Biochemistry: a short course
Gespeichert in:
| Hauptverfasser: | , , |
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| Format: | Buch |
| Sprache: | English |
| Veröffentlicht: |
New York, NY
Freeman
2015
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| Ausgabe: | 3. ed. |
| Schlagworte: | |
| Online-Zugang: | Inhaltsverzeichnis |
| Beschreibung: | Getr. Zählung Ill., graph. Darst. |
| ISBN: | 9781464126130 1464126135 |
Internformat
MARC
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| 020 | |a 9781464126130 |9 978-1-4641-2613-0 | ||
| 020 | |a 1464126135 |9 1-4641-2613-5 | ||
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| 100 | 1 | |a Tymoczko, John L. |d 1948-2019 |e Verfasser |0 (DE-588)124601103 |4 aut | |
| 245 | 1 | 0 | |a Biochemistry |b a short course |c John L. Tymoczko ; Jeremy M. Berg ; Lubert Stryer |
| 250 | |a 3. ed. | ||
| 264 | 1 | |a New York, NY |b Freeman |c 2015 | |
| 300 | |a Getr. Zählung |b Ill., graph. Darst. | ||
| 336 | |b txt |2 rdacontent | ||
| 337 | |b n |2 rdamedia | ||
| 338 | |b nc |2 rdacarrier | ||
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| 700 | 1 | |a Berg, Jeremy M. |d 1958- |e Verfasser |0 (DE-588)12460109X |4 aut | |
| 700 | 1 | |a Stryer, Lubert |d 1938-2024 |e Verfasser |0 (DE-588)124601197 |4 aut | |
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Datensatz im Suchindex
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Brief Contents
PART I
The Molecular Design of Life
SECTION 1 Biochemistry Helps Us Understand
Our World 1
Chapter 1 Biochemistry and the Unity of Life 3
Chapter 2 Water, Weak Bonds, and the
Generation of Order Out of Chaos 17
SECTION 2 Protein Composition and Structure 35
Chapter 3 Amino Acids 37
Chapter 4 Protein Three-Dimensional Structure 47
Chapter 5 Techniques in Protein Biochemistry 69
SECTION 3 Basic Concepts and Kinetics of
Enzymes 95
Chapter 6 Basic Concepts of Enzyme Action 97
Chapter 7 Kinetics and Regulation 111
Chapter 8 Mechanisms and Inhibitors 131
Chapter 9 Hemoglobin, an Allosteric Protein 149
SECTION 4 Carbohydrates and Lipids 165
Chapter 10 Carbohydrates 167
Chapter 11 Lipids 189
SECTION 5 Cell Membranes, Channels,
Pumps, and Receptors 203
Chapter 12 Membrane Structure and Function 205
Chapter 13 Signal-Transduction Pathways 225
PART II
Transducing and Storing Energy
SECTION 6 Basic Concepts and Design
of Metabolism 245
Chapter 14 Digestion: Turning a Meal into
Cellular Biochemicals 247
Chapter 15 Metabolism: Basic Concepts
and Design 257
SECTION 7 Glycolysis and Gluconeogenesis 281
Chapter 16 Glycolysis 283
Chapter 17 Gluconeogenesis 313
SECTION 8 The Citric Acid Cycle 329
Chapter 18 Preparation for the Cycle 331
Chapter 19 Harvesting Electrons from the
Cycle 343
SECTION 9 Oxidative Phosphorylation 361
Chapter 20 The Electron-Transport Chain 363
Chapter 21 The Proton-Motive Force 383
SECTION 10 The Light Reactions of
Photosynthesis and the Calvin Cycle 405
Chapter 22 The Light Reactions 407
Chapter 23 The Calvin Cycle 427
SECTION 11 Glycogen Metabolism and
the Pentose Phosphate Pathway 443
Chapter 24 Glycogen Degradation 445
Chapter 25 Glycogen Synthesis 459
Chapter 26 The Pentose Phosphate
Pathway 473
SECTION 12 Fatty Acid and Lipid Metabolism 487
Chapter 27 Fatty Acid Degradation 489
Chapter 28 Fatty Acid Synthesis 507
Chapter 29 Lipid Synthesis: Storage Lipids,
Phospholipids, and Cholesterol 523
SECTION 13 The Metabolism of Nitrogen-
Containing Molecules 549
Chapter 30 Amino Acid Degradation and
the Urea Cycle 551
Chapter 31 Amino Acid Synthesis 571
Chapter 32 Nucleotide Metabolism 585
PART III
Synthesizing the Molecules of Life
SECTION 14 Nucleic Acid Structure
and DNA Replication 605
Chapter 33 The Structure of Informational
Macromolecules: DNA and RNA 607
Chapter 34 DNA Replication 627
Chapter 35 DNA Repair and Recombination 643
SECTION 15 RNA Synthesis, Processing,
and Regulation 657
Chapter 36 RNA Synthesis and Regulation
in Bacteria 659
Chapter 37 Gene Expression in Eukaryotes 675
Chapter 38 RNA Processing in Eukaryotes 691
SECTION 16 Protein Synthesis and
Recombinant DNA Techniques 705
Chapter 39 The Genetic Code 707
Chapter 40 The Mechanism of Protein
Synthesis 721
Chapter 41 Recombinant DNA Techniques 743
XV
Contents
PARTI
The Molecular Design of Life
SECTION 1
Biochemistry Helps Us to Understand Our World 1
Chapter t Biochemistry and the Unity of Life 3
1.1 Living Systems Require a L imited Variety of Atoms
and Molecules 4
1.2 There Are Four Major Classes of Biomolecules 5
Proteins Are Highly Versatile Biomolecules 5
Nucleic Adds Are the Information Molecules of the Cel) 6
Lipids Are a Storage form of Fuel and Serve as a Barrier 6
Carbohydrates Are Fuels and Informational Molecules 7
1.3 The Central Dogma Describes the Basic Principles
of Biological Information Transfer 7
1.4 Membranes Define the Cell and Carry Out Cellular
Functions 8
Biochemical Functions Are Sequestered in Cellular
Compartments 11
Some Organelles Process and Sort Proteins and Exchange
Material with the Environment 12
Clinical Insight Defects in Organelle Function
May Lead to Disease 14
Chapter 2 Water, Weak Bonds, and the
Generation of Order Out of Chaos 17
2.1 Thermal Motions Power Biological Interactions 18
2.2 Biochemical Interactions Take Place
in an Aqueous Solution 18
2.3 Weak interactions Are Important Biochemical
Properties 20
Electrostatic Interactions Are Between Electrical Charges 20
Hydrogen Bonds Form Between an Electronegative
Atom and Hydrogen 21
van der Waals Interactions Depend on Transient
Asymmetry in Electrical Charge 21
Weak Bonds Permit Repeated Interactions 22
2.4 Hydrophobic Molecules Cluster Together 22
Membrane Formation Is Powered by the
Hydrophobic Effect 23
Protein Folding Is Powered by the Hydrophobic Effect 24
Functional Groups Have Specific Chemical Properties 24
2.5 pH Is an Important Parameter of
Biochemical Systems 26
Water Ionizes to a Small Extent 26
An Acid Is a Proton Donor, Whereas a Base Is a
Proton Acceptor 27
Adds Have Differing Tendencies to Ionize 27
Buffers Resist Changes in pH 28
Buffers Are Crucial in Biological Systems 29
Making Buffers Is a Common Laboratory Practice 30
SECTION 2
Protein Composition and Structure 35
Chapter 3 Amino Acids 37
Two Different Ways of Depicting Biomolecules
Will Be Used 38
3.1 Proteins Are Built from a Repertoire of
20 Amino Acids 38
Most Amino Acids Exist in Two Mirror-Image Forms 38
All Amino Acids Have at Least Two Charged Groups 38
3.2 Amino Acids Contain a Wide Array of
Functional Groups 39
Hydrophobic Amino Acids Have Mainly
Hydrocarbon Side Chains 39
Polar Amino Acids Have Side Chains That Contain an
Electronegative Atom 41
Positively Charged Amino Acids Are Hydrophilic 42
Negatively Charged Amino Acids Have Acidic
Side Chains 43
The lonizable Side Chains Enhance Reactivity and
Bonding 43
3.3 Essential Amino Acids Must Be Obtained from
the Diet 44
PjJ Clinical Insight Pathological Conditions Result
If Protein Intake Is Inadequate 44
Chapter 4 Protein Three-Dimensional
Structure 47
4.1 Primary Structure: Amino Acids Are Linked
