Front Matter

Front Matter Introduction

About the Authors

Preface

New to this Edition

Teaching and Learning Tools

Media and Supplements

Acknowledgments

SECTION 1: Biochemistry Helps Us to Understand Our World

Section Introduction

1: Biochemistry and the Unity of Life

Chapter Introduction

1.1 Living Systems Require a Limited Variety of Atoms and Molecules

1.2 There Are Four Major Classes of Biomolecules

1.3 The Central Dogma Describes the Basic Principles of Biological Information Transfer

1.4 Membranes Define the Cell and Carry Out Cellular Functions

SUMMARY

KEY TERMS

PROBLEMS

2: Water, Weak Bonds, and the Generation of Order Out of Chaos

Chapter Introduction

2.1 Thermal Motions Power Biological Interactions

2.2 Biochemical Interactions Take Place in an Aqueous Solution

2.3 Weak Interactions Are Important Biochemical Properties

2.4 Hydrophobic Molecules Cluster Together

2.5 pH Is an Important Parameter of Biochemical Systems

SUMMARY

KEY TERMS

PROBLEMS

SECTION 2: Protein Composition and Structure

Section Introduction

3: Amino Acids

Chapter Introduction

3.1 Proteins Are Built from a Repertoire of 20 Amino Acids

3.2 Amino Acids Contain a Wide Array of Functional Groups

3.3 Essential Amino Acids Must Be Obtained from the Diet

SUMMARY

KEY TERMS

PROBLEMS

4: Protein Three-Dimensional Structure

Chapter Introduction

4.1 Primary Structure: Amino Acids Are Linked by Peptide Bonds to Form Polypeptide Chains

4.2 Secondary Structure: Polypeptide Chains Can Fold into Regular Structures

4.3 Tertiary Structure: Water-Soluble Proteins Fold into Compact Structures

4.4 Quaternary Structure: Multiple Polypeptide Chains Can Assemble into a Single Protein

4.5 The Amino Acid Sequence of a Protein Determines Its Three-Dimensional Structure

SUMMARY

KEY TERMS

PROBLEMS

5: Techniques in Protein Biochemistry

Chapter Introduction

5.1 The Proteome Is the Functional Representation of the Genome

5.2 The Purification of a Protein Is the First Step in Understanding Its Function

5.3 Immunological Techniques Are Used to Purify and Characterize Proteins

5.4 Determination of Primary Structure Facilitates an Understanding of Protein Function

SUMMARY

KEY TERMS

PROBLEMS

SECTION 3: Basic Concepts and Kinetics of Enzymes

Section Introduction

6: Basic Concepts of Enzyme Action

Chapter Introduction

6.1 Enzymes Are Powerful and Highly Specific Catalysts

6.2 Many Enzymes Require Cofactors for Activity

6.3 Gibbs Free Energy Is a Useful Thermodynamic Function for Understanding Enzymes

6.4 Enzymes Facilitate the Formation of the Transition State

SUMMARY

KEY TERMS

PROBLEMS

7: Kinetics and Regulation

Chapter Introduction

7.1 Kinetics Is the Study of Reaction Rates

7.2 The Michaelis–Menten Model Describes the Kinetics of Many Enzymes

7.3 Allosteric Enzymes Are Catalysts and Information Sensors

7.4 Enzymes Can Be Studied One Molecule at a Time

SUMMARY

APPENDIX: Derivation of the Michaelis–Menten Equation

KEY TERMS

PROBLEMS

8: Mechanisms and Inhibitors

Chapter Introduction

8.1 A Few Basic Catalytic Strategies Are Used by Many Enzymes

8.2 Enzyme Activity Can Be Modulated by Temperature, pH, and Inhibitory Molecules

