Lehninger, The New Edition

LEHNINGER

PRINCIPLES OF BIOCHEMISTRY

international edition

Sixth edition

David L. Nelson

University of Wisconsin–Madison

Michael M. Cox

University of Wisconsin–Madison

February 2013Hardback 978-1-4641-0962-1

in this edition

new art

the new edition features a dramatically

re-envisioned art program with the

following advantages:

n new renditions of classic figures are now easier

to interpret and learn from

n Figures that pair molecular models with

schematic cartoons, generated specifically for this

book use consistent shapes and color schemes

n Figures with numbered, annotated steps walk

students through complex processes

n Summary figures help students to keep the big

picture in mind while learning the specifics.

Chaperonins in protein folding.

F i G U r e 4 - 3 1

Clear writing and illustrations…Clear explanations of difficult

concepts…Clear communication of the ways that biochemistry is

currently understood and practiced. For over 35 years, in edition after

bestselling edition, Principles of Biochemistry has put those defining

principles into practice, guiding students through a coherent introduction

to the essentials of biochemistry without overwhelming them.

the new edition brings this remarkable text into a new era. like its

predecessors, Lehninger Principles of Biochemistry, Sixth edition strikes a

careful balance of current science and enduring concepts, incorporating

many new research findings, but focusing on those that help illustrate

important principles of biochemistry. With this edition, students will

encounter new information emerging from high throughput dna

sequencing, x-ray crystallography, and the manipulation of genes and gene

expression, and other techniques. in addition, students will see how

contemporary biochemistry has shifted away from exploring metabolic

pathways in isolation to focusing on interactions among pathways. they

will also get an updated understanding of the relevance of biochemistry to

the study of human disease (especially diabetes) as well as the important

role of evolutionary theory in biochemical research.

these extensive content changes, as well as new artwork, make this edition

of Lehninger Principles of Biochemistry the most impressive yet.

New Sixth Edition

Updated GenomicsChapter 9, DNA-Based Information Technologies, now

incorporates the latest genomic methods, such as:

n next generation dna sequencing, including the illumina and 454 sequencing

methods and platforms

n applications of genomics, including the use of haplotypes to trace human

migrations and phylogenetics to locate human genes associated with

inherited disease conditions

n Forensic genotyping and the use of personalized genomics in medicine

numerous other chapters have been updated to reflect advances gained from

new genomic methods.

New Scienceevery chapter has been thoroughly revised and updated to include both the

most important advances in biochemistry and information needed in a modern

biochemistry text. among the new and/or thoroughly

updated topics in this edition are:

n Prebiotic evolution, black smokers, and the rna world (Chapter 1)

n intrinsically disordered proteins (Chapter 4)

n transition state analogs and irreversible inhibition (Chapter 6)

n Blood coagulation pathways in the context of enzymatic regulation

(Chapter 6)

n asymmetric lipid distribution in bilayer (Chapter 11)

n role of Bar superfamily proteins in membrane curvature (Chapter 11)

n Scaffold proteins (aKaPS and others) and their regulatory roles

(Chapter 12)

n reactive oxygen species as by-products and as signals (Chapter 19)

n Structure and function of the oxygen-evolving metal cluster in PSii

(Chapter 19)

n Formation, transport of lipoproteins in mammals, including the roles of

SreBP, SCaP, insig in cholesterol regulation (Chapter 21)

n integration of carbohydrate and lipid metabolism by PPars, SreBPs,

mtorC1, and lxr (Chapters 21, 23)

n Creatine phosphate and the role of creatine kinase in moving atP to

cytosol (Chapter 23)

n Microbial symbionts in the gut and their influence on energy

metabolism and adipogenesis (Chapter 23)

n nucleosomes: their modification and positioning and higher order

chromatin structure (Chapter 24)

n dna polymerases and homologous recombination (Chapter 25)

n loading eukaryotic rna polymerase ii (Chapter 26)

n Mutation-resistant nature of the genetic code (Chapter 27)

n ribosome structure and the initiation of translation in eukaryotes

(Chapter 27)

n dna looping, combinatorial control, chromatin remodeling, and positive

regulation in eukaryotes (Chapter 28)

n regulation of the initiation of transcription in eukaryotes (Chapter 28)

n Steroid-binding nuclear receptors (Chapter 28)

