Kalthoff / Second Edition Contents
ANALYSIS OF BIOLOGICAL DEVELOPMENT, second edition
Klaus Kalthoff / University of Texas
DETAILED TABLE OF CONTENTS
( Basic Principles)
( Key Strategies)
CHAPTER 1 /ANALYSIS OF DEVELOPMENT
1.1
1.2
1.3
1.4
1.5
The Principle of Epigenesis
Developmental Periods and Stages in the Life Cycle
Embryonic Development Begins with Fertilization and Ends
with the Completion of Histogenesis
Postembryonic Development Can Be Direct or Indirect
Adulthood Begins with the Onset of Reproduction and Ends with Death
Classical Analytical Strategies in Developmental Biology
Embryonic Cells Have Predictable Fates in Development
Analysis of Development Requires a Strategy of Controlled Interference
 Isolation, Removal and Transplantation of Embryonic Parts Are Key
Strategies of Embryologists
Genetic Analysis of Development
Reductionist and Synthetic Analyses of Development
CHAPTER 2 / THE ROLE OF CELLS IN DEVELOPMENT
2.1
2.2
2.3
2.4
2.5
2.6
2.7
2.8
The Principle of Cellular Continuity
The Cell and Its Organelles
Cell Shape and the Cytoskeleton
Cells Change Their External Shape As Well As Their Internal Order
Microtubules Maintain Cell Shape and Mediate Intracellular Transport
Microfilaments Generate Contracting Forces and Stabilize the
Cell Surface
Intermediate Filaments Vary Among Different Cell Types
The Cell Cycle and Its Control
Chromosomes Are Duplicated During S Phase
Chromosome Duplicates Are Split between Daughter Cells During Mitosis
The Cell Cytoplasm Is Divided During Cytokinesis
The Cyclic Activity of a Protein Complex Controls the Cell Cycle
Cell Membranes
Cellular Movement
Cell Junctions in Epithelia and Mesenchyme
Cellular Signaling
Intercellular Signals Vary with Respect to Distance, Speed of
Action, and Complexity
Membrane Receptors Initiate Different Signaling Pathways
Adenylate Cyclase Generates cAMP as a Second Messenger
Phospholipase C- Generates Diacylglycerol, Inositol
Trisphosphate, and Calcium Ions as Second Messengers
The RTK-Ras-ERK Pathway Activates Transcription Factors
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2.9
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Signal Transduction Pathways Are Linked with One Another
and With Cell Adhesion
Conclusion and Outlook
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CHAPTER 3 / GAMETOGENESIS
3.1
3.2
3.3
The Discovery of the Mammalian Egg
The Germ Line Concept and the Dual Origin of Gonads
Meiosis
Homologous Chromosomes Are Separated during Meiosis
The Timing of Meiosis Differs between Males and Females
Meiosis Promotes Genetic Variation, Helps to Establish
Homozygous Mutant Alleles, and Eliminates Bad Genes
3.4
Spermatogenesis
Male Germ Cells Develop in Seminiferous Tubules
Spermiogenesis
Spermatogonia Behave as Stem Cells
3.5
Oogenesis
Oocytes Are Supplied with Large Amounts of RNA
Yolk Proteins for Oocytes Are Synthesized in the Liver or Fat Body
The Ovarian Autonomy Imposes Polarity on the Oocyte
Maturation Processes Prepare the Oocyte for Ovulation and
Fertilization
Eggs Are Protected by Different Types of Envelopes
*Method 3.1 Autoradiography
CHAPTER 4 / FERTILIZATION
4.1
4.2
4.3
4.4
4.5
Interactions Before Sperm-Egg Adhesion
Many Sperm Are Attracted to Eggs by Chemical Signals
Some Sperm Must Undergo Capacitation Before They Can
Fertilize Eggs
Fertilization in Sea Urchins
Sea Urchin Sperm Undergo the Acrosome Reaction Before
They Adhere to the Vitelline Envelope
Sea Urchin Sperm Adhere to Eggs with an Acrosomal Protein
Called Bindin
Gamete Fusion Leads to the Formation of a Fertilization Cone
Fertilization in Mammals
Mouse Sperm Undergo the Acrosome Reaction After They
Adhere to the Sona Pellucida
 A Bioassay Is a Powerful Strategy to Reveal a Biologically
Active Component
Mouse Sperm Adhere to a Specific Zona Pellucida Protein
Antibodies to Sperm-Egg Adhesion Proteins Can Act as Contraceptives
Egg Activation
Egg Activation May Be Triggered by Different Signaling Mechanisms
