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biochemi str y
seventh edition
Reginald H. Garrett | Charles M. Grisham

University of Virginia
With molecular graphic images

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Reginald H. Garrett, Charles M. Grisham
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About the Authors
Reginald H. Garrett was educated in the Baltimore city public Charles M. Grisham was born and raised in Minneapolis,
schools and at the Johns Hopkins University, where he earned his Minnesota, and educated at Benilde High School. He received
Ph.D. in biology in 1968. Since then, he has been at the University his B.S. in chemistry from the Illinois Institute of Technology in
of Virginia, where he is a Professor Emeritus of Biology. He is the 1969 and his Ph.D. in chemistry from the University of Minne-
author of previous editions of Biochemistry as well as Principles sota in 1973. Following a postdoctoral appointment at the Insti-
of Biochemistry (Cengage, Brooks/Cole) and numerous papers tute for Cancer Research in Philadelphia, he joined the faculty
and review articles on the biochemical, genetic, and molecular of the University of Virginia, where he is Professor of Chemis-
biological aspects of inorganic nitrogen metabolism. His research try. He is the author of previous editions of Biochemistry and
interests focused on the pathway of nitrate assimilation in Principles of Biochemistry (Cengage, Brooks/Cole) and numer-
filamentous fungi. His investigations contributed substantially ous papers and review articles on active transport of sodium,
to our understanding of the enzymology, genetics, and regula- potassium, and calcium in mammalian systems, on protein
tion of this major pathway of biological nitrogen acquisition. kinase C, and on the applications of NMR and EPR spectros-
More recently, he has collaborated in systems approaches to the copy to the study of biological systems. He also authored Inter-
metabolic basis of nutrition-related diseases. His research active Biochemistry CD-ROM and Workbook, a tutorial CD for
has been supported by the National Institutes of Health, the students. His work has been supported by the National Insti-
National Science Foundation, and private industry. He is a tutes of Health, the National Science Foundation, the Muscular
former Fulbright Scholar at the Universität für Bodenkultur in Dystrophy Association of America, the Research Corporation,
Vienna, Austria and served as Visiting Scholar at the University the American Heart Association, and the American Chemical
of Cambridge on two separate occasions. During the second, he Society. He is a Research Career Development Awardee of the
was the Thomas Jefferson Visiting Fellow in Downing College. National Institutes of Health, and in 1983 and 1984 he was a
In 2003, he was Professeur Invité at the Université Paul Sabatier/ Visiting Scientist at the Aarhus University Institute of Physiol-
Toulouse III and the Centre National de la Recherche ogy Denmark. In 1999, he was Knapp Professor of Chemistry at
Scientifique, Institute for Pharmacology and Structural Biology the University of San Diego. He has taught biochemistry, intro-
in France. He taught biochemistry at the University of Virginia ductory chemistry, and physical chemistry at the University of
for 46 years. He is a member of the American Society for Virginia for 48 years. He is a member of the American Society
Biochemistry and Molecular Biology. for Biochemistry and Molecular Biology.
Gene Runion
Reginald H. Garrett and Charles M. Grisham

iv
Contents in Brief
Part I Molecular Components of Cells 1

1 The Facts of Life: Chemistry Is the Logic of Biological Phenomena 1
2 Water: The Medium of Life 29
3 Thermodynamics of Biological Systems 51
4 Amino Acids and the Peptide Bond 77
5 Proteins: Their Primary Structure and Biological Functions 105
6 Proteins: Secondary, Tertiary, and Quaternary Structure 145
7 Carbohydrates and the Glycoconjugates of Cell Surfaces 193
8 Lipids 237
9 Membranes and Membrane Transport 265
10 Nucleotides and Nucleic Acids 317
11 Structure of Nucleic Acids 345
12 Recombinant DNA, Cloning, Gene Editing, and Synthetic Biology—An Introduction 393
Part II Protein Dynamics 433

13 Enzymes—Kinetics and Specificity 433
14 Mechanisms of Enzyme Action 475
15 Enzyme Regulation 511
16 Molecular Motors 543
Part III Metabolism and Its Regulation 573

17 Metabolism: An Overview 573
18 Glycolysis 603
19 The Tricarboxylic Acid Cycle 635
20 Electron Transport and Oxidative Phosphorylation 671
21 Photosynthesis 713
22 Gluconeogenesis, Glycogen Metabolism, and the Pentose Phosphate Pathway 751
23 Fatty Acid Catabolism 791
24 Lipid Biosynthesis 821
25 Nitrogen Acquisition and Amino Acid Metabolism 871
26 Synthesis and Degradation of Nucleotides 921
27 Metabolic Integration and Organ Specialization 951
Part IV Information Transfer 981

28 DNA Metabolism: Replication, Recombination, and Repair 981
29 Transcription and the Regulation of Gene Expression 1033
30 Protein Synthesis 1089
31 Completing the Protein Life Cycle: Folding, Processing, and Degradation 1131
32 The Reception and Transmission of Extracellular Information 1161
Abbreviated Answers to Problems A-1

Index I-1
v
Detailed Contents
Part I Molecular Components of Cells 1 eak Forces Restrict Organisms to a Narrow Range
W
of Environmental Conditions 16
1
Enzymes Catalyze Metabolic Reactions 17
Biomolecular Phenomena across the Time Scale of Life 17
1.5 What Are the Organization and Structure of
The Facts of Life: Chemistry Is the Logic of Cells? 17
The Eukaryotic Cell Likely Emerged from an Archaeal
Biological Phenomena 1 Lineage 18
1.1 What Are the Distinctive Properties of Living How Many Genes Does a Cell Need? 18
Systems? 1 Critical Developments in Biochemistry: Synthetic Life 19
1.2 What Kinds of Molecules Are Biomolecules? 4 Archaea and Bacteria Have a Relatively Simple
Biomolecules Are Carbon Compounds 5 Structural Organization 20
1.3 What Is the Structural Organization of Complex The Structural Organization of Eukaryotic Cells Is More
Biomolecules? 5 Complex Than That of Prokaryotic Cells 21
etabolites Are Used to Form the Building Blocks
M 1.6 What Are Viruses? 22
of Macromolecules 5 Foundational Biochemistry 25
Organelles Represent a Higher Order in Biomolecular Problems 25
Organization 8
Further Reading 27
Membranes Are Supramolecular Assemblies That Define
2
the Boundaries of Cells 9
The Unit of Life Is the Cell 9
1.4 How Do the Properties of Biomolecules Reflect Their
Fitness to the Living Condition? 10
Water: The Medium of Life 29
iological Macromolecules and Their Building Blocks
B
Have a “Sense” or Directionality 10 2.1 What Are the Properties of Water? 29
Biological Macromolecules Are Informational 11 Water Has Unusual Properties 30
Biomolecules Have Characteristic Hydrogen Bonding in Water Is Key to Its Properties 30
Three-Dimensional Architecture 12 The Structure of Ice Is Based on H-Bond
Weak Forces Maintain Biological Structure and Formation 30
Determine Biomolecular Interactions 12 Molecular Interactions in Liquid Water Are Based on
Van der Waals Forces Are Short-Range Interactions H Bonds 31
between Atoms in Nearby Molecules 12 The Solvent Properties of Water Derive from Its Polar
Hydrogen Bonds Form between H Atoms and Nature 32
Electronegative Atoms Like N and O 14 Water Can Ionize to Form H1 and OH2 36
Ionic Interactions Form between Oppositely Charged 2.2 What Is pH? 37
Groups or Molecules 14
Strong Electrolytes Dissociate Completely in Water 38
Hydrophobic Interactions Are the Result of Favorable
Weak Electrolytes Are Substances That Dissociate
Interactions between Water Molecules 14
Only Slightly in Water 39
The Defining Concept of Biochemistry Is
The Henderson–Hasselbalch Equation Describes the
“Molecular Recognition through Structural
Dissociation of a Weak Acid in the Presence of Its
Complementarity” 14
Conjugate Base 39
Biomolecular Recognition Is Mediated by Weak Chemical
Forces 15
vi
Detailed Contents vii
itration Curves Illustrate the Progressive

T tandard Reduction Potentials Are Measured
S
Dissociation of a Weak Acid 41 in Reaction Half-Cells 66
Phosphoric Acid Has Three Dissociable H1 42 %o9 Values Can Be Used to Predict the Direction
2.3 What Are Buffers, and What Do They Do? 43 of Redox Reactions 67
he Phosphate Buffer System Is a Major Intracellular
T %o9 Values Can Be Used to Analyze Energy Changes
Buffering System 43 in Redox Reactions 67
The Imidazole Group of Histidine Also Serves The Reduction Potential Depends on Concentration 69
as an Intracellular Buffering System 44 3.7 Why Are Coupled Processes Important to Living
The Bicarbonate Buffer System of Blood Plasma 44 Things? 69
Buffers Useful within Physiological pH Ranges 45 A Deeper Look: Coupling a Reaction to ATP Hydrolysis
Human Biochemistry: Blood pH and Respiration 46 Changes Its Keq by a Factor of 108 71
2.4 What Properties of Water Give It a Unique 3.8 What Is the Daily Human Requirement for ATP? 71
Role in the Environment? 46 Foundational Biochemistry 72
Foundational Biochemistry 47 Problems 73
Problems 47 Further Reading 75
Further Reading 49
3 4
Amino Acids and the Peptide Bond 77
Thermodynamics of Biological Systems 51 4.1 What Are the Structures and Properties of Amino Acids? 77
3.1 What Are the Basic Concepts of Thermodynamics? 51 Typical Amino Acids Contain a Central Tetrahedral
hree Quantities Describe the Energetics
T Carbon Atom 77
of Biochemical Reactions 52 Amino Acids Can Join via Peptide Bonds 78
All Reactions and Processes Follow There Are 20 Common Amino Acids 78
the Laws of Thermodynamics 53 Are There Other Ways to Classify Amino Acids? 82
A Deeper Look: Entropy, Information, and the A Deeper Look: Why Nature Chose Selenium 82
Importance of “Negentropy” 54 Amino Acids 21 and 22—and More? 83
Free Energy Provides a Simple Criterion for Equilibrium 54 Several Amino Acids Occur Only Rarely in Proteins 83
3.2 What Is the Effect of Concentration on Net 4.2 What Are the Acid–Base Properties of Amino Acids? 83
Free Energy Changes? 55 Amino Acids Are Weak Polyprotic Acids 83
3.3 How May Standard State, Equilibrium, and Cellular Critical Developments in Biochemistry: Adding New
Conditions Be Compared? 56 Chemistry to Proteins with Unnatural Amino Acids 84
3.4 What Are the Characteristics of High-Energy Side Chains of Amino Acids Undergo Characteristic
Biomolecules? 57 Ionizations 86
High Energy Phosphate Compounds 58 4.3 What Reactions Do Amino Acids Undergo? 89
A Deeper Look: Why Nature Chose Phosphates 59 4.4 What Are the Optical and Stereochemical Properties
ATP Is an Intermediate Energy-Shuttle Molecule 59 of Amino Acids? 89
The Hydrolysis of Phosphoric Acid Anhydrides Amino Acids Are Chiral Molecules 89
Is Highly Favorable 60 Chiral Molecules Are Described by the d,l and (R,S)
The Hydrolysis DG° of ATP and ADP Is Greater Than Naming Conventions 90
That of AMP 61 Critical Developments in Biochemistry: Green
Acetyl Phosphate and 1,3-Bisphosphoglycerate Fluorescent Protein—The “Light Fantastic”—from
Are Phosphoric-Carboxylic Anhydrides 61 Jellyfish to Gene Expression 91
Enol Phosphates Are Potent Phosphorylating Agents 62 4.5 What Are the Spectroscopic Properties of Amino
The Complex Equilibria Involved in ATP Hydrolysis 64 Acids? 91
3.5 How Do Group Transfer Potentials Quantify the henylalanine, Tyrosine, and Tryptophan
P
Reactivity of Functional Groups? 65 Absorb Ultraviolet Light 92
3.6 What Are Reduction Potentials, and How Are They Amino Acids Can Be Characterized by Nuclear Magnetic
Resonance 92
Used to Account for Free Energy Changes in Redox
Reactions? 66 Critical Developments in Biochemistry: Rules for
Description of Chiral Centers in the (R,S) System 93
viii Detailed Contents
4.6 How Are Amino Acid Mixtures Step 6. Reconstruction of the Overall Amino Acid
Separated and Analyzed? 94 Sequence 120
Amino Acids Can Be Separated by Chromatography 94 The Amino Acid Sequence of a Protein Can Be
Determined by Mass Spectrometry 122
4.7 What Is the Fundamental Structural
Pattern in Proteins? 94 Sequence Databases Contain the Amino Acid Sequences
of Millions of Different Proteins 125
The Peptide Bond Has Partial Double-Bond Character 95
The Polypeptide Backbone Is Relatively Polar 97 5.4 What Is the Nature of Amino Acid Sequences? 126
Peptides Can Be Classified According to How Many omologous Proteins from Different Organisms Have
H
Amino Acids They Contain 97 Homologous Amino Acid Sequences 127
The Amino Acids in Polypeptides Can Be Released Computer Programs Can Align Sequences and Discover
by Acid Hydrolysis 98 Homology between Proteins 127
Proteins Are Composed of One or More Polypeptide Related Proteins Share a Common Evolutionary
Chains 98 Origin 130
Apparently Different Proteins May Share a Common
Foundational Biochemistry 100
Ancestry 131
Problems 101
A Mutant Protein Is a Protein with a Slightly Different
Further Reading 103 Amino Acid Sequence 131
5.5 How Are Polypeptides Synthesized
5 in the Laboratory? 131