by Peptide Bonds to Form Polypeptide Chains 48
Proteins Have Unique Ammo Acid Sequences
Specified by Genes 49
Polypeptide Chains Are Flexible Vet Conformationally
Restricted 50
4.2 Secondary Structure: Polypeptide Chains
Can Fold into Regular Structures 52
The Alpha Helix Is a Coiled Structure Stabilized by
Intrachain Hydrogen Bonds 52
Beta Sheets Are Stabilized by Hydrogen Bonding
Between Polypeptide Strands 53
Polypeptide Chains Can Change Direction by
Making Reverse Turns and Loops 55
Fibrous Proteins Provide Structural Support for
Cells and Tissues 55
Ü Clinical Insight Defects in Collagen Structure
Result in Pathological Conditions 57
4.3 Tertiary Structure: Water-Soluble Proteins Fold
into Compact Structures 57
Myoglobin Illustrates the Principles of Tertiary Structure 57
The Tertiary Structure of Many Proteins Can Be
Divided into Structural and Functional Units 59
4.4 Quaternary Structure: Multiple Polypeptide
Chains Can Assemble into a Single Protein 59
XVI
Contents XVII
4.5 The Amino Acid Sequence of a Protein 6.4 Enzymes Facilitate the Formation of
Determines Its Three-Dimensional Structure 60 the Transition State 103
Proteins Fold by the Progressive Stabilization of Intermediates Rather Than by Random Search 61 The Formation of an Enzyme-Substrate Complex Is the First Step in Enzymatic Catalysis 103
Some Proteins Are Inherently Unstructured arid Can The Active Sites of Enzymes Elave Some Common Features 104
Exist in Multiple Conformations ЕпЭ 62 The Binding Energy Between Enzyme and Substrate Is
|fi| Clinical Insight Protein Misfolding and Aggregation Important for Catalysis 105
Are Associated with Some Neurological Diseases 63 Transition-State Analogs Are Potent Inhibitors of Enzyme 106
Chapter 5 Techniques in Protein Biochemistry 69 Chapter 7 Kinetics and Regulation m
5.1 The Proteome Is the Functional Representation of the Genome 70 7.1 Kinetics Is the Study of Reaction Rates 112
5.2 The Purification of a Protein Is the First Step in Understanding Its Function 70 7.2 The Michaelis-Menten Model Describes the Kinetics of Many Enzymes 113
Proteins Can Be Purified on the Basis of Differences in Their Chemical Properties 71 Clinical Insight Variations in KM Can Flave Physiological Consequences 114
Proteins Must Be Removed from the Cell to Be Purified 71 KM and l/max Values Can Be Determined by Several Means 115
Proteins Can Be Purified According to Solubility, Size, KM and Vm3x Values Are Important Enzyme Characteristics 115
Charge, and Binding Affinity 72 kat/KM С a Measure of Catalytic Efficiency 116
Proteins Can Be Separated by Cel Electrophoresis and Most Biochemical Reactions Include Multiple Substrates 117
Displayed 74 7.3 Allosteric Enzymes Are Catalysts and Information
A Purification Scheme Can Be Quantitatively Evaluated 77 Sensors 118
5.3 Immunological Techniques Are Used to Purify and Characterize Proteins 78 Allosteric Enzymes Are Regulated by Products of the Pathways Under Their Control 120
Centrifugation Is a Means of Separating Proteins 78 Allosterically Regulated Enzymes Do Not Conform to
Gradient Centrifugation Provides an Assay for the Michaelis-Menten Kinetics 121
Estradiol-Receptor Complex 79 Allosteric Enzymes Depend on Alterations in Quaternary
Antibodies to Specific Proteins Can Be Generated 80 Structure 121
Monoclonal Antibodies with Virtually Any Desired Regulator Molecules Modulate the R s — T Equilibrium 122
Specificity Can Be Readily Prepared 81 The Sequential Model Also Can Account for
The Estrogen Receptor Can Be Purified by Allosteric Effects StfS 123
Immunoprécipitation 83 Clinical Insight Loss of Allosteric Control May
Proteins Can Be Detected and Quantified with Result in Pathological Conditions 123
the. Use of an Enzyme-Linked Immunosorbent Assay 84 7.4 Enzymes Can Be Studied One Molecule at a Time 123
Western Blotting Permits the Detection of Proteins Separated by Cel Electrophoresis 84 Chapter 8 Mechanisms and Inhibitors 131
5.4 Determination of Primary Structure Facilitates an Understanding of Protein Function 86 8.1 A Few Basic Catalytic Strategies Are Used by Many Enzymes 131
Mass Spectrometry Can Be Used to Determine a Protein's Mass, Identity, and Sequence 88 8.2 Enzyme Activity Can Be Modulated by Temperature, pH, and Inhibitory Molecules 132
Amino Acids Are Sources of Many Kinds of Insight 90 Temperature Enhances the Rate of Enzyme-Catalyzed
SECTION 3 Reactions 132
Basic Concepts and Kinetics of Enzymes 95 Most Enzymes Have an Optimal pH 133
Chapter 6 Basic Concepts of Enzyme Action 97 Enzymes Can Be Inhibited by Specific Molecules Reversible Inhibitors Are Kinetically Distinguishable 134 135
6.1 Enzymes Are Powerful and Highly Specific Catalysts 97 Irreversible Inhibitors Can Be Used to Map
Proteolytic Enzymes Illustrate the Range of Enzyme the Active Site 137
Specificity 98 If Clinical Insight Penicillin Irreversibly Inactivates a
There Are Six Major Classes of Enzymes 98 Key Enzyme in Bacterial Cell-Wall Synthesis 138
6.2 Many Enzymes Require Cofactors for Activity 99 8.3 Chymotrypsin Illustrates Basic Principles of
6.3 Gibbs Free Energy Is a Useful Thermodynamic Catalysis and Inhibition 140
Function for Understanding Enzymes 100 Serine 195 Is Required for Chymotrypsin Activity 140
The Free-Energy Change Provides Information About the Spontaneity but Not the Rate of a Reaction 100 Chymotrypsin Action Proceeds In Two Steps Linked by a Covalently Bound Intermediate 141
The Standard Free-Energy Change of a Reaction Is Related to the Equilibrium Constant 101 The Catalytic Role of Histidine 57 Was Demonstrated by Affinity Labeling 142
Enzymes Alter the Reaction Rate but Not the Reaction Equilibrium 102 Serine Is Part of a CatalyticTriad That Includes Histidine and AsparticAcid 142
XViii Contents
Chapter 9 Hemoglobin, an Allosteric Protein
9.1 Hemoglobin Displays Cooperative Behavior
9.2 Myoglobin and Hemoglobin Bind Oxygen in
Heme Croups
H Clinical Insight functional Magnetic Resonance
Imaging Reveals Regions of the Brain Processing
Sensory Information
9.3 Hemoglobin Binds Oxygen Cooperatively
9.4 An Allosteric Regulator Determines the
Oxygen Affinity of Hemoglobin
I Clinical Insight Hemoglobin's Oxygen Affinity Is
Adjusted to Meet Environmental Needs
H Biological Insight Hemoglobin Adaptations Allow
Oxygen Transport in Extreme Environments
9.5 Hydrogen Ions and Carbon Dioxide Promote the
Release of Oxygen
9.6 Mutations in Genes Encoding Hemoglobin
Subunits Can Result in Disease
I Clinical Insight Sickle-Cell Anemia Is a Disease
Caused by a Mutation in Hemoglobin
I NEW Clinical Insight Thalassemia is Caused by an
Imbalanced Production of Hemoglobin Chains
SECTION 4
Carbohydrates and Lipids
Chapter 10 Carbohydrates
10.1 Monosaccharides Are the Simplest Carbohydrates
Many Common Sugars Exist in Cyclic Forms
NEW Pyranose and Furanose Rings Can Assume
Different Conformations
BM NEW Clinical Insight Glucose Is a Reducing Sugar
Monosaccharides Are Joined to Alcohols and
Amines Through Glycosidic Bonds
Q Biological Insight Glucosinolates Protect Plants