A Temporary Rise in Ca2+ Concentration is Followed by
Activation of Protein Kinase C
Activation Accelerates The Egg’s Metabolism in Preparation
for Cleavage
Blocks to Polyspermy
The Fertilization Potential Serves as a Fast Block to Polyspermy
The Cortical Reaction Causes a Slow Block to Polyspermy
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4.6
The Principle of Synergistic Mechanisms
4.7
Parthenogenesis
*Method 4.3 Immunostaining
CHAPTER 5 / CLEAVAGE
5.1
5.2
5.3
5.4
Yolk Distribution and Cleavage Pattern
Cleavage Patterns of Representative Animals
Sea Urchins Have Isolecithal Eggs and Undergo Holoblastic
Cleavage
Amphibians Have Mesolecithal Eggs but Still Cleave
Holoblastically
Snails Have Isolecithal Eggs and Follow a Spiral Cleavage
Pattern
The Ascidian Cleavage Pattern Is Bilaterally Symmetrical
Mammalian Eggs Show Rotational Cleavage
Eggs With Variable Cleavage Show Regulative Development
Birds, Reptiles, and Many Fishes Have Telolecithal Eggs and
Undergo Discoidal Cleavage
Insects Have Centrolecithal Eggs and Undergo Superficial
Cleavage
Spatial Control of Cleavage: Positioning and Orientation of Mitotic Spindles
Actin and Myosin Form the Contractile Ring in Cytokinesis
The Mitotic Spindle Axis Determines the Orientation of the
Cleavage Plane
Mechanical Constraints May Orient Mitotic Spindles
Centrosomes Organize Mitotic Spindles in Regular Ways
During Cleavage
Specific Sites in the Egg Cortex Attract and Anchor
Centrosomes
Maternal Gene Products May Orient Mitotic Spindles
The Timing of Cleavage Divisions
The Cell Cycle Slows Down during the Midblastula Transition
(MBT)
Stage- and Region- Specific Gene Activities Modulate the
Basic Cell Cycle After MBT
The Nucleocytoplasmic Ratio May Trigger MBT According to
a Titration Model
CHAPTER 6 / CELL FATE, POTENCY, AND DETERMINATION
6.1
6.2
6.3
6.4
Fate Mapping
Conal Analysis
Potency of Embryonic Cells
Determination of Embryonic Cells
Cell Determination Is Discovered Through Operational Criteria
Drosophila Blastoderm Cells Are Determined to Form
Structures within a Single Segment
Drosophila Cells Are Born with a Bias that is Honed by
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Subsequent Interactions
Prospective Neural Plate Cells of Amphibians Are Determined
During Gastrulation
Mouse Embryonic Cells Are Not Determined Until the
Blastocyst Stage
6.5
Properties of the Determined State
Determination Is a Stepwise Process of Instruction and
Commitment
Cell Determination Occurs as Part of Embryonic Pattern
Formation
The Determined State Is (Almost) Stably Passed On during
Mitosis
6.6
Regulation in Development
*Method 6.1 Labeling Cells by Somatic Crossover
CHAPTER 7 / GENOMIC EQUIVALENCE AND THE CYPTOPLASMIC ENVIRONMENT
7.1
7.2
7.3
7.4
7.5
7.6
7.7
7.8
7.9
Theories of Cell Differentiation
Observation on Cells
Cells May Carry Out Different Functions at Different Times
Cells Change Their Differentiated State During Regeneration
Observations on Chromosomes
Different Cells from the Same Individual Show the Same Set of
Chromosomes
Polytene Chromosomes Show the Same Banding Pattern in
Different Tissues
Chromosome Elimination Is Associated with the Dichotomy
between Germ Line and Somatic Cells
Molecular Data on Genomic Equivalence
Totipotency of Differentiated Plant Cells
Totipotency of Nuclei from Embryonic Animal Cells
Newt Blastomeres Develop Normally after Delayed Nucleation
Nuclei from Embryonic Cells Are Still Totipotent
Pluripotency of Nuclei from Differentiated Animal Cells
Nuclei from Older Donor Cells Show a Decreasing Ability to
Promote the Development of a New Organism
Nuclei from Mature Cells Are Unprepared for the Fast Mitotic
Cycles of Early Frog Embryos
Some Differentiated Cells Contain Highly Pluripotent Nuclei
Mammals Can Be Cloned by Fusing Fetal or Adults Cells with
Enucleated Eggs
Nuclear Transfer Experiments with Mammals Have Important
Applications in Science, Agriculture, and Medicine
Will Humans be Cloned in the Future?