Orthogonal Synthesis of Peptides 132
Solid-Phase Methods Are Very Useful in Peptide
Proteins: Their Primary Structure and Biological Synthesis 132
Functions 105 5.6 Do Proteins Have Chemical Groups Other Than
5.1 What Architectural Arrangements Amino Acids? 132
Characterize Protein Structure? 105 5.7 What Are the Many Biological Functions of
roteins Fall into Three Basic Classes According to Shape
P Proteins? 134
and Solubility 106 Proteins Are the Agents of Biological Function 135
Protein Structure Is Described in Terms of Four Levels of All Proteins Function through Specific Recognition and
Organization 107 Binding of Some Target Molecule 135
Noncovalent Forces Drive Formation of the Higher 5.8 What Is the Proteome and What Does It Tell Us? 137
Orders of Protein Structure 109
The Proteome Is Large and Dynamic 137
A Protein’s Conformation Can Be Described as Its Overall
Determining the Proteome of a Cell 137
Three-Dimensional Structure 109
Critical Developments in Biochemistry: Two New Suffixes
5.2 How Are Proteins Isolated and Purified from in Molecular Biology and Biochemistry: “-ome” and
Cells? 110 “-omics” 138
Number of Protein Separation Methods Exploit
A A Deeper Look: The Human Proteome Project and the
Differences in Size and Charge 110 Human Protein Atlas 139
A Deeper Look: Estimation of Protein Concentrations in Foundational Biochemistry 139
Solutions of Biological Origin 111
Problems 140
A Typical Protein Purification Scheme Uses a Series of
Separation Methods 111 Further Reading 142
A Deeper Look: Techniques Used in Protein
5.3
Purification 112
How Is the Primary Structure of a Protein
Determined? 115
6
Sanger Was the First to Determine the Sequence of a Proteins: Secondary, Tertiary, and Quaternary
Protein 116 Structure 145
Both Chemical and Enzymatic Methodologies Are Used 6.1 What Noncovalent Interactions Stabilize the Higher
in Protein Sequencing 116
Levels of Protein Structure? 146
Step 1. Separation of Polypeptide Chains 116
Hydrogen Bonds Are Formed Whenever Possible 146
A Deeper Look: The Virtually Limitless Number of
Hydrophobic Interactions Drive Protein Folding 146
Different Amino Acid Sequences 117
Ionic Interactions Usually Occur on the Protein
Step 2. Cleavage of Disulfide Bridges 117
Surface 147
Step 3. N- and C-Terminal Analysis 117
Van der Waals Interactions Are Ubiquitous 147
Steps 4 and 5. Fragmentation of the Polypeptide Chain 119
Detailed Contents ix
6.2 What Role Does the Amino Acid Sequence Play rtificial Intelligence Algorithms Predict Protein
A
in Protein Structure? 147 Structures Accurately 180
6.3 What Are the Elements of Secondary Structure in Critical Developments in Biochemistry: NMR and
Proteins, and How Are They Formed? 148 Cryo-EM — Revolutionary Methods that Probe Protein
Structure and Dynamics 180
he Amide Plane Is a Fundamental Determinant of
T
All Protein Structure 148 6.5 How Do Protein Subunits Interact at the Quaternary
The Alpha-Helix Is a Key Secondary Structure 150 Level of Protein Structure? 181
A Deeper Look: Knowing What the Right Hand and Left Human Biochemistry: Diseases of Protein Folding 182
Hand Are Doing 150 Human Biochemistry: Structural Genomics 183
Critical Developments in Biochemistry: X-ray There Is Symmetry in Quaternary Structures 183
Crystallography of Proteins 151 Quaternary Association Is Driven by Weak Forces 183
The b-Pleated Sheet Is a Core Structure in Proteins 153 Open Quaternary Structures Can Polymerize 185
Critical Developments in Biochemistry: In Bed with a There Are Structural and Functional Advantages to
Cold, Pauling Stumbles onto the α-Helix and a Nobel Quaternary Association 185
Prize 154 Human Biochemistry: Faster-Acting Insulin: Genetic
Helix–Sheet Composites Provide Flexibility and Engineering Solves a Quaternary Structure Problem 186
Strength in Spider Silk 156 Foundational Biochemistry 188
b-Turns Allow the Protein Strand to Change Direction 156
Problems 188
6.4 How Do Polypeptides Fold into Three-Dimensional Further Reading 190
Protein Structures? 157
7
A Deeper Look: The Coiled-Coil Motif in Proteins 158
Fibrous Proteins Usually Play a Structural Role 159
Globular Proteins Mediate Cellular Function 162
Helices and Sheets Make Up the Core of Most Globular Carbohydrates and the Glycoconjugates of Cell
Proteins 162
Critical Developments in Biochemistry: The Protein Data
Surfaces 193
Bank (PDB) 162 7.1 How Are Carbohydrates Named? 194
Water Molecules on the Protein Surface Stabilize the 7.2 What Are the Structure and Chemistry
Structure 163 of Monosaccharides? 194
Packing Considerations 163 Monosaccharides Are Classified as Aldoses and
Human Biochemistry: Collagen-Related Diseases 164 Ketoses 194
Protein Domains Are Nature’s Modular Strategy for Stereochemistry Is a Prominent Feature of
Protein Design 165 Monosaccharides 195
Denaturation Leads to Loss of Protein Structure and Monosaccharides Exist in Cyclic and Anomeric Forms 197
Function 165 Haworth Projections Are a Convenient Device for
Human Biochemistry: Domain-Based Engineering Drawing Sugars 198
of Proteins Forms the Basis of a Novel Cancer Monosaccharides Can Be Converted to Several Derivative
Treatment 167 Forms 200
Anfinsen’s Classic Experiment Proved That Sequence A Deeper Look: Honey—An Ancestral Carbohydrate
Determines Structure 169 Treat 202
Is There a Single Mechanism for Protein Folding? 169
7.3 What Are the Structure and Chemistry
What Is the Thermodynamic Driving Force for Protein of Oligosaccharides? 204
Folding? 172
Disaccharides Are the Simplest Oligosaccharides 204
Marginal Stability of the Tertiary Structure Makes Proteins
A Deeper Look: Trehalose—A Natural Protectant for
Flexible 172
Bugs 205
Motion in Globular Proteins 172
A Variety of Higher Oligosaccharides Occur in
The Folding Tendencies and Patterns of Globular Nature 206
Proteins 173
Human Biochemistry: Alpha-Gal and Red Meat
A Deeper Look: Metamorphic Proteins—A Consequence Allergy 207
of Dynamism and Marginal Stability 175
7.4 What Are the Structure and Chemistry
Proteins Can Be Classified on the Basis of Folding
Patterns 176 of Polysaccharides? 208
Molecular Chaperones Are Proteins That Help Other omenclature for Polysaccharides Is Based on Their
N
Proteins to Fold 177 Composition and Structure 208
Some Proteins Are Intrinsically Unstructured 178 Polysaccharides Serve Energy Storage, Structure, and
Protection Functions 209
x Detailed Contents
Polysaccharides Provide Stores of Energy 209 8.3 What Are the Structures and Chemistry
Polysaccharides Provide Physical Structure and Strength of Glycerophospholipids? 242
to Organisms 210 Glycerophospholipids Are the Most Common
A Deeper Look: Ruth Benerito and Wrinkle-Free Cotton Phospholipids 243
Fabrics 213 Ether Glycerophospholipids Include PAF and
Polysaccharides Provide Strength and Rigidity to Bacterial Plasmalogens 245
Cell Walls 214 Human Biochemistry: Platelet-Activating Factor: A
A Deeper Look: A Complex Polysaccharide in Red Potent Ether Glycerophospholipid Mediator 246
Wine—The Strange Story of Rhamnogalacturonan II 215 8.4 What Are Sphingolipids, and How Are They Important
Animals Display a Variety of Cell Surface for Higher Animals? 246
Polysaccharides 218
8.5 What Are Waxes, and How Are They Used? 246
7.5 What Are Glycoproteins, and How Do They Function A Deeper Look: Novel Lipids with Valuable
in Cells? 218 Properties 248
Carbohydrates on Proteins Can Be O-Linked or
8.6 What Are Terpenes, and What Is Their Relevance
N-Linked 218
to Biological Systems? 249
O-GlcNAc Signaling Is Altered in Diabetes and
A Deeper Look: Why Do Plants Emit Isoprene? 251
Cancer 220
Human Biochemistry: Coumadin or Warfarin—Agent of
O-Linked Saccharides Form Rigid Extended Extracellular
Life or Death 251
Structures 220
Polar Fish Depend on Antifreeze Glycoproteins 220 8.7 What Are Steroids, and What Are Their Cellular
N-Linked Oligosaccharides Can Affect the Physical Functions? 252
Properties and Functions of a Protein 221 Cholesterol 252
Human Biochemistry: Drug Research Finds a Sweet Steroid Hormones Are Derived from Cholesterol 252
Spot 222 Human Biochemistry: Plant Sterols and Stanols—
Sialic Acid Terminates the Oligosaccharides of Natural Cholesterol Fighters 254
Glycoproteins and Glycolipids 222 8.8 How Do Lipid Metabolites Act
Sialic Acid Cleavage Can Serve as a Timing Device for as Biological Signals? 254
Protein Degradation 223 Human Biochemistry: Beating COVID-19: Lipid
7.6 How Do Proteoglycans Modulate Processes Nanoparticles Make RNA Vaccines Practical and
in Cells and Organisms? 224 Effective 255
Functions of Proteoglycans Involve Binding to Other Human Biochemistry: The Endocannabinoid
Proteins 225 Signaling System: A Target for Next-Generation
Proteoglycans May Modulate Cell Growth Processes 226 Therapeutics 256
Proteoglycans Make Cartilage Flexible and Resilient 226 Human Biochemistry: Fingolimod—a Sphingosine-1-P
Mimic Is an Oral Drug for Multiple Sclerosis 258
7.7 How Do Carbohydrates Provide a Structural
Code? 226 8.9 What Can Lipidomics Tell Us about Cell, Tissue,
and Organ Physiology? 259
Lectins Translate the Sugar Code 228
Selectins, Rolling Leukocytes, and the Inflammatory
Response 228 Problems 261
Galectins—Mediators of Inflammation, Immunity, and Further Reading 263
Cancer 229
Problems 231
Further Reading 234
9
Membranes and Membrane Transport 265
8 9.1 What Are the Chemical and Physical