and Add Flavor to Our Diets
10.2 Monosaccharides Are Linked to Form Complex
Carbohydrates
Specific Enzymes Are Responsible for Oligosaccharide
Assembly
Sucrose, Lactose, and Maltose Are the Common
Disacchandes
Glycogen and Starch Are Storage Forms of Glucose
Cellulose, a Structural Component of Plants, Is
Made of Chains of Glucose
10.3 Carbohydrates Are Attached to Proteins to
Form Glycoproteins
Carbohydrates May Be Linked to Asparagine, Serine, or
Threonine Residues of Proteins
Eg Clinical Insight The Hormone Erythropoietin Is a
Glycoprotein
Proteoglycans, Composed of Polysaccharides and
Protein, Have Important Structural Roles
Bä Clinical Insight Proteoglycans Are Important
Components of Cartilage
| Clinical Insight Mucins Are Glycoprotein
Components of Mucus
149 B Biological Insight Blood Groups Are Based on
150 Protein Glycosylation Patterns 181
150 | Clinical Insight Lack of Glycosylation Can Result in Pathological Conditions 182
10.4 Lectins Are Specific Carbohydrate-Binding Proteins 182
152 Lectins Promote Interactions Between Cells 183
152 | Clinical Insight Lectins Facilitate Embryonic
Development 183
154 | Clinical Insight Influenza Virus Binds to
Sialic Acid Residues 183
154 Chapter 11 lipids 189
155 11.1 Fatty Acids Are a Main Source of Fuel 190
155 Fatty Acids Vary in Chain Length and Degree of Un saturation 191
156 157 The Degree and Type of Unsaturation Are Important to Health 192
112 Triacylgiycerols Are the Storage Form of Fatty Acids 193
159 11.3 There Are Three Common Types of Membrane Lipids 194
Phospholipids Are the Major Class of Membrane Lipids 194
165 Membrane Lipids Can Include Carbohydrates 196
167 Steroids Are Lipids That Have a Variety of Roles 196
168 | Biological Insight Membranes of Extremophiles
Are Built from Ether Lipids with Branched Chains 197
169 171 171 Membrane Lipids Contain a Hydrophilic and a Hydrophobic Moiety 197
Some Proteins Are Modified by the Covalent Attachment of HydropFiobicGroups 198
172 | Clinical Insight Premature Aging Can Result from the Improper Attachment of a Hydrophobic Group to a Protein 199
175 173 SECTION 5 Cell Membranes, Channels, Pumps, and
173 Receptors 203
174 Chapter 12 Membrane Structure and
175 Function 205
175 12.1 Phospholipids and Glycolipids Form Bimolecular Sheets 206
177 | Clinical Insight Lipid Vesicles Can Be Formed from Phospholipids 207
177 Lipid Bilayers Are Highly Impermeable to Ions and Most Polar Molecules 207
178 12.2 Membrane Fluidity Is Controlled by Fatty Acid Composition and Cholesterol Content 208
12.3 Proteins Carry Out Most Membrane Processes 209
178 Proteins Associate with the Lipid Bilayer in
179 a Variety of Ways 209
■ Clinical Insight The Association of Prostaglandin H2
180 Synthase-I with the Membrane Accounts for the Action of Aspirin 211
12.4 Lipids and Many Membrane Proteins Diffuse
Laterally in the Membrane
12.5 A Major Role of Membrane Proteins Is to
Function As Transporters
The Na+-K+ ATPase Is an Important Pump in
Many Cells
Q Clinical Insight Multidrug Resistance Highlights
a Family of Membrane Pumps with ATP-Binding
Domains
O Clinical Insight Harlequin Ichthyosis Is a Dramatic
Result of a Mutation in an ABC Transporter Protein
SecondaryTransporters Use One Concentration
Gradient to Power the Formation of Another
|H Clinical Insight Digitalis Inhibits the Na+֊Kf Pump
by Blocking Its Déphosphorylation
Specific Channels Can RapidlyTransport Ions Across
Membranes
Biological Insight Venomous Pit Vipers Use Ion
Channels to Generate a Thermal Image
The Structure of the Potassium Ion Channel Reveals
the Basis of Ion Specificity
The Structure of the Potassium Ion Channel Explains
Its Rapid Rate ofTransport
Chapter 13 Signal-Transduction Pathways
13.1 Signal Transduction Depends on Molecular
Circuits
13.2 Receptor Proteins Transmit Information into
the Cell
Seven-Transmembrane-Helix Receptors Change
Conformation in Response to Ligand Binding and
Activate G Proteins
L igand Binding to 7TM Receptors Leads to the
Activation of G Proteins
Activated G Proteins Transmit Signals by Binding to
Other Proteins
Cyclic AMP Stimulates the Phosphorylation of
Many Target Proteins by Activating Protein Kinase A
11 NEW Clinical Insight Mutations in Protein Kinase A
Can Cause Cushing/s Syndrome
G Proteins Spontaneously Reset Themselves
Through GTP Hydrolysis
H Clinical Insight Cholera and Whooping Cough Are
Due to Altered G֊Protein Activity
The Hydrolysis of Phosphatidylinositol Bisphosphate by
Phospholipase C Generates Two Second Messengers
13.3 Some Receptors Dimerize in Response to Ligand
Binding and Recruit Tyrosine Kinases
Receptor Dimerization May Result in Tyrosine Kinase
Recruitment
k! Clinical Insight Some Receptors Contain Tyrosine
Kinase Domains Within Their Covalent Structures
Ras Belongs to Another Class of Signaling G Proteins
13.4 Metabolism in Context: Insulin Signaling
Regulates Metabolism
The Insulin Receptor Is a Dimer That Closes Around
a Bound Insulin Molecule
Contents The Activated Insulin-Receptor Kinase Initiates a Kinase XIX
211 Cascade Insulin Signaling Is Terminated by the Action of 237
212 Phosphatases 13.5 Calcium Ion Is a Ubiquitous Cytoplasmic 238
213 Messenger 13.6 Defects in Signaling Pathways Can Lead to 238
Diseases 239
214 11 Clinical Insight The Conversion of Proto-oncogenes into Oncogenes Disrupts
214 the Regulation of Cell Growth HI Clinical Insight Protein Kinase Inhibitors May 239
214 Be Effective Anticancer Drugs 240
215 PÂRTII
216 Transducing and Storing Energy SECTION 6
216 Basic Concepts and Design of Metabolism 245
216 Chapter 14 Digestion: Turninga Meal into
Cellular Biochemicals 247
218 14.1 Digestion Prepares Large Biomolecules for
Use in Metabolism 247
225 Most Digestive Enzymes Are Secreted as Inactive
Precursors 248
225 14.2 Proteases Digest Proteins into Amino Acids and
Peptides 248
227 HI NEW Clinical Insight Protein Digestion Begins in the
Stomach 248
NEW Protein Digestion Continues in the Intestine 249
227 0 MEW Clinical Insight Celiac Disease Results from
the Inability to Properly Digest Certain Proteins 251
228 14.3 Dietary Carbohydrates Are Digested by
Alpha-Amylase 251
229 14.4 The Digestion of Lipids Is Complicated by
229 Their Hydrophobicity 252
Q Biological Insight Snake Venoms Digest from
230 the Inside Out Chapter 15 Metabolism: Basic Concepts and 254
230 Design 15.1 Energy Is Required to Meet Three 257
231 NEW Fundamental Needs 15.2 Metabolism Is Composed of Many 258
232 Interconnecting Reactions Metabolism Consists of Energy-Yielding Reactions and 258
233 Energy-Requiring Reactions A Thermodynamically Unfavorable Reaction Can 259
233 Be Driven by a Favorable Reaction 260
235 15.3 ATP Is the Universal Currency of Free Energy 260
ATP Hydrolysis Is Exergonic 261
236 ATP Hydrolysis Drives Metabolism by Shifting
the Equilibrium of Coupled Reactions 261
236 The High Phosphory 1-Transfer Potential of ATP Results from Structural Differences Between
236 ATP and Its Hydrolysis Products 263
XX Contents
Phosphoryl-Transfer Potential Is an Important Form of