Control of Nuclear Activities by the Cytoplasmic Environment
Gene Expression Changes upon Transplantation of Nuclei to
New Cytoplasmic Environments
Cell Fusion Exposes Nuclei to New Cytoplasmic Signals
Conclusion and Outlook
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CHAPTER 8 / LOCALIZED CYTOPLASMIC DETERMINANTS
8.1
8.2
8.3
8.4
8.5
8.6
8.7
8.8
8.9
The Principle of Cytoplasmic Localization
Polar Lobe Formation as a Means of Cytoplasmic Localization
Germ Cell Determinants in Insects Eggs
Rescue Experiments Can Restore Defective Embryos to
Normal Development
Heterotopic Transplantation Tests Cytoplasmic Determinants
for Activity in Abnormal Locations
Localization of Polar Granules Requires RNA Transport Along
Microtubules
Bicoid mRNA in Drosophila Eggs
Myoplasm in Ascidian Eggs
Cytoplasmic Components of Ascidian Eggs Are Segregated
Upon Fertilization
Myoplasm is Necessary and Sufficient for Tail Muscle
Formation
Myoplasm Segregation Involves a Plasma Membrane Lamina
Myoplasm is Associated with Localized Maternal RNAs
Cytoplasmic Localization at Advanced Embryonic Stages
Bioassays for Localized Cytoplasmic Determinants
The Principle of Default Programs
Properties of Localized Cytoplasmic Determinants
Cytoplasmic Determinants Control Certain Cell Lineages or
Entire Body Regions
Localized Cytoplasmic Determinants May Be Activating or
Inhibitory
Cytoplasmic Localization May or May Not Be Associated with
Visible Markers
Most Localized Cytoplasmic Determinants Are Maternal
mRNAs
Cytoplasmic Organization Occurs at Different Stages of
Development Using Various Cellular Mechanisms
CHAPTER 9 / AXIS FORMATION AND MESODERM INDUCTION
9.1
9.2
9.3
9.4
9.5
Body Axes and Planes
Generation of Rhizoid-Thallus Axis in Fucus
Determination of the Animal-Vegetal Axis in Amphibians
The Animal-Vegetal Axis Originates by Oriented Transport
during Oogenesis
The Animal-Vegetal Polarity Determines the Spatial Order of
the Germ Layers
Vegetal Blastomeres Induce Their Animal Neighbors to Form
Mesoderm
The Principle of Induction
Determination of the Dorsoventral Axis in Amphibians
Deep Cytoplasm Undergoes Regular Movements during Egg
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9.6
9.7
9.8
Activation
Cytoplasmic Rearrangements Following Fertilization Involve
Cortical Rotation
Microtubules Move Cytoplasmic Components Dorsally
Beyond Cortical Displacement
A Dorsalizing Activity Moves from the Vegetal Pole to the
Dorsal Side During Cortical Rotation
Dorsal Vegetal and Equatorial Blastomeres Rescue Ventralized
Embryos
Effect of Dorsoventral Polarity on Mesoderm Induction in Xenopus
Mesoderm is Induced with a Rudimentary Dorsoventral Pattern
Dorsal Marginal Cells Induce an Array of Mesodermal Organ
Rudiments
Molecular Mechanisms of Dorsoventral Axis Formation and Mesoderm Induction
-Catenin May Specify Dorsoventral Polartiy
Several Growth Factors Have Mesoderm-Inducing Activity
TGF- Members and B-Catenin May Act Synergistically in
Inducing Spemann’s Organizer
Determination of Left-Right Asymmetry
Chapter 10 / Gastulation
10.1
10.2
10.3
10.4