New Biochemical Methods

new methods or updates described in this edition include:

n Modern Sanger protein sequencing and mass spectrometry (Chapter 3)

n Mass spectrometry applied to proteomics, glycomics, lipidomics, and

metabolomics (Chapters 3, 7, 10)

n oligosaccharide microarrays to explore protein-oligosaccharide

interactions and the “carbohydrate code” (Chapter 7)

n Modern genomic methods, including next-generation dna sequencing

(Chapter 9)

n the use of genomics to locate genes involved in human disease

(Chapter 9)

n Quantitative PCr and reverse transcriptase PCr (Chapter 9)

n Genetic engineering of photosynthetic organisms (Chapter 20)

n Use of Positron emission tomography (Pet) to visualize tumors and

brown adipose tissue (Chapter 23)

n development of bacterial strains with altered genetic codes for

site-specific insertion of novel amino acids into proteins (Chapter 27)

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Binding of the intrinsically disordered carboxyl terminus of p53 protein to its binding partners.

Next generation reversible terminator sequencing.

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New Sixth Edition

New Medical ApplicationsThis icon is used throughout the book to denote material of special

medical interest. Many sections explore what we know about the molecular mechanisms of disease. A few of the new or

revised medical applications in this edition are:

n Box 4-6, death by Misfolding: the Prior diseases (Chapter 4)

n Paganini and ehlers-danlos Syndrome (Chapter 4)

n hiV protease inhibitors and how basic enzymatic principles influenced

their design (Chapter 6)

n Blood coagulation cascade and hemophilia (Chapter 6)

n Curing african Sleeping Sickness with an enzymatic suicide inhibitor

(Chapter 6)

n how researchers locate human genes involved in inherited diseases

(Chapter 9)

n Multidrug resistance transporters and their importance in clinical

medicine (Chapter 11)

n Multistep progression to colorectal cancer (Chapter 12)

n Cholesterol metabolism, cardiovascular disease, and mechanism of

plaque formation in atherosclerosis (Chapter 21)

n P450 and drug interactions (Chapter 21) hMG-Coa reductase

(Chapter 21) and Box 21–3, the lipid hypothesis and the development

of Statins

n Box 24–1, Curing disease by inhibiting topoisomerases, describing the

use of topoisomerase inhibitors in the treatment of bacterial infections and

cancer, including material on ciprofloxacin (the antibiotic effective for

anthrax) Stem cells (Chapter 28)

Special Theme: Understanding Metabolism through Obesity and DiabetesThe urgent world health issue of diabetes provides a defining theme for the book’s chapters on metabolism and its control. Sections and boxes

highlighting the interplay of metabolism, obesity, and diabetes are:

n Untreated diabetes Produces life-threatening acidosis (Chapter 2)

n Box 7–1, Blood Glucose Measurements in the diagnosis and treatment of

diabetes, introducing hemoglobin glycation and aGes and their role in the

pathology of advanced diabetes

n Glucose Uptake is deficient in type 1 diabetes Mellitus (Chapter 14)

n Ketone Bodies are overproduced in diabetes and during Starvation

(Chapter 17)

n Some Mutations in Mitochondrial Genomes Cause disease (Chapter 19)

n diabetes Can result from defects in the Mitochondria of Pancreatic Cells

(Chapter 19)

n adipose tissue Generates Glycerol 3-phosphate by Glyceroneogenesis

(Chapter 21)

n diabetes Mellitus arises from defects in insulin Production or action

(Chapter 23)

n Section 23.4, obesity and the regulation of Body Mass, includes a new

discussion of the regulatory roles of torC1 in regulating cell growth

n Section 23.5, obesity, the Metabolic Syndrome, and type 2 diabetes,

discusses the role of ectopic lipids and inflammation in the development of

insulin resistance, and the management of type 2 diabetes with exercise,

diet, and medication

Overloading adipocytes with triglycerides triggers inflammation in fat tissue, and ectopic lipid deposition and insulin resistance.