Cellular Energy Transformation 264
Clinical Insight Exercise Depends on Various
Means of Generating ATP 265
Phosphates Play a Prominent Role in
Biochemical Processes 266
15.4 The Oxidation of Carbon Fuels Is an
Important Source of Cellular Energy 266
Carbon Oxidation Is Paired with a Reduction 266
Compounds with High Phosphoryl-Transfer Potential
Can Couple Carbon Oxidation to ATP Synthesis 267
15.5 Metabolic Pathways Contain Many Recurring
Motifs 268
Activated Carriers Exemplify the Modular Design and
Economy of Metabolism 268
Clinical Insight Lack of Activated Pantothenate
Results in Neurological Problems 271
Many Activated Carriers Are Derived from Vitamins 271
15.6 Metabolic Processes Are Regulated in
Three Principal Ways 273
The Amounts of Enzymes Are Control led 274
Catalytic Activity Is Regulated 274
The Accessibility of Substrates Is Regulated 275
SECTION 7
Glycolysis and Gluconeogenesis 281
Chapter 16 Glycolysis 285
16.1 Glycolysis Is an Energy-Conversion Pathway 284
Hexokinase Traps Glucose in the Cell and
Begins Glycolysis 284
Fructose 1,6-bisphosphate Is Generated from Glucose
6֊phosphate 286
Clinical Insight The Six Carbon Sugar Is Cleaved
into Two Three-Carbon Fragments 287
The Oxidation of an Aldehyde Powers the Formation
of a Compound Having High Phosphoryl-Transfer
Potential 288
ATP Is Formed by Phosphoryi Transfer from
1,3-Bisphosphoglycerate 289
Additional ATP Is Generated with the Formation
of Pyruvate 2.90
Two ATP Molecules Are Formed in the Conversion
of Glucose into Pyruvate 291
16.2 NAD+ Is Regenerated from the Metabolism of
Pyruvate 291
Fermentations Are a Means of Oxidizing NADH 292
ЦП Biological Insight Fermentations Provide Usable
Energy in the Absence of Oxygen 294
16.3 Fructose and Galactose Are Converted into
Glycolytic Intermediates 294
NEW Fructose Is Converted into Glycolytic Intermediates
by Fructokinase 295
O NEW Clinical Insight Excessive Fructose
Consumption Can Lead to Pathological Conditions 295
NEW Galactose Js Converted into Glucose 6-phosphate 296
НЦ Clinical Insight Many Adults Are Intolerant of Milk
Because They Are Deficient in Lactase 297
НЯ Clinical Insight Galactose Is Highly Toxic If the
Transferase Is Missing 298
16.4 The Glycolytic Pathway Is Tightly Controlled 299
Glycolysis in Muscle Is Regulated by Feedback
Inhibition to Meet the Need for ATP 299
The Regulation of Glycolysis in the Liver
Corresponds to the Biochemical Versatility of
the Liver 300
A Family of Transporters Enables Glucose to Enter and
Leave Animal Cells 303
H NEW Clinical Insight Aerobic Glycolysis Is a
Property of Rapidly Growing Cells 304
ИЯ Clinical Insight Cancer and Exercise Training Affect
Glycolysis in a Similar Fashion 305
16.5 Metabolism in Context: Glycolysis Helps
Pancreatic Beta Cells Sense Glucose 305
Chapter 17 Gluconeogenesis 313
17.1 Glucose Can Be Synthesized from
Noncarbohydrate Precursors 314
Gluconeogenesis Is Not a Complete Reversal of
Glycolysis 314
The Conversion of Pyruvate into Phosphoenolpyruvate
Begins with the Formation of Oxaloacetate 316
Oxaloacetate Is Shuttled into the Cytoplasm and
Converted into Phosphoenolpyruvate 317
The Conversion of Fructose 1,6-bisphosphate into
Fructose 6-phosphate and Orthophosphate Is an
Irreversible Step 318
The Generation of Free Glucose Fs an Important
Control Point 319
Six High-Transfer-Potential Phosphoryi Groups Are
Spent in Synthesizing Glucose from Pyruvate 319
17.2 Gluconeogenesis and Glycolysis Are
Reciprocally Regulated 320
Energy Charge Determines Whether Glycolysis or
Gluconeogenesis Will Be More Active 320
The Balance Between Glycolysis and Gluconeogenesis
in the Liver Is Sensitive to Blood-Glucose
Concentration 321
ЦР Clinical Insight Insulin Fails to Inhibit
Gluconeogenesis in Type 2 Diabetes 323
Ц Clinical Insight Substrate Cycles Amplify
Metabolic Signals 323
17.3 Metabolism in Context: Precursors Formed by
Muscle Are Used by Other Organs 324
SECTION 8
The Citric Acid Cycle 329
Chapter 18 Preparation for the Cycle 331
18.1 Pyruvate Dehydrogenase Forms Acetyl
Coenzyme A from Pyruvate 332
The Synthesis of Acetyl Coenzyme A from
Pyruvate Requires Three Enzymes and
Five Coenzymes 333
Flexible Linkages Allow Lipoamide to Move Between
Different Active Sites 335
18.2 The Pyruvate Dehydrogenase Complex Is
Regulated by Two Mechanisms 337
Contents XXI
§U Clinical Insight Defective Regulation of Pyruvate
Dehydrogenase Results in Lactic Acidosis 338
Ü Clinical Insight Enhanced Pyruvate
Dehydrogenase Kinase Activity Facilitates the
Development of Cancer 339
II Clinical Insight The Disruption of Pyruvate
Metabolism Is the Cause of Beriberi 339
Chapter 19 Harvesting Electrons from
the Cycle 343
19.1 The Citric Acid Cycle Consists of Two Stages 344
19.2 Stage One Oxidizes Two Carbon Atoms to
Gather Energy-Rich Electrons 344
Citrate Synthase. Forms Citrate from Oxaioacetate and
Acetyl Coenzyme A 344
The Mechanism of Citrate Synthase Prevents
Undesirable Reactions 345
Citrate Is Isomerized into Isocitrate 346
Isocitrate Is Oxidtzed and Decarboxyl a ted to
Alpha-Ketogluta rate 346
Succmy! Coenzyme A Is Formed by the Oxidative
Decarboxylation of Alpha-Ketoglutarate 347
19.3 Stage Two Regenerates Oxaioacetate and
Harvests Energy-Rich Electrons 347
A Compound with High Phosphoryl· Transfer
Potential Is Generated from Succinyl Coenzyme A 347
Succmyl Coenzyme A Synthetase transforms
types of Biochemical Energy 348
Oxaioacetate Is Regenerated by the Oxidation of
Succinate 349
Fhe Citric Acid Cycle Produces High-Fransfer-Potential
Electrons, an ATP, and Carbon Dioxide 349
19.4 The Citric Acid Cycle Is Regulated 352
The. Citric. Acid Cycle Is Controlled at Several Points 352
The Citric Acid Cycle Is a Source of Biosynthetic
Precursors 353
The Citric Acid Cycle Must Be Capable of Being Rapidly
Replenished 353
§(§ Clinical Insight Defects in the Citric Acid Cycle
Contribute to the Development of Cancer 354
19.5 The Glyoxylate Cycle Enables Plants and
Bacteria to Convert Fats into Carbohydrates 355
SECTION 9
Oxidative Phosphorylation 361
Chapter 20 The Electron-Transport Chain 363
20.1 Oxidative Phosphorylation in Eukaryotes
Takes Place in Mitochondria 364
Mitochondria Are Bounded by a Double Membrane 364
E3 Biological Insight Mitochondria Are the
Result of an Endosymbjotic Event 365
20.2 Oxidative Phosphorylation Depends on Electron
Transfer 366
The Electron-Transfer Potential of an Electron Is
Measured as Redox Potential 366
Electron Flow Through the Electron-Transport
Chain Creates a Proton Gradient 367
The Electron-Transport Chain Is a Series of
Coupled Oxidation-Reduction Reactions 368
Ü NEW Clinical Insight Loss of Iron-Sulfur Cluster
Results in Friedreich's Ataxia 371
20.3 The Respiratory Chain Consists of Proton
Pumps and a Physical Link to the Citric
Acid Cycle 371
The High-Potential Electrons of NADH Enter
the Respiratory Chain at NADH-Q Oxidoreductase 371
Ubiquinol Is the Entry Point for Electrons from
FADH2 of Flavoproteins 373
Electrons Flow from Ubiquinol to Cytochrome c
Through Q-Cytochrome c Oxidoreductase 373
The Q Cycle Funnels Electrons from a Two-Electron