Properties of Membranes? 266
The Composition of Membranes Suits Their
Lipids 237 Functions 266
Lipids Form Ordered Structures Spontaneously in
8.1 What Are the Structures and Chemistry of Fatty
Water 267
Acids? 237
The Fluid Mosaic Model Describes Membrane
8.2 What Are the Structures and Chemistry Dynamics 269
of Triacylglycerols? 241
9.2 What Are the Structure and Chemistry of Membrane
A Deeper Look: Polar Bears Prefer Nonpolar Food 242
Proteins? 271
Detailed Contents xi
eripheral Membrane Proteins Associate Loosely

P 10.2 What Are Nucleosides? 319
with the Membrane 271 Human Biochemistry: Adenosine: A Nucleoside
Integral Membrane Proteins Are Firmly Anchored with Physiological Activity 320
in the Membrane 271 10.3 What Are the Structure and Chemistry
Lipid-Anchored Membrane Proteins Can Act as Switching of Nucleotides? 320
Devices 280
Cyclic Nucleotides Are Cyclic Phosphodiesters 322
9.3 How Are Biological Membranes Organized? 282 Nucleoside Diphosphates and Triphosphates Are
9.4 What Are the Dynamic Processes Nucleotides with Two or Three Phosphate Groups 322
That Modulate Membrane Function? 283 Human Biochemistry: cGAMP, a Cyclic Dinucleotide
L ipids and Proteins Undergo a Variety That Triggers a Response to Infection 322
of Movements in Membranes 283 NDPs and NTPs Are Polyprotic Acids 323
Membrane Lipids Can Be Ordered to Different Nucleoside 59-Triphosphates Are Carriers of Chemical
Extents 284 Energy 323
9.5 How Does Transport Occur across Biological 10.4 What Are Nucleic Acids? 324
Membranes? 293 The Base Sequence of a Nucleic Acid Is Its Defining
9.6 What Is Passive Diffusion? 294 Characteristic 325
9.7 How Does Facilitated Diffusion Occur? 294 10.5 What Are the Different Classes of Nucleic
Membrane Channel Proteins Facilitate Diffusion 295 Acids? 326
The Bacillus cereus NaK Channel Uses a Variation The Fundamental Structure of DNA Is a Double
on the K1 Selectivity Filter 297 Helix 326
CorA Is a Pentameric Mg21 Channel 298 Various Forms of RNA Serve Different Roles in
Cells 330
Chloride, Water, Glycerol, and Ammonia
Flow through Single-Subunit Pores 299 A Deeper Look: The RNA World and Early
Evolution 334
9.8 How Does Energy Input Drive Active Transport
The Chemical Differences between DNA and RNA Have
Processes? 300 Biological Significance 334
ll Active Transport Systems Are Energy-Coupling
A
Devices 300
10.6 Are Nucleic Acids Susceptible to Hydrolysis? 335
RNA Is Susceptible to Hydrolysis by Base, but DNA
Many Active Transport Processes Are Driven by
Is Not 335
ATP 301
The Enzymes That Hydrolyze Nucleic Acids Are
A Deeper Look: Cardiac Glycosides: Potent Drugs from
Phosphodiesterases 335
Ancient Times 305
Nucleases Differ in Their Specificity for Different Forms
ABC Transporters Use ATP to Drive Import and Export
of Nucleic Acid 335
Functions and Provide Multidrug Resistance 306
Restriction Enzymes Are Nucleases That Cleave
9.9 How Are Certain Transport Processes Driven by Light Double-Stranded DNA Molecules 337
Energy? 308
Type II Restriction Endonucleases Are Useful
9.10 How Is Secondary Active Transport Driven by Ion for Manipulating DNA in the Lab 337
Gradients? 310 Restriction Endonucleases Can Be Used to Map
Na1 and H1 Drive Secondary Active Transport 310 the Structure of a DNA Fragment 339
AcrB Is a Secondary Active Transport System 310 Foundational Biochemistry 341
Foundational Biochemistry 311 Problems 341
Problems 312 Further Reading 343
Further Reading 314
10 11
Structure of Nucleic Acids 345
Nucleotides and Nucleic Acids 317
11.1 How Do Scientists Determine the Primary Structure
10.1 What Are the Structure and Chemistry of Nucleic Acids? 345
of Nitrogenous Bases? 318 he Nucleotide Sequence of DNA Can Be Determined
T
Three Pyrimidines and Two Purines Are Commonly from the Electrophoretic Migration of a Defined Set of
Found in Cells 318 Polynucleotide Fragments 346
The Properties of Pyrimidines and Purines Can Be Traced Sanger’s Chain Termination or Dideoxy Method Uses
to Their Electron-Rich Nature 319 DNA Replication to Generate a Defined Set of
Polynucleotide Fragments 346
xii Detailed Contents
ext-Generation Sequencing by Synthesis 348