10.5
The Analysis of Morphogenesis
Morphogenesis Involves Typical Epithelial Movements
Morphogenesis Is Based on a Small Repertoire of Cell
Activities
Gastrulation in Sea Urchins
Gastrulation in Amphibians
Different Gastrula Areas Show Distinct Cellular Behaviors
Bottle Cells Generate the Initial Depression of the Blastopore
Deep Marginal Zone Cells Are Necessary for Involution
Deep Zone Cells and Involuting Marginal Zone Cells Migrate
on the Inside of the Blastocoel Roof
Convergent Extension Is Especially Strong in the Dorsal
Marginal Zone
Animal Cap and Noninvoluting Marginal Zone Undergo
Epiboly
How Are Patterns of Cell Behavior Related to Gene
Expression?
Gastrulation in Fishes and Birds
Zebrafish Embryos Develop from Two Cell Layers Mostly by
Convergent Extension
Chicken Embryos Develop From One Cell Layer Mostly by
Ingression
Gastrulation in Humans
CHAPTER 11 / CELL ADHESION AND MORPHOGENESIS
11.1
Cell Aggregation Studies in Vitro
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Cells from Different Tissues Adhere to One Another Selectively
Tissues Form Hierarchies of Adhesiveness
11.2
Cell Adhesion Molecules
Immunoglobulinlike CAMs May Allow or Hinder Cell
Adhesion
Cadherins Mediate CA2+ – Dependent Cell Adhesion
Lectins Bind Heterotupically to Sugar Residues
11.3
ECM Molecules and Their Cellular Receptors
Glycosaminopglycans and Proteoglycans Form and
Amorphous, Hydrophilic Ground Substance
Fibrous Glycoproteins Make Up the Dynamic Meshwork of
the ECM
Integrins Mediate Cell Adhesion to ECM Molecules
11.4
The Role of Cell and Substrate Adhesion Molecules in Morphogenesis
CAM Expression is Correlated with Cell Fates
Cell Adhesiveness Changes during Sea Urchin Gastrulation
CAMs Facilitate the Formation of Cell Junctions
Fibrous ECM Components Provide Contact Guidance to Cells
Amphibian Gastrulation Requires Fibronectin on the Inner
Surface of the Blastocoel Roof
11.5
Morphoregulatory Roles of Cell and Substrate Adhesion Molecules
CAM and SAM Genes Are Controlled by Selector Genes
Cell-Cell Adhesion and Cell-Substratum Interactions Affect
Gene Activity
* Methods 11.1 Isolating Cell Adhesion Molecules and Their Genes with Antibodies
CHAPTER 12 / NEURULATION AND AXIS INDUCTION
12.1
12.2
12.3
12.4
Neurulation as an Example of Organogenesis
Neurulation Is of Scientific and Medical Interest
Neurulation in Amphibians Occurs in Two Phases
Neural Tubes of Bird Embryos Have Hinges
In Humans, Neural Tube Closure Begins in the Neck Region
In Fish, the Neural Tube Originates as a Solid Rod
Mechanisms of Neurulation in Amphibians
Neurulation Depends on Tissues Adjacent to the Neural Plate
Neural Plate Cells Undergo Columnization
Intercalation of Nerual Plate Cells Causes Convergent
Extension
Both Columnarization and Cell Intercalation Contribute to
Generating the Keyhole Shape
Neural Tube Closure Is Associated with Apical Constriction,
Rapid Anteroposterior Extension, and Cell Crawling
The Role of Induction in Axis Formation
The Dorsal Blastopore Lip Organizes the Formation of an
Entire Embryo
Does the Organizer Have Structure?