Carrier to a One-Electron Carrier and Pumps Protons 374
Cytochrome c Oxidase Catalyzes the Reduction of
Molecular Oxygen to Water 375
Ü3 Biological Insight The Dead /one: Too Much
Respiration 377
Toxic Derivatives of Molecular Oxygen Such As Superoxide
Radical Are Scavenged by Protective Enzymes 377
Chapter 21 The Proton Motive Force 383
21.1 A Proton Gradient Powers the Synthesis of ATP 384
ATP Synthase Is Composed of a Proton-Conducting
Unit and a Catalytic Unit 385
Proton Flow Through AFP Synthase Leads to the
Release of Tightly Bound ATP 386
Rotational Catalysis Is the World's Smallest
Molecular Motor 387
Proton Flow Around the c Ring Powers ATP
Synthesis 388
21.2 Shuttles Allow Movement Across
Mitochondrial Membranes 390
Electrons from Cytoplasmic NADH Enter
Mitochondria by Shuttles 390
The Entry of ADP into Mitochondria Is Coupled to
the Exit of ATP 392
Mitochondrial Transporters Allow Metabolite Exchange
Between the Cytoplasm and Mitochondria 393
21.3 Cellular Respiration Is Regulated by
the Need for ATP 393
The Complete Oxidation of Glucose Yields About
30 Molecules of ATP 393
The Rate of Oxidative Phosphorylation Is Determined
by the Need for ATP 395
O NEW Clinical Insight ATP Synthase Can Be
Regulated 395
El Biological Insight Regulated Uncoupling Leads to
the Generation of Heat 396
O Clinical Insight Oxidative Phosphorylation Can Be
Inhibited at Many Stages 398
H Clinical Insight Mitochondrial Diseases Are Being
Discovered in Increasing Numbers 399
Power Transmission by Proton Gradients Is a
Central Motif of Bioenergetics 400
XXli Contents
SECTION 10
The Light Reactions of Photosynthesis and
the Calvin Cycle 405
Chapter 22 The Light Reactions 407
22.1 Photosynthesis Takes Place in Chloroplasts 408
^ Biological Insight Chloroplasts, Like Mitochondria,
Arose from an Endosymbiotic Event 409
22.2 Photosynthesis Transforms Light Energy into
Chemical Energy 409
Chlorophyll Is the Primary Receptor in Most
Photosynthetic Systems 410
Light-Harvesting Complexes Enhance the Efficiency
of Photosynthesis 411
Q Biological Insight Chlorophyll in Potatoes
Suggests the Presence of a Toxin 413
22.3 Two Photosystems Generate a Proton
Gradient and NADPH 413
Photosystem I Uses L ight Energy to Generate Reduced
Ferredoxin, a Powerful Redudant 414
Photosystem H Transfers Electrons to
Photosystem I and Generates a Proton Gradient 415
Cytochrome Links Photosystem II to
Photosystem I 416
The Oxidation of Water Achieves Oxidation-Reduction
Balance and Contributes Protons to the Proton
Gradient 416
22.4 A Proton Gradient Drives ATP Synthesis 418
1 he ATP Synthase of Chloroplasts Closely Resembles
That of Mitochondria 418
NEW The Activity of Chloroplast ATP Synthase Is
Regulated 419
Cyclic Electron Flow Through Photosystem I Leads
to the Production of ATP Instead of NADPH 419
The Absorption of Eight Photons Yields One 02,
Two NADPH, and Three ATP Molecules 420
The Components of Photosynthesis Are
Highly Organized 421
Biological Insight Many Herbicides Inhibit the
Light Reactions of Photosynthesis 421
Chapter 23 The Calvin Cycle 427
23.1 The Calvin Cycle Synthesizes Hexoses from
Carbon Dioxide and Water 428
Carbon Dioxide Reacts with Ribuiose
1,5-bisphosphate to Form Two Molecules of
3-Pho5phoglycerate 429
Hexose Phosphates Are Made from
Phosphoglycerate, and Ribuiose 1,5-bisphosphate
Is Regenerated 430
Three Molecules of ATP and Two Molecules of
NADPH Are Used to Bring Carbon Dioxide to
the Level of a Hexose 430
El Biological Insight A Volcanic Eruption Can Affect
Photosynthesis Worldwide 432
Starch and Sucrose Are the Major Carbohydrate
Stores in Plants 433
Q Biological Insight Why Bread Becomes Stale:
The Role of Starch 434
23.2 The Calvin Cycle Is Regulated by the Environment 434
Thioredoxin Plays a Key Role in Regulating
the Calvin Cycle 435
Rubisco Also Catalyzes a Wasteful Oxygenase Reaction 436
The C4 Pathway of Tropical Plants Accelerates
Photosynthesis by Concentrating Carbon Dioxide 436
Crassulacean Acid Metabolism Permits Growth in Arid
Ecosystems 438
SECTION 11
Glycogen Metabolism and the Pentose
Phosphate Pathway 443
Chapter 24 Glycogen Degradation 445
24.1 Glycogen Breakdown Requires Several Enzymes 446
Phospborylase Cleaves Glycogen to Release Glucose
1-phosphate 446
A Debranching Enzyme Also Is Needed for
the Breakdown of Glycogen 447
Phosphoglucomutase Converts Glucose 1-phosphate
into Glucose 6-phosphate 448
Liver Contains Glucose 6-phosphatase,
a Hydrolytic Enzyme Absent from Muscle 448
24.2 Phospborylase Is Regulated by Allosteric
Interactions and Reversible Phosphorylation 449
Liver Phosphorylase Produces Glucose for Use by
OtherTissues 449
Muscle Phosphorylase Is Regulated by
the Intracellular Energy Charge 450
Biochemical Characteristics of Muscle Fiber Types Differ 451
NEW Phosphorylation Promotes the Conversion of
Phosphorylase b to Phosphorylase a 451
Phosphorylase Kinase Is Activated by
Phosphorylation and Calcium Ions 452
H Clinical Insight Hers Disease Is Due to a
Phosphorylase Deficiency 453
24.3 Epinephrine and Glucagon Signal
the Need for Glycogen Breakdown 455
C Proteins Transmit the Signal for the Initiation
of Glycogen Breakdown 453
Glycogen Breakdown Must Be RapidlyTurned
Off When Necessary 455
H Biological Insight Glycogen Depletion Coincides
with the Onset of Fatigue 455
Chapter 25 Glycogen Synthesis 459
25.1 Glycogen Is Synthesized and Degraded by
Different Pathways 459
UDP-Clucose Is an Activated Form of Glucose 460
Glycogen Synthase Catalyzes the Transfer of
Glucose from UDP-Giucose to a Growing Cham 460
A Branching Enzyme Forms Alpha֊!,6 Linkages 461
Glycogen Synthase Is the Key Regulatory Enzyme
in Glycogen Synthesis 461
Glycogen Is an Efficient Storage Form of Glucose 462
25.2 Metabolism in Context: Glycogen Breakdown
and Synthesis Are Reciprocally Regulated 462
Protein Phosphatase 1 Reverses the Regulatory
Effects of Kinases on Glycogen Metabolism 462
Insulin Stimulates Glycogen Synthesis by Inactivating
Glycogen Synthase Kinase 464
Contents XXIII
Glycogen Metabolism in the Liver Regulates
the Blood -Glucose Concentration
§H Clinical Insight Diabetes Mellitus Results from
Insulin Insufficiency and Glucagon Excess
H Clinical Insight A Biochemical Understanding of
Glycogen-Storage Diseases Is Possible
Chapter 26 The Pentose Phosphate
Pathway
26.1 The Pentose Phosphate Pathway Yields
NADPH and Five-Carbon Sugars
Two Molecules of NADPH Are Generated in
the Conversion of Glucose 6-phosphate into
Ribulose 5-phosphate
The Pentose Phosphate Pathway and Glycolysis
Are Linked by Transketolase and Transaldolase
26.2 Metabolism in Context: Glycolysis and
the Pentose Phosphate Pathway Are
Coordinately Controlled
The Rate of the Pentose Phosphate Pathway Is
Controlled by the Level of NADP+
The Fate of Glucose 6-phosphate Depends on
the Need for NADPH, Ribose 5-phosphate, and ATP
Į-jĮ NEW Clinical Insight The Pentose Phosphate
Pathway Is Required For Rapid Cell Growth
26.3 Glucose 6֊phosphate Dehydrogenase
Lessens Oxidative Stress
O Clinical Insight Glucose 6 phosphate.