8.3 Chymotrypsin Illustrates Basic Principles of Catalysis and Inhibition

SUMMARY

KEY TERMS

PROBLEMS

9: Hemoglobin, an Allosteric Protein

Chapter Introduction

9.1 Hemoglobin Displays Cooperative Behavior

9.2 Myoglobin and Hemoglobin Bind Oxygen in Heme Groups

9.3 Hemoglobin Binds Oxygen Cooperatively

9.4 An Allosteric Regulator Determines the Oxygen Affinity of Hemoglobin

9.5 Hydrogen Ions and Carbon Dioxide Promote the Release of Oxygen

9.6 Mutations in Genes Encoding Hemoglobin Subunits Can Result in Disease

SUMMARY

KEY TERMS

PROBLEMS

SECTION 4: Carbohydrates and Lipids

Section Introduction

10: Carbohydrates

Chapter Introduction

10.1 Monosaccharides Are the Simplest Carbohydrates

10.2 Monosaccharides Are Linked to Form Complex Carbohydrates

10.3 Carbohydrates Are Attached to Proteins to Form Glycoproteins

10.4 Lectins Are Specific Carbohydrate-Binding Proteins

SUMMARY

KEY TERMS

PROBLEMS

11: Lipids

Chapter Introduction

11.1 Fatty Acids Are a Main Source of Fuel

11.2 Triacylglycerols Are the Storage Form of Fatty Acids

11.3 There Are Three Common Types of Membrane Lipids

SUMMARY

KEY TERMS

PROBLEMS

SECTION 5: Cell Membranes, Channels, Pumps, and Receptors

Section Introduction

12: Membrane Structure and Function

Chapter Introduction

12.1 Phospholipids and Glycolipids Form Bimolecular Sheets

12.2 Membrane Fluidity Is Controlled by Fatty Acid Composition and Cholesterol Content

12.3 Proteins Carry Out Most Membrane Processes

12.4 Lipids and Many Membrane Proteins Diffuse Laterally in the Membrane

12.5 A Major Role of Membrane Proteins Is to Function as Transporters

SUMMARY

KEY TERMS

PROBLEMS

13: Signal-Transduction Pathways

Chapter Introduction

13.1 Signal Transduction Depends on Molecular Circuits

13.2 Receptor Proteins Transmit Information into the Cell

13.3 Some Receptors Dimerize in Response to Ligand Binding and Recruit Tyrosine Kinases

13.4 Metabolism in Context: Insulin Signaling Regulates Metabolism

13.5 Calcium Ion Is a Ubiquitous Cytoplasmic Messenger

13.6 Defects in Signaling Pathways Can Lead to Diseases

SUMMARY

KEY TERMS

PROBLEMS

SECTION 6: Basic Concepts and Design of Metabolism

Section Introduction

14: Digestion: Turning a Meal into Cellular Biochemicals

Chapter Introduction

14.1 Digestion Prepares Large Biomolecules for Use in Metabolism

14.2 Proteases Digest Proteins into Amino Acids and Peptides

14.3 Dietary Carbohydrates Are Digested by Alpha-Amylase

14.4 The Digestion of Lipids Is Complicated by Their Hydrophobicity

SUMMARY

KEY TERMS

PROBLEMS

15: Metabolism: Basic Concepts and Design

Chapter Introduction

15.1 Energy Is Required to Meet Three Fundamental Needs

15.2 Metabolism Is Composed of Many Interconnecting Reactions

15.3 ATP Is the Universal Currency of Free Energy

15.4 The Oxidation of Carbon Fuels Is an Important Source of Cellular Energy

15.5 Metabolic Pathways Contain Many Recurring Motifs

15.6 Metabolic Processes Are Regulated in Three Principal Ways

SUMMARY

KEY TERMS

PROBLEMS

SECTION 7: Glycolysis and Gluconeogenesis

Section Introduction

16: Glycolysis

Chapter Introduction

16.1 Glycolysis Is an Energy-Conversion Pathway

16.2 NAD+ Is Regenerated from the Metabolism of Pyruvate