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Nelson | Cox L E H N I N G E R P R I N C I P L E S O F B I O C H E M I S T R Y I N T E R N A T I O N A L E D I T I O N S I x T H E D I T I O N

Special Theme: Evolution Evolution is presented as a foundational principle of biochemistry

throughout this edition, including:

n Section 1.5, evolutionary Foundations, discusses how life may have evolved

and canvasses some of the early milestones in the evolution of eukaryotic

cells (Chapter 1)

n Genome Sequencing informs Us about our humanity (Chapter 9)

n Genome Comparisons help locate Genes involved in disease (Chapter 9)

n Genome Sequences inform Us about our Past and Provide opportunities

for the Future (Chapter 9)

n Box 9–3, Getting to Know the neanderthals (Chapter 9)

n aBC transporters Use atP to drive the active transport of a Wide Variety

of Substrates (Chapter 11)

n Signaling Systems of Plants have Some of the Same Components Used by

Microbes and Mammals (Chapter 12)

n the β-oxidation enzymes of different organelles have diverged during

evolution (Chapter 17)

n Section 19.10, the evolution of oxygenic Photosynthesis

n Mitochondria and Chloroplasts evolved from endosymbiotic Bacteria

(Chapter 19)

n Photosystems i and ii evolved from Bacterial Photosystems (Chapter 19)

n rna Synthesis offers important Clues to Biochemical evolution (Chapter 26)

n Box 27–1, exceptions that Prove the rule: natural

Variations in the Genetic Code (Chapter 27)

n Box 27–2, From an rna World to a Protein World

(Chapter 27)

n Box 28-1, of Fins, Wings, Beaks, and things

(Chapter 28)

Lehninger Teaching HallmarksStudents encountering biochemistry for the first time often have difficulty with two key aspects of the course: approaching quantitative problems and drawing on their previous studies of organic chemistry. Those same students must also learn a complex language, with conventions that are often unstated. As always, Principles of Biochemistry

helps students cope with these challenges:

Focus on Chemical Logicn Section 13.2, Chemical logic and common

biochemical reactions, discusses the common

biochemical reaction types that underlie all

metabolic reactions, helping student to connect

organic chemistry with biochemistry.

n NEW chemical logic figures highlight the

conservation of mechanism and illustrate patterns

that make learning pathways easier. Chemical

logic figures are provided for each of the central

metabolic pathways: glycolysis (Fig 14-3), citric

acid cycle (Fig. 16-7), and fatty acid oxidation (

Fig 17-9).

n Mechanism figures feature step-by-step descriptions to help students

understand the reaction process. these figures use a consistent set of

conventions introduced and explained in detail with the first enzyme

mechanism encountered.

Problem-Solving Tools

n In-text Worked Examples take students through some of the most

difficult equations step by step. new worked examples appear in

Chapters 1, 2, and 19.

n More than 600 end-of-chapter problems (over 75 of them new) give

students further opportunity to practice what they have learned.

n Data Analysis Problems (one at the end of each chapter), contributed

by Brian White of the University of Massachusetts–Boston, encourage

students interpret data from the literature.

Key Conventions

the text highlights clear statements of many assumptions and

conventions that students are often expected to assimilate without

being told (for example, peptide sequences are written from amino-to

carboxyl-terminal end, left to right; nucleotide sequences are written

from 5’ to 3’ end, left to right).

