Frontmatter

About the Authors

Molecular Cell Biology, Eighth Edition

Preface

Media and Supplements

Acknowledgments

Chapter 1: Molecules, Cells, and Model Organisms

Chapter Introduction

1.1 The Molecules of Life

Proteins Give Cells Structure and Perform Most Cellular Tasks

Nucleic Acids Carry Coded Information for Making Proteins at the Right Time and Place

Phospholipids Are the Conserved Building Blocks of All Cellular Membranes

1.2 Prokaryotic Cell Structure and Function

Prokaryotes Comprise Two Kingdoms: Archaea and Eubacteria

Escherichia coli Is Widely Used in Biological Research

1.3 Eukaryotic Cell Structure and Function

The Cytoskeleton Has Many Important Functions

The Nucleus Contains the DNA Genome, RNA Synthetic Apparatus, and a Fibrous Matrix

Eukaryotic Cells Contain a Large Number of Internal Membrane Structures

Mitochondria Are the Principal Sites of ATP Production in Aerobic Cells

Chloroplasts Contain Internal Compartments in Which Photosynthesis Takes Place

All Eukaryotic Cells Use a Similar Cycle to Regulate Their Division

1.4 Unicellular Eukaryotic Model Organisms

Yeasts Are Used to Study Fundamental Aspects of Eukaryotic Cell Structure and Function

Mutations in Yeast Led to the Identification of Key Cell Cycle Proteins

Studies in the Alga Chlamydomonas reinhardtii Led to the Development of a Powerful Technique to Study Brain Function

The Parasite That Causes Malaria Has Novel Organelles That Allow It to Undergo a Remarkable Life Cycle

1.5 Metazoan Structure, Differentiation, and Model Organisms

Multicellularity Requires Cell-Cell and Cell-Matrix Adhesions

Epithelia Originated Early in Evolution

Tissues Are Organized into Organs

Genomics Has Revealed Important Aspects of Metazoan Evolution and Cell Function

Embryonic Development Uses a Conserved Set of Master Transcription Factors

Planaria Are Used to Study Stem Cells and Tissue Regeneration

Invertebrates, Fish, Mice, and Other Organisms Serve as Experimental Systems for Study of Human Development and Disease

Genetic Diseases Elucidate Important Aspects of Cell Function

The Following Chapters Present Much Experimental Data That Explains How We Know What We Know About Cell Structure and Function

Chapter 2: Chemical Foundations

Chapter Introduction

2.1 Covalent Bonds and Noncovalent Interactions

The Electronic Structure of an Atom Determines the Number and Geometry of the Covalent Bonds It Can Make

Electrons May Be Shared Equally or Unequally in Covalent Bonds

Covalent Bonds Are Much Stronger and More Stable Than Noncovalent Interactions

Ionic Interactions Are Attractions Between Oppositely Charged Ions

Hydrogen Bonds Are Noncovalent Interactions That Determine the Water Solubility of Uncharged Molecules

Van der Waals Interactions Are Weak Attractive Interactions Caused by Transient Dipoles

The Hydrophobic Effect Causes Nonpolar Molecules to Adhere to One Another

Molecular Complementarity Due to Noncovalent Interactions Leads to a Lock-and-Key Fit Between Biomolecules

Key Concepts of Section 2.1

2.2 Chemical Building Blocks of Cells

Amino Acids Differing Only in Their Side Chains Compose Proteins

Five Different Nucleotides Are Used to Build Nucleic Acids

Monosaccharides Covalently Assemble into Linear and Branched Polysaccharides

Phospholipids Associate Noncovalently to Form the Basic Bilayer Structure of Biomembranes

Key Concepts of Section 2.2

2.3 Chemical Reactions and Chemical Equilibrium

A Chemical Reaction Is in Equilibrium When the Rates of the Forward and Reverse Reactions Are Equal

The Equilibrium Constant Reflects the Extent of a Chemical Reaction

Chemical Reactions in Cells Are at Steady State

Dissociation Constants of Binding Reactions Reflect the Affinity of Interacting Molecules

Biological Fluids Have Characteristic pH Values

Hydrogen Ions Are Released by Acids and Taken Up by Bases

Buffers Maintain the pH of Intracellular and Extracellular Fluids

Key Concepts of Section 2.3

2.4 Biochemical Energetics

Several Forms of Energy Are Important in Biological Systems

Cells Can Transform One Type of Energy into Another

The Change in Free Energy Determines If a Chemical Reaction Will Occur Spontaneously

The ΔG°′ of a Reaction Can Be Calculated from Its Keq

The Rate of a Reaction Depends on the Activation Energy Necessary to Energize the Reactants into a Transition State

Life Depends on the Coupling of Unfavorable Chemical Reactions with Energetically Favorable Ones

Hydrolysis of ATP Releases Substantial Free Energy and Drives Many Cellular Processes

ATP Is Generated During Photosynthesis and Respiration

NAD+ and FAD Couple Many Biological Oxidation and Reduction Reactions

Key Concepts of Section 2.4

Key Terms

Review the Concepts

References

Chapter 3: Protein Structure and Function

Chapter Introduction

3.1 Hierarchical Structure of Proteins

The Primary Structure of a Protein Is Its Linear Arrangement of Amino Acids

Secondary Structures Are the Core Elements of Protein Architecture

Tertiary Structure Is the Overall Folding of a Polypeptide Chain

There Are Four Broad Structural Categories of Proteins

Different Ways of Depicting the Conformation of Proteins Convey Different Types of Information

Structural Motifs Are Regular Combinations of Secondary Structures

Domains Are Modules of Tertiary Structure

Multiple Polypeptides Assemble into Quaternary Structures and Supramolecular Complexes

Comparing Protein Sequences and Structures Provides Insight into Protein Function and Evolution

Key Concepts of Section 3.1

3.2 Protein Folding

Planar Peptide Bonds Limit the Shapes into Which Proteins Can Fold

The Amino Acid Sequence of a Protein Determines How It Will Fold

Folding of Proteins in Vivo Is Promoted by Chaperones

Protein Folding Is Promoted by Proline Isomerases

Abnormally Folded Proteins Can Form Amyloids That Are Implicated in Diseases

Key Concepts of Section 3.2

3.3 Protein Binding and Enzyme Catalysis

Specific Binding of Ligands Underlies the Functions of Most Proteins

Enzymes Are Highly Efficient and Specific Catalysts

An Enzyme’s Active Site Binds Substrates and Carries Out Catalysis

Serine Proteases Demonstrate How an Enzyme’s Active Site Works

Enzymes in a Common Pathway Are Often Physically Associated with One Another

Key Concepts of Section 3.3

3.4 Regulating Protein Function

Regulated Synthesis and Degradation of Proteins Is a Fundamental Property of Cells

The Proteasome Is a Molecular Machine Used to Degrade Proteins

Ubiquitin Marks Cytosolic Proteins for Degradation in Proteasomes

Noncovalent Binding Permits Allosteric, or Cooperative, Regulation of Proteins

Noncovalent Binding of Calcium and GTP Are Widely Used as Allosteric Switches to Control Protein Activity

Phosphorylation and Dephosphorylation Covalently Regulate Protein Activity

Ubiquitinylation and Deubiquitinylation Covalently Regulate Protein Activity

Proteolytic Cleavage Irreversibly Activates or Inactivates Some Proteins

Higher-Order Regulation Includes Control of Protein Location

Key Concepts of Section 3.4

3.5 Purifying, Detecting, and Characterizing Proteins

Centrifugation Can Separate Particles and Molecules That Differ in Mass or Density

Electrophoresis Separates Molecules on the Basis of Their Charge-to-Mass Ratio

Liquid Chromatography Resolves Proteins by Mass, Charge, or Affinity

Highly Specific Enzyme and Antibody Assays Can Detect Individual Proteins

Radioisotopes Are Indispensable Tools for Detecting Biological Molecules

Mass Spectrometry Can Determine the Mass and Sequence of Proteins

Protein Primary Structure Can Be Determined by Chemical Methods and from Gene Sequences

Protein Conformation Is Determined by Sophisticated Physical Methods

Key Concepts of Section 3.5

3.6 Proteomics

Proteomics Is the Study of All or a Large Subset of Proteins in a Biological System

Advanced Techniques in Mass Spectrometry Are Critical to Proteomic Analysis

Key Concepts of Section 3.6

Key Terms

Review the Concepts

Extended References

Perspectives for the Future

Chapter 4: Culturing and Visualizing Cells

Chapter Introduction

4.1 Growing and Studying Cells in Culture

Culture of Animal Cells Requires Nutrient-Rich Media and Special Solid Surfaces

Primary Cell Cultures and Cell Strains Have a Finite Life Span

Transformed Cells Can Grow Indefinitely in Culture

Flow Cytometry Separates Different Cell Types

Growth of Cells in Two-Dimensional and Three-Dimensional Culture Mimics the In Vivo Environment

Hybridomas Produce Abundant Monoclonal Antibodies

A Wide Variety of Cell Biological Processes Can Be Studied with Cultured Cells

Drugs Are Commonly Used in Cell Biological Research

Key Concepts of Section 4.1

4.2 Light Microscopy: Exploring Cell Structure and Visualizing Proteins Within Cells

The Resolution of the Conventional Light Microscope Is About 0.2 µm

Phase-Contrast and Differential-Interference-Contrast Microscopy Visualize Unstained Live Cells

Imaging Subcellular Details Often Requires That Specimens Be Fixed, Sectioned, and Stained

Fluorescence Microscopy Can Localize and Quantify Specific Molecules in Live Cells

Intracellular Ion Concentrations Can Be Determined with Ion-Sensitive Fluorescent Dyes

Immunofluorescence Microscopy Can Detect Specific Proteins in Fixed Cells

Tagging with Fluorescent Proteins Allows the Visualization of Specific Proteins in Live Cells

Deconvolution and Confocal Microscopy Enhance Visualization of Three-Dimensional Fluorescent Objects

Two-Photon Excitation Microscopy Allows Imaging Deep into Tissue Samples

TIRF Microscopy Provides Exceptional Imaging in One Focal Plane

FRAP Reveals the Dynamics of Cellular Components

FRET Measures Distance Between Fluorochromes

Super-Resolution Microscopy Can Localize Proteins to Nanometer Accuracy

Light-Sheet Microscopy Can Rapidly Image Cells in Living Tissue

Key Concepts of Section 4.2

4.3 Electron Microscopy: High-Resolution Imaging

Single Molecules or Structures Can Be Imaged Using a Negative Stain or Metal Shadowing

Cells and Tissues Are Cut into Thin Sections for Viewing by Electron Microscopy

Immunoelectron Microscopy Localizes Proteins at the Ultrastructural Level

Cryoelectron Microscopy Allows Visualization of Specimens Without Fixation or Staining