16.3 Fructose and Galactose Are Converted into Glycolytic Intermediates

16.4 The Glycolytic Pathway Is Tightly Controlled

16.5 Metabolism in Context: Glycolysis Helps Pancreatic Beta Cells Sense Glucose

SUMMARY

KEY TERMS

PROBLEMS

17: Gluconeogenesis

Chapter Introduction

17.1 Glucose Can Be Synthesized from Noncarbohydrate Precursors

17.2 Gluconeogenesis and Glycolysis Are Reciprocally Regulated

17.3 Metabolism in Context: Precursors Formed by Muscle Are Used by Other Organs

SUMMARY

KEY TERMS

PROBLEMS

SECTION 8: The Citric Acid Cycle

Section Introduction

18: Preparation for the Cycle

Chapter Introduction

18.1 Pyruvate Dehydrogenase Forms Acetyl Coenzyme A from Pyruvate

18.2 The Pyruvate Dehydrogenase Complex Is Regulated by Two Mechanisms

SUMMARY

KEY TERMS

PROBLEMS

19: Harvesting Electrons from the Cycle

Chapter Introduction

19.1 The Citric Acid Cycle Consists of Two Stages

19.2 Stage One Oxidizes Two Carbon Atoms to Gather Energy-Rich Electrons

19.3 Stage Two Regenerates Oxaloacetate and Harvests Energy-Rich Electrons

19.4 The Citric Acid Cycle Is Regulated

19.5 The Glyoxylate Cycle Enables Plants and Bacteria to Convert Fats into Carbohydrates

SUMMARY

KEY TERMS

PROBLEMS

SECTION 9: Oxidative Phosphorylation

Section Introduction

20: The Electron-Transport Chain

Chapter Introduction

20.1 Oxidative Phosphorylation in Eukaryotes Takes Place in Mitochondria

20.2 Oxidative Phosphorylation Depends on Electron Transfer

20.3 The Respiratory Chain Consists of Proton Pumps and a Physical Link to the Citric Acid Cycle

SUMMARY

KEY TERMS

PROBLEMS

21: The Proton-Motive Force

Chapter Introduction

21.1 A Proton Gradient Powers the Synthesis of ATP

21.2 Shuttles Allow Movement Across Mitochondrial Membranes

21.3 Cellular Respiration is Regulated by the Need for ATP

SUMMARY

KEY TERMS

PROBLEMS

SECTION 10: The Light Reactions of Photosynthesis and the Calvin Cycle

Section Introduction

22: The Light Reactions

Chapter Introduction

22.1 Photosynthesis Takes Place in Chloroplasts

22.2 Photosynthesis Transforms Light Energy into Chemical Energy

22.3 Two Photosystems Generate a Proton Gradient and NADPH

22.4 A Proton Gradient Drives ATP Synthesis

SUMMARY

KEY TERMS

PROBLEMS

23: The Calvin Cycle

Chapter Introduction

23.1 The Calvin Cycle Synthesizes Hexoses from Carbon Dioxide and Water

23.2 The Calvin Cycle Is Regulated by the Environment

SUMMARY

KEY TERMS

PROBLEMS

SECTION 11: Glycogen Metabolism and the Pentose Phosphate Pathway

Section Introduction

24: Glycogen Degradation

Chapter Introduction

24.1 Glycogen Breakdown Requires Several Enzymes

24.2 Phosphorylase Is Regulated by Allosteric Interactions and Reversible Phosphorylation

24.3 Epinephrine and Glucagon Signal the Need for Glycogen Breakdown

SUMMARY

KEY TERMS

PROBLEMS

25: Glycogen Synthesis

Chapter Introduction

25.1 Glycogen Is Synthesized and Degraded by Different Pathways

25.2 Metabolism in Context: Glycogen Breakdown and Synthesis Are Reciprocally Regulated

SUMMARY

KEY TERMS

PROBLEMS

26: The Pentose Phosphate Pathway

Chapter Introduction

26.1 The Pentose Phosphate Pathway Yields NADPH and Five-Carbon Sugars