N MC Proteins Establish Chromosome Organization and
S
High-Throughput DNA Sequencing Based on Proton Mediate Chromosome Dynamics 375
Detection 349 11.6 How Can Nucleic Acids Be Synthesized
Single-Molecule Real-Time Sequencing 351 Chemically? 375
Human Biochemistry: The Human Genome hosphoramidite Chemistry Is Used to Form
P
Project 352 Oligonucleotides from Nucleotides 376
Oxford Nanopore Technologies (ONT) Real-Time Genes and Even Chromosomes Can Be Synthesized
Sequencing 352 Chemically 376
Bioinformatics—Functional Genomics, Proteomics, and 11.7 What Are the Secondary and Tertiary
More 353 Structures of RNA? 378
11.2 What Sorts of Secondary Structures Can ransfer RNA Adopts Higher-Order Structure Through
T
Double-Stranded DNA Molecules Adopt? 354 Intrastrand Base Pairing 380
Conformational Variation in Polynucleotide Messenger RNA Adopts Higher-Order Structure Through
Strands 354 Intrastrand Base Pairing 381
DNA Usually Occurs in the Form of Double-Stranded Ribosomal RNA Also Adopts Higher-Order Structure
Helices 355 Through Intrastrand Base Pairing 382
Watson–Crick Base Pairs Have Virtually Identical Aptamers Are Oligonucleotides Specifically Selected for
Dimensions 355 Their Ligand-Binding Ability 384
The DNA Double Helix Is a Stable Structure 355 Foundational Biochemistry 386
Double Helical Structures Can Adopt a Number of Stable Problems 387
Conformations 357
Further Reading 389
A-Form DNA Is an Alternative Form of Right-Handed
12
DNA 357
Z-DNA Is a Conformational Variation in the Form of a
Left-Handed Double Helix 358
The Double Helix Is Flexible 361
Human Biochemistry: DNA Methylation, CpG Islands, Recombinant DNA, Cloning, Gene Editing, and
and Epigenetics 362 Synthetic Biology—An Introduction 393
Alternative Hydrogen-Bonding Interactions Give Rise to 12.1 What Is DNA Cloning? 394
Novel DNA Structures: Cruciforms, Triplexes, and
Plasmids Are Very Useful in Cloning Genes 394
Quadruplexes 362
Shuttle Vectors Are Plasmids That Can Propagate in Two
11.3 Can the Secondary Structure of DNA Different Organisms 399
Be Denatured and Renatured? 366 Artificial Chromosomes Can Be Created from
hermal Denaturation of DNA Can Be Observed by
T Recombinant DNA 399
Changes in UV Absorbance 367
12.2 What Is a DNA Library? 400
pH Extremes or Strong H-Bonding Solutes also Denature
Genomic Libraries Are Prepared from the Total DNA in an
DNA Duplexes 367
Organism 400
Single-Stranded DNA Can Renature to Form DNA
Critical Developments in Biochemistry: Combinatorial
Duplexes 367
Libraries 400
A Deeper Look: The Buoyant Density of DNA 368
Libraries Can Be Screened for the Presence of Specific
The Rate of DNA Renaturation Is an Index of DNA Genes 401
Sequence Complexity 368
Probes for Screening Libraries Can Be Prepared
Nucleic Acid Hybridization: Different DNA Strands of in a Variety of Ways 402
Similar Sequence Can Form Hybrid Duplexes 369
PCR Is Used to Clone and Amplify Specific
11.4 How Does DNA Adopt Structures of Genes 402
Higher Complexity? 370 Critical Developments in Biochemistry: Identifying
Supercoils Are One Kind of Structural Complexity in Specific DNA Sequences by Southern Blotting
DNA 370 (Southern Hybridization) 404
Supercoiled DNA Is Defined by Its Linking Number 370 cDNA Libraries Are DNA Libraries Prepared from
Enzymes Can Add or Remove Supercoils 371 mRNA 405
11.5 What Is the Structure of Eukaryotic DNA Microarrays (Gene Chips) Are Arrays of Different
Chromosomes? 372 Oligonucleotides Immobilized on a Chip 407
Nucleosomes Are the Fundamental Structural Unit in 12.3 Can the Genes in Libraries Be Transcribed
Chromatin 373 and Translated? 408
Higher-Order Structural Organization of Chromatin Gives Expression Vectors Are Engineered So That the RNA or
Rise to Chromosomes 374 Protein Products of Cloned Genes Can Be Synthesized 408
Detailed Contents xiii
Reporter Gene Constructs Are Chimeric DNA Molecules Coenzymes and Cofactors Are Nonprotein Components
Composed of Gene Regulatory Sequences Positioned Essential to Enzyme Activity 437
Next to an Easily Expressible Gene Product 411 13.2 How Can the Rate of an Enzyme-Catalyzed Reaction
Specific Protein–Protein Interactions Can Be Identified Be Defined in a Mathematical Way? 438
Using the Yeast Two-Hybrid System 412
Chemical Kinetics Provides a Foundation for Exploring
In Vitro Mutagenesis—Introducing Changes into Enzyme Kinetics 438
Expressed Proteins 413
Bimolecular Reactions Are Reactions Involving Two
12.4 How Is RNA Interference Used to Reveal the Function Reactant Molecules 439
of Genes? 413 Catalysts Lower the Free Energy of Activation for a
12.5 Gene Therapy: Is It Possible for Scientists to Alter or Reaction 439
Rewrite the Genome of an Organism? 414 Decreasing DG‡ Increases Reaction Rate 440
Human Gene Therapy Can Repair Genetic 13.3 What Equations Define the Kinetics of Enzyme‑
Deficiencies 415 Catalyzed Reactions? 441
Viruses as Vectors in Human Gene Therapy 416 The Substrate Binds at the Active Site of an Enzyme 441
Genome Engineering 416 The Michaelis–Menten Equation Is the Fundamental
Genome Editing with CRISPR-Cas9 417 Equation of Enzyme Kinetics 442
A Deeper Look: The Biological Role of CRISPR-Cas Rate Calculations Assume That [ES] Remains Constant
Systems: Adaptive immunity in Bacteria 418 During an Enzymatic Reaction 442
Base Editing with CRISPR-Cas9 420 A Calculations Are Simplified If Velocity Measurements
Prime Editing with CRISPR-Cas9 421 Are Made Immediately after Adding S 442
Gene Silencing with CRISPR-Cas9 421 The Michaelis Constant, Km, Is Defined as
(k21 1 k2)/k1 443
12.6 How Does High-Throughput Technology Allow Global
When [S] 5 Km, v 5 Vmax/2 444
Study of Millions of Genes or Molecules at Once? 421
Plots of v versus [S] Illustrate the Relationships between
High-Throughput Screening 422
Vmax, Km, and Reaction Order 444
High-Throughput RNAi Screening of Mammalian
Turnover Number Defines the Activity of One Enzyme
Genomes 422
Molecule 445
High-Throughput Protein Screening 422
The Ratio, kcat/Km, Defines the Catalytic Efficiency of an
12.7 What Is the Field of Synthetic Biology? 423 Enzyme 445
iGEM and BioBricks (Registry of Standard Biological Linear Plots Can Be Derived from the Michaelis–Menten
Parts) 423 Equation 446
Metabolic Engineering 424 A Deeper Look: An Example of the Effect of Amino Acid
Synthetic Genomes 425 Substitutions on Km and kcat: Wild-Type and Mutant
Foundational Biochemistry 427 Forms of Human Sulfite Oxidase 448
Problems 428 Nonlinear Lineweaver–Burk or Hanes–Woolf Plots Are a
Property of Regulatory Enzymes 448
Further Reading 429
Enzymatic Activity Is Strongly Influenced by pH 448
The Response of Enzymatic Activity to Temperature Is
Complex 448
Part II Protein Dynamics 433
13.4 What Can Be Learned from the Inhibition of Enzyme
13
Activity? 449
Enzymes May Be Inhibited Reversibly or Irreversibly 449
Reversible Inhibitors May Bind at the Active Site or at
Some Other Site 450
Enzymes—Kinetics and Specificity 433
A Deeper Look: The Equations of Competitive
Enzymes Are the Agents of Metabolic Function 434 Inhibition 451
13.1 What Characteristic Features Define Enzymes? 434 Enzymes Also Can Be Inhibited in an Irreversible
Catalytic Power Is Defined as the Ratio of the Enzyme- Manner 453
Catalyzed Rate of a Reaction to the Uncatalyzed Rate 435 13.5 What Is the Kinetic Behavior of Enzymes Catalyzing
Specificity Is the Term Used to Define the Selectivity of Bimolecular Reactions? 455
Enzymes for Their Substrates 435
Human Biochemistry: Viagra—An Unexpected Outcome
Regulation of Enzyme Activity Ensures That the Rate of in a Program of Drug Design 456
Metabolic Reactions Is Appropriate to Cellular
The Conversion of AEB to PEQ Is the Rate-Limiting Step
Requirements 435
in Random, Single-Displacement Reactions 457
Enzyme Nomenclature Provides a Systematic Way of
In an Ordered, Single-Displacement Reaction, the
Naming Metabolic Reactions 435
Leading Substrate Must Bind First 458
xiv Detailed Contents
Double-Displacement (Ping-Pong) Reactions Proceed Via Noncatalytic Residues May Still Play a Role in Catalysis in
Formation of a Covalently Modified Enzyme Enzyme Active Sites 490
Intermediate 459 14.6 What Can Be Learned from the Mechanisms of Some
Exchange Reactions Are One Way to Diagnose Typical Enzymes? 491
Bisubstrate Mechanisms 461
Serine Proteases 491
Multisubstrate Reactions Can Also Occur in Cells 461
The Digestive Serine Proteases 491
13.6 How Can Enzymes Be So Specific? 461 The Chymotrypsin Mechanism in Detail: Kinetics 493
The Lock-and-Key Hypothesis Was the First Explanation The Serine Protease Mechanism in Detail: Events at the
for Specificity 462 Active Site 493
The Induced-Fit Hypothesis Provides a More Accurate The Aspartic Proteases 496
Description of Specificity 462
The Mechanism of Action of Aspartic Proteases 497
Induced Fit Favors Formation of the Transition State 462
The HIV-1 Protease Is an Aspartic Protease 497
Specificity and Reactivity 462
Critical Developments in Biochemistry: Frances Arnold
13.7 Are All Enzymes Proteins? 463 and Directed Evolution of Enzymes 499
RNA Molecules That Are Catalytic Have Been Termed Chorismate Mutase: A Model for Understanding Catalytic
Ribozymes 463 Power and Efficiency 499
Antibody Molecules Can Have Catalytic Activity 464 Human Biochemistry: Protease Inhibitors-Game
13.8 Is It Possible to Design an Enzyme to Catalyze Any Changers for AIDS Patients 502
Desired Reaction? 466 Foundational Biochemistry 504
A Designer Enzyme for Bioremediation 467 Problems 505
A Case Study for Designer Enzyme Development — Kemp Further Reading 508
Eliminases 467
15
Problems 469
Further Reading 471
Enzyme Regulation 511
14 15.1 What Factors Influence Enzymatic Activity? 511

he Availability of Substrates and Cofactors Usually
T
Determines How Fast the Reaction Goes 512
Mechanisms of Enzyme Action 475 As Product Accumulates, the Apparent Rate
14.1 What Are the Magnitudes of Enzyme-Induced Rate of the Enzymatic Reaction Will Decrease 512
Accelerations? 475 Genetic Regulation of Enzyme Synthesis Determines
the Amount of Enzyme Present at Any Moment 512
14.2 What Role Does Transition-State Stabilization Play in
Enzyme Activity Can Be Regulated Allosterically 512
Enzyme Catalysis? 477
Enzyme Activity Can Be Regulated through Covalent
14.3 How Does Destabilization of the Enzyme-Substrate Modification 513
Complex Affect Enzyme Catalysis? 478 Zymogens Are Inactive Precursors of Enzymes 513
14.4 What Are Transition State Analogs and How Tightly Isozymes Are Enzymes with Slightly Different Subunits
Do They Bind to the Active Sites of Enzymes? 479 and Kinetic Properties 514
A Deeper Look: Transition-State Analogs Make Our 15.2 What Are the General Features of Allosteric
World Better 480 Regulation? 516
14.5 What Are the Mechanisms of Enzyme Catalysis? 483 egulatory Enzymes Have Certain Exceptional
R
Enzymes Facilitate Formation of Near-Attack Properties 516
Conformations 483 15.3 Can Allosteric Regulation Be Explained by Conforma-
A Deeper Look: How to Read and Write Mechanisms 484 tional Changes in Proteins? 517
Protein Motions Are Essential to Enzyme Catalysis 484 he Symmetry Model for Allosteric Regulation Is Based
T
Covalent Catalysis 486 on Two Conformational States for a Protein 517
General Acid-Base Catalysis 487 The Sequential Model for Allosteric Regulation Is Based
Low-Barrier Hydrogen Bonds 488 on Ligand-Induced Conformational Changes 517
Quantum Mechanical Tunneling in Electron and Proton The Notable Difference between the MWC and KNF
Transfers 489 Models 519
Human Biochemistry: Antibiotic Resistance by Changes in the Oligomeric State of a Protein
Superbugs 489 Can Also Give Allosteric Behavior 519
Metal Ion Catalysis 490
Detailed Contents xv
15.4 What Kinds of Covalent Modification Regulate Human Biochemistry: Hemoglobin and Nitric Oxide 536
the Activity of Enzymes? 519 Sickle-Cell Disease Is Characterized by Abnormal Red
ovalent Modification through Reversible
C Blood Cells 537
Phosphorylation 519 Sickle-Cell Disease Has Been Characterized as a
Protein Kinases: Target Recognition and Intrasteric Molecular Disease 537
Control 519 Foundational Biochemistry 538
Phosphorylation Is Not the Only Form of Covalent Problems 539
Modification That Regulates Enzyme Function 522
Further Reading 541
Acetylation Is a Prominent Modification
16
for the Regulation of Metabolic Enzymes 522
15.5 Is the Activity of Some Enzymes Controlled by Both
Allosteric Regulation and Covalent Modification? 523
he Glycogen Phosphorylase Reaction Converts
T
Glycogen into Readily Usable Fuel in the Form of
Molecular Motors 543
Glucose-1-Phosphate 523 16.1 What Are the General Features of Molecular
Glycogen Phosphorylase Is a Homodimer 523 Motors? 543
Glycogen Phosphorylase Activity Is Regulated 16.2 What Is the Molecular Mechanism
Allosterically 524 of Muscle Contraction? 544
Covalent Modification of Glycogen Phosphorylase uscle Contraction Is Triggered by Ca 21 Release
M
Overrides Allosteric Regulation 526 from Intracellular Stores 544
Enzyme Cascades Regulate Glycogen Phosphorylase Human Biochemistry: Medical Applications of Smooth
Covalent Modification 526 Muscle Effectors 545
15.6 What Is the Relationship between Quaternary Structure The Molecular Structure of Skeletal Muscle
and Allosteric Regulation? 528 Is Based on Actin and Myosin 545
he Comparative Biochemistry of Myoglobin and
T The Mechanism of Muscle Contraction Is Based on
Hemoglobin Reveals Insights into Allostery 528 Sliding Filaments 548
Myoglobin Is an Oxygen-Storage Protein 529 A Deeper Look: The P-Loop NTPases—Energy to Run
O2 Binds to the Mb Heme Group 529 the Motors 549
O2 Binding Alters Mb Conformation 529 16.3 What Are the Molecular Motors That Orchestrate the
A Deeper Look: The Oxygen-Binding Curves of Mechanochemistry of Microtubules? 551
Myoglobin and Hemoglobin 530 ilaments of the Cytoskeleton Are Highways
F
Cooperative Binding of Oxygen by Hemoglobin That Move Cellular Cargo 552
Has Important Physiological Significance 531 Three Classes of Motor Proteins Move Intracellular
Hemoglobin Has an a2b2 Tetrameric Structure 531 Cargo 553
Oxygenation Markedly Alters the Quaternary Dyneins Move Organelles in a Plus-to-Minus Direction;
Structure of Hb 532 Kinesins, in a Minus-to-Plus Direction 554
Movement of the Heme Iron by Less Than 0.04 nm Human Biochemistry: Effectors of Microtubule
Induces the Conformational Change in Hemoglobin 532 Polymerization as Therapeutic Agents 555
The Physiological Significance of the Hb:O2 Cytoskeletal Motors Are Highly Processive 555
Interaction 532 ATP Binding and Hydrolysis Drive Hand-over-Hand
The Oxy and Deoxy Forms of Hemoglobin Represent Movement of Kinesin 557
Two Different Conformational States 533 Conformation Changes in a Hexameric Ring
The Allosteric Behavior of Hemoglobin Has Both Drive the Dynein Motor 558
Symmetry (MWC) Model and Sequential (KNF) 16.4 How Do Molecular Motors Unwind DNA? 558
Model Components 533 egative Cooperativity Facilitates Hand-over-Hand
N
H1 Promotes the Dissociation of Oxygen from Movement 561
Hemoglobin 533 Papillomavirus E1 Helicase Moves along DNA as on a
CO2 Also Promotes the Dissociation of O2 from Spiral Staircase 562
Hemoglobin 534
16.5 How Do Bacterial Flagella Use a Proton Gradient
2,3-Bisphosphoglycerate Is an Important Allosteric
to Drive Rotation? 564
Effector for Hemoglobin 535
The Flagellar Rotor Is a Complex Structure 565
BPG Binding to Hb Has Important Physiological
Significance 535 Gradients of H1 and Na1 Drive Flagellar Rotors 565
Fetal Hemoglobin Has a Higher Affinity for O2 The Flagellar Rotor Self-Assembles in a
Because It Has a Lower Affinity for BPG 535 Spontaneous Process 565
xvi Detailed Contents
lagellar Filaments Are Composed of Protofilaments of