Dehydrogenase Deficiency Causes a Drug-Induced
Hemolytic Anemia
B3 Biological Insight A Deficiency of Glucose
6֊phosphate Dehydrogenase Confers an
Evolutionary Advantage in Some Circumstances
SECTION 12
Fatty Acid and Lipid Metabolism
Chapter 27 Fatty Acid Degradation
27.1 Fatty Adds Are Processed in Three Stages
KB·
ip Clinical Insight Triacylglycerols Are Hydrolyzed
by Hormone-Stimulated Lipases
NEW Free Fatty Acids and Glycerol Are Released into
the Blood
Fatty Acids Are Linked to Coenzyme A Before
They Are Oxidized
lp Clinical Insight Pathological Conditions Result if
Fatty Acids Cannot Enter the Mitochondria
Acetyl CoĄ NADH, and FADH2Are Generated by
Fatty Acid Oxidation
The Complete Oxidation of Palmitate Yields
106 Molecules of ATP
27.2 The Degradation of Unsaturated and
Odd-Chain Fatty Acids Requires Additional Steps
An Isomerase and a Reductase Are Required for the
Oxidation of Unsaturated Fatty Adds
Odd-Chain Fatty Acids Yield Propionyl CoA in the
Final Thiolysis Step
27.3 Ketone Bodies Are Another Fuel Source
Derived from Fats
Ketone-Body Synthesis Takes Place in the Liver
NEW Clinical Insight Ketogenic Diets May Have
465 Therapeutic Properties 498
Animals Cannot Convert Fatty Acids into Glucose 498
466 27.4 Metabolism in Context: Fatty Acid Metabolism
467 Is a Source of Insight into Various Physiological States '499
473 H Clinical Insight Diabetes Can Lead to a Life-Threatening Excess of Ketone-Body Production 499
Ü Clinical Insight Ketone Bodies Are a Crucial Fuel Source During Starvation 500
474 11 NEW Clinical Insight Some Fatty Acids May Contribute
to the Development of Pathological Conditions 501
474
Chapter 28 Fatty Add Synthesis 507
474 28.1 Fatty Acid Synthesis Takes Place in Three Stages 507
Citrate Carries Acetyl Groups from Mitochondria to the Cytoplasm 508
478 Several Sources Supply NADPH for Fatty Acid Synthesis 508
478 The Formation of Malonyl CoA Is the Committed Step in Fatty Acid Synthesis 509
478 Fatty Add Synthesis Consists of a Senes of Condensation, Reduction, Dehydration, and
Reduction Reactions 510
481 The Synthesis of Palmitate Requires 8 Molecules of
481 Acetyl CoA, 14 Molecules of NADPH, and 7 Molecules of ATP 512
Fatty Acids Are Synthesized by a Multifunctional Enzyme Complex in Animals 512
481 j^j Clinical Insight Fatty Add Metabolism Is Altered in
Tumor Cells 513
4^ 03 U-i O Clinical Insight A Small Fatty Acid That Causes Big Problems 513
487 28.2 Additional Enzymes Elongate and Desaturate Fatty Acids 514
Membrane-Bound Enzymes Generate Unsaturated
489 Fatty Acids 514
489 Eicosanoid Hormones Are Derived from Polyunsaturated Fatty Acids 514
490 H Clinical Insight Aspirin Exerts Its Effects by Covalently Modifying a Key Enzyme 515
491 28.3 Acetyl CoA Carboxylase Is a Key Regulator of Fatty Acid Metabolism 516
491 Acetyl CoA Carboxylase Is Regulated by Conditions in the Cell 516
495 Acetyl CoA Carboxylase Is Regulated by a Variety of Hormones 516
493 28.4 Metabolism in Context: Ethanol Alters Energy Metabolism in the Liver 517
495 495 Chapter 29 Lipid Synthesis: Storage Lipids, Phospholipids, and Cholesterol 523
495 29.1 Phosphatidate Is a Precursor of Storage Lipids and Many Membrane Lipids 523
497 Triacylglycerol Is Synthesized from Phosphatidate in Two Steps 524
Phospholipid Synthesis Requires Activated Precursors 524
497 11 NEW Clinical Insight Phosphatidylcholine Is an
497 Abundant Phospholipid 526
xxiv Contents
Sphingoiipids Are Synthesized from Ceramide
Q Clinical Insight Gangliosides Serve as Binding
Sites for Pathogens
II Clinical Insight Disrupted Lipid Metabolism
Results in Respiratory Distress Syndrome and
Tay Sachs Disease
PhosphatidicAcid Phosphatase Is a Key
Regulatory Enzyme in Lipid Metabolism
29.2 Cholesterol Is Synthesized from Acetyl
Coenzyme A in Three Stages
The Synthesis of Mevalonate Initiates the Synthesis of
Cholesterol
Squalene (C30) Is Synthesized from Six Molecules of
Isopentenyl Pyrophosphate (C5)
Squalene Cyclizes to Form Cholesterol
29.3 The Regulation of Cholesterol Synthesis
Takes Place at Several Levels
29.4 Lipoproteins Transport Cholesterol and
Triacylglycerols Throughout the Organism
Low-Density Lipoproteins Play a Central Role in
Cholesterol Metabolism
Clinical Insight The Absence of the LDL Receptor
Leads to Familial Hypercholesterolemia and
Atherosclerosis
^1 NEW Clinical Insight Cycling of the LDL Receptor
Is Regulated
ff Clinical Insight HDL Seems to Protect Against
Atherosclerosis
m NEW Clinical Insight The Clinical Management of
Cholesterol Levels Can Be Understood at a
Biochemical Level
29.5 Cholesterol Is the Precursor of Steroid
Hormones
О NEW Clinical Insight Bile Salts Facilitate Lipid
Absorption
Steroid Hormones Are Crucial Signal Molecules
Vitamin D Is Derived from Cholesterol by
the Energy of Sunlight
Clinical Insight Vitamin D Is Necessary for Bone
Development
Q Clinical Insight Androgens Can Be Used to
Artificially Enhance Athletic Performance
Oxygen Atoms Are Added to Steroids by
Cytochrome P450 Monooxygenases
Metabolism in Context: Ethanol Also Is Processed by
the Cytochrome P450 System
SECTION 13
The Metabolism of Nitrogen-Containing
Molecules
Chapter 30 Amino Acid Degradation and
the Urea Cycle
30.1 Nitrogen Removal Is the First Step in
the Degradation of Amino Acids
Alpha-Amino Croups Are Converted into Ammonium
Ions by the Oxidative Deamination of Glutamate
Щ NEW Clinical insight Blood Levels of
Amonitransferases Serve a Diagnostic Function
NEW Serine and Threonine Can Be Directly Deaminated
Peripheral Tissues Transport Nitrogen to the Liver
30.2 Ammonium Ion Is Converted into Urea in Most
Terrestrial Vertebrates
NEW Carbamoyl Phosphate Synthetase Is the Key
Regulatory Enzyme for Urea Synthesis
NEW Carbamoyl Phosphate Reacts with Ornithine to Begin
Urea Cycle
The Urea Cycle Is Linked to Gluconeogenesis
il Clinical Insight Metabolism in Context:
Inherited Defects of the Urea Cycle Cause
Hyperammonemia
Biological Insight Hibernation Presents Nitrogen
Disposal Problems
Q Biological Insight Urea Is Not the Only Means of
Disposing of Excess Nitrogen
30.3 Carbon Atoms of Degraded Amino Acids
Emerge as Major Metabolic Intermediates
Pyruvate Is a Point of Entry into Metabolism
Oxaloacetate Is Another Point of Entry into
Metabolism
Alpha-Ketoglutarate Is Yet Another Point of Entry into
Metabolism
Succinyl Coenzyme A Is a Point of Entry for
Several Nonpolar Amino Acids
The Branched-Cham Amino Acids Yield Acetyl
Co enzyme Ą A ceto acetate, or Succinyl Co enzyme A
Oxygenases Are Required for the Degradation of
Aromatic Amino Acids
Methionine Is Degraded into Succinyl Coenzyme A
^1 Clinical Insight Inborn Errors of Metabolism
Can Disrupt Amino Acid Degradation
^1 NEW Clinical Insight Determining the Basis of the
Neurological Symptoms of Phenylketonuria
Is an Active Area of Research
Chapter 31 Amino Add Synthesis
31.1 The Nitrogenase Complex Fixes Nitrogen
The Molybdenum-Iron Cofactor of Nitrogenase
Binds and Reduces Atmospheric Nitrogen
Ammonium Ion Is Incorporated into an Amino
Acid Through Glutamate and Glutamine
31.2 Amino Acids Are Made from Intermediates
of Major Pathways
Human Beings Can Synthesize Some Amino Adds
but Must Obtain Others from the Diet
Some Amino Acids Can Be Made by Simple
Transamination Reactions
Serine, Cysteine, and Glycine Are Formed from
3֊Phosphoglycerate
H Clinical Insight Tetra hydrofolate Carries
Activated One-Carbon Units
S-Adenosylmethionme Is the Major Donor of
Methyl Croups
11 Clinical Insight High Homocysteine Levels
Correlate with Vascular Disease
31.3 Feedback Inhibition Regulates Amino Acid
Biosynthesis
The Committed Step Is the Common Site
of Regulation
Branched Pathways Require Sophisticated
Regulation
526
527
528
529
529
530
530
532
532
534
535