Scanning Electron Microscopy of Metal-Coated Specimens Reveals Surface Features

Key Concepts of Section 4.3

4.4 Isolation of Cell Organelles

Disruption of Cells Releases Their Organelles and Other Contents

Centrifugation Can Separate Many Types of Organelles

Organelle-Specific Antibodies Are Useful in Preparing Highly Purified Organelles

Proteomics Reveals the Protein Composition of Organelles

Key Concepts of Section 4.4

Key Terms

Review the Concepts

Extended References

Perspectives for the Future

Classic Experiment 4-1: Separating Organelles

Chapter 5: Fundamental Molecular Genetic Mechanisms

Chapter Introduction

5.1 Structure of Nucleic Acids

A Nucleic Acid Strand Is a Linear Polymer with End-to-End Directionality

Native DNA Is a Double Helix of Complementary Antiparallel Strands

DNA Can Undergo Reversible Strand Separation

Torsional Stress in DNA Is Relieved by Enzymes

Different Types of RNA Exhibit Various Conformations Related to Their Functions

Key Concepts of Section 5.1

5.2 Transcription of Protein-Coding Genes and Formation of Functional mRNA

A Template DNA Strand Is Transcribed into a Complementary RNA Chain by RNA Polymerase

Organization of Genes Differs in Prokaryotic and Eukaryotic DNA

Eukaryotic Precursor mRNAs Are Processed to Form Functional mRNAs

Alternative RNA Splicing Increases the Number of Proteins Expressed from a Single Eukaryotic Gene

Key Concepts of Section 5.2

5.3 The Decoding of mRNA by tRNAs

Messenger RNA Carries Information from DNA in a Three-Letter Genetic Code

The Folded Structure of tRNA Promotes Its Decoding Functions

Nonstandard Base Pairing Often Occurs Between Codons and Anticodons

Amino Acids Become Activated When Covalently Linked to tRNAs

Key Concepts of Section 5.3

5.4 Stepwise Synthesis of Proteins on Ribosomes

Ribosomes Are Protein-Synthesizing Machines

Methionyl-tRNAiMet Recognizes the AUG Start Codon

Eukaryotic Translation Initiation Usually Occurs at the First AUG Downstream from the 5′ End of an mRNA

During Chain Elongation Each Incoming Aminoacyl-tRNA Moves Through Three Ribosomal Sites

Translation Is Terminated by Release Factors When a Stop Codon Is Reached

Polysomes and Rapid Ribosome Recycling Increase the Efficiency of Translation

GTPase-Superfamily Proteins Function in Several Quality-Control Steps of Translation

Nonsense Mutations Cause Premature Termination of Protein Synthesis

Key Concepts of Section 5.4

5.5 DNA Replication

DNA Polymerases Require a Primer to Initiate Replication

Duplex DNA Is Unwound, and Daughter Strands Are Formed at the DNA Replication Fork

Several Proteins Participate in DNA Replication

DNA Replication Occurs Bidirectionally from Each Origin

Key Concepts of Section 5.5

5.6 DNA Repair and Recombination

DNA Polymerases Introduce Copying Errors and Also Correct Them

Chemical and Radiation Damage to DNA Can Lead to Mutations

High-Fidelity DNA Excision-Repair Systems Recognize and Repair Damage

Base Excision Repairs T-G Mismatches and Damaged Bases

Mismatch Excision Repairs Other Mismatches and Small Insertions and Deletions

Nucleotide Excision Repairs Chemical Adducts that Distort Normal DNA Shape

Two Systems Use Recombination to Repair Double-Strand Breaks in DNA

Homologous Recombination Can Repair DNA Damage and Generate Genetic Diversity

Key Concepts of Section 5.6

5.7 Viruses: Parasites of the Cellular Genetic System

Most Viral Host Ranges Are Narrow

Viral Capsids Are Regular Arrays of One or a Few Types of Protein

Viruses Can Be Cloned and Counted in Plaque Assays

Lytic Viral Growth Cycles Lead to Death of Host Cells

Viral DNA Is Integrated into the Host-Cell Genome in Some Nonlytic Viral Growth Cycles

Key Concepts of Section 5.7

Key Terms

Review the Concepts

Extended References

Perspectives for the Future

Chapter 6: Molecular Genetic Techniques

Chapter Introduction

6.1 Genetic Analysis of Mutations to Identify and Study Genes

Recessive and Dominant Mutant Alleles Generally Have Opposite Effects on Gene Function

Segregation of Mutations in Breeding Experiments Reveals Their Dominance or Recessivity

Conditional Mutations Can Be Used to Study Essential Genes in Yeast

Recessive Lethal Mutations in Diploids Can Be Identified by Inbreeding and Maintained in Heterozygotes

Complementation Tests Determine Whether Different Recessive Mutations Are in the Same Gene

Double Mutants Are Useful in Assessing the Order in Which Proteins Function

Genetic Suppression and Synthetic Lethality Can Reveal Interacting or Redundant Proteins

Genes Can Be Identified by Their Map Position on the Chromosome

Key Concepts of Section 6.1

6.2 DNA Cloning and Characterization

Restriction Enzymes and DNA Ligases Allow Insertion of DNA Fragments into Cloning Vectors

Isolated DNA Fragments Can Be Cloned into E. coli Plasmid Vectors

Yeast Genomic Libraries Can Be Constructed with Shuttle Vectors and Screened by Functional Complementation

cDNA Libraries Represent the Sequences of Protein-Coding Genes

The Polymerase Chain Reaction Amplifies a Specific DNA Sequence from a Complex Mixture

Cloned DNA Molecules Can Be Sequenced Rapidly by Methods Based on PCR

Key Concepts of Section 6.2

6.3 Using Cloned DNA Fragments to Study Gene Expression

Hybridization Techniques Permit Detection of Specific DNA Fragments and mRNAs

DNA Microarrays Can Be Used to Evaluate the Expression of Many Genes at One Time

Cluster Analysis of Multiple Expression Experiments Identifies Co-regulated Genes

E. coli Expression Systems Can Produce Large Quantities of Proteins from Cloned Genes

Plasmid Expression Vectors Can Be Designed for Use in Animal Cells

Key Concepts of Section 6.3

6.4 Locating and Identifying Human Disease Genes

Monogenic Diseases Show One of Three Patterns of Inheritance

DNA Polymorphisms Are Used as Markers for Linkage Mapping of Human Mutations

Linkage Studies Can Map Disease Genes with a Resolution of About 1 Centimorgan

Further Analysis Is Needed to Locate a Disease Gene in Cloned DNA

Many Inherited Diseases Result from Multiple Genetic Defects

Key Concepts of Section 6.4

6.5 Inactivating the Function of Specific Genes in Eukaryotes

Normal Yeast Genes Can Be Replaced with Mutant Alleles by Homologous Recombination

Genes Can Be Placed Under the Control of an Experimentally Regulated Promoter

Specific Genes Can Be Permanently Inactivated in the Germ Line of Mice

Somatic Cell Recombination Can Inactivate Genes in Specific Tissues

Dominant-Negative Alleles Can Inhibit the Function of Some Genes

RNA Interference Causes Gene Inactivation by Destroying the Corresponding mRNA

Engineered CRISPR–Cas9 Systems Allow Precise Genome Editing

Key Concepts of Section 6.5

Key Terms

Review the Concepts

Extended References

Perspectives for the Future

Chapter 7: Biomembrane Structure

Chapter Introduction

7.1 The Lipid Bilayer: Composition and Structural Organization

Phospholipids Spontaneously Form Bilayers

Phospholipid Bilayers Form a Sealed Compartment Surrounding an Internal Aqueous Space

Biomembranes Contain Three Principal Classes of Lipids

Most Lipids and Many Proteins Are Laterally Mobile in Biomembranes

Lipid Composition Influences the Physical Properties of Membranes

Lipid Composition Is Different in the Exoplasmic and Cytosolic Leaflets

Cholesterol and Sphingolipids Cluster with Specific Proteins in Membrane Microdomains

Cells Store Excess Lipids in Lipid Droplets

Key Concepts of Section 7.1

7.2 Membrane Proteins: Structure and Basic Functions

Proteins Interact with Membranes in Three Different Ways

Most Transmembrane Proteins Have Membrane-Spanning α Helices

Multiple β Strands in Porins Form Membrane-Spanning “Barrels”

Covalently Attached Lipids Anchor Some Proteins to Membranes

All Transmembrane Proteins and Glycolipids Are Asymmetrically Oriented in the Bilayer

Lipid-Binding Motifs Help Target Peripheral Proteins to the Membrane

Proteins Can Be Removed from Membranes by Detergents or High-Salt Solutions

Key Concepts of Section 7.2

7.3 Phospholipids, Sphingolipids, and Cholesterol: Synthesis and Intracellular Movement

Fatty Acids Are Assembled from Two-Carbon Building Blocks by Several Important Enzymes

Small Cytosolic Proteins Facilitate Movement of Fatty Acids

Fatty Acids Are Incorporated into Phospholipids Primarily on the ER Membrane

Flippases Move Phospholipids from One Membrane Leaflet to the Opposite Leaflet

Cholesterol Is Synthesized by Enzymes in the Cytosol and ER Membrane

Cholesterol and Phospholipids Are Transported Between Organelles by Several Mechanisms

Key Concepts of Section 7.3

Key Terms

Review the Concepts

References

Perspectives for the Future

Chapter 8: Genes, Genomics, and Chromosomes

Chapter Introduction

8.1 Eukaryotic Gene Structure

Most Eukaryotic Genes Contain Introns and Produce mRNAs Encoding Single Proteins

Simple and Complex Transcription Units Are Found in Eukaryotic Genomes

Protein-Coding Genes May Be Solitary or Belong to a Gene Family

Heavily Used Gene Products Are Encoded by Multiple Copies of Genes

Nonprotein-Coding Genes Encode Functional RNAs

Key Concepts of Section 8.1

8.2 Chromosomal Organization of Genes and Noncoding DNA

Genomes of Many Organisms Contain Nonfunctional DNA

Most Simple-Sequence DNAs Are Concentrated in Specific Chromosomal Locations

DNA Fingerprinting Depends on Differences in Length of Simple-Sequence DNAs

Unclassified Intergenic DNA Occupies a Significant Portion of the Genome

Key Concepts of Section 8.2

8.3 Transposable (Mobile) DNA Elements

Movement of Mobile Elements Involves a DNA or an RNA Intermediate

DNA Transposons Are Present in Prokaryotes and Eukaryotes

LTR Retrotransposons Behave Like Intracellular Retroviruses

Non-LTR Retrotransposons Transpose by a Distinct Mechanism

Other Retroposed RNAs Are Found in Genomic DNA

Mobile DNA Elements Have Significantly Influenced Evolution

Key Concepts of Section 8.3

8.4 Genomics: Genome-Wide Analysis of Gene Structure and Function

Stored Sequences Suggest Functions of Newly Identified Genes and Proteins

Comparison of Related Sequences from Different Species Can Give Clues to Evolutionary Relationships Among Proteins