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Lehninger Contents

1 The Foundations of Biochemistry1.1 Cellular Foundations 1.2 Chemical Foundations Box 1–1 Molecular Weight, Molecular Mass, and their Correct Units Box 1–2 louis Pasteur and optical activity: In Vino, Veritas 1.3 Physical Foundations Box 1–3 entropy: things Fall apart 1.4 Genetic Foundations 1.5 evolutionary Foundations

2 Water 2.1 Weak interactions in aqueous Systems 2.2 ionization of Water, Weak acids, and Weak Bases 2.3 Buffering against ph Changes in Biological Systems Box 2–1 Medicine: on Being one’s own rabbit [don’t trythis at home!] 2.4 Water as a reactant 2.5 the Fitness of the aqueous environment for living organisms

3 Amino Acids, Peptides, and Proteins 3.1 amino acids Box 3–1 Methods: absorption of light by Molecules: the lambert-Beer law 3.2 Peptides and Proteins 3.3 Working with Proteins 3.4 the Structure of Proteins: Primary Structure Box 3–2 Consensus Sequences and Sequence logos

4 The Three-Dimensional Structure of Proteins 4.1 overview of Protein Structure 4.2 Protein Secondary Structure Box 4–1 Methods: Knowing the right hand from the left 4.3 Protein tertiary and Quaternary Structures Box 4–2 Permanent Waving is Biochemical engineering Box 4–3 Why Sailors, explorers, and College Students Should eat their Fresh Fruits and Vegetables Box 4–4 the Protein data Bank Box 4–5 Methods: Methods for determining the three-dimensional Structure of a Protein 4.4 Protein denaturation and Folding Box 4–6 Medicine: death by Misfolding: the Prion diseases

5 Protein Function 5.1 reversible Binding of a Protein to a ligand: oxygen-Binding Proteins Box 5–1 Medicine: Carbon Monoxide: a Stealthy Killer 5.2 Complementary interactions between Proteins and ligands: the immune System and immunoglobulins

5.3 Protein interactions Modulated by Chemical energy: actin, Myosin, and Molecular Motors

6 Enzymes 6.1 an introduction to enzymes 6.2 how enzymes Work 6.3 enzyme Kinetics as an approach to Understanding Mechanism Box 6–1 transformations of the Michaelis-Menten equation: the double-reciprocal PlotBox 6–2 Kinetic tests for determining inhibition Mechanisms Box 6–3 Curing african Sleeping Sickness with a Biochemical trojan horse 6.4 examples of enzymatic reactions 6.5 regulatory enzymes

7 Carbohydrates and Glycobiology 7.1 Monosaccharides and disaccharides Box 7–1 Medicine: Blood Glucose Measurements in the diagnosis and treatment of diabetes Box 7–2 Sugar is Sweet, and So are . . . a Few other things 7.2 Polysaccharides 7.3 Glycoconjugates: Proteoglycans, Glycoproteins, and Glycolipids 7.4 Carbohydrates as informational Molecules: the Sugar Code 7.5 Working with Carbohydrates

8 Nucleotides and Nucleic Acids 8.1 Some Basics 8.2 nucleic acid Structure 8.3 nucleic acid Chemistry 8.4 other Functions of nucleotides

9 DNA-Based Information Technologies 9.1 Studying Genes and their Products Box 9–1 a Powerful tool in Forensic Medicine 9.2 Using dna-Based Methods to Understand Protein Function 9.3 Genomics and the human Story Box 9–2 Medicine: Personalized Genomic Medicine Box 9–3 Getting to Know the neanderthals

10 Lipids 10.1 Storage lipids 10.2 Structural lipids in Membranes Box 10–1 Medicine: abnormal accumulations of Membrane lipids: Some inherited human diseases 10.3 lipids as Signals, Cofactors, and Pigments 10.4 Working with lipids

Nelson | Cox L E H N I N G E R P R I N C I P L E S O F B I O C H E M I S T R Y I N T E R N A T I O N A L E D I T I O N S I x T H E D I T I O N

11 Biological Membranes and Transport 11.1 the Composition and architecture of Membranes 11.2 Membrane dynamics 11.3 Solute transport across Membranes Box 11–1 Medicine: defective Glucose and Water transport in two Forms of diabetesBox 11–2 Medicine: a defective ion Channel in Cystic Fibrosis