Axis Induction Shows Regional Specificity
Mechanisms of Neural Induction
There Are Two Signaling Pathways—Planar and Vertical—for
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12.5
Neural Induction
Planar Induction Plays a Major Role in Xenopus Embryos
Neural Induction is a Multistep Process
Axis Induction by Disinhibition
Dorsal Development Occurs as a Default Program in Xenopus
Spemann’s Organizer Inactivates a Ventralizing Signal
Basic Questions are Still Unresolved
CHAPTER 13 / ECTODERMAL ORGANS
13.1
13.2
13.3
13.4
Neural Tube
Nervous Tissue Consists of Neurons and Glial Cells
The Spinal Cord Is Patterned by Signal from Adjacent Tissues
The Basic Organization of the Spinal Cord Is Modified in the
Brain
The Peripheral Nervous System is of Diverse Origin
Neural Crest
NC Cells Arise at the Boundary Between Neural Plate and
Epidermis
NC Cells Have Different Migration Routes and a Wide Range
of Fates
 The Strategy of Clonal Analysis Shows that NC Cells Are a
Heterogeneous Population of Pluripotent Cells
The Strategies of Heterotopic and Heterochronic
Transplantation Reveal Spatial and Temporal Restrictions to NC Cell
Migration
Extracellular Matrix Affects the Determination of NC Cells
Region-Specific Growth Factors Are Involved in NC Cell
Determination
Ectodermal Placodes
The Otic Placode Forms the Inner Ear
The Lens Placode Develops Together with the Retina
Nasal Placodes Form Olfactory Sensory Epithelia
Epidermis
CHAPTER 14 / ENDODERMAL AND MESODERMAL ORGANS
14.1
14.2
14.3
Endodermal Derivatives
The Embryonic Pharynx Contains a Series of Arches
Embryos Pass through a Phylotypic Stage after Organogenesis
The Endoderm Lines the Inside of the Intestine and Its
Appendages
Axial and Paraxial Mesoderm
Axial Mesoderm Forms the Prechordal Plate and the
Notochord
Paraxial Mesoderm Forms the Presomitic Plates and Somites
Somites Are Patterned by Signal from Surrounding Organ
Rudiments
Connective Tissue and Skeletal Muscle
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14.4
14.5
14.6
14.7
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Connective Tissue Contains Large Amounts of Extracellular
Matrix
Skeletal Muscle Fibers Arise Through Cell Fusion
Intermediate Mesoderm
The Principle of Reciprocal Interaction
Lateral Plates
The Lateral Plates Surround the Coelomic Cavities
The Cardiovascular System Develops from Various
Mesodermal Precursors
Cardiovascular System Development Recapitulates the
Phylotopic Stage
Smooth Muscle Consists of Single Cells
Extraembryonic Membranes
The Amnion and Chorion Are Formed by Layers of Ectoderm
and Mesoderm
The Yolk Sac and Allantois Are Formed by Layers of
Endoderm and Mesoderm
The Mammalian Placenta Is Formed from the Embryonic
Trophoblast and the Uterine Endometrium
CHAPTER 15 / THE USE OF MUTANTS AND TRANSGENIC ORGANISMS IN THE ANALYSIS OF
DEVELOPMENT
15.1
15.2
15.3
15.4
15.5
15.6
The Historical Separation of Genetics from Developmental Biology
Modern Genetic Analysis of Development
Mutants Reveal the Hidden “Logic” of Embryonic Development
Drosophila Mutants Allow the Analysis of a Complex Body
Pattern
Caenorhabditis elegans Mutants Uncover Gene Activities
Controlling the Cell Lineages
Genetic Analysis of the Mouse Mus musculus Uses Embryonic
Stem Cells
The Zebra Fish Danio rerio is Genetically Tractable and
Suitable for Cell Transplantation
Genetic Analysis Reveals Similarity of Plant and Animal
Development
DNA Cloning and Sequencing
DNA Can Be Replicated, Transcribed, and ReverseTranscribed In Vitro
Nucleic Acid Hybridization Allows the Detection of Specific
Nucleotide Sequences
DNA Can Be Cut and Spliced Enzymatically
DNA Sequencing May Reveal the Biochemical Activity of a
Gene Product
Transfection and Genetic Transformation of Cells
The Strategies of Gene Overexpression, Dominant Interference and Gene Knockout.