Genes Can Be Identified Within Genomic DNA Sequences

The Number of Protein-Coding Genes in an Organism’s Genome Is Not Directly Related to Its Biological Complexity

Key Concepts of Section 8.4

8.5 Structural Organization of Eukaryotic Chromosomes

Chromatin Exists in Extended and Condensed Forms

Modifications of Histone Tails Control Chromatin Condensation and Function

Nonhistone Proteins Organize Long Chromatin Loops

Additional Nonhistone Proteins Regulate Transcription and Replication

Key Concepts of Section 8.5

8.6 Morphology and Functional Elements of Eukaryotic Chromosomes

Chromosome Number, Size, and Shape at Metaphase Are Species-Specific

During Metaphase, Chromosomes Can Be Distinguished by Banding Patterns and Chromosome Painting

Chromosome Painting and DNA Sequencing Reveal the Evolution of Chromosomes

Interphase Polytene Chromosomes Arise by DNA Amplification

Three Functional Elements Are Required for Replication and Stable Inheritance of Chromosomes

Centromere Sequences Vary Greatly in Length and Complexity

Addition of Telomeric Sequences by Telomerase Prevents Shortening of Chromosomes

Key Concepts of Section 8.6

Key Terms

Review the Concepts

Extended References

Perspectives for the Future

Chapter 9: Transcriptional Control of Gene Expression

Chapter Introduction

9.1 Control of Gene Expression in Bacteria

Transcription Initiation by Bacterial RNA Polymerase Requires Association with a Sigma Factor

Initiation of lac Operon Transcription Can Be Repressed or Activated

Small Molecules Regulate Expression of Many Bacterial Genes via DNA-Binding Repressors and Activators

Transcription Initiation from Some Promoters Requires Alternative Sigma Factors

Transcription by σ54-RNA Polymerase Is Controlled by Activators That Bind Far from the Promoter

Many Bacterial Responses Are Controlled by Two-Component Regulatory Systems

Expression of Many Bacterial Operons Is Controlled by Regulation of Transcriptional Elongation

Key Concepts of Section 9.1

9.2 Overview of Eukaryotic Gene Control

Regulatory Elements in Eukaryotic DNA Are Found Both Close to and Many Kilobases Away from Transcription Start Sites

Three Eukaryotic RNA Polymerases Catalyze Formation of Different RNAs

The Largest Subunit in RNA Polymerase II Has an Essential Carboxy-Terminal Repeat

Key Concepts of Section 9.2

9.3 RNA Polymerase II Promoters and General Transcription Factors

RNA Polymerase II Initiates Transcription at DNA Sequences Corresponding to the 5′ Cap of mRNAs

The TATA Box, Initiators, and CpG Islands Function as Promoters in Eukaryotic DNA

General Transcription Factors Position RNA Polymerase II at Start Sites and Assist in Initiation

Elongation Factors Regulate the Initial Stages of Transcription in the Promoter-Proximal Region

Key Concepts of Section 9.3

9.4 Regulatory Sequences in Protein-Coding Genes and the Proteins Through Which They Function

Promoter-Proximal Elements Help Regulate Eukaryotic Genes

Distant Enhancers Often Stimulate Transcription by RNA Polymerase II

Most Eukaryotic Genes Are Regulated by Multiple Transcription-Control Elements

DNase I Footprinting and EMSA Detect Protein-DNA Interactions

Activators Are Composed of Distinct Functional Domains

Repressors Are the Functional Converse of Activators

DNA-Binding Domains Can Be Classified into Numerous Structural Types

Structurally Diverse Activation and Repression Domains Regulate Transcription

Transcription Factor Interactions Increase Gene-Control Options

Multiprotein Complexes Form on Enhancers

Key Concepts of Section 9.4

9.5 Molecular Mechanisms of Transcription Repression and Activation

Formation of Heterochromatin Silences Gene Expression at Telomeres, near Centromeres, and in Other Regions

Repressors Can Direct Histone Deacetylation at Specific Genes

Activators Can Direct Histone Acetylation at Specific Genes

Chromatin-Remodeling Complexes Help Activate or Repress Transcription

Pioneer Transcription Factors Initiate the Process of Gene Activation During Cellular Differentiation

The Mediator Complex Forms a Molecular Bridge Between Activation Domains and Pol II

Key Concepts of Section 9.5

9.6 Regulation of Transcription-Factor Activity

DNase I Hypersensitive Sites Reflect the Developmental History of Cellular Differentiation

Nuclear Receptors Are Regulated by Extracellular Signals

All Nuclear Receptors Share a Common Domain Structure

Nuclear-Receptor Response Elements Contain Inverted or Direct Repeats

Hormone Binding to a Nuclear Receptor Regulates Its Activity as a Transcription Factor

Metazoans Regulate the RNA Polymerase II Transition from Initiation to Elongation

Termination of Transcription Is Also Regulated

Key Concepts of Section 9.6

9.7 Epigenetic Regulation of Transcription

DNA Methylation Represses Transcription

Methylation of Specific Histone Lysines Is Linked to Epigenetic Mechanisms of Gene Repression

Epigenetic Control by Polycomb and Trithorax Complexes

Long Noncoding RNAs Direct Epigenetic Repression in Metazoans

Key Concepts of Section 9.7

9.8 Other Eukaryotic Transcription Systems

Transcription Initiation by Pol I and Pol III Is Analogous to That by Pol II

Key Concepts of Section 9.8

Key Terms

Review the Concepts

Extended References

Perspectives for the Future

Chapter 10: Post-transcriptional Gene Control

Chapter Introduction

10.1 Processing of Eukaryotic Pre-mRNA

The 5′ Cap Is Added to Nascent RNAs Shortly After Transcription Initiation

A Diverse Set of Proteins with Conserved RNA-Binding Domains Associate with Pre-mRNAs

Splicing Occurs at Short, Conserved Sequences in Pre-mRNAs via Two Transesterification Reactions

During Splicing, snRNAs Base-Pair with Pre-mRNA

Spliceosomes, Assembled from snRNPs and a Pre-mRNA, Carry Out Splicing

Chain Elongation by RNA Polymerase II Is Coupled to the Presence of RNA-Processing Factors

SR Proteins Contribute to Exon Definition in Long Pre-mRNAs

Self-Splicing Group II Introns Provide Clues to the Evolution of snRNAs

3′ Cleavage and Polyadenylation of Pre-mRNAs Are Tightly Coupled

Nuclear Exoribonucleases Degrade RNA That Is Processed Out of Pre-mRNAs

RNA Processing Solves the Problem of Pervasive Transcription of the Genome in Metazoans

Key Concepts of Section 10.1

10.2 Regulation of Pre-mRNA Processing

Alternative Splicing Generates Transcripts with Different Combinations of Exons

A Cascade of Regulated RNA Splicing Controls Drosophila Sexual Differentiation

Splicing Repressors and Activators Control Splicing at Alternative Sites

RNA Editing Alters the Sequences of Some Pre-mRNAs

Key Concepts of Section 10.2

10.3 Transport of mRNA Across the Nuclear Envelope

Phosphorylation and Dephosphorylation of SR Proteins Imposes Directionality on mRNP Export Across the Nuclear Pore Complex

Balbiani Rings in Insect Larval Salivary Glands Allow Direct Visualization of mRNP Export Through NPCs

Pre-mRNAs in Spliceosomes Are Not Exported from the Nucleus

HIV Rev Protein Regulates the Transport of Unspliced Viral mRNAs

Key Concepts of Section 10.3

10.4 Cytoplasmic Mechanisms of Post-transcriptional Control

Degradation of mRNAs in the Cytoplasm Occurs by Several Mechanisms

Adenines in mRNAs and lncRNAs May Be Post-transcriptionally Modified by N6 Methylation

Micro-RNAs Repress Translation and Induce Degradation of Specific mRNAs

Alternative Polyadenylation Increases miRNA Control Options

RNA Interference Induces Degradation of Precisely Complementary mRNAs

Cytoplasmic Polyadenylation Promotes Translation of Some mRNAs

Protein Synthesis Can Be Globally Regulated

Sequence-Specific RNA-Binding Proteins Control Translation of Specific mRNAs

Surveillance Mechanisms Prevent Translation of Improperly Processed mRNAs

Localization of mRNAs Permits Production of Proteins at Specific Regions Within the Cytoplasm

Key Concepts of Section 10.4

10.5 Processing of rRNA and tRNA

Pre-rRNA Genes Function as Nucleolar Organizers

Small Nucleolar RNAs Assist in Processing Pre-rRNAs

Self-Splicing Group I Introns Were the First Examples of Catalytic RNA

Pre-tRNAs Undergo Extensive Modification in the Nucleus

Nuclear Bodies Are Functionally Specialized Nuclear Domains

Key Concepts of Section 10.5

Key Terms

Review the Concepts

Extended References

Perspectives for the Future

Chapter 11: Transmembrane Transport of Ions and Small Molecules

Chapter Introduction

11.1 Overview of Transmembrane Transport

Only Gases and Small Uncharged Molecules Cross Membranes by Simple Diffusion

Three Main Classes of Membrane Proteins Transport Molecules and Ions Across Cellular Membranes

Key Concepts of Section 11.1

11.2 Facilitated Transport of Glucose and Water

Uniport Transport Is Faster and More Specific than Simple Diffusion

The Low Km of the GLUT1 Uniporter Enables It to Transport Glucose into Most Mammalian Cells

The Human Genome Encodes a Family of Sugar-Transporting GLUT Proteins

Transport Proteins Can Be Studied Using Artificial Membranes and Recombinant Cells

Osmotic Pressure Causes Water to Move Across Membranes

Aquaporins Increase the Water Permeability of Cellular Membranes

Key Concepts of Section 11.2

11.3 ATP-Powered Pumps and the Intracellular Ionic Environment

There Are Four Main Classes of ATP-Powered Pumps

ATP-Powered Ion Pumps Generate and Maintain Ionic Gradients Across Cellular Membranes

Muscle Relaxation Depends on Ca2+ ATPases That Pump Ca2+ from the Cytosol into the Sarcoplasmic Reticulum