12 Biosignaling 12.1 General Features of Signal transduction Box 12–1 Methods: Scatchard analysis Quantifies the receptor-ligand interaction 12.2 Protein–Coupled receptors and Second Messengers Box 12–2 Medicine: G Proteins: Binary Switches in health and disease Box 12–3 Methods: Fret: Biochemistry Visualized in a living Cell 12.3 receptor tyrosine Kinases 12.4 receptor Guanylyl Cyclases, cGMP, and Protein Kinase G 12.5 Multivalent adaptor Proteins and Membrane rafts 12.6 Gated ion Channels 12.7 integrins: Bidirectional Cell adhesion receptors 12.8 regulation of transcription by nuclear hormone receptors 12.9 Signaling in Microorganisms and Plants 12.10 Sensory transduction in Vision, olfaction, and Gustation Box 12–4 Medicine: Color Blindness: John dalton’s experiment from the Grave12.11 regulation of the Cell Cycle by Protein Kinases 12.12 oncogenes, tumor Suppressor Genes, and Programmed Cell death Box 12–5 Medicine: development of Protein Kinase inhibitors for Cancer treatment

13 Bioenergetics and Biochemical Reaction Types 13.1 Bioenergetics and thermodynamics 13.2 Chemical logic and Common Biochemical reactions 13.3 Phosphoryl Group transfers and atP Box 13–1 Firefly Flashes: Glowing reports of atP 13.4 Biological oxidation-reduction reactions

14 Glycolysis, Gluconeogenesis, and the Pentose Phosphate Pathway 14.1 Glycolysis Box 14–1 Medicine: high rate of Glycolysis in tumors Suggests targets for Chemotherapy and Facilitates diagnosis 14.2 Feeder Pathways for Glycolysis .

14.3 Fates of Pyruvate under anaerobic Conditions:Fermentation Box 14–2 athletes, alligators, and Coelacanths: Glycolysis at limiting Concentrations of oxygen Box 14–3 ethanol Fermentations: Brewing Beer and Producing Biofuels14.4 Gluconeogenesis 14.5 Pentose Phosphate Pathway of Glucose oxidation Box 14–4 Medicine: Why Pythagoras Wouldn’t eat Falafel: Glucose 6-Phosphate dehydrogenase deficiency

15 Principles of Metabolic Regulation 15.1 regulation of Metabolic Pathways 15.2 analysis of Metabolic Control Box 15–1 Methods: Metabolic Control analysis: Quantitative aspects 15.3 Coordinated regulation of Glycolysis and Gluconeogenesis Box 15–2 isozymes: different Proteins that Catalyze the Same reaction Box 15–3 Medicine: Genetic Mutations that lead to rare Forms of diabetes 15.4 the Metabolism of Glycogen in animals Box 15–4 Carl and Gerty Cori: Pioneers in Glycogen Metabolism and disease 15.5 Coordinated regulation of Glycogen Synthesis and Breakdown

16 The Citric Acid Cycle 16.1 Production of acetyl-Coa (activated acetate) 16.2 reactions of the Citric acid Cycle Box 16–1 Moonlighting enzymes: Proteins with More than one Job Box 16–2 Synthases and Synthetases; ligases and lyases; Kinases, Phosphatases, and Phosphorylases: Yes, the names are Confusing! Box 16–3 Citrate: a Symmetric Molecule that reacts asymmetrically 16.3 regulation of the Citric acid Cycle 16.4 the Glyoxylate Cycle

17 Fatty Acid Catabolism 17.1 digestion, Mobilization, and transport of Fats 17.2 oxidation of Fatty acids Box 17–1 Fat Bears Carry out oxidation in their Sleep Box 17–2 Coenzyme B12: a radical Solution to a Perplexing Problem 17.3 Ketone Bodies