536
537
537
538
539
539
539
540
541
542
542
543
549
551
552
552
553
553
554
Chapter 32 Nucleotide Metabolism
32.1 An Overview of Nucleotide Biosynthesis and
Nomenclature
32.2 The Pyrimidine Ring Is Assembled and
Then Attached to a Ribose Sugar
CTP Is Formed by the Amination of UTP
Kinases Convert Nucleoside Monophosphates into
Nucleoside Triphosphates
0 NEW Clinical Insight Salvage Pathways Recycle
Pyrimidine Bases
32.3 The Purine Ring Is Assembled on Ribose
Phosphate
AMP and CMP Are Formed from IMP
0 Clinical Insight Enzymes of the Purine Synthesis
Pathway Are Associated with One Another in Vivo
Bases Can Be Recycled by Salvage Pathways
32.4 Ribonucleotides Are Reduced to
Deoxyribonucleotides
Thymidylate Is Formed by the Méthylation of
Deoxyuridylate
El Clinical Insight Several Valuable. Anticancer Drugs
Block the Synthesis of Thymidylate
32.5 Nucleotide Biosynthesis Is Regulated by
Feedback Inhibition
Pyrimidine Biosynthesis Is Regulated by Aspartate
Transcarbamoylase
The Synthesis of Purine Nucleotides is Controlled by
Feedback Inhibition at Several Sites
Q NEW Clinical Insight The Synthesis of
Deoxyribonucleotides Is Controlled by the
Regulation of Ribonucleotide Reductase
32.6 Disruptions in Nucleotide Metabolism Can
Cause Pathological Conditions
P Clinical Insight The Loss of Adenosine Deaminase
Activity Results in Severe Combined
Immunodeficiency
0 Clinical Insight Gout Is Induced by High Serum
Levels of Urate
0 Clinical Insight Lesch-Nyhan Syndrome Is a
Dramatic Consequence of Mutations in a
Salvage-Pathway Enzyme
0 Clinical Insight Folic Add Deficiency Promotes
Birth Defects Such As Spina Bifida
PART III
Synthesizing the Molecules of Life
Section 14
Nucleic Acid Structure and DNA Replication
Chapter 33 The Structure of Informational
Macromolecules: DNA and RNA
33.1 A Nucleic Acid Consists of Bases Linked to
a Sugar-Phosphate Backbone
DNA and RNA Differ in the Sugar Component and
One of the Bases
Nucleotides Are the Monomeric Units of Nucleic Acids
DNA Molecules Are Very Long and Eiave Directionality
Contents XXV
585 33.2 Nucleic Acid Strands Can Form a Double-Helical
Structure 611
586 The Double Helix Is Stabilized by Hydrogen
Bonds and the Hydrophobic Effect 611
587 The Double Helix Facilitates the Accurate
589 Transmission of Hereditary Information Meselson and Stahl Demonstrated That Replication Is 613
589 Semiconservative The Strands of the Double Helix Can Be Reversibly 614
589 Separated 615
33.3 DNA Double Helices Can Adopt Multiple Forms Z-DNA Is a Left-Handed Double Helix in Which 615
590 Backbone Phosphoryl Groups Zigzag 616
590 The Major and Minor Grooves Are Lined by
592 Sequence-Specific Hydrogen-Bonding Croups 616
Double-Stranded DNA Can Wrap Around Itself to Form
593 Supercoiled Structures 33.4 Eukaryotic DNA Is Associated with Specific 617
593 Proteins 619
Nucleosomes Are Complexes of DNA and Histones 619
594 Eukaryotic DNA Is Wrapped Around Histones to Form
Nucleosomes 620
595 ЕЯ Clinical Insight Damaging DNA Can Inhibit
Cancer-Cell Growth 622
596 33.5 RNA Can Adopt Elaborate Structures 622
596 Chapter 34 DNA Replication 627
596 34.1 DNA Is Replicated by Polymerases 628
DNA Polymerase Catalyzes Phosphodiester-Linkage
597 Formation The Specificity of Replication Is Dictated by the 628
Complementarity of Bases Ol Clinical Insight The Separation of DNA Strands 630
598 Requires Specific Helicases and ATP Hydrolysis Topoisomerases Prepare the Double Helix for 630
Unwinding 632
598 0 Clinical Insight Bacterial Topoisomerase Is a
599 Therapeutic Target 632
Many Polymerases Proofread the Newly Added
Bases and Excise Errors 633
600 600 34.2 DNA Replication Is Highly Coordinated 633
DNA Replication in E. coll Begins at a Unique Site An RNA Primer Synthesized by Primase Enables DNA 634
Synthesis to Begin One Strand of DNA Is Made Continuously and 634
the Other Strand Is Synthesized in Fragments DNA Replication Requires Highly Processive 635
Polymerases The Leading and l aggingStrands Are Synthesized in a 635
605 Coordinated Fashion DNA Synthesis Is More Complex in Eukaryotes 636
607 Than in Bacteria Telomeres Are Unique Structures at the Ends of 638
Linear Chromosomes 0| Clinical Insight Telomeres Are Replicated by 638
608 Telomerase, a Specialized Polymerase That
608 Carries Its Own RNA Template 639
609 Chapter 35 DNA Repair and Recombination 643
610 35.1 Errors Can Arise in DNA Replication 644
XXVI Contents
El Clinical Insight Some Genetic Diseases Are Caused
by the Expansion of Repeats of Three Nucleotides
Bases Can Be Damaged by Oxidizing Agents,
Alkylating Agents, and Light
35.2 DNA Damage Can Be Detected and Repaired
The Presence of Thymine Instead of Uracil in
DNA Permits the Repair of Deaminated Cytosine
Clinical Insight Many Cancers Are Caused by
the Defective Repair of DNA
11 Clinical Insight Many Potential Carcinogens
Can Be Detected by Their Mutagenic Action
on Bacteria
35.3 DNA Recombination Plays Important Roles in
Replication and Repair
Double Strand Breaks Can Be Repaired by
Recombination
DNA Recombination Is Important in a Variety of
Biological Processes
SECTION 15
RNA Synthesis, Processing, and Regulation
Chapter 36 RNA Synthesis and Regulation
in Bacteria
36.1 Cellular RNA Is Synthesized by RNA Polymerases
Genes Are the Transcriptional Units
RNA Polymerase Is Composed of Multiple Subunits
36.2 RNA Synthesis Comprises Three Stages
Transcription Is Initiated at Promoter Sites on the DNA
Template
Sigma Subunits of RNA Polymerase Recognize
Promoter Sites
RNA Strands Grow in the 5'--to-3' Direction
Elongation Takes Place at Transcription Bubbles
That Move Along the DNA Template
An RNA Hairpin Followed by Several Uracil Residues
Terminates the Transcription of Some Genes
The Rho Protein Helps Terminate the Transcription of
Some Genes
Precursors of Transfer and Ribosomal RNA Are
Cleaved and Chemically Modified After Transcription
61 Clinical Insight Some Antibiotics Inhibit
Transcription
563 The lac Operon Illustrates the Control of
Bacterial Gene Expression
An Operon Consists of Regulatory Elements and
Protein-Encoding Genes
Ligand Binding Can Induce Structural Changes in
Regulatory Proteins
Transcription Can Be Stimulated by Proteins
That Contact RNA Polymerase
Clinical and Biological Insight Many Bacterial Cells
Release Chemical Signals That Regulate Gene
Expression in Other Cells
Some Messenger RNAs Directly Sense Metabolite
Concentrations
Chapter 37 Gene Expression in Eukaryotes 675
644 37.1 Eukaryotic Cells Have Three Types of RNA
Polymerases 676
645 37.2 RNA Polymerase II Requires Complex
647 Regulation 678
649 The Transcription Factor IID Protein Complex Initiates the /\ssembiy of the Active Transcription Complex 679
650 Enhancer Sequences Can Stimulate Transcription at Start Sites Thousands of Bases Away 679
El Clinical Insight Inappropriate Enhancer Use May Cause Cancer 680
650 Multiple Transcription Factors Interact with Eukaryotic
Promoters and Enhancers 680
651 E Clinical Insight Induced Pluripotent Stem Cells
652 Can Be Generated by Introducing Four Transcription Factors into Differentiated Cells 680
373 Gene Expression Is Regulated by Hormones 681
652 Nuclear Hormone Receptors Have Similar Domain
Structures 681
Nuclear Hormone Receptors Recruit Coactivators and Co rep ressors 682
657 E Clinical Insight Steroid-Hormone Receptors Are Targets for Drugs 683
659 37.4 Histone Acetylation Results in Chromatin Remodeling 684
659 Metabolism in Context: Acetyl CoA Plays a Key Role in
660 the Regulation of Transcription 684
661 Histone Deacetylases Contribute to Transcriptional Repression 686
661 Chapter 38 RNA Processing in Eukaryotes 691
661 38.1 Mature Ribosomal RNA Is Generated by
the Cleavage of a Precursor Molecule 692