The Mechanism of Action of the Ca2+ Pump Is Known in Detail

Calmodulin Regulates the Plasma-Membrane Pumps That Control Cytosolic Ca2+ Concentrations

The Na+/K+ ATPase Maintains the Intracellular Na+ and K+ Concentrations in Animal Cells

V-Class H+ ATPases Maintain the Acidity of Lysosomes and Vacuoles

ABC Proteins Export a Wide Variety of Drugs and Toxins from the Cell

Certain ABC Proteins “Flip” Phospholipids and Other Lipid-Soluble Substrates from One Membrane Leaflet to the Other

The ABC Cystic Fibrosis Transmembrane Regulator Is a Chloride Channel, Not a Pump

Key Concepts of Section 11.3

11.4 Nongated Ion Channels and the Resting Membrane Potential

Selective Movement of Ions Creates a Transmembrane Electric Gradient

The Resting Membrane Potential in Animal Cells Depends Largely on the Outward Flow of K+ Ions Through Open K+ Channels

Ion Channels Are Selective for Certain Ions by Virtue of a Molecular “Selectivity Filter”

Patch Clamps Permit Measurement of Ion Movements Through Single Channels

Novel Ion Channels Can Be Characterized by a Combination of Oocyte Expression and Patch Clamping

Key Concepts of Section 11.4

11.5 Cotransport by Symporters and Antiporters

Na+ Entry into Mammalian Cells Is Thermodynamically Favored

Na+-Linked Symporters Enable Animal Cells to Import Glucose and Amino Acids Against High Concentration Gradients

A Bacterial Na+/Amino Acid Symporter Reveals How Symport Works

A Na+-Linked Ca2+ Antiporter Regulates the Strength of Cardiac Muscle Contraction

Several Cotransporters Regulate Cytosolic pH

An Anion Antiporter Is Essential for Transport of CO2 by Erythrocytes

Numerous Transport Proteins Enable Plant Vacuoles to Accumulate Metabolites and Ions

Key Concepts of Section 11.5

11.6 Transcellular Transport

Multiple Transport Proteins Are Needed to Move Glucose and Amino Acids Across Epithelia

Simple Rehydration Therapy Depends on the Osmotic Gradient Created by Absorption of Glucose and Na+

Parietal Cells Acidify the Stomach Contents While Maintaining a Neutral Cytosolic pH

Bone Resorption Requires the Coordinated Function of a V-Class Proton Pump and a Specific Chloride Channel

Key Concepts of Section 11.6

Key Terms

Review the Concepts

Extended References

Perspectives for the Future

Classic Experiment 11-1: Stumbling upon Active Transport

Chapter 12: Cellular Energetics

Chapter Introduction

12.1 First Step of Harvesting Energy from Glucose: Glycolysis

During Glycolysis (Stage I), Cytosolic Enzymes Convert Glucose to Pyruvate

The Rate of Glycolysis Is Adjusted to Meet the Cell’s Need for ATP

Glucose Is Fermented When Oxygen Is Scarce

Key Concepts of Section 12.1

12.2 The Structure and Functions of Mitochondria

Mitochondria Are Multifunctional Organelles

Mitochondria Have Two Structurally and Functionally Distinct Membranes

Mitochondria Contain DNA Located in the Matrix

The Size, Structure, and Coding Capacity of mtDNA Vary Considerably Among Organisms

Products of Mitochondrial Genes Are Not Exported

Mitochondria Evolved from a Single Endosymbiotic Event Involving a Rickettsia-Like Bacterium

Mitochondrial Genetic Codes Differ from the Standard Nuclear Code

Mutations in Mitochondrial DNA Cause Several Genetic Diseases in Humans

Mitochondria Are Dynamic Organelles That Interact Directly with One Another

Mitochondria Are Influenced by Direct Contacts with the Endoplasmic Reticulum

Key Concepts of Section 12.2

12.3 The Citric Acid Cycle and Fatty Acid Oxidation

In the First Part of Stage II, Pyruvate Is Converted to Acetyl CoA and High-Energy Electrons

In the Second Part of Stage II, the Citric Acid Cycle Oxidizes the Acetyl Group in Acetyl CoA to CO2 and Generates High-Energy Electrons

Transporters in the Inner Mitochondrial Membrane Help Maintain Appropriate Cytosolic and Matrix Concentrations of NAD+ and NADH

Mitochondrial Oxidation of Fatty Acids Generates ATP

Peroxisomal Oxidation of Fatty Acids Generates No ATP

Key Concepts of Section 12.3

12.4 The Electron-Transport Chain and Generation of the Proton-Motive Force

Oxidation of NADH and FADH2 Releases a Significant Amount of Energy

Electron Transport in Mitochondria Is Coupled to Proton Pumping

Electrons Flow “Downhill” Through a Series of Electron Carriers

Four Large Multiprotein Complexes Couple Electron Transport to Proton Pumping Across the Inner Mitochondrial Membrane

The Reduction Potentials of Electron Carriers in the Electron-Transport Chain Favor Electron Flow from NADH to O2

The Multiprotein Complexes of the Electron-Transport Chain Assemble into Supercomplexes

Reactive Oxygen Species Are By-Products of Electron Transport

Experiments Using Purified Electron-Transport Chain Complexes Established the Stoichiometry of Proton Pumping

The Proton-Motive Force in Mitochondria Is Due Largely to a Voltage Gradient Across the Inner Membrane

Key Concepts of Section 12.4

12.5 Harnessing the Proton-Motive Force to Synthesize ATP

The Mechanism of ATP Synthesis Is Shared Among Bacteria, Mitochondria, and Chloroplasts

ATP Synthase Comprises F0 and F1 Multiprotein Complexes

Rotation of the F1 γ Subunit, Driven by Proton Movement Through F0, Powers ATP Synthesis

Multiple Protons Must Pass Through ATP Synthase to Synthesize One ATP

F0 c Ring Rotation Is Driven by Protons Flowing Through Transmembrane Channels

ATP-ADP Exchange Across the Inner Mitochondrial Membrane Is Powered by the Proton-Motive Force

The Rate of Mitochondrial Oxidation Normally Depends on ADP Levels

Mitochondria in Brown Fat Use the Proton-Motive Force to Generate Heat

Key Concepts of Section 12.5

12.6 Photosynthesis and Light-Absorbing Pigments

Thylakoid Membranes in Chloroplasts Are the Sites of Photosynthesis in Plants

Chloroplasts Contain Large DNAs Often Encoding More Than a Hundred Proteins

Three of the Four Stages in Photosynthesis Occur Only During Illumination

Photosystems Comprise a Reaction Center and Associated Light-Harvesting Complexes

Photoelectron Transport from Energized Reaction-Center Chlorophyll a Produces a Charge Separation

Internal Antennas and Light-Harvesting Complexes Increase the Efficiency of Photosynthesis

Key Concepts of Section 12.6

12.7 Molecular Analysis of Photosystems

The Single Photosystem of Purple Bacteria Generates a Proton-Motive Force but No O2

Chloroplasts Contain Two Functionally and Spatially Distinct Photosystems

Linear Electron Flow Through Both Plant Photosystems Generates a Proton-Motive Force, O2, and NADPH

An Oxygen-Evolving Complex Is Located on the Luminal Surface of the PSII Reaction Center

Multiple Mechanisms Protect Cells Against Damage from Reactive Oxygen Species During Photoelectron Transport

Cyclic Electron Flow Through PSI Generates a Proton-Motive Force but No NADPH or O2

Relative Activities of Photosystems I and II Are Regulated

Key Concepts of Section 12.7

12.8 CO2 Metabolism During Photosynthesis

Rubisco Fixes CO2 in the Chloroplast Stroma

Synthesis of Sucrose Using Fixed CO2 Is Completed in the Cytosol

Light and Rubisco Activase Stimulate CO2 Fixation

Photorespiration Competes with Carbon Fixation and Is Reduced in C4 Plants

Key Concepts of Section 12.8

Key Terms

Review the Concepts

Extended References

Perspectives for the Future

Chapter 13: Moving Proteins into Membranes and Organelles

Chapter Introduction

13.1 Targeting Proteins To and Across the ER Membrane

Pulse-Chase Experiments with Purified ER Membranes Demonstrated That Secreted Proteins Cross the ER Membrane

A Hydrophobic N-Terminal Signal Sequence Targets Nascent Secretory Proteins to the ER

Cotranslational Translocation Is Initiated by Two GTP-Hydrolyzing Proteins

Passage of Growing Polypeptides Through the Translocon Is Driven by Translation

ATP Hydrolysis Powers Post-translational Translocation of Some Secretory Proteins in Yeast

Key Concepts of Section 13.1

13.2 Insertion of Membrane Proteins into the ER

Several Topological Classes of Integral Membrane Proteins Are Synthesized on the ER

Internal Stop-Transfer Anchor and Signal-Anchor Sequences Determine Topology of Single-Pass Proteins

Multipass Proteins Have Multiple Internal Topogenic Sequences

A Phospholipid Anchor Tethers Some Cell-Surface Proteins to the Membrane

The Topology of a Membrane Protein Can Often Be Deduced from Its Sequence

Key Concepts of Section 13.2

13.3 Protein Modifications, Folding, and Quality Control in the ER

A Preformed N-Linked Oligosaccharide Is Added to Many Proteins in the Rough ER

Oligosaccharide Side Chains May Promote Folding and Stability of Glycoproteins

Disulfide Bonds Are Formed and Rearranged by Proteins in the ER Lumen

Chaperones and Other ER Proteins Facilitate Folding and Assembly of Proteins

Improperly Folded Proteins in the ER Induce Expression of Protein-Folding Catalysts

Unassembled or Misfolded Proteins in the ER Are Often Transported to the Cytosol for Degradation

Key Concepts of Section 13.3

13.4 Targeting of Proteins to Mitochondria and Chloroplasts

Amphipathic N-Terminal Targeting Sequences Direct Proteins to the Mitochondrial Matrix

Mitochondrial Protein Import Requires Outer-Membrane Receptors and Translocons in Both Membranes

Studies with Chimeric Proteins Demonstrate Important Features of Mitochondrial Protein Import

Three Energy Inputs Are Needed to Import Proteins into Mitochondria

Multiple Signals and Pathways Target Proteins to Submitochondrial Compartments

Import of Chloroplast Stromal Proteins Is Similar to Import of Mitochondrial Matrix Proteins

Proteins Are Targeted to Thylakoids by Mechanisms Related to Bacterial Protein Translocation

Key Concepts of Section 13.4

13.5 Targeting of Peroxisomal Proteins

A Cytosolic Receptor Targets Proteins with an SKL Sequence at the C-Terminus to the Peroxisomal Matrix