Lehninger Contents

18 Amino Acid Oxidation and the Production of Urea 18.1 Metabolic Fates of amino Groups 18.2 nitrogen excretion and the Urea Cycle Box 18–1 Medicine: assays for tissue damage 18.3 Pathways of amino acid degradation Box 18–2 Medicine: Scientific Sleuths Solve a Murder Mystery

19 Oxidative Phosphorylation and Photophosphorylation

Oxidative Phosphorylation

19.1 electron-transfer reactions in Mitochondria Box 19–1 hot, Stinking Plants and alternative respiratory Pathways19.2 atP Synthesis Box 19–2 Methods: atomic Force Microscopy to Visualize Membrane Proteins 19.3 regulation of oxidative Phosphorylation 19.4 Mitochondria in thermogenesis, Steroid Synthesis, and apoptosis 19.5 Mitochondrial Genes: their origin and the effects of Mutations

Photosynthesis: Harvesting Light Energy

19.6 General Features of Photophosphorylation 19.7 light absorption 19.8 the Central Photochemical event: light-driven electron Flow 19.9 atP Synthesis by Photophosphorylation 19.10 the evolution of oxygenic Photosynthesis

20 Carbohydrate Biosynthesis in Plants and Bacteria

20.1 Photosynthetic Carbohydrate Synthesis 20.2 Photorespiration and the C4 and CaM PathwaysBox 20–1 Will Genetic engineering of Photosynthetic organisms increase their efficiency? 20.3 Biosynthesis of Starch and Sucrose 20.4 Synthesis of Cell Wall Polysaccharides: Plant Cellulose and Bacterial Peptidoglycan 20.5 integration of Carbohydrate Metabolism in the Plant Cell

21 Lipid Biosynthesis

21.1 Biosynthesis of Fatty acids and eicosanoids Box 21–1 Medicine: Mixed-Function oxidases, Cytochrome P-450s and drug overdoses 21.2 Biosynthesis of triacylglycerols 21.3 Biosynthesis of Membrane Phospholipids 21.4 Cholesterol, Steroids, and isoprenoids: Biosynthesis, regulation, and transport Box 21–2 Medicine: apoe alleles Predict incidence of alzheimer’s disease

Box 21–3 Medicine: the lipid hypothesis and the development of Statins

22 Biosynthesis of Amino Acids, Nucleotides, and Related Molecules

22.1 overview of nitrogen Metabolism Box 22–1 Unusual lifestyles of the obscure but abundant 22.2 Biosynthesis of amino acids 22.3 Molecules derived from amino acids Box 22–2 on Kings and Vampires 22.4 Biosynthesis and degradation of nucleotides

23 Hormonal Regulation and Integration of Mammalian Metabolism

23.1 hormones: diverse Structures for diverse Functions Box 23–1 Medicine: how is a hormone discovered? the arduous Path to Purified insulin 23.2 tissue-Specific Metabolism: the division of labor Box 23–2 Creatine and Creatine Kinase: invaluable diagnostic aids and the Muscle Builder’s Friends 23.3 hormonal regulation of Fuel Metabolism 23.4 obesity and the regulation of Body Mass 23.5 obesity, the Metabolic Syndrome, and type 2 diabetes

24 Genes and Chromosomes

24.1 Chromosomal elements 24.2 dna Supercoiling Box 24–1 Medicine: Curing disease by inhibiting topoisomerases 24.3 the Structure of Chromosomes Box 24–2 Medicine: epigenetics, nucleosome Structure, and histone Variants

25 DNA Metabolism

25.1 dna replication 25.2 dna repair Box 25–1 Medicine: dna repair and Cancer 25.3 dna recombination Box 25–2 Medicine: Why Proper Chromosomal Segregation Matters

26 RNA Metabolism

26.1 dna-dependent Synthesis of rna Box 26–1 Methods: rna Polymerase leaves its Footprint on a Promoter 26.2 rna Processing 26.3 rna-dependent Synthesis of rna and dna Box 26–2 Medicine: Fighting aidS with inhibitors of hiV reverse transcriptase Box 26–3 Methods: the Selex Method for Generating rna Polymers with new Functions Box 26–4 an expanding rna Universe Filled with tUF rnas

F i G U r e 2 3 - 3 0 Fuel metabolism in the liver

during prolonged fasting or in

uncontrolled diabetes mellitus.