F Mutations Create Specific Metabolic Blocks 588
Flagellin 566 Isotopic Tracers Can Be Used as Metabolic Probes 588
Motor Reversal Involves Conformation Switching NMR Spectroscopy Is a Noninvasive Metabolic Probe 589
of Motor and Filament Proteins 566 Cell Fractionation Reveals Which Compartment Contains
Foundational Biochemistry 568 a Metabolic Pathway 589
Problems 568 Metabolic Pathways Are Also Time-Dependent 591
Further Reading 570 17.5 What Can the Metabolome Tell Us about a Biological
System? 592
Part III Metabolism and Its Regulation 573 17.6 What Food Substances Form the Basis
of Human Nutrition? 596
17
Humans Require Protein 596
Carbohydrates Provide Metabolic Energy 597
Lipids Are Essential, but in Moderation 597
Metabolism: An Overview 573 Foundational Biochemistry 597

Problems 598
17.1 Is Metabolism Similar in Different Organisms? 573
Further Reading 600
Living Things Vary in Their Carbon and Energy
Requirements 574
Oxygen Is Essential to Life for Aerobes 574
The Flow of Energy in the Biosphere and the Carbon and
Oxygen Cycles Are Intimately Related 574
18
A Deeper Look: Calcium Carbonate—A Biological Sink Glycolysis 603
for CO2 575 18.1 What Are the Essential Features of Glycolysis? 603
17.2 What Can Be Learned from Metabolic Maps? 575 18.2 Why Are Coupled Reactions Important in
The Metabolic Map Can Be Viewed as a Set of Dots and Glycolysis? 605
Lines 575
18.3 What Are the Chemical Principles and Features of the
Alternative Metabolic Pathway Models Can Provide New
First Phase of Glycolysis? 606
Insights into Pathways 577
eaction 1: Glucose Is Phosphorylated by Hexokinase
R
Metabolism Is a Network 578
or Glucokinase—The First Priming Reaction 606
Metabolic Pathways with Multiple Enzymes May Take
A Deeper Look: Glucokinase—An Enzyme with Different
Different Forms 579
Roles in Different Cells 609
17.3 How Do Anabolic and Catabolic Processes Form the Reaction 2: Phosphoglucoisomerase Catalyzes
Core of Metabolic Pathways? 580 the Isomerization of Glucose-6-Phosphate 609
Anabolism Is Biosynthesis 580 Reaction 3: ATP Drives a Second Phosphorylation
Anabolism and Catabolism Are Not Mutually Exclusive 580 by Phosphofructokinase—The Second Priming
The Pathways of Catabolism Converge to a Few End Reaction 610
Products 581 Reaction 4: Cleavage by Fructose Bisphosphate
Anabolic Pathways Diverge, Synthesizing an Astounding Aldolase Creates Two 3-Carbon Intermediates 612
Variety of Biomolecules from a Limited Set of Building Reaction 5: Triose Phosphate Isomerase
Blocks 581 Completes the First Phase of Glycolysis 613
Amphibolic Intermediates Play Dual Roles 583 Human Biochemistry: Methylglyoxal and Advanced
Corresponding Pathways of Catabolism and Anabolism Glycation End-Products—The Dark Side of
Differ in Important Ways 583 Glycolysis 614
Metabolic Regulation Requires Different Pathways for 18.4 What Are the Chemical Principles and Features of the
Oppositely Directed Metabolic Sequences 583 Second Phase of Glycolysis? 615
ATP Is Produced and Consumed in a Cellular Energy eaction 6: Glyceraldehyde-3-Phosphate Dehydrogenase
R
Cycle 583 Creates a High-Energy Intermediate 615
NAD1 Collects Electrons Released in Catabolism 584 Reaction 7: Phosphoglycerate Kinase Is the Break-Even
NADPH Provides the Reducing Power for Anabolic Reaction 615
Processes 585 Reaction 8: Phosphoglycerate Mutase Catalyzes a
Coenzymes and Vitamins Provide Unique Chemistry and Phosphoryl Transfer 617
Essential Nutrients to Pathways 586 Reaction 9: Dehydration by Enolase Creates PEP 618
17.4 What Experiments Can Be Used to Elucidate Reaction 10: Pyruvate Kinase Yields More ATP 618
Metabolic Pathways? 587 Human Biochemistry: Pyruvate Kinase M2 Moonlights
nzyme Inhibitors Can Be Used to Reveal Individual
E as a Protein Kinase in Cancer 620
Metabolic Reactions 587
Detailed Contents xvii
18.5 What Are the Metabolic Fates of NADH and Pyruvate 19.5 What Are the Energetic Consequences
Produced in Glycolysis? 621 of the TCA Cycle? 651
Anaerobic Metabolism of Pyruvate Leads to Lactate or he Carbon Atoms of Acetyl-CoA Have Different Fates in
T
Ethanol 621 the TCA Cycle 652
Lactate Accumulates Under Anaerobic Conditions in A Deeper Look: Steric Preferences in NAD1-Dependent
Animal Tissues 622 Dehydrogenases 654
Critical Developments in Biochemistry: The Warburg 19.6 Can the TCA Cycle Provide Intermediates
Effect and Cancer 623 for Biosynthesis? 654
18.6 How Do Cells Regulate Glycolysis? 624 Human Biochemistry: Leber’s Hereditary Optic
18.7 Are Substrates Other Than Glucose Neuropathy Arises from Mitochondrial DNA
Used in Glycolysis? 624 Mutations 655
ructose Catabolism in Liver Is Unregulated—
F 19.7 What Are the Anaplerotic, or Filling Up,
and Potentially Harmful 624 Reactions? 656
Mannose Enters Glycolysis in Two Steps 626 A Deeper Look: Anaplerosis Plays a Critical Role in
Galactose Enters Glycolysis via the Leloir Pathway 626 Insulin Secretion 657
Human Biochemistry: Tumor Diagnosis Using Positron A Deeper Look: Fool’s Gold and the Reductive Citric
Emission Tomography (PET) 626 Acid Cycle—The First Metabolic Pathway? 658
Glycerol Can Also Enter Glycolysis 628 19.8 How Is the TCA Cycle Regulated? 658
18.8 How Do Cells Respond to Hypoxic Stress? 628 yruvate Dehydrogenase Is Regulated
P
by Phosphorylation/Dephosphorylation 659
Human Biochemistry: Lactose and Lactose Intolerance:
From Mother’s Milk to Yogurt 629 Isocitrate Dehydrogenase Is Strongly Regulated 660
Foundational Biochemistry 631 Regulation of TCA Cycle Enzymes by Acetylation 660
Human Biochemistry: TCA Metabolites Play Roles in
Problems 631
Many Pathways via Post-Translational Modifications 661
Further Reading 633
Two Covalent Modifications Regulate E. coli Isocitrate
Dehydrogenase 661
19
TCA Metabolites Can Act as Cellular Signals 662
The TCA Cycle Operates as a Metabolon 662
19.9 Can Any Organisms Use Acetate as Their Sole Carbon
The Tricarboxylic Acid Cycle 635 Source? 662
he Glyoxylate Cycle Operates in Specialized
T
19.1 What Is the Chemical Logic of the TCA Cycle? 636
Organelles 663
19.2 How Is Pyruvate Oxidatively Decarboxylated Isocitrate Lyase Short-Circuits the TCA Cycle
to Acetyl-CoA? 639 by Producing Glyoxylate and Succinate 664
A Deeper Look: The Coenzymes of the Pyruvate The Glyoxylate Cycle Helps Plants Grow in the Dark 665
Dehydrogenase Complex 642
Glyoxysomes Must Borrow Three Reactions from
19.3 How Are Two CO2 Molecules Produced Mitochondria 665
from Acetyl-CoA? 644 Foundational Biochemistry 666
he Citrate Synthase Reaction Initiates the
T Problems 666
TCA Cycle 644
Further Reading 668
Citrate Is Isomerized by Aconitase to Form
Isocitrate 645
Isocitrate Dehydrogenase Catalyzes the First
Oxidative Decarboxylation in the Cycle 647
α-Ketoglutarate Dehydrogenase Catalyzes the Second
20
Oxidative Decarboxylation of the TCA Cycle 648 Electron Transport and Oxidative
19.4 How Is Oxaloacetate Regenerated to Complete Phosphorylation 671
the TCA Cycle? 648 20.1 Where in the Cell Do Electron Transport and Oxidative
uccinyl-CoA Synthetase Catalyzes Substrate-Level
S Phosphorylation Occur? 671
Phosphorylation 649 itochondrial Functions Are Localized in Specific
M
Succinate Dehydrogenase Is FAD-Dependent 650 Compartments 672
Fumarase Catalyzes the Trans-Hydration Human Biochemistry: Mitochondrial Dynamics in
of Fumarate to Form l-Malate 650 Human Diseases 673
Malate Dehydrogenase Completes the Cycle The Mitochondrial Matrix Contains the Enzymes of the
by Oxidizing Malate to Oxaloacetate 651 TCA Cycle 674
xviii Detailed Contents
20.2 How Is the Electron-Transport Chain Organized? 674 20.7 How Do Mitochondria Mediate Apoptosis? 704
The Electron-Transport Chain Can Be Isolated in Four Cytochrome c Triggers Apoptosome Assembly 706
Complexes 675 Foundational Biochemistry 707
Complex I Oxidizes NADH and Reduces Coenzyme Q 676 Problems 707
Human Biochemistry: Solving a Medical Mystery
Further Reading 710
Revolutionized Our Treatment of Parkinson’s Disease 679
21
Complex II Oxidizes Succinate and Reduces Coenzyme
Q 680
Complex III Mediates Electron Transport from Coenzyme
Q to Cytochrome c 681
Complex IV Transfers Electrons from Cytochrome c Photosynthesis 713
to Reduce Oxygen on the Matrix Side 684 21.1 What Are the General Properties of
Proton Transport Across Cytochrome c Oxidase Photosynthesis? 714
Is Coupled to Oxygen Reduction 686 Photosynthesis Occurs in Membranes 714
Reversal of Complex II May Short-Circuit Photosynthesis Consists of Both Light Reactions
the Electron Transfer Pathway 687 and Dark Reactions 715
The Complexes of Electron Transport Water Is the Ultimate Electron Donor
May Function as Supercomplexes 687 for Photosynthetic NADP1 Reduction 716
Electron Transfer Energy Stored in a Proton
21.2 How Is Solar Energy Captured by Chlorophyll? 717
Gradient: The Mitchell Hypothesis 689
hlorophylls and Accessory Light-Harvesting Pigments
C
20.3 What Are the Thermodynamic Implications of Absorb Light of Different Wavelengths 717
Chemiosmotic Coupling? 690 The Light Energy Absorbed by Photosynthetic
20.4 How Does a Proton Gradient Drive Pigments Has Several Possible Fates 718
the Synthesis of ATP? 690 The Transduction of Light Energy into Chemical
ATP Synthase Is Composed of F1 and F0 Complexes 691 Energy Involves Oxidation–Reduction 719
he Catalytic Sites of ATP Synthase Adopt
T Photosynthetic Units Consist of Many Chlorophyll
Three Different Conformations 693 Molecules but Only a Single Reaction Center 720
Boyer’s 18O Exchange Experiment Identified the 21.3 What Kinds of Photosystems Are Used to Capture
Energy-Requiring Step 693 Light Energy? 721
The Binding Change Mechanism Describes hlorophyll Exists in Plant Membranes in Association
C
the Events of Rotational Catalysis 694 with Proteins 721
Proton Flow Through F0 Drives Rotation PSI and PSII Participate in the Overall Process of
of the Motor and Synthesis of ATP 694 Photosynthesis 721
How Many Protons Are Required to Make an ATP? The Pathway of Photosynthetic Electron Transfer Is Called
It Depends on the Organism 696 the Z Scheme 722
The Mitchell Model Has Been Confirmed Oxygen Evolution Requires the Accumulation of Four
in a Reconstitution Experiment 696 Oxidizing Equivalents in PSII 722
Inhibitors of Oxidative Phosphorylation Electrons Are Taken from H2O to Replace Electrons Lost
Reveal Insights About the Mechanism 697 from P680 724
Uncouplers Disrupt the Coupling of Electron Electrons from PSII Are Transferred to PSI via the
Transport and ATP Synthase 698 Cytochrome b6 f Complex 724
ATP–ADP Translocase Mediates the Movement of ATP Plastocyanin Transfers Electrons
and ADP Across the Mitochondrial Membrane 698 from the Cytochrome b6 f Complex to PSI 724
Human Biochemistry: Endogenous Uncouplers—Novel 21.4 What Is the Molecular Architecture of Photosynthetic
Proteins with Many Beneficial Effects 699
Reaction Centers? 725
20.5 What Is the P/O Ratio for Mitochondrial The R. viridis Photosynthetic Reaction Center Is an
Oxidative Phosphorylation? 700 Integral Membrane Protein 725
20.6 How Are the Electrons of Cytosolic NADH Photosynthetic Electron Transfer by the R. viridis
Fed into Electron Transport? 701 Reaction Center Leads to ATP Synthesis 726
Glycerophosphate Shuttle Ensures Efficient Use of
The The Molecular Architecture of PSII Resembles
Cytosolic NADH 701 the R. viridis Reaction Center Architecture 727
The Malate–Aspartate Shuttle Is Reversible 702 How Does PSII Generate O2 from H2O? 729
The Net Yield of ATP from Glucose Oxidation The Molecular Architecture of PSI Resembles the
Depends on the Shuttle Used 703 R. viridis Reaction Center and PSII Architecture 730
3.5 Billion Years of Evolution Have Resulted How Do Green Plants Carry Out Photosynthesis? 731
in a Very Efficient System 704
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Vaunut vierivät rämisten ylös mäkeä Ernstin tuvalle.
Hän ihmetteli, mitä se mahtoi olla, meni etehisen kuistiin ja asetti