Peroxisomal Membrane and Matrix Proteins Are Incorporated by Different Pathways

Key Concepts of Section 13.5

13.6 Transport Into and Out of the Nucleus

Large and Small Molecules Enter and Leave the Nucleus via Nuclear Pore Complexes

Nuclear Transport Receptors Escort Proteins Containing Nuclear-Localization Signals into the Nucleus

A Second Type of Nuclear Transport Receptor Escorts Proteins Containing Nuclear-Export Signals Out of the Nucleus

Most mRNAs Are Exported from the Nucleus by a Ran-Independent Mechanism

Key Concepts of Section 13.6

Key Terms

Review the Concepts

References

Perspectives for the Future

Chapter 14: Vesicular Traffic, Secretion, and Endocytosis

Chapter Introduction

14.1 Techniques for Studying the Secretory Pathway

Transport of a Protein Through the Secretory Pathway Can Be Assayed in Live Cells

Yeast Mutants Define Major Stages and Many Components in Vesicular Transport

Cell-Free Transport Assays Allow Dissection of Individual Steps in Vesicular Transport

Key Concepts of Section 14.1

14.2 Molecular Mechanisms of Vesicle Budding and Fusion

Assembly of a Protein Coat Drives Vesicle Formation and Selection of Cargo Molecules

A Conserved Set of GTPase Switch Proteins Controls the Assembly of Different Vesicle Coats

Targeting Sequences on Cargo Proteins Make Specific Molecular Contacts with Coat Proteins

Rab GTPases Control Docking of Vesicles on Target Membranes

Paired Sets of SNARE Proteins Mediate Fusion of Vesicles with Target Membranes

Dissociation of SNARE Complexes After Membrane Fusion Is Driven by ATP Hydrolysis

Key Concepts of Section 14.2

14.3 Early Stages of the Secretory Pathway

COPII Vesicles Mediate Transport from the ER to the Golgi

COPI Vesicles Mediate Retrograde Transport Within the Golgi and from the Golgi to the ER

Anterograde Transport Through the Golgi Occurs by Cisternal Maturation

Key Concepts of Section 14.3

14.4 Later Stages of the Secretory Pathway

Vesicles Coated with Clathrin and Adapter Proteins Mediate Transport from the trans-Golgi

Dynamin Is Required for Pinching Off of Clathrin-Coated Vesicles

Mannose 6-Phosphate Residues Target Soluble Proteins to Lysosomes

Study of Lysosomal Storage Diseases Revealed Key Components of the Lysosomal Sorting Pathway

Protein Aggregation in the trans-Golgi May Function in Sorting Proteins to Regulated Secretory Vesicles

Some Proteins Undergo Proteolytic Processing After Leaving the trans-Golgi

Several Pathways Sort Membrane Proteins to the Apical or Basolateral Region of Polarized Cells

Key Concepts of Section 14.4

14.5 Receptor-Mediated Endocytosis

Cells Take Up Lipids from the Blood in the Form of Large, Well-Defined Lipoprotein Complexes

Receptors for Macromolecular Ligands Contain Sorting Signals That Target Them for Endocytosis

The Acidic pH of Late Endosomes Causes Most Receptor-Ligand Complexes to Dissociate

The Endocytic Pathway Delivers Iron to Cells Without Dissociation of the Transferrin–Transferrin Receptor Complex in Endosomes

Key Concepts of Section 14.5

14.6 Directing Membrane Proteins and Cytosolic Materials to the Lysosome

Multivesicular Endosomes Segregate Membrane Proteins Destined for the Lysosomal Membrane from Proteins Destined for Lysosomal Degradation

Retroviruses Bud from the Plasma Membrane by a Process Similar to Formation of Multivesicular Endosomes

The Autophagic Pathway Delivers Cytosolic Proteins or Entire Organelles to Lysosomes

Key Concepts of Section 14.6

Key Terms

Review the Concepts

References

Classic Experiment 14-1: Following a Protein Out of the Cell

Chapter 15: Signal Transduction and G Protein–Coupled Receptors

Chapter Introduction

15.1 Signal Transduction: From Extracellular Signal to Cellular Response

Signaling Molecules Can Act Locally or at a Distance

Receptors Bind Only a Single Type of Hormone or a Group of Closely Related Hormones

Protein Kinases and Phosphatases Are Employed in Many Signaling Pathways

GTP-Binding Proteins Are Frequently Used in Signal Transduction Pathways as On/Off Switches

Intracellular “Second Messengers” Transmit Signals from Many Receptors

Signal Transduction Pathways Can Amplify the Effects of Extracellular Signals

Key Concepts of Section 15.1

15.2 Studying Cell-Surface Receptors and Signal Transduction Proteins

The Dissociation Constant Is a Measure of the Affinity of a Receptor for Its Ligand

Binding Assays Are Used to Detect Receptors and Determine Their Affinity and Specificity for Ligands

Near-Maximal Cellular Response to a Signaling Molecule Usually Does Not Require Activation of All Receptors

Sensitivity of a Cell to External Signals Is Determined by the Number of Cell-Surface Receptors and Their Affinity for Ligand

Hormone Analogs Are Widely Used as Drugs

Receptors Can Be Purified by Affinity Chromatography Techniques

Immunoprecipitation Assays and Affinity Techniques Can Be Used to Study the Activity of Signal Transduction Proteins

Key Concepts of Section 15.2

15.3 G Protein–Coupled Receptors: Structure and Mechanism

All G Protein–Coupled Receptors Share the Same Basic Structure

Ligand-Activated G Protein–Coupled Receptors Catalyze Exchange of GTP for GDP on the α Subunit of a Heterotrimeric G Protein

Different G Proteins Are Activated by Different GPCRs and In Turn Regulate Different Effector Proteins

Key Concepts of Section 15.3

15.4 G Protein–Coupled Receptors That Regulate Ion Channels

Acetylcholine Receptors in the Heart Muscle Activate a G Protein That Opens K+ Channels

Light Activates Rhodopsin in Rod Cells of the Eye

Activation of Rhodopsin by Light Leads to Closing of cGMP-Gated Cation Channels

Signal Amplification Makes the Rhodopsin Signal Transduction Pathway Exquisitely Sensitive

Rapid Termination of the Rhodopsin Signal Transduction Pathway Is Essential for the Temporal Resolution of Vision

Rod Cells Adapt to Varying Levels of Ambient Light by Intracellular Trafficking of Arrestin and Transducin

Key Concepts of Section 15.4

15.5 G Protein–Coupled Receptors That Activate or Inhibit Adenylyl Cyclase

Adenylyl Cyclase Is Stimulated and Inhibited by Different Receptor-Ligand Complexes

Structural Studies Established How Gαs·GTP Binds to and Activates Adenylyl Cyclase

cAMP Activates Protein Kinase A by Releasing Inhibitory Subunits

Glycogen Metabolism Is Regulated by Hormone-Induced Activation of PKA

cAMP-Mediated Activation of PKA Produces Diverse Responses in Different Cell Types

Signal Amplification Occurs in the cAMP-PKA Pathway

CREB Links cAMP and PKA to Activation of Gene Transcription

Anchoring Proteins Localize Effects of cAMP to Specific Regions of the Cell

Multiple Mechanisms Suppress Signaling from the GPCR/cAMP/PKA Pathway

Key Concepts of Section 15.5

15.6 G Protein–Coupled Receptors That Trigger Elevations in Cytosolic and Mitochondrial Calcium

Calcium Concentrations in the Mitochondrial Matrix, ER, and Cytosol Can Be Measured with Targeted Fluorescent Proteins

Activated Phospholipase C Generates Two Key Second Messengers Derived from the Membrane Lipid Phosphatidylinositol 4,5-Bisphosphate

The Ca2+-Calmodulin Complex Mediates Many Cellular Responses to External Signals

DAG Activates Protein Kinase C

Integration of Ca2+ and cAMP Second Messengers Regulates Glycogenolysis

Signal-Induced Relaxation of Vascular Smooth Muscle Is Mediated by a Ca2+-Nitric Oxide-cGMP-Activated Protein Kinase G Pathway

Key Concepts of Section 15.6

Key Terms

Review the Concepts

References

Perspectives for the Future

Classic Experiment 15-1: The Infancy of Signal Transduction Studies: GTP Stimulation of cAMP Synthesis

Chapter 16: Signaling Pathways That Control Gene Expression

Chapter Introduction

16.1 Receptor Serine Kinases That Activate Smads

TGF-β Proteins Are Stored in an Inactive Form in the Extracellular Matrix

Three Separate TGF-β Receptor Proteins Participate in Binding TGF-β and Activating Signal Transduction

Activated TGF-β Receptors Phosphorylate Smad Transcription Factors

The Smad3/Smad4 Complex Activates Expression of Different Genes in Different Cell Types

Negative Feedback Loops Regulate TGF-β/Smad Signaling

Key Concepts of Section 16.1

16.2 Cytokine Receptors and the JAK/STAT Signaling Pathway

Cytokines Influence the Development of Many Cell Types

Binding of a Cytokine to Its Receptor Activates One or More Tightly Bound JAK Protein Tyrosine Kinases

Phosphotyrosine Residues Are Binding Surfaces for Multiple Proteins with Conserved Domains

SH2 Domains in Action: JAK Kinases Activate STAT Transcription Factors

Multiple Mechanisms Down-Regulate Signaling from Cytokine Receptors

Key Concepts of Section 16.2

16.3 Receptor Tyrosine Kinases

Binding of Ligand Promotes Dimerization of an RTK and Leads to Activation of Its Intrinsic Tyrosine Kinase

Homo- and Hetero-oligomers of Epidermal Growth Factor Receptors Bind Members of the Epidermal Growth Factor Family

Activation of the EGF Receptor Results in the Formation of an Asymmetric Active Kinase Dimer

Multiple Mechanisms Down-Regulate Signaling from RTKs

Key Concepts of Section 16.3

16.4 The Ras/MAP Kinase Pathway

Ras, a GTPase Switch Protein, Operates Downstream of Most RTKs and Cytokine Receptors

Genetic Studies in Drosophila Identified Key Signal-Transducing Proteins in the Ras/MAP Kinase Pathway

Receptor Tyrosine Kinases Are Linked to Ras by Adapter Proteins

Binding of Sos to Inactive Ras Causes a Conformational Change That Triggers an Exchange of GTP for GDP

Signals Pass from Activated Ras to a Cascade of Protein Kinases Ending with MAP Kinase

Phosphorylation of MAP Kinase Results in a Conformational Change That Enhances Its Catalytic Activity and Promotes Its Dimerization

MAP Kinase Regulates the Activity of Many Transcription Factors Controlling Early Response Genes

