynamics involve transferring energy by chemical bonds.

• Energy for phosphorylation comes from oxidation of carbohydrate, lipid, and protein macronutrients.

Page 30: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Cellular Oxidation

• Oxidation reactionOxidation reaction: an element loses electrons (e-); a compound loses electrons, often accompanied by hydrogen ions (H+), or it gains oxygen.

• Reduction reactionReduction reaction: an element gains electrons (e-); a compound gains electrons, or it loses oxygen.

Page 31: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Cellular Oxidation

• Oxidation-reduction reactions are coupled. Every oxidation coincides with a reduction.

• OOIILL RRIIGG:

OOxidation IInvolves LLoss

RReduction IInvolves GGain• LLEEOO the lion says GGEERR:

LLose EElectrons OOxidation

GGain EElectrons RReduction

Page 32: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Cellular Oxidation

• Oxidation-reduction reactions are coupled. Every reduction coincides with a oxidation (redox).

• Cellular oxidation-reduction constitutes mechanism for energy metabolism.

• Carbohydrate, fat, and protein provide hydrogen atoms for this process.

Page 33: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Cellular Oxidation

• Hydrogens released from food molecules picked up by coenzyme NAD+ & sometimes FAD in cytosol.

• Substrate oxidizes & loses hydrogens (electrons), NAD+ gains a hydrogen & 2 electrons and reduces to NADH, the other H+ in fluid

• FAD also catalyzes dehydro-genations and accepts pairs of electrons.

Page 34: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Cellular Oxidation

• NADH and FADH2 formed in macronutrient breakdown in cytosol carry electrons to cytochromes in mitochondrial membrane. Animation: chemical rxns mitochondrion

• Cytochromes, a series of iron-protein electron carriers, pass pairs of electrons in bucket brigade fashion on the inner membranes of mitochondrion.

Page 35: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Transport of electrons by specific carriers constitutes the respiratory chain or electron transport chain.

Cellular Oxidation

Page 36: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Cellular Oxidation

• The final electron acceptor (oxidizer) in the respiratory chain is oxygen which forms water.

• Without oxygen as final oxidizer, respiratory chain cannot proceed and H remain in cellular cytosol. Animation: Electron Transport

Page 37: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Oxidative Phosphorylation

• Oxidative phosphorylation: refers to the phosphorylation of ADP during the electron transport from NADH and FADH2 to oxygen.

• Electrochemical energy generated in the ETC is harnessed and transferred to ATP synthase which phosphorylates ADP to ATP.

Page 38: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Energy Release from Food

Page 39: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Energy Release from Carbohydrate

Energy Release from Carbohydrate

• What is the primary function of CHO?• The only macronutrient whose potential

energy generates ATP anaerobically• During light & moderate intensity activity

CHO supplies about ????? body’s needs.• A continual breakdown of CHO is required

so lipid can be used for energy.

Carbohydrate

Page 40: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Energy Release from Carbohydrate

• Compare rate of aerobic breakdown of CHO to lipid.

• Net energy yield per molecule of glucose is ?? moles of ATP.

• Of 689 kCal from one mole of glucose, only 38% (263 kCals) of the energy is conserved within ATP bonds; the remainder is dissipated as heat

Carbohydrate

Page 41: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Carbohydrate

Glycolysis

Glycolysis: a series of 10 enzymatically controlled chemical reactions involving breakdown of glucose to two molecules of pyruvate.Anaerobic Glycolysis: breakdown of glucose to 2 lactates.Aerobic Glycolysis: breakdown of glucose to 2 pyruvates which then continue catabolism to acetyl coenzyme A.

Page 42: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Carbohydrate

• Glycolysis occurs in cell’s cytoplasm.

• What is energy yield in glycolysis?

• Energy from glycolysis is useful performing what exercise?

• How many pairs of hydrogen are released during glycolysis?

Page 43: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

AnaerobiAnaerobic c GlycolysiGlycolysiss

Carbohydrate

Formation of lactic acid occurs when excess H from NADH temporarily combine with pyruvic acid.

Page 44: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Carbohydrate

Body disposes of lactic acid in two ways:1. When sufficient O2

available within muscles, lactate returns H to pyruvic acid for aerobic metabolism.

2. Cori cycle in liver: gluconeogenesis – make glucose from lactate. Animation: Biochemical Cori Cycle

Page 45: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Stage Two of energy release from carbohydrate takes place in the mitochondrion. Two coenzyme A’s pick up the 2 two-carbon acetyl group and transport into mitochondria, releasing carbon dioxide molecules. Coenzyme A then releases the acetyl to begin TCA cycle.

Carbohydrate

Page 46: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Carbohydrate

Page 47: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Carbohydrate

The citric acid cycle, or Kreb’s cycle, or tricarboxylate cycle is a series of 10 enzymatically controlled chemical reactions which begins with oxaloacetate and ends with oxaloacetate. Animation: citric acid cycleWhat is the most important function of the citric acid cycle?How many ATP are formed directly from substrates?

Page 48: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Carbohydrate

What is the net energy transfer from the complete catabolism of glucose?

Page 49: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Energy Release from FatEnergy Release from Fat

• Adipocytes– Site of fat storage and mobilization– 95% of an adipocyte’s volume is

occupied by triacylglycerol (TG) fat droplets

– Lipolysis splits TG molecules into glycerol and three water-soluble free fatty acids (FFA)

– Catalyzed by hormone-sensitive lipase

Fat

Page 50: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Transport and Uptake of Free Fatty Acids

Transport and Uptake of Free Fatty Acids

• After diffusing into the circulation, FFA are transported within the circulation bound to albumin

• FFA are then taken up by active skeletal muscle in proportion to their flow and concentration

Fat

Page 51: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Breakdown of Glycerol and Fatty Acids

Breakdown of Glycerol and Fatty Acids

• Glycerol– Is converted to 3-

phosphoglyceraldehyde, an intermediate glycolytic metabolite

• FFA– Are transformed into acetyl–CoA in the

mitochondria during -oxidation– A process that successively releases 2-

carbon acetyl fragments split from long fatty acid chains

Fat

Page 52: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Fat

Fuel reserves from fat represents about 60,000 to 100,000 kcals in fat cells and 30,000 in intramuscular triglycerides. Carbohydrate reserves is <2,000 kcal.

Before energy release from fat: lypolysis.FA diffuse from adipocyte into blood, bind to albumin & delivered to tissues.

Page 53: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Beta Oxidation

Fat

Page 54: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Did You Know?Did You Know?

• As carbohydrate levels decrease, the availability of oxaloacetate may become inadequate, which impairs fat catabolism.

• Fats burn in a carbohydrate flame.

Fat

Page 55: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

• Excess macronutrients convert to fat.

• Any excess carbohydrate, lipid, or protein readily converted to fatty acid.

Page 56: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Energy Release from Energy Release from ProteinProtein

• Protein can be used as energy substrate during endurance-type exercise.

• To provide energy a.a. must be converted to usable form.

Page 57: Copyright © 2006 Lippincott Williams & Wilkins. Fundamentals of Human Energy Transfer Chapter 5 Section 3: Energy Transfer.

Copyright © 2006 Lippincott Williams & Wilkins.

Metabolic Mill

Molecules degraded to few simple units, mostly acetyl CoA.Notice reversibility except pyruvic acid to acetyl CoA.

Each pathway has rate-limiting enzyme.Cellular [ADP] is most important factor that affects enzymes controlling energy metabolism.