kätensä auringon varjoksi.
— Herranen aika, vallesmanni!
Hän kalpeni yht'äkkiä. Hänen ensi ajatuksensa oli: vaimo, lapsi! —

Hän kuuli vaimonsa laulelevan pienokaiselle…
Mutta sitten tyyntyi hän taas heti; ehkei se ollut mitään vaarallista.
Mutta nyt olivat vaunut hänen ovensa edustalla. Eräs talonpoika

istui kyytimiehen vierellä. Nimismies astui alas.
— Hyvää päivää, Ernst! — Se kuului onnettomuutta ennustavalta,

tyyneltä.
— Herra vallesmanni suo minulle sen kunnian.
Ernst kumarsi. Sydän sylkytti.
— Tahtoisin puhua hiukan sinun kanssasi. Ja minulla on todistajat,

kuten näet.
— Todistajat!?
Nyt selveni hänelle kaikki. Mutta hän nieleksi, tyynennäköisenä —

hän ei antaisi itseään ilmi.
— Sinut on ilmiannettu — arveli nimismies.
— Vai niin!
Ernst koetti nauraa.
— Sinä et näy sen siitään paranneen.
Ernst tuli veripunaiseksi.
— Älkää puhuko siitä!
— Minä puhun uudesta. Sinut on ilmiannettu…
— Kenenkä toimesta, jos saan luvan kysyä?
— Sitä minun ei tarvitse sanoa.
— Silloin voin minä sen sanoa. Se ei ole tietysti kukaan muu kuin
tuo, joka hiipii tuolla viidakossa ja tähystelee tänne.
Hän osoitti. Toiset katsoivat osoitettuun suuntaan. Siellä puikki

Hans pensaiden suojaan.
— Tule sinä, Hans — kirkui Ernst — tule katsomaan, mitä olet

aikaan saanut. Mutta puukkosahaa minä en ole ottanut sen sinä
kyllä tiedät.
— Mitä puukkosahaa? — kysyi nimismies. — Tässä ei ole
kysymys mistään puukkosahasta. Mutta sinä olet tehnyt väärää
rahaa.
— Herra jumala!
Ernst koetti käsillään päätään.
Hans oli tullut nyt saapuville. Hän asettui vaunujen taakse. Hän
pelkäsi Ernstin nyrkkejä.
— No, Ernst, nyt tiedät sinä, kuka on ilmiantajasi. Se oli oikein

tehty, sillä minulla on todistus kädessäni.
— Mikä todistus?
Näytettiin kaksikruunuista.
Ernst tuijotti.
— Mistä vallesmanni on tuon saanut, jos saan olla nenäkäs?
— Sen tiedät parhaiden itse.
— Tahtovatko siis kaikki ihmiset tehdä minut onnettomaksi?
Samalla osoitti hän alas tielle.
— Katsokaa, ettekö usko, että hänkin tulee tänne! Niin niin,

udellaan, kun nimismies kulkee pitäjän läpi.
Eriksson läheni, vähän arkaillen. Mutta hän tahtoi nähdä

kohtauksen.
Nimismies seisoi raha kädessä, ja piti sitä ylhäällä auringon
paisteessa.
Tunnustatko väärentäneesi tämän?
— Ernst! — kuului kirahdus puolipimeästä etehisestä.
Kaikki kääntyivät katsomaan. Siellä seisoi vaimo pelästyneenä.

Mies ei ollut huomannut, että tuutilaulu oli vaiennut.
— Tule, Anna-Reetta!
Vaimo juoksi hänen luokseen.
— Sinä et ole tehnyt mitään pahaa — se ei ole totta! Sano, että se

ei ole totta…!
— Kas niin, älä itke. Anna-Reetta!
— Tunnustatko? — kysyi taas nimismies, joka seisoi kuin

kuvapatsas hänen edessään.
— Sano, että se on valhe, Ernst!
Mutta silloin sai hänen äänensä lujuutta.
— Valhe? Miksi valehteleisin minä? Enkö minä ole tehnyt sitä

sinun tähtesi! Minä näin kuinka kalpeana sinä makasit sängyssä
pienokaisen kanssa. Ei ollut palaistakaan ruokaa koko talossa.
Minun täytyi hankkia rahaa. En voinut sallia sinun nälkään kuolevan.
Minä istuin valveilla koko yön. Minun korvissani suhisi ja lauloi koko
ajan kun työskentelin, lauloi sinusta. Luulin tekeväni hyvän työn.
Uskoin, Herramme auttaisi meitä… Sinä olit raukka, Eriksson, kun et
viskannut rahaa vuonoon!
Kauppias tahtoi vastata, mutta oikeuden valvoja sanoi:
— Sinä siis tunnustat?
— Ei, ei — huudahti vaimo.
— Kyllä, Anna-Reetta!
Hans'in kasvot loistivat. Eriksson puri huultaan, todistajat katsoivat

maahan. Nimismies nyökytti päätään ikäänkuin valitellen. Mutta
vaimo heittäytyi rajusti miehensä kaulaan:
— Ernst, Ernst!
Syntyi hetken äänettömyys.
— Mihinkä minut nyt viedään?
— Vankeuteen, tietysti.
— Kuinka käy minun ja lapsen — uikutti vaimo.
— Kysy Hansilta!
Lapsen kirkuna kuului sisältä.
Saanhan käydä häntä katsomassa ennenkuin minut viedään.
— Kyllä, mene!
Mies ja vaimo menivät sisään.
Samassa juoksi muudan poika viidakosta.
— Mitä se on? — kysäsi Hans.

— Puukkosaha! — sanoi poika. — Minulta se hävisi — mutta
sitten — en sitä löytänyt — pelkäsin selkäsaunaa — en uskaltanut
sanoa — mutta sitten sen löysin — karttakopasta.
— Lurjus! — mutisi Hans ja otti työkalun poikansa kädestä. —

Juokse takaisin kotiin! Saat selkääsi, jahka tulen.
Poika alkoi itkeä tillittää ja lähti juoksemaan.
— Näetkös, etten minä ole sitä ottanut — sanoi Ernst.
— Ethän sinäkään voi kaikkea pahaa tehdä.
— Kitas kiinni!
— Jumalan käsi! — aprikoi Hans. — Lapsen kautta keksittiin rikos.
— Tule nyt! kehoitti nimismies.
Niin sai Ernst istua kuskilaudalle, talonpoika istui rattaiden

etupuolelle.
Vaunut vierivät tiehensä. Ernst ei tulisi näkemään tupaansa pitkiin

aikoihin.
Vaimo vaipui portaille, nosti esiliinan silmilleen ja nyyhkytti…
Hans käveli kotiinsa. Hän vihelteli.
Eriksson vaelteli hiljaa puodilleen. Hän ei saanut päästään pois

ajatusta, jonka Ernst oli lausunut:
— Sinä olit raukka, Eriksson, kun et viskannut rahaa vuonoon!