G Protein–Coupled Receptors Transmit Signals to MAP Kinase in Yeast Mating Pathways

Scaffold Proteins Separate Multiple MAP Kinase Pathways in Eukaryotic Cells

Key Concepts of Section 16.4

16.5 Phosphoinositide Signaling Pathways

Phospholipase Cγ Is Activated by Some RTKs and Cytokine Receptors

Recruitment of PI-3 Kinase to Activated Receptors Leads to Synthesis of Three Phosphorylated Phosphatidylinositols

Accumulation of PI 3-Phosphates in the Plasma Membrane Leads to Activation of Several Kinases

Activated Protein Kinase B Induces Many Cellular Responses

The PI-3 Kinase Pathway Is Negatively Regulated by PTEN Phosphatase

Key Concepts of Section 16.5

16.6 Signaling Pathways Controlled by Ubiquitinylation and Protein Degradation: Wnt, Hedgehog, and NF-κB

Wnt Signaling Triggers Release of a Transcription Factor from a Cytosolic Protein Complex

Concentration Gradients of Wnt Protein Are Essential for Many Steps in Development

Hedgehog Signaling Relieves Repression of Target Genes

Hedgehog Signaling in Vertebrates Requires Primary Cilia

Degradation of an Inhibitor Protein Activates the NF-κB Transcription Factor

Polyubiquitin Chains Serve as Scaffolds Linking Receptors to Downstream Proteins in the NF-κB Pathway

Key Concepts of Section 16.6

16.7 Signaling Pathways Controlled by Protein Cleavage: Notch/Delta, SREBP, and Alzheimer’s Disease

On Binding Delta, the Notch Receptor Is Cleaved, Releasing a Component Transcription Factor

Matrix Metalloproteases Catalyze Cleavage of Many Signaling Proteins from the Cell Surface

Inappropriate Cleavage of Amyloid Precursor Protein Can Lead to Alzheimer’s Disease

Regulated Intramembrane Proteolysis of SREBPs Releases a Transcription Factor That Acts to Maintain Phospholipid and Cholesterol Levels

Key Concepts of Section 16.7

16.8 Integration of Cellular Responses to Multiple Signaling Pathways: Insulin Action

Insulin and Glucagon Work Together to Maintain a Stable Blood Glucose Level

A Rise in Blood Glucose Triggers Insulin Secretion from the β Islet Cells

In Fat and Muscle Cells, Insulin Triggers Fusion of Intracellular Vesicles Containing the GLUT4 Glucose Transporter to the Plasma Membrane

Insulin Inhibits Glucose Synthesis and Enhances Storage of Glucose as Glycogen

Multiple Signal Transduction Pathways Interact to Regulate Adipocyte Differentiation Through PPARγ, the Master Transcriptional Regulator

Inflammatory Hormones Cause Derangement of Adipose Cell Function in Obesity

Key Concepts of Section 16.8

Key Terms

Review the Concepts

References

Perspectives for the Future

Chapter 17: Cell Organization and Movement I: Microfilaments

Chapter Introduction

17.1 Microfilaments and Actin Structures

Actin Is Ancient, Abundant, and Highly Conserved

G-Actin Monomers Assemble into Long, Helical F-Actin Polymers

F-Actin Has Structural and Functional Polarity

Key Concepts of Section 17.1

17.2 Dynamics of Actin Filaments

Actin Polymerization In Vitro Proceeds in Three Steps

Actin Filaments Grow Faster at (+) Ends Than at (−) Ends

Actin Filament Treadmilling Is Accelerated by Profilin and Cofilin

Thymosin-β4 Provides a Reservoir of Actin for Polymerization

Capping Proteins Block Assembly and Disassembly at Actin Filament Ends

Key Concepts of Section 17.2

17.3 Mechanisms of Actin Filament Assembly

Formins Assemble Unbranched Filaments

The Arp2/3 Complex Nucleates Branched Filament Assembly

Intracellular Movements Can Be Powered by Actin Polymerization

Microfilaments Function in Endocytosis

Toxins That Perturb the Pool of Actin Monomers Are Useful for Studying Actin Dynamics

Key Concepts of Section 17.3

17.4 Organization of Actin-Based Cellular Structures

Cross-Linking Proteins Organize Actin Filaments into Bundles or Networks

Adapter Proteins Link Actin Filaments to Membranes

Key Concepts of Section 17.4

17.5 Myosins: Actin-Based Motor Proteins

Myosins Have Head, Neck, and Tail Domains with Distinct Functions

Myosins Make Up a Large Family of Mechanochemical Motor Proteins

Conformational Changes in the Myosin Head Couple ATP Hydrolysis to Movement

Myosin Heads Take Discrete Steps Along Actin Filaments

Key Concepts of Section 17.5

17.6 Myosin-Powered Movements

Myosin Thick Filaments and Actin Thin Filaments in Skeletal Muscle Slide Past Each Other During Contraction

Skeletal Muscle Is Structured by Stabilizing and Scaffolding Proteins

Contraction of Skeletal Muscle Is Regulated by Ca2+ and Actin-Binding Proteins

Actin and Myosin II Form Contractile Bundles in Nonmuscle Cells

Myosin-Dependent Mechanisms Regulate Contraction in Smooth Muscle and Nonmuscle Cells

Myosin V–Bound Vesicles Are Carried Along Actin Filaments

Key Concepts of Section 17.6

17.7 Cell Migration: Mechanism, Signaling, and Chemotaxis

Cell Migration Coordinates Force Generation with Cell Adhesion and Membrane Recycling

The Small GTP-Binding Proteins Cdc42, Rac, and Rho Control Actin Organization

Cell Migration Involves the Coordinate Regulation of Cdc42, Rac, and Rho

Migrating Cells Are Steered by Chemotactic Molecules

Key Concepts of Section 17.7

Key Terms

Review the Concepts

Extended References

Perspectives for the Future

Classic Experiment 17-1: Looking at Muscle Contraction

Classic Experiment 17-2: Sensing Chemotactic Gradients

Chapter 18: Cell Organization and Movement II: Microtubules and Intermediate Filaments

Chapter Introduction

18.1 Microtubule Structure and Organization

Microtubule Walls Are Polarized Structures Built from αβ-Tubulin Dimers

Microtubules Are Assembled from MTOCs to Generate Diverse Configurations

Key Concepts of Section 18.1

18.2 Microtubule Dynamics

Individual Microtubules Exhibit Dynamic Instability

Localized Assembly and “Search and Capture” Help Organize Microtubules

Drugs Affecting Tubulin Polymerization Are Useful Experimentally and in Treatment of Diseases

Key Concepts of Section 18.2

18.3 Regulation of Microtubule Structure and Dynamics

Microtubules Are Stabilized by Side-Binding Proteins

+TIPs Regulate the Properties and Functions of the Microtubule (+) End

Other End-Binding Proteins Regulate Microtubule Disassembly

Key Concepts of Section 18.3

18.4 Kinesins and Dyneins: Microtubule-Based Motor Proteins

Organelles in Axons Are Transported Along Microtubules in Both Directions

Kinesin-1 Powers Anterograde Transport of Vesicles Down Axons Toward the (+) Ends of Microtubules

The Kinesins Form a Large Protein Superfamily with Diverse Functions

Kinesin-1 Is a Highly Processive Motor

Dynein Motors Transport Organelles Toward the (−) Ends of Microtubules

Kinesins and Dyneins Cooperate in the Transport of Organelles Throughout the Cell

Tubulin Modifications Distinguish Different Classes of Microtubules and Their Accessibility to Motors

Key Concepts of Section 18.4

18.5 Cilia and Flagella: Microtubule-Based Surface Structures

Eukaryotic Cilia and Flagella Contain Long Doublet Microtubules Bridged by Dynein Motors

Ciliary and Flagellar Beating Are Produced by Controlled Sliding of Outer Doublet Microtubules

Intraflagellar Transport Moves Material Up and Down Cilia and Flagella

Primary Cilia Are Sensory Organelles on Interphase Cells

Defects in Primary Cilia Underlie Many Diseases

Key Concepts of Section 18.5

18.6 Mitosis

Centrosomes Duplicate Early in the Cell Cycle in Preparation for Mitosis

Mitosis Can Be Divided into Six Stages

The Mitotic Spindle Contains Three Classes of Microtubules

Microtubule Dynamics Increase Dramatically in Mitosis

Mitotic Asters Are Pushed Apart by Kinesin-5 and Oriented by Dynein

Chromosomes Are Captured and Oriented During Prometaphase

Duplicated Chromosomes Are Aligned by Motors and Microtubule Dynamics

The Chromosomal Passenger Complex Regulates Microtubule Attachment at Kinetochores

Anaphase A Moves Chromosomes to Poles by Microtubule Shortening

Anaphase B Separates Poles by the Combined Action of Kinesins and Dynein

Additional Mechanisms Contribute to Spindle Formation

Cytokinesis Splits the Duplicated Cell in Two

Plant Cells Reorganize Their Microtubules and Build a New Cell Wall in Mitosis

Key Concepts of Section 18.6

18.7 Intermediate Filaments

Intermediate Filaments Are Assembled from Subunit Dimers

Intermediate Filaments Are Dynamic

Cytoplasmic Intermediate Filament Proteins Are Expressed in a Tissue-Specific Manner

Lamins Line the Inner Nuclear Envelope To Provide Organization and Rigidity to the Nucleus

Lamins Are Reversibly Disassembled by Phosphorylation During Mitosis

Key Concepts of Section 18.7

18.8 Coordination and Cooperation Between Cytoskeletal Elements

Intermediate Filament–Associated Proteins Contribute to Cellular Organization

Microfilaments and Microtubules Cooperate to Transport Melanosomes

Cdc42 Coordinates Microtubules and Microfilaments During Cell Migration

Advancement of Neural Growth Cones Is Coordinated by Microfilaments and Microtubules

Key Concepts of Section 18.8

Key Terms

Review the Concepts

Extended References

Perspectives for the Future

Chapter 19: The Eukaryotic Cell Cycle

Chapter Introduction

19.1 Overview of the Cell Cycle and Its Control

The Cell Cycle Is an Ordered Series of Events Leading to Cell Replication

Cyclin-Dependent Kinases Control the Eukaryotic Cell Cycle

Several Key Principles Govern the Cell Cycle

Key Concepts of Section 19.1

19.2 Model Organisms and Methods of Studying the Cell Cycle

Budding and Fission Yeasts Are Powerful Systems for Genetic Analysis of the Cell Cycle

Frog Oocytes and Early Embryos Facilitate Biochemical Characterization of the Cell Cycle Machinery