26.2 Metabolism in Context: Glycolysis and the Pentose Phosphate Pathway Are Coordinately Controlled

26.3 Glucose 6-phosphate Dehydrogenase Lessens Oxidative Stress

SUMMARY

KEY TERMS

PROBLEMS

SECTION 12: Fatty Acid and Lipid Metabolism

Section Introduction

27: Fatty Acid Degradation

Chapter Introduction

27.1 Fatty Acids Are Processed in Three Stages

27.2 The Degradation of Unsaturated and Odd-Chain Fatty Acids Requires Additional Steps

27.3 Ketone Bodies Are Another Fuel Source Derived from Fats

27.4 Metabolism in Context: Fatty Acid Metabolism Is a Source of Insight into Various Physiological States

SUMMARY

KEY TERMS

PROBLEMS

28: Fatty Acid Synthesis

Chapter Introduction

28.1 Fatty Acid Synthesis Takes Place in Three Stages

28.2 Additional Enzymes Elongate and Desaturate Fatty Acids

28.3 Acetyl CoA Carboxylase Is a Key Regulator of Fatty Acid Metabolism

28.4 Metabolism in Context: Ethanol Alters Energy Metabolism in the Liver

SUMMARY

KEY TERMS

PROBLEMS

29: Lipid Synthesis: Storage Lipids, Phospholipids, and Cholesterol

Chapter Introduction

29.1 Phosphatidate Is a Precursor of Storage Lipids and Many Membrane Lipids

29.2 Cholesterol Is Synthesized from Acetyl Coenzyme A in Three Stages

29.3 The Regulation of Cholesterol Synthesis Takes Place at Several Levels

29.4 Lipoproteins Transport Cholesterol and Triacylglycerols Throughout the Organism

29.5 Cholesterol Is the Precursor of Steroid Hormones

SUMMARY

KEY TERMS

PROBLEMS

SECTION 13: The Metabolism of Nitrogen-Containing Molecules

Section Introduction

30: Amino Acid Degradation and the Urea Cycle

Chapter Introduction

30.1 Nitrogen Removal Is the First Step in the Degradation of Amino Acids

30.2 Ammonium Ion Is Converted into Urea in Most Terrestrial Vertebrates

30.3 Carbon Atoms of Degraded Amino Acids Emerge As Major Metabolic Intermediates

SUMMARY

KEY TERMS

PROBLEMS

31: Amino Acid Synthesis

Chapter Introduction

31.1 The Nitrogenase Complex Fixes Nitrogen

31.2 Amino Acids Are Made from Intermediates of Major Pathways

31.3 Feedback Inhibition Regulates Amino Acid Biosynthesis

SUMMARY

KEY TERMS

PROBLEMS

32: Nucleotide Metabolism

Chapter Introduction

32.1 An Overview of Nucleotide Biosynthesis and Nomenclature

32.2 The Pyrimidine Ring Is Assembled and Then Attached to a Ribose Sugar

32.3 The Purine Ring Is Assembled on Ribose Phosphate

32.4 Ribonucleotides Are Reduced to Deoxyribonucleotides

32.5 Nucleotide Biosynthesis Is Regulated by Feedback Inhibition

32.6 Disruptions in Nucleotide Metabolism Can Cause Pathological Conditions

SUMMARY

KEY TERMS

PROBLEMS

SECTION 14: Nucleic Acid Structure and DNA Replication

Section Introduction

33: The Structure of Informational Macromolecules: DNA and RNA

Chapter Introduction

33.1 A Nucleic Acid Consists of Bases Linked to a Sugar–Phosphate Backbone

33.2 Nucleic Acid Strands Can Form a Double-Helical Structure

33.3 DNA Double Helices Can Adopt Multiple Forms

33.4 Eukaryotic DNA Is Associated with Specific Proteins

33.5 RNA Can Adopt Elaborate Structures

SUMMARY

KEY TERMS

PROBLEMS

34: DNA Replication

Chapter Introduction

34.1 DNA Is Replicated by Polymerases

34.2 DNA Replication Is Highly Coordinated