Jos hän olisi tehnyt sen! Nyt ei hän voisi nukkua tänä yönä.
Ehkäpä sitten, jos hän ottaa oikein aimo todin.
Kosinta.
Iltapuolipäivä keskikesällä. Ukko Olli on istuutunut muutamalle

kivelle tupansa edustalla. Hän imee piippunysäänsä ja hymyilee
leveästi. Hymynsä ei ole auringolle, vaan siksi että hän ajattelee: Ei
tahdo saada kuvan ruokaa, kun ei ole naisväkeä talossa.
Hän on äskettäin menettänyt eukkonsa — Jumala hälle anteeksi

antakoon, hän oli häijy. Ja hän asuu nyt yksin poikansa kanssa, joka
parastaikaa puuhailee kuokka kädessä perunamaalla. Olli on
seitsemänkymmentäkolme ja poika viisikymmentä vuotta vanha.
Vanhuksen siinä niin istuessa ja sylkiessä haikujen välillä ja

leskimiehen pulmallista elämää, mietiskellessä asettaa Herramme
asiat niin, että vanha Riitta tulee tietä myöten. Olli varjostaa
kädellään auringonpaistetta ja tuumii itsekseen:
— Kas vaan, hän on liikkeellä.
Hän tuntee Riitan hyvin — hän katseli nuoruudessaan Riittaa

silmällä kuin helmellä, sanotaan. Mutta Riitta meni naimisiin toisen
kanssa ja Olli toisen kanssa, ja niin joutuivat he erilleen. Mutta sitten
kuoli Riitan mies, ja kun Olli jäi leskeksi, ovat nuoruuden ajatukset
hänessä taas valveutuneet. Ja siksi hän nyt niin kirkastuu, kun saa
nähdä Riitan.
Hän pistää sammuneen piipun liivin taskuun, sivasee käden
selkämyställä suutaan ja valmistautuu kohteliaasen vastaanottoon.
Kun Riitta näkee Ollin siinä seisovan noin jotakin tarkoittavana ja

muhoilevana, kirkastuvat hänenkin kasvonsa — näyttää siltä, kuin
näkisi hän mielellään ukon tuollaisena.
Olli nyökyttelee ja tohuilee ja tervehtii:
— Hyvää iltapäivää!
Riitta pysähtyy. Olli näkee rypyt hänen kasvoissaan. Mutta hän ei

välitä vaan sanoo:
— Että hän on lähtenyt liikkeelle. Pitkästä matkasta on hän

varmaan väsynyt, sillä hän tulee kai kotoaan, luulen ma.
— Niin onkin, piti kauppamiehen luo.
— Silloin voi hän hyvin istua ja levätä hiukan. Siitä on jo aikoja kun
olemme viimeksi puhelleet.
— On kyllä, on. — Ja. Riitta istuu kivelle.
— Ho hoi, niin niin, — kiikkuu hän edestakasin — sitä käy jo niin

vanhaksi, että tuskin voi kävellä.
— Niin käy, niin. Mutta miten hän näyttää ripsakalta ja reippaalta!
— Miten hän sanoo! Ja toinen on jo seitsemänkymmentä vuotta!
— Hän tulee kyllä vanhemmaksi, jos hän saa elää.
— Niin, vanhemmaksihan sitä tulee.

Ja sitten he eivät puhu mitään.
Mutta se alkaa käydä Ollista sietämättömäksi. Hän pöyrii

tukkaansa ja katsoo likaisia kynsiään.
— Minä ajattelin sanoa ikään jotakin.
— Ho hoi, niin niin — huokaa Riitta ikäänkuin ei olisi kuunnellut.
— Niin, näes, minä olen nyt yksinäinen, ja se on niin ikävää.
— Hän saa hankkia jonkun, joka hoitaa ja huolehtii talossa.
— Sitäpä minä juuri tässä ajattelin.
— Niin käy se hyvin, niin.
Ja sitten he vaikenevat.
— Näes, minussa on vielä vanhaa rakkautta häneen — alkaa Olli

taas.
— Oh, siitä on niin pitkä aika, siitä.
Riitta nyökyttelee päätään ja vaipuu muistoihinsa.
— Niinpä voisi nyt olla aika, että me tulisimme yhteen, ajattelen

minä.
— Hän ei toki koskaan tarkoita, että me… — Tuo ihmeellinen

ajatus saa
Riitan melkein pelokkaaksi.
-— Kyllä, juuri sitä niinä tarkoitan. Niin kysyn minä nyt kunniani
kautta rehellisesti, tahtooko hän ottaa minut aviomiehekseen myötä-
ja vastoinkäymisessä.
— Oh, herranen aika! — Eukon katse oikein vilkastuu. — Niin, se

riippuu siitä, mitä silloin saa tehdä.
— Hoitaa talouttani — ja poikaa, tietysti.
»Poika» on viisikymmenvuotias ja seisoo hän perunamaalla ja

kuokkii. Kun hän kuulee heidän keskustelunsa, katsahtaa hän ylös,
seisauttaa työnsä ja menee heidän luokseen.
Riitta nauraa:
— Poikaa! Sitä on kuitenkin koko vanha veitikka, tuota Ollia.
— Onko niin, niin pidä kiin'! — sanoo Olli reippaasti.
— Eihän voi muuta, kun kuin pyytää niin kauniisti. Minäkin olen
ajatellut sitä samaa, jos sanon niinkuin asia on.
— Hän on kerrassaan mokoma.
Niin vaikenevat he. Sillä »poika» seisoo nyt heidän edessään. Hän
kysyy eukolta:
— Mitäpä se isä hänelle oikeastaan puhuikaan?
— Niin, hän tahtoo hankkia sinulle äitipuolen, Pekka.
— Voi sun perhana!

— Niin, näetkös — sanoo Olli hieman hämillään — minulla on
taipumusta naisiin.
— Oletteko hullu, tehän olette jo seitsemänkymmentä ja kolme

vuotta vanha!
— Sitä nuorisoa, se ei ajattele mitään muuta kuin rakkautta ja

sellaista.
— Ho hoi, niin niin — huokailee eukko.
— Mutta emmekö mene tupaan nyt? — arvelee Olli.
— Mennään vaan. — Riitta nousee, he menevät tupaan päin. —

Siellä ei näytä niinkään siistiltä, luulen minä.
— Aiotteko te maata myöskin oikein aviosängyssä? — kysyy

»poika» ja katsoa tirrittää.
— Ole vaiti, se ei kuulu lapsille! — nauraa Olli.
Ja niin menevät he sisään.
Vappuyö.
Viileä oli yö toukokuun ensimäistä päivää vasten. Ilma oli täynnä

niityn ja metsän tuoksuja. Kautta ilman kävi keväälle omituinen
suhina. Posket ja sydän tulivat punaisiksi, kun hengitti sen yön ilmaa,
mieli alkoi rakkausasioita kaipailla.
Puolihämärässä seisoi Frans kädet taskussa tuvan edessä,
äänetönnä, katsellen maita, joilla äsken kynnetyistä va'oista nousi
vehmas tuoksu.
Hän uneksi kuiluista, jota hänellä ei ollut. Hän oli suuri ja vahva,
kaksikymmenvuotias.
Hän katseli kauvas vuorille, jotka valtavina varjoina kuvastuivat

vappuyön puolivaloisaa taivasta vasten. Itsepintaisesti kääntyi hänen
katseensa Uusevuorelle, sen metsättömälle, sileälle ja valaskalan
selkää muistuttavalle selänteelle. Vuoren keskikohdalla roihusi
lieskuva tuli, joka valaisi vuoren selkämää siksi, että hän sen valossa
voi eroittaa ihmisvarjoja juoksentelevina sinne tänne.
Se oli kylän poikien vappukokko. He olivat siellä yön helmassa

hempukoineen, sillä kun Frans oli tarkasti tähystellyt, voi hän
varjojen joukosta nähdä ei ainoastaan poikia, vaan myöskin
helmaväkeä.
Ja silloin leimusi hänen silmissään, kuin olisi sielläkin kokko

palanut. Hän väänsi ruumistaan, kaivoi kädet syvemmälle
taskuihinsa, sylkäsi ja sanoi:
— S—tana!
Sitten asettui hän taas miettimään, kiinnittäen silmänsä kylän

poikien kokkotuleen.
Hän virnistelihe hyvillä mielin. Kas, joskaan hänellä ei ollutkaan

omaa hempukkaansa, niin tiesi hän kuitenkin, kenenkä hän olisi
tahtonut saada. Se oli Vanhankylän Amalia. Mutta ei ikinä elämässä
tulisi hänen vanha ärtyisä äitinsä sallimaan, että hän Amalian saisi,
sen hän tiesi. Siksipä hän juuri kirosi.
Koettaakseen mitä äiti sanoisi moisesta pariskunnasta, oli hän

kerran maininnut Amaliaa talon pojan vaimoksi — samalla kuin
tuskallisesti oli äitiä katsellut.
Mutta silloin oli eukko pauhun nostanut, niin vanha kuin olikin,
vaaleat, laihat kasvot näyttivät suuttumuksesta vavahtelevan, ja
puolisokeat silmät olivat muuttuneet melkein vihreiksi. Hän oli
huutanut:
— Sen köyhän luutun! Ei koskaan elämässäni!
Silloin ymmärsi Frans, että se oli mahdotonta, ja vaikeni. Eukko oli

ahnas kuin juutalainen ja varma kuin vallesmanni eikä hän koskaan
tulisi sallimaan, että torpan Amalia ottaisi hänen kolikkonsa arkun
pohjalta ja saisi talonpojan pojan.
Frans piti sitä konstailemisena, hän olisi melkein voinut itkeä.

Kylän pojat, jotka olivat kaikki köyhiä, olivat saaneet jokainen oman
kullan — heillä on sydämelliset vanhemmat.
Hän polki maata ja repi tukkaansa.
— Että pitääkin olla sellainen äiti!
Samassa kuului äidin käheä, kimeä ääni tuvasta:
— Frans!
Hän vastasi kärsimättömästi:
— Niin!
Äiti kutsui häntä sisään. — Mitähän hän taas tahtoi? Aina olisi
hänen pitänyt jotakin tehdä, kiitosta hän ei saanut koskaan. Hän hoiti
koko talouden, heinän kulotuksesta sikolättiin saakka, mutta hänellä
oli sellainen äiti, ettei hän voinut saada armastansa omakseen.
Äiti tahtoi, että heidän pitäisi mennä myllyyn jauhoja hakemaan.
— Näin myöhään! — murisi Frans. — Voimmehan sen tehdä

huomennakin.
— Laiskuri! — huusi äiti. — Tuos' on säkki!
— Pekka ei ole myllyllä — koetti Frans vastustella.
— Tuuleehan nyt.
Siinä hän oli oikeassa. Frans oli kyllä nähnyt Pekan myllyn siipien
huiskavan metsänreunan yli, kun hän äsken oli seisonut mäellä. Äitiä
ei käynyt petkuttaminen — hän käsitti, että Pekka oli myllyssä,
vaikka näkikin niin huonosti, ettei voinut myllyn siipiä eroittaa.
Frans jukkaili, mutta hänen täytyi kuitenkin ottaa säkki toiseen

käteensä, äiti kulki toiseen nojaten, ja niin he menivät.
Astellessaan siinä äitinsä vieressä tunsi Frans itsensä hyvin

kaukaiseksi äitistään, joka aina oli ollut ankara häntä kohtaan ja
karkoittanut sillä tavalla pojanrakkauden hänestä. Jo pienenä
poikana oli hän saanut itkeä tätä. Isä ei välittänyt hänestä mitään —
hän rakasti kartanoa ja maata, oli vaiti ja kuoli. Sitä mukaa kuin
Frans kasvoi, muuttui hänen surunsa kovakiskoisuudeksi. Äiti ei ollut
tehnyt hänelle muuta hyvää kuin synnyttänyt hänet maailmaan, eikä
se ollut hyvin, koska hän sen kautta oli tottunut viilaamaan sitä, jota
hänen piti rakastaa…
Joka yön eli hän kädet nyrkkiin puristettuina!
Hän pelkäsi tehdä äidilleen mitään pahaa, pelkäsi olla yhdessä

hänen kanssaan.
He olivat vaiti tiellä…
Lopuksi kysyi Frans ikäänkuin toisten onnella itseään suututtaen:
— Näettekö vapputulia tuolta vuorelta?
Äiti pysähtyi ja katsoi sinne, varjostaen käsillään silmiään.
— Sieltä kajastaa heikko valo. Ketä he ovat, jotka sytyttävät tulen

metsään?
— Eihän siellä mitään metsää ole. Ne ovat kylänpoikia, jotka

hempukoineen vappuyötä viettävät, ja kyllä heillä on hauska.
— Niin niin, siellä he nyt yöllä keikkuvat ja niin siitä tulee heti häitä
jälestä — sanoi äiti.
Hän ei voinut koskaan kärsiä nuorten iloa. Hän ei muistanut omaa

nuoruuttaan.
— Teillä ei ole mitään tekemistä sen kanssa mitä jälestä tapahtuu!