Fruit Flies Reveal the Interplay Between Development and the Cell Cycle

The Study of Tissue Culture Cells Uncovers Cell Cycle Regulation in Mammals

Researchers Use Multiple Tools to Study the Cell Cycle

Key Concepts of Section 19.2

19.3 Regulation of CDK Activity

Cyclin-Dependent Kinases Are Small Protein Kinases That Require a Regulatory Cyclin Subunit for Their Activity

Cyclins Determine the Activity of CDKs

Cyclin Levels Are Primarily Regulated by Protein Degradation

CDKs Are Regulated by Activating and Inhibitory Phosphorylation

CDK Inhibitors Control Cyclin-CDK Activity

Genetically Engineered CDKs Led to the Discovery of CDK Functions

Key Concepts of Section 19.3

19.4 Commitment to the Cell Cycle and DNA Replication

Cells Are Irreversibly Committed to Division at a Cell Cycle Point Called START or the Restriction Point

The E2F Transcription Factor and Its Regulator Rb Control the G1–S Phase Transition in Metazoans

Extracellular Signals Govern Cell Cycle Entry

Degradation of an S Phase CDK Inhibitor Triggers DNA Replication

Replication at Each Origin Is Initiated Once and Only Once During the Cell Cycle

Duplicated DNA Strands Become Linked During Replication

Key Concepts of Section 19.4

19.5 Entry into Mitosis

Precipitous Activation of Mitotic CDKs Initiates Mitosis

Mitotic CDKs Promote Nuclear Envelope Breakdown

Mitotic CDKs Promote Mitotic Spindle Formation

Chromosome Condensation Facilitates Chromosome Segregation

Key Concepts of Section 19.5

19.6 Completion of Mitosis: Chromosome Segregation and Exit from Mitosis

Separase-Mediated Cleavage of Cohesins Initiates Chromosome Segregation

APC/C Activates Separase Through Securin Ubiquitinylation

Mitotic CDK Inactivation Triggers Exit from Mitosis

Cytokinesis Creates Two Daughter Cells

Key Concepts of Section 19.6

19.7 Surveillance Mechanisms in Cell Cycle Regulation

Checkpoint Pathways Establish Dependencies and Prevent Errors in the Cell Cycle

The Growth Checkpoint Pathway Ensures That Cells Enter the Cell Cycle Only After Sufficient Macromolecule Biosynthesis

The DNA Damage Response System Halts Cell Cycle Progression When DNA Is Compromised

The Spindle Assembly Checkpoint Pathway Prevents Chromosome Segregation Until Chromosomes Are Accurately Attached to the Mitotic Spindle

The Spindle Position Checkpoint Pathway Ensures That the Nucleus Is Accurately Partitioned Between Two Daughter Cells

Key Concepts of Section 19.7

19.8 Meiosis: A Special Type of Cell Division

Extracellular and Intracellular Cues Regulate Germ Cell Formation

Several Key Features Distinguish Meiosis from Mitosis

Recombination and a Meiosis-Specific Cohesin Subunit Are Necessary for the Specialized Chromosome Segregation in Meiosis I

Co-orienting Sister Kinetochores Is Critical for Meiosis I Chromosome Segregation

DNA Replication Is Inhibited Between the Two Meiotic Divisions

Key Concepts of Section 19.8

Key Terms

Review the Concepts

References

Perspectives for the Future

Classic Experiment 19-1: How Cyclins Were Discovered

Classic Experiment 19-2: Synthesis and Degradation of Mitotic Cyclin Are Required for Progression through Mitosis

Classic Experiment 19-3: The Formulation of the Checkpoint Concept

Chapter 20: Integrating Cells into Tissues

Chapter Introduction

20.1 Cell-Cell and Cell–Extracellular Matrix Adhesion: An Overview

Cell-Adhesion Molecules Bind to One Another and to Intracellular Proteins

The Extracellular Matrix Participates in Adhesion, Signaling, and Other Functions

The Evolution of Multifaceted Adhesion Molecules Made Possible the Evolution of Diverse Animal Tissues

Cell-Adhesion Molecules Mediate Mechanotransduction

Key Concepts of Section 20.1

20.2 Cell-Cell and Cell–Extracellular Junctions and Their Adhesion Molecules

Epithelial Cells Have Distinct Apical, Lateral, and Basal Surfaces

Three Types of Junctions Mediate Many Cell-Cell and Cell-ECM Interactions

Cadherins Mediate Cell-Cell Adhesions in Adherens Junctions and Desmosomes

Integrins Mediate Cell-ECM Adhesions, Including Those in Epithelial-Cell Hemidesmosomes

Tight Junctions Seal Off Body Cavities and Restrict Diffusion of Membrane Components

Gap Junctions Composed of Connexins Allow Small Molecules to Pass Directly Between the Cytosols of Adjacent Cells

Key Concepts of Section 20.2

20.3 The Extracellular Matrix I: The Basal Lamina

The Basal Lamina Provides a Foundation for Assembly of Cells into Tissues

Laminin, a Multi-adhesive Matrix Protein, Helps Cross-Link Components of the Basal Lamina

Sheet-Forming Type IV Collagen Is a Major Structural Component of the Basal Lamina

Perlecan, a Proteoglycan, Cross-Links Components of the Basal Lamina and Cell-Surface Receptors

Key Concepts of Section 20.3

20.4 The Extracellular Matrix II: Connective Tissue

Fibrillar Collagens Are the Major Fibrous Proteins in the ECM of Connective Tissues

Fibrillar Collagen Is Secreted and Assembled into Fibrils Outside the Cell

Type I and II Collagens Associate with Nonfibrillar Collagens to Form Diverse Structures

Proteoglycans and Their Constituent GAGs Play Diverse Roles in the ECM

Hyaluronan Resists Compression, Facilitates Cell Migration, and Gives Cartilage Its Gel-Like Properties

Fibronectins Connect Cells and ECM, Influencing Cell Shape, Differentiation, and Movement

Elastic Fibers Permit Many Tissues to Undergo Repeated Stretching and Recoiling

Metalloproteases Remodel and Degrade the Extracellular Matrix

Key Concepts of Section 20.4

20.5 Adhesive Interactions in Motile and Nonmotile Cells

Integrins Mediate Adhesion and Relay Signals Between Cells and Their Three-Dimensional Environment

Regulation of Integrin-Mediated Adhesion and Signaling Controls Cell Movement

Connections Between the ECM and Cytoskeleton Are Defective in Muscular Dystrophy

IgCAMs Mediate Cell-Cell Adhesion in Neural and Other Tissues

Leukocyte Movement into Tissues Is Orchestrated by a Precisely Timed Sequence of Adhesive Interactions

Key Concepts of Section 20.5

20.6 Plant Tissues

The Plant Cell Wall Is a Laminate of Cellulose Fibrils in a Matrix of Glycoproteins

Loosening of the Cell Wall Permits Plant Cell Growth

Plasmodesmata Directly Connect the Cytosols of Adjacent Cells

Tunneling Nanotubes Resemble Plasmodesmata and Transfer Molecules and Organelles Between Animal Cells

Only a Few Adhesion Molecules Have Been Identified in Plants

Key Concepts of Section 20.6

Key Terms

Review the Concepts

Extended References

Perspectives for the Future

Chapter 21: Stem Cells, Cell Asymmetry, and Cell Death

Chapter Introduction

21.1 Early Mammalian Development

Fertilization Unifies the Genome

Cleavage of the Mammalian Embryo Leads to the First Differentiation Events

Key Concepts of Section 21.1

21.2 Embryonic Stem Cells and Induced Pluripotent Stem Cells

The Inner Cell Mass Is the Source of ES Cells

Multiple Factors Control the Pluripotency of ES Cells

Animal Cloning Shows That Differentiation Can Be Reversed

Somatic Cells Can Generate iPS Cells

ES and iPS Cells Can Generate Functional Differentiated Human Cells

Key Concepts of Section 21.2

21.3 Stem Cells and Niches in Multicellular Organisms

Adult Planaria Contain Pluripotent Stem Cells

Multipotent Somatic Stem Cells Give Rise to Both Stem Cells and Differentiating Cells

Stem Cells for Different Tissues Occupy Sustaining Niches

Germ-Line Stem Cells Produce Sperm or Oocytes

Intestinal Stem Cells Continuously Generate All the Cells of the Intestinal Epithelium

Hematopoietic Stem Cells Form All Blood Cells

Rare Types of Cells Constitute the Niche for Hematopoietic Stem Cells

Meristems Are Niches for Stem Cells in Plants

A Negative Feedback Loop Maintains the Size of the Shoot Apical Stem-Cell Population

The Root Meristem Resembles the Shoot Meristem in Structure and Function

Key Concepts of Section 21.3

21.4 Mechanisms of Cell Polarity and Asymmetric Cell Division

The Intrinsic Polarity Program Depends on a Positive Feedback Loop Involving Cdc42

Cell Polarization Before Cell Division Follows a Common Hierarchy of Steps

Polarized Membrane Traffic Allows Yeast to Grow Asymmetrically During Mating

The Par Proteins Direct Cell Asymmetry in the Nematode Embryo

The Par Proteins and Other Polarity Complexes Are Involved in Epithelial-Cell Polarity

The Planar Cell Polarity Pathway Orients Cells Within an Epithelium

The Par Proteins Are Involved in Asymmetric Division of Stem Cells

Key Concepts of Section 21.4

21.5 Cell Death and Its Regulation

Most Programmed Cell Death Occurs Through Apoptosis

Evolutionarily Conserved Proteins Participate in the Apoptotic Pathway

Caspases Amplify the Initial Apoptotic Signal and Destroy Key Cellular Proteins

Neurotrophins Promote Survival of Neurons

Mitochondria Play a Central Role in Regulation of Apoptosis in Vertebrate Cells

The Pro-apoptotic Proteins Bax and Bak Form Pores and Holes in the Outer Mitochondrial Membrane

Release of Cytochrome c and SMAC/DIABLO Proteins from Mitochondria Leads to Formation of the Apoptosome and Caspase Activation

Trophic Factors Induce Inactivation of Bad, a Pro-apoptotic BH3-Only Protein

Vertebrate Apoptosis Is Regulated by BH3-Only Pro-apoptotic Proteins That Are Activated by Environmental Stresses

Two Types of Cell Murder Are Triggered by Tumor Necrosis Factor, Fas Ligand, and Related Death Signals