662 38.2 Transfer RNA Is Extensively Processed 692
663 383 Messenger RNA Is Modified and Spliced 693
664 Sequences at the Ends of Introns Specify Splice Sites in mRNA Precursors 694
664 Small Nuclear RNAs in Spliceosomes Catalyze the Splicing of mRNA Precursors 695
665 jrj Clinical Insight Mutations That Affect Pre-mRNA Splicing Cause Disease 696
666 Clinical Insight Most Human Pre-mRNAs Can Be Spliced in Alternative Ways to Yield Different
Proteins 697
667 The Transcription and Processing of mRNA
Are Coupled 698
668 E Biological Insight RNA Editing Changes
the Proteins Encoded by mRNA 698
668 38.4 RNA Can Function as a Catalyst 699
669 SECTION 16
669 Protein Synthesis and Recombinant DNA Techniques 705
Chapter 39 The Genetic Code 707
670 39.1 The Genetic Code Links Nucleic Acid and
Protein Information 708
670 The Genetic Code Is Nearly Universal 708
Contents
XXVII
Transfer RNA Molecules Have a Common Design
Some Transfer RNA Molecules Recognize More
Than One Codon Because of Wobble
in Base-Pairing
The Synthesis of Long Proteins Requires a Low Error
Frequency
39.2 Amino Acids Are Activated by Attachment to
Transfer RNA
Amino Acids Are First Activated by Adenylation
Aminoacyl-tRNA Synthetases Have Highly
Discriminating Ammo Acid Activation Sites
Proofreading byAminoacyf tRNA Synthetases
Increases the Fidelity of Protein Synthesis
Synthetases Recognize the Anticodon Loops and
Acceptor Stems of Transfer RNA Molecules
39.3 A Ribosome Is a Ribonucleoprotein Particle
Made of Two Subunits
Ribosomai RNAs Play a Central Role in Protein
Synthesis
Messenger RNA Is Translated in the 5Чо-3'
Direction
Chapter 40 The Mechanism of
Protein Synthesis
40.1 Protein Synthesis Decodes the Information in
Messenger RNA
Ribosomes Have Three tRNA-Binding Sites That
Bridge the 30S and SOS Subunits
The Start Signal Is AUG Preceded by
Several Bases That Pair with 16S Ribosomai RNA
Bacterial Protein Synthesis Is Initiated by
Formylmethionyl Transfer RNA
Formylmethionyl-tRNAf Is Placed in the PSite of
the Ribosome in the Formation of the 70S Initiation
Complex
Elongation Factors Deliver Aminoacyl-tRNA to
the Ribosome
40.2 Peptidy! Transferase Catalyzes Peptide-Bond
Synthesis
The Formation of a Peptide Bond Is Followed by the
GTP-Driven Translocation of tRNAs and mRNA
Protein Synthesis Is Terminated by Release Factors
That Read Stop Codons
40.3 Bacteria and Eukaryotes Differ in the Initiation
of Protein Synthesis
Ж Clinical Insight Mutations in Initiation Factor 2
Cause a Curious Pathological Condition
40.4 A Variety of Biomolecules Can Inhibit Protein
Synthesis
Ж Clinical Insight Some Antibiotics Inhibit Protein
Synthesis
Ж Clinical Insight Diphtheria Toxin Blocks Protein
Synthesis in Eukaryotes by Inhibiting Translocation
Ц Clinical Insight Ricin Fatally Modifies
28S Ribosomai RNA
40.5 Ribosomes Bound to the Endoplasmic Reticulum
Manufacture Secretory and Membrane Proteins
Protein Synthesis Begins on Ribosomes That Are
Free in the Cytoplasm 733
Signal Sequences Mark Proteins for translocation
Across the. Endoplasmic Reticulum Membrane 733
40.6 Protein Synthesis Is Regulated by a Number of
Mechanisms 735
Messenger RNA Use Is Subject to Regulation 735
The Stability of Messenger RNA Also Can Be
Regulated 736
Small RNAs Can Regulate mRNA Stability and Use 736
Chapter 41 Recombinant DNA Techniques 743
41.1 Nucleic Acids Can Be Synthesized from
Protein-Sequence Data 744
Protein Sequence Is a Cu.de to NudeicAcid
Information 744
DNA Probes Can Be Synthesized by Automated
Methods 744
41.2 Recombinant DNA Technology Has
Revolutionized All Aspects of Biology 745
Restriction Enzymes Split DNA into Specific
Fragments 745
Restriction Fragments Can Be Separated by Cel
Electrophoresis and Visualized 746
Restriction Enzymes and DNA Ligase Are Key Tools for
Forming Recombinant DNA Molecules 747
41.3 Eukaryotic Genes Can Be Manipulated with
Considerable Precision 748
Complementary DNA Prepared from mRNA Can Be
Expressed in Host Cells 748
Estrogen-Receptor cDNA Can Be Identified by
Screening a cDNA Library 749
Complementary DNA Libraries Can Be Screened for
Synthesized Protein 750
Specific Genes Can Be Cloned from Digests of
Genomic DNA 750
DNA Can Be Sequenced by the Controlled
Termination of Replication 751
Ж Clinical and Biological Insight Next-Generation
Sequencing Methods Enable the Rapid
Determination of a Complete Genome Sequence 753
Selected DNA Sequences Can Be Greatly Amplified by
the Polymerase Chain Reaction 754
Ж Clinical and Biological Insight PCR Is a Powerful
Technique in Medical Diagnostics, Forensics, and
Studies of Molecular Evolution 756
Gene-Expression Levels Can Be Comprehensively
Examined 756
Appendices A1
Glossary B1
Answers to Problems Cl
Index D1
Selected Readings
(online atwww.whfreeman.com/tymoczko3e) El
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| any_adam_object | 1 |
| author | Tymoczko, John L. 1948-2019 Berg, Jeremy M. 1958- Stryer, Lubert 1938-2024 |
| author_GND | (DE-588)124601103 (DE-588)12460109X (DE-588)124601197 |
| author_facet | Tymoczko, John L. 1948-2019 Berg, Jeremy M. 1958- Stryer, Lubert 1938-2024 |
| author_role | aut aut aut |
| author_sort | Tymoczko, John L. 1948-2019 |
| author_variant | j l t jl jlt j m b jm jmb l s ls |
| building | Verbundindex |
| bvnumber | BV042382369 |
| classification_rvk | WD 4010 |
| classification_tum | CHE 800f |
| ctrlnum | (OCoLC)906074493 (DE-599)BVBBV042382369 |
| dewey-full | 572 |
| dewey-hundreds | 500 - Natural sciences and mathematics |
| dewey-ones | 572 - Biochemistry |
| dewey-raw | 572 |
| dewey-search | 572 |
| dewey-sort | 3572 |
| dewey-tens | 570 - Biology |
| discipline | Biologie Chemie |
| edition | 3. ed. |
| format | Book |
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| indexdate | 2024-08-06T00:21:51Z |
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| spelling | Tymoczko, John L. 1948-2019 Verfasser (DE-588)124601103 aut Biochemistry a short course John L. Tymoczko ; Jeremy M. Berg ; Lubert Stryer 3. ed. New York, NY Freeman 2015 Getr. Zählung Ill., graph. Darst. txt rdacontent n rdamedia nc rdacarrier Biochemie (DE-588)4006777-4 gnd rswk-swf 1\p (DE-588)4123623-3 Lehrbuch gnd-content Biochemie (DE-588)4006777-4 s DE-604 Berg, Jeremy M. 1958- Verfasser (DE-588)12460109X aut Stryer, Lubert 1938-2024 Verfasser (DE-588)124601197 aut Digitalisierung UB Regensburg - ADAM Catalogue Enrichment application/pdf http://bvbr.bib-bvb.de:8991/F?func=service&doc_library=BVB01&local_base=BVB01&doc_number=027818424&sequence=000002&line_number=0001&func_code=DB_RECORDS&service_type=MEDIA Inhaltsverzeichnis 1\p cgwrk 20201028 DE-101 https://d-nb.info/provenance/plan#cgwrk |
| spellingShingle | Tymoczko, John L. 1948-2019 Berg, Jeremy M. 1958- Stryer, Lubert 1938-2024 Biochemistry a short course Biochemie (DE-588)4006777-4 gnd |
| subject_GND | (DE-588)4006777-4 (DE-588)4123623-3 |
| title | Biochemistry a short course |
| title_auth | Biochemistry a short course |
| title_exact_search | Biochemistry a short course |
| title_full | Biochemistry a short course John L. Tymoczko ; Jeremy M. Berg ; Lubert Stryer |
| title_fullStr | Biochemistry a short course John L. Tymoczko ; Jeremy M. Berg ; Lubert Stryer |
| title_full_unstemmed | Biochemistry a short course John L. Tymoczko ; Jeremy M. Berg ; Lubert Stryer |
| title_short | Biochemistry |
| title_sort | biochemistry a short course |
| title_sub | a short course |
| topic | Biochemie (DE-588)4006777-4 gnd |
| topic_facet | Biochemie Lehrbuch |
| url | http://bvbr.bib-bvb.de:8991/F?func=service&doc_library=BVB01&local_base=BVB01&doc_number=027818424&sequence=000002&line_number=0001&func_code=DB_RECORDS&service_type=MEDIA |
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