SUMMARY

KEY TERMS

PROBLEMS

35: DNA Repair and Recombination

Chapter Introduction

35.1 Errors Can Arise in DNA Replication

35.2 DNA Damage Can Be Detected and Repaired

35.3 DNA Recombination Plays Important Roles in Replication and Repair

SUMMARY

KEY TERMS

PROBLEMS

SECTION 15: RNA Synthesis, Processing, and Regulation

Section Introduction

36: RNA Synthesis and Regulation in Bacteria

Chapter Introduction

36.1 Cellular RNA Is Synthesized by RNA Polymerases

36.2 RNA Synthesis Comprises Three Stages

36.3 The lac Operon Illustrates the Control of Bacterial Gene Expression

SUMMARY

KEY TERMS

PROBLEMS

37: Gene Expression in Eukaryotes

Chapter Introduction

37.1 Eukaryotic Cells Have Three Types of RNA Polymerases

37.2 RNA Polymerase II Requires Complex Regulation

37.3 Gene Expression Is Regulated by Hormones

37.4 Histone Acetylation Results in Chromatin Remodeling

SUMMARY

KEY TERMS

PROBLEMS

38: RNA Processing in Eukaryotes

Chapter Introduction

38.1 Mature Ribosomal RNA Is Generated by the Cleavage of a Precursor Molecule

38.2 Transfer RNA Is Extensively Processed

38.3 Messenger RNA Is Modified and Spliced

38.4 RNA Can Function as a Catalyst

SUMMARY

KEY TERMS

PROBLEMS

SECTION 16: Protein Synthesis and Recombinant DNA Techniques

Section Introduction

39: The Genetic Code

Chapter Introduction

39.1 The Genetic Code Links Nucleic Acid and Protein Information

39.2 Amino Acids Are Activated by Attachment to Transfer RNA

39.3 A Ribosome Is a Ribonucleoprotein Particle Made of Two Subunits

SUMMARY

KEY TERMS

PROBLEMS

40: The Mechanism of Protein Synthesis

Chapter Introduction

40.1 Protein Synthesis Decodes the Information in Messenger RNA

40.2 Peptidyl Transferase Catalyzes Peptide-Bond Synthesis

40.3 Bacteria and Eukaryotes Differ in the Initiation of Protein Synthesis

40.4 A Variety of Biomolecules Can Inhibit Protein Synthesis

40.5 Ribosomes Bound to the Endoplasmic Reticulum Manufacture Secretory and Membrane Proteins

40.6 Protein Synthesis Is Regulated by a Number of Mechanisms

SUMMARY

KEY TERMS

PROBLEMS

41: Recombinant DNA Techniques

Chapter Introduction

41.1 Nucleic Acids Can Be Synthesized from Protein-Sequence Data

41.2 Recombinant DNA Technology Has Revolutionized All Aspects of Biology

41.3 Eukaryotic Genes Can Be Manipulated with Considerable Precision

SUMMARY

KEY TERMS

PROBLEMS

APPENDIX A: Physical Constants and Conversion of Units

APPENDIX A: Physical Constants and Conversion of Units

APPENDIX B: Acidity Constants

APPENDIX B: Acidity Constants

APPENDIX C: Standard Bond Lengths

APPENDIX C: Standard Bond Lengths

APPENDIX D: Water-Soluble Vitamins

APPENDIX D: Water-Soluble Vitamins

Answers to Problems

Chapter 1

Chapter 2

Chapter 3

Chapter 4

Chapter 5

Chapter 6

Chapter 7

Chapter 8

Chapter 9

Chapter 10

Chapter 11

Chapter 12

Chapter 13

Chapter 14

Chapter 15

Chapter 16

Chapter 17

Chapter 18

Chapter 19

Chapter 20

Chapter 21

Chapter 22

Chapter 23

Chapter 24

Chapter 25

Chapter 26

Chapter 27

Chapter 28

Chapter 29

Chapter 30

Chapter 31

Chapter 32

Chapter 33

Chapter 34

Chapter 35

Chapter 36

Chapter 37

Chapter 38

Chapter 39

Chapter 40

Chapter 41