Nelson | Cox L E H N I N G E R P R I N C I P L E S O F B I O C H E M I S T R Y I N T E R N A T I O N A L E D I T I O N S I x T H E D I T I O N

27 Protein Metabolism 27.1 the Genetic Code Box 27–1 exceptions that Prove the rule: natural Variations in the Genetic Code 27.2 Protein Synthesis Box 27–2 From an rna World to a Protein World Box 27–3 natural and Unnatural expansion of the Genetic Code Box 27–4 induced Variation in the Genetic Code: nonsense Suppression 27.3 Protein targeting and degradation

28 Regulation of Gene Expression

28.1 Principles of Gene regulation 28.2 regulation of Gene expression in Bacteria 28.3 regulation of Gene expression in eukaryotes Box 28–1 of Fins, Wings, Beaks, and things

Appendix A Common Abbreviations in the Biochemical Research Literature Appendix B Abbreviated Solutions to Problems Glossary Credits Index

F O R I N S T R U C T O R S

Instructor’s Resource DVD ( ISBN: 978-1-4641-0969-0 ) the dVd includes fully optimized JPeG files of text art, animated enzyme mechanisms, animated biochemical techniques, protein data bank ids, living Graphs, and the test Bank.

n Fully optimized JPEG files—every figure, photo, and table in the text is available as a separate jpeg or in PowerPoint, with colors, resolution and labels enhanced. these files have been reviewed by course instructors and tested in a large lecture hall to ensure maximum clarity and visibility.

n Animated Enzyme Mechanisms and Animated Biochemical Techniques—available in Flash files and preloaded into PowerPoint, in both PC and Macintosh formats.

n Protein Data Bank IDs for the structures in the text and the Jmol interactive Web browser applet, arranged by figure number.

n Living Graphs—with these dynamic versions of key equations from the textbook, you and your students can see the graphic results of changing parameters.

n A comprehensive Test Bank ( in PDF and editable Word formats)—150 multiple-choice and short-answer problems per chapter, rated by level of difficulty.

F O R S T U D E N T S

FREE Book Companion Site, www.whfreeman.com/lehninger6eprovides tudents with media designed to enhance their understanding of biochemical principles and improve their problem-solving ability.

n Molecular Structure Tutorials—students use the Jmol interactive Web browser to explore the molecular structures in the textbook, including:

▶ Protein architecture

▶ Bacteriorhodopsin

▶ lac repressor

▶ nucleotides

▶ MhC Molecules

▶ trimeric G Proteins

▶ oxygen-Binding Proteins

▶ restriction endonucleases

▶ hammerhead ribozyme

n Protein Data Bank IDs for the structures in the text and the Jmol interactive Web browser applet, arranged by figure number.

n Living Graphs—with these dynamic versions of key equations from the textbook, students can see the graphic results of changing parameters.

The Absolute, Ultimate Guide to Lehninger Principles of Biochemistry, Sixth Edition, Study Guide and Solutions ManualMarcy Osgood, University of New Mexico School of MedicineKaren Ocorr, Sanford-Burnham Medical Research Institute Pape

iSBn 978-1-4292-9476-8

The Absolute, Ultimate Guide combines an innovative study guide with a

reliable solutions manual (providing extended solutions to end-of-chapter

problems) in one convenient volume. thoroughly class-tested, the Study

Guide includes for each chapter:

▶ Major Concepts: a roadmap through the chapter

▶ What to review: questions that recap key points from previous chapters

▶ discussion Questions: provided for each section; designed for individual review, study groups, or classroom discussion

▶ a Self-test: “do you know the terms?”; crossword puzzles; multiple-choice, fact-driven questions; and questions that ask students to apply their new knowledge in new directions—plus answers!

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