Key Concepts of Section 21.5

Key Terms

Review the Concepts

References

Perspectives for the Future

Chapter 22: Cells of the Nervous System

Chapter Introduction

22.1 Neurons and Glia: Building Blocks of the Nervous System

Information Flows Through Neurons from Dendrites to Axons

Information Moves Along Axons as Pulses of Ion Flow Called Action Potentials

Information Flows Between Neurons via Synapses

The Nervous System Uses Signaling Circuits Composed of Multiple Neurons

Glial Cells Form Myelin Sheaths and Support Neurons

Neural Stem Cells Form Nerve and Glial Cells in the Central Nervous System

Key Concepts of Section 22.1

22.2 Voltage-Gated Ion Channels and the Propagation of Action Potentials

The Magnitude of the Action Potential Is Close to ENa and Is Caused by Na+ Influx Through Open Na+ Channels

Sequential Opening and Closing of Voltage-Gated Na+ and K+ Channels Generate Action Potentials

Action Potentials Are Propagated Unidirectionally Without Diminution

Nerve Cells Can Conduct Many Action Potentials in the Absence of ATP

All Voltage-Gated Ion Channels Have Similar Structures

Voltage-Sensing S4 α Helices Move in Response to Membrane Depolarization

Movement of the Channel-Inactivating Segment into the Open Pore Blocks Ion Flow

Myelination Increases the Velocity of Impulse Conduction

Action Potentials “Jump” from Node to Node in Myelinated Axons

Two Types of Glia Produce Myelin Sheaths

Light-Activated Ion Channels and Optogenetics

Key Concepts of Section 22.2

22.3 Communication at Synapses

Formation of Synapses Requires Assembly of Presynaptic and Postsynaptic Structures

Neurotransmitters Are Transported into Synaptic Vesicles by H+-Linked Antiport Proteins

Three Pools of Synaptic Vesicles Loaded with Neurotransmitter Are Present in the Presynaptic Terminal

Influx of Ca2+ Triggers Release of Neurotransmitters

A Calcium-Binding Protein Regulates Fusion of Synaptic Vesicles with the Plasma Membrane

Fly Mutants Lacking Dynamin Cannot Recycle Synaptic Vesicles

Signaling at Synapses Is Terminated by Degradation or Reuptake of Neurotransmitters

Opening of Acetylcholine-Gated Cation Channels Leads to Muscle Contraction

All Five Subunits in the Nicotinic Acetylcholine Receptor Contribute to the Ion Channel

Nerve Cells Integrate Many Inputs to Make an All-or-None Decision to Generate an Action Potential

Gap Junctions Allow Direct Communication Between Neurons and Between Glia

Key Concepts of Section 22.3

22.4 Sensing the Environment: Touch, Pain, Taste, and Smell

Mechanoreceptors Are Gated Cation Channels

Pain Receptors Are Also Gated Cation Channels

Five Primary Tastes Are Sensed by Subsets of Cells in Each Taste Bud

A Plethora of Receptors Detect Odors

Each Olfactory Receptor Neuron Expresses a Single Type of Odorant Receptor

Key Concepts of Section 22.4

22.5 Forming and Storing Memories

Memories Are Formed by Changing the Number or Strength of Synapses Between Neurons

The Hippocampus Is Required for Memory Formation

Multiple Molecular Mechanisms Contribute to Synaptic Plasticity

Formation of Long-Term Memories Requires Gene Expression

Key Concepts of Section 22.5

Key Terms

Review the Concepts

Extended References

Perspectives for the Future

Chapter 23: Immunology

Chapter Introduction

23.1 Overview of Host Defenses

Pathogens Enter the Body Through Different Routes and Replicate at Different Sites

Leukocytes Circulate Throughout the Body and Take Up Residence in Tissues and Lymph Nodes

Mechanical and Chemical Boundaries Form a First Layer of Defense Against Pathogens

Innate Immunity Provides a Second Line of Defense

Inflammation Is a Complex Response to Injury That Encompasses Both Innate and Adaptive Immunity

Adaptive Immunity, the Third Line of Defense, Exhibits Specificity

Key Concepts of Section 23.1

23.2 Immunoglobulins: Structure and Function

Immunoglobulins Have a Conserved Structure Consisting of Heavy and Light Chains

Multiple Immunoglobulin Isotypes Exist, Each with Different Functions

Each Naive B Cell Produces a Unique Immunoglobulin

Immunoglobulin Domains Have a Characteristic Fold Composed of Two β Sheets Stabilized by a Disulfide Bond

An Immunoglobulin’s Constant Region Determines Its Functional Properties

Key Concepts of Section 23.2

23.3 Generation of Antibody Diversity and B-Cell Development

A Functional Light-Chain Gene Requires Assembly of V and J Gene Segments

Rearrangement of the Heavy-Chain Locus Involves V, D, and J Gene Segments

Somatic Hypermutation Allows the Generation and Selection of Antibodies with Improved Affinities

B-Cell Development Requires Input from a Pre-B-Cell Receptor

During an Adaptive Response, B Cells Switch from Making Membrane-Bound Ig to Making Secreted Ig

B Cells Can Switch the Isotype of Immunoglobulin They Make

Key Concepts of Section 23.3

23.4 The MHC and Antigen Presentation

The MHC Determines the Ability of Two Unrelated Individuals of the Same Species to Accept or Reject Grafts

The Killing Activity of Cytotoxic T Cells Is Antigen Specific and MHC Restricted

T Cells with Different Functional Properties Are Guided by Two Distinct Classes of MHC Molecules

MHC Molecules Bind Peptide Antigens and Interact with the T-Cell Receptor

Antigen Presentation Is the Process by Which Protein Fragments Are Complexed with MHC Products and Posted to the Cell Surface

The Class I MHC Pathway Presents Cytosolic Antigens

The Class II MHC Pathway Presents Antigens Delivered to the Endocytic Pathway

Key Concepts of Section 23.4

23.5 T Cells, T-Cell Receptors, and T-Cell Development

The Structure of the T-Cell Receptor Resembles the F(ab) Portion of an Immunoglobulin

TCR Genes Are Rearranged in a Manner Similar to Immunoglobulin Genes

Many of the Variable Residues of TCRs Are Encoded in the Junctions Between V, D, and J Gene Segments

Signaling via Antigen-Specific Receptors Triggers Proliferation and Differentiation of T and B Cells

T Cells Capable of Recognizing MHC Molecules Develop Through a Process of Positive and Negative Selection

T Cells Commit to the CD4 or CD8 Lineage in the Thymus

T Cells Require Two Types of Signals for Full Activation

Cytotoxic T Cells Carry the CD8 Co-receptor and Are Specialized for Killing

T Cells Produce an Array of Cytokines That Provide Signals to Other Immune-System Cells

Helper T Cells Are Divided into Distinct Subsets Based on Their Cytokine Production and Expression of Surface Markers

Leukocytes Move in Response to Chemotactic Cues Provided by Chemokines

Key Concepts of Section 23.5

23.6 Collaboration of Immune-System Cells in the Adaptive Response

Toll-Like Receptors Perceive a Variety of Pathogen-Derived Macromolecular Patterns

Engagement of Toll-Like Receptors Leads to Activation of Antigen-Presenting Cells

Production of High-Affinity Antibodies Requires Collaboration Between B and T cells

Vaccines Elicit Protective Immunity Against a Variety of Pathogens

The Immune System Defends Against Cancer

Key Concepts of Section 23.6

Key Terms

Review the Concepts

Extended References

Perspectives for the Future

Classic Experiment 23-1: Two Genes Become One: Somatic Recombination of Immunoglobulin Genes

Chapter 24: Cancer

Chapter Introduction

24.1 How Tumor Cells Differ from Normal Cells

The Genetic Makeup of Most Cancer Cells Is Dramatically Altered

Cellular Housekeeping Functions Are Fundamentally Altered in Cancer Cells

Uncontrolled Proliferation Is a Universal Trait of Cancer

Cancer Cells Escape the Confines of Tissues

Tumors Are Heterogeneous Organs That Are Sculpted by Their Environment

Tumor Growth Requires Formation of New Blood Vessels

Invasion and Metastasis Are Late Stages of Tumorigenesis

Key Concepts of Section 24.1

24.2 The Origins and Development of Cancer

Carcinogens Induce Cancer by Damaging DNA

Some Carcinogens Have Been Linked to Specific Cancers

The Multi-hit Model Can Explain the Progress of Cancer

Successive Oncogenic Mutations Can Be Traced in Colon Cancers

Cancer Development Can Be Studied in Cultured Cells and in Animal Models

Key Concepts of Section 24.2

24.3 The Genetic Basis of Cancer

Gain-of-Function Mutations Convert Proto-oncogenes into Oncogenes

Cancer-Causing Viruses Contain Oncogenes or Activate Cellular Proto-oncogenes

Loss-of-Function Mutations in Tumor-Suppressor Genes Are Oncogenic

Inherited Mutations in Tumor-Suppressor Genes Increase Cancer Risk

Epigenetic Changes Can Contribute to Tumorigenesis

Micro-RNAs Can Promote and Inhibit Tumorigenesis

Researchers Are Identifying Drivers of Tumorigenesis

Molecular Cell Biology Is Changing How Cancer Is Diagnosed and Treated

Key Concepts of Section 24.3

24.4 Misregulation of Cell Growth and Death Pathways in Cancer

Oncogenic Receptors Can Promote Proliferation in the Absence of External Growth Factors

Many Oncogenes Encode Constitutively Active Signal-Transducing Proteins

Inappropriate Production of Nuclear Transcription Factors Can Induce Transformation

Aberrations in Signaling Pathways That Control Development Are Associated with Many Cancers

Genes That Regulate Apoptosis Can Function as Proto-oncogenes or Tumor-Suppressor Genes

Key Concepts of Section 24.4

24.5 Deregulation of the Cell Cycle and Genome Maintenance Pathways in Cancer

Mutations That Promote Unregulated Passage from G1 to S Phase Are Oncogenic

Loss of p53 Abolishes the DNA Damage Checkpoint

Loss of DNA-Repair Systems Can Lead to Cancer

Key Concepts of Section 24.5

Key Terms

Review the Concepts

References

Perspectives for the Future

Classic Experiment 24-1: Identification of the RAS Oncogene

Glossary

Glossary - A

Glossary - B

Glossary - C

Glossary - D

Glossary - E

Glossary - F

Glossary - G

Glossary - H

Glossary - I

Glossary - J

Glossary - K

Glossary - L

Glossary - M

Glossary - N

Glossary - O

Glossary - P

Glossary - Q

Glossary - R

Glossary - S

Glossary - T

Glossary - U

Glossary - V

Glossary - W

Glossary - X, Y, Z

Index

Index - A

Index - B

Index - C

Index - D

Index - E

Index - F

Index - G

Index - H

Index - I

Index - J

Index - K

Index - L

Index - M

Index - N

Index - O

Index - P

Index - Q

Index - R

Index - S

Index - T

Index - U

Index - V

Index - W

Index - X, Y, Z