huudahti Frans. Pitääkö teidän haukkua heitäkin? Eikö siinä kyllin,
että minua…?
— Sinua? Enkö minä ole ollut äiti sinulle? Oletko kärsinyt vilua ja
nälkää, hä?
— En niin juuri. Mutta te kiellätte minulta sen, josta minä enin

pidän.
— Joko taas vetelet sitä hompsukkaa esiin? Hyi tokin! Ei ikinä tule
hän minun kattoni alle.
— Te olette kova äiti. Mutta ensi vuonna minä olen täysi-ikäinen.
— Mitä aiot silloin tehdä? Naidako ja ajaa minut maantielle? Älä

sinä — minua et saa minnekään! On oikein häpeää, kun sinä saisit
varakkaitakin talonpojan tyttäriä, sekä Ottilian että Jeanetten, jos
vain sanoisit sanankaan.
— Mutta sitä sanaa minä en sano.
— Sinä olet häväistykseni, Frans. Sinä saatat minut hautaan.
74
— Minä en voi odottaa kauvemmin. Minun pitää saada Malin,

kuuletteko!
Se on niin, että siitä voi tulla aivan hulluksi, kun on nuori.
— Nuorten täytyy totella.
— Yö tuntuu niin lämpimältä, etten tunne itseäni.
— Ei minusta.
— Jos te estätte minut saamasta Amalian, en tiedä mitä minä

teen.
— No, mitä sitten? Olisi hauska tietää.
— Äiti!!
He vaikenivat… ja jatkoivat matkaa.

Pian saapuivat he myllylle, jonka siipien suhina ja kivien jauhava
jymy kuului heidän korviinsa.
— Näetkös. Pekka jauhaa kuitenkin.
Frans ei kuullut, mitä äiti sanoi. Hänellä oli ainoastaan yksi ajatus:
Malin!
Ja toinen ajatus: Äiti! Jos hän olisi kuollut!
Herra Jumala, oliko hän niin huono ihminen, että sellaisia ajatteli!
Hänen käsivartensa vapisivat, ja äiti kysyi:
— Onko sinulla vilu?
— Täällä, jossa on niin kuuma! vastasi hän. — Mutta hänen

huulensa vapisivat.
— Sinähän vapiset kuin vilussa. Sinä ajattelet jotakin pahaa. No,

ilkeä sinä oletkin aina ollut.
Tämän sanoi äiti hänelle, joka aina oli tahtonut häntä rakastaa!
— No, olemmeko jo perillä? — kysyi äiti.
— Kyllä, nyt menemme sillan yli.
He kulkivat yli pienen sillan, jonka alitse keväisellä vauhdilla riensi

syvä vesipaljous, ja saapuivat myllymäelle.
Pekka lauleli ylhäällä myllynkivien jymisevään tahtiin, pauhasi niin,

että sitä kuullessa olisi voinut tulla aivan kuuroksi. Siinä oli jotakin
loihdittua tuossa kumeassa pauhussa ja siipien viuhinassa… huss…
uss…
Ne leikkasivat ilmaa niin, että se pilvenä kasvoihin painui. Kaiken

tämän seasta voi eroittaa silloin tällöin Pekan yksinäisen, kimeän
rallatuksen…
— Nyt menemme sitten ylös myllyyn, sanoi äiti.
Hän hapuili toisella kädellään.
— Näettekö siipiä? — kysyi Frans ja kävi ruosteenkarvaiseksi.
— En.
Frans päästi äidin käsivarrestaan ja meni siipien sivuitse myllyn

portaille.
— Mihin sinä menit? — kysyi äiti ja pysähtyi.
— Minä avaan myllyn oven teille.
— Minä kuulen, missä sinä olet. Minä tulen.
Hän läksi kävelemään myllyn ovelle, jossa Frans oli.
Frans näki, että hän kulki suoraan myllyn siipiä kohti.
Hän ei huutanut hänelle…
Frans ei liikkunut. Hän torkkui…
Hänen piti pitää kiinni avatusta myllyn ovesta, ettei putoaisi…

Toinen myllyn siipi paukautti ankarasti äitiä päähän ja heitti hänet
alas mäen rinteelle, johon hän jäi makaamaan…
Frans, joka tunsi elämän ja kuoleman tuskan kuristavan

kuikkaansa, sai tuskin kähistyksi:
— Apua!
Sitten riensi hän äitinsä luo, heittäytyi hänen ylitsensä ja huusi:
— Äiti, äiti!
Pekan rallatus loppui. Hän näyttäytyi myllyn aukosta ja kysyi

alhaalta:
— Mitä se on?
— Hän on kuollut…? — huusi Frans ylös.
Samassa kuului myllyn portailta Pekan puukenkien kolina, ja pian

oli hän maassa, valkeana kuin aave.
— Kuinka se tapahtui? Kuinka se tapahtui? — voivotteli Pekka.
— Hän meni suoraan siipiä kohti. Hänhän oli niin sokea. Minun piti
juuri huutaa hänelle, mutta se oli liian myöhäistä.
— Sellainen onnettomuus! Ja juuri minun myllylläni!…
— Meidän pitää viedä hänet sinun luoksesi.
— Minä menen ja seisotan myllyn.
Sen Pekka teki ja yhdessä kantoivat he ruumiin Pekan kotiin.

Frans tunsi kuin kevään rinnassaan. Ihmeellistä, kuinka äiti oli
kevyt kantaa; verta kyllä vuoti käsille.
Sydän löi ankarasti, ja veri kiehui suonissa. Hän ei tiennyt, pitikö

hänen itkeä vai nauraa. Se oli synti äidin tähden. Oli synti hänen
itsensä tähden.
Mutta sitten ajatteli hän Malin'ia, joka istui Vanhassakylässä ja

ikävöitsi häntä, kaipaus sai hänet hikoilemaan, ja lopuksi hän itki
lämmintä itkuaan tuossa kevään ja rakkauden yössä…
Ja vapputuli leimusi ylhäällä vuorella ja näytti nuoleksivan taivaan

vaalenevia avaruuksia, nuoren väen suudellessa ja nauraessa…
Keulapuolella.
Olimme huviksemme purjehtimassa.
Jahti piti suuntaansa suoraan Nokiaukean yli.
Aikomuksemme oli käydä lähellä Nokisaarta — äärimmäinen

merelle päin. Mutta yhä lisääntyvässä aallokossa päätimme me,
naisten tähden, kääntyä maihin.
Juuri nyt käännyttyä, jahdin kiitäessä myötätuulta kotia kohti, näin

minä Ninuksen, nuoren laivurin seisovan keulalla ja katsovan meihin
puoliksi surkeana, sillä aikaa kuin hänen vanhempi toverinsa
mukavasti istui ruorissa.
— Mitäs Ninus hymyilee? — kysyin minä.

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Biochemistry 7th Edition Reginald H.

BioChemistry 6th Edition By Reginald H. Garrett

Lehninger Principles of Biochemistry 7th Edition eBook

Textbook of Biochemistry with Clinical Correlations,

Biochemistry, 7th ed: The Molecular basis of life 7th

Lippincott Illustrated Reviews: Biochemistry

Dalzielov Duh Reginald Hill

BRS Biochemistry, Molecular Biology, and Genetics

atlas of human anatomy 7th Edition Frank H Netter

Reginald H. Garrett | Charles M. Grisham

With molecular graphic images

Australia • Brazil • Canada • Mexico • Singapore • United Kingdom • United States

Learning Designer: Mona Zeftel

Content Program Manager: Ivan Corriher

Senior IP Project Manager: Betsy Hathaway

Production Service: Lumina Datamatics, Ltd.

Cover Image: CRISPR-Cas9, by Tina Chai

Printed in the United States of America

To our grandchildren Jackson, Bella, Reggie, Ricky, Charlotte

Reginald H. Garrett and Charles M. Grisham

Part I Molecular Components of Cells 1

Part II Protein Dynamics 433

Part III Metabolism and Its Regulation 573

Part IV Information Transfer 981

Abbreviated Answers to Problems A-1

 itration Curves Illustrate the Progressive

5 in the Laboratory? 131

8 9.1 What Are the Chemical and Physical

 eripheral Membrane Proteins Associate Loosely

 ext-Generation Sequencing by Synthesis 348

14 15.1 What Factors Influence Enzymatic Activity? 511

 lagellar Filaments Are Composed of Protofilaments of

Metabolism: An Overview 573 Foundational Biochemistry 597

— Minä möin vuodan.

Vaunut vierivät rämisten ylös mäkeä Ernstin tuvalle.

Hän ihmetteli, mitä se mahtoi olla, meni etehisen kuistiin ja asetti

— Herranen aika, vallesmanni!

Hän kalpeni yht'äkkiä. Hänen ensi ajatuksensa oli: vaimo, lapsi! —

Mutta nyt olivat vaunut hänen ovensa edustalla. Eräs talonpoika

— Hyvää päivää, Ernst! — Se kuului onnettomuutta ennustavalta,

— Herra vallesmanni suo minulle sen kunnian.

Ernst kumarsi. Sydän sylkytti.

— Tahtoisin puhua hiukan sinun kanssasi. Ja minulla on todistajat,

Nyt selveni hänelle kaikki. Mutta hän nieleksi, tyynennäköisenä —

— Sinut on ilmiannettu — arveli nimismies.

Ernst koetti nauraa.

— Sinä et näy sen siitään paranneen.

Ernst tuli veripunaiseksi.

— Älkää puhuko siitä!

— Minä puhun uudesta. Sinut on ilmiannettu…

— Kenenkä toimesta, jos saan luvan kysyä?

— Sitä minun ei tarvitse sanoa.

Hän osoitti. Toiset katsoivat osoitettuun suuntaan. Siellä puikki

— Tule sinä, Hans — kirkui Ernst — tule katsomaan, mitä olet

Ernst koetti käsillään päätään.

— No, Ernst, nyt tiedät sinä, kuka on ilmiantajasi. Se oli oikein

— Mistä vallesmanni on tuon saanut, jos saan olla nenäkäs?

— Sen tiedät parhaiden itse.

— Tahtovatko siis kaikki ihmiset tehdä minut onnettomaksi?

Samalla osoitti hän alas tielle.

— Katsokaa, ettekö usko, että hänkin tulee tänne! Niin niin,

Eriksson läheni, vähän arkaillen. Mutta hän tahtoi nähdä

Tunnustatko väärentäneesi tämän?

— Ernst! — kuului kirahdus puolipimeästä etehisestä.

itration Curves Illustrate the Progressive

eripheral Membrane Proteins Associate Loosely

ext-Generation Sequencing by Synthesis 348

lagellar Filaments Are Composed of Protofilaments of