Germ Line Transformation
The Genetic Transformation of Drosophila Utilizes
Transposable DNA Elements
Mammals Can Be Transformed by Injecting Transgenes
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Directly into an Egg Pronucleus
Transgenic Mice Can Be Raised from Transformed Embryonic
Stem Cells
DNA Insertion Can Be Used for Mutagenesis and Promoter
Trapping
Transgenic Organisms Have Many Uses in Basic and Applied
Science
* Methods 15.1 The Generation and Maintenance of Mutants
* Methods 15.2 Saturation Mutagenesis Screens
* Methods 15.3 Northern Blotting and Southern Blotting
* Methods 15.4 Recombinant DNA Techniques
CHAPTER 16 / TRANSCRIPTIONAL CONTROL
The Principle of Differential Gene Expression
Evidence for Transcriptional Control
DNA Sequences Controlling Transcription
Regulatory DNA Sequences Are Studied with Fusion Genes
Promoters and Enhancers Are Regulatory DNA Regions with
Different Properties
16.4
Transcription Factors and Their Role in Development
General Transcription Factors Bind to All Promoters
Transcriptional Activators and Repressors Associate with
Restricted Sets of Genes and Occur Only in Certain Cells
Transcriptional Activators Have Highly Conserved DNA
Binding Domains
The bicoid Protein Acts as a Transcriptional Activator on the
hunchback+ Gene in the Drosophila Embryo
The Activity of Transcription Factors Themselves May Be
Regulated
16.5
Chromatin Structure and Transcription
Heterchromatic Chromosome Regions Are Not Transcribed
Puffs in Polytene Chromosome Are Actively Transcribed
DNA in Transcribed Chromatin Is Sensitive to DNase I
Digestion
Transcriptional Control Depends on Histone Acetylation and
Deacetylation
16.6
Transcriptional Control and Cell Determination
Combinatorial Action of Transcription Factors Explains the
Stepwise Process of Cell Determination
Cell Determination May Be Based on Bistable Control Circuit
of Switch Genes
Drosophila Homeotic Genes Show Switch Gene
Characteristics
DNA Methylation Maintains Patterns of Gene Expression
* Method 16.1 In Situ Hybridization
16.1
16.2
16.3
CHAPTER 17 / RNA PROCESSING
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17.1
17.2
Posttranscriptional Modifications of Pre-Messenger RNA
Control of Development by Alternative Splicing
A Cascade of Alternative Splicing Steps Controls Sex
Development in Drosophila
Alternative Splicing of Calcitonin and Neuropeptide mRNA Is
Regulated by Blockage of the Calcitonin-Specific Splice Acceptor Site
17.3
Messenger RNP Transport from Nucleus to Cytoplasm
Nuclear Pore Complexes Are Controlled Gates for the
Transport of RNPs to the Cytoplasm
Experiments with Cloned cDNAs Indicate Differential mRNP
Retention
17.4
Messenger RNA Degradation
The Half-Life of Messenger RNAs in Cells Is Regulated
Selectively
The Degradation of mRNAs in Cells Is Controlled by Proteins
Binding to Specific RNA Motifs
* Method 17.1 Ribonuclease Protection Assay
* Method 17.2 Pulse Labeling of Molecules and Their Half-life in Cells
CHAPTER 18 / TRANSLATIONAL CONTROL AND POSTRANSLATIONAL MODIFICATIONS
18.1
18.2
18.3
18.4
18.5
Formation of Polysomes and Nontranslated mRNP Particles
Most mRNAs Are Immediately Recruited into Polysomes and
Translated
Some mRNAs Are Stored as Nontranslated mRNP Particles
Mechanisms of Translational Control
Calcium Ions and pH May Regulate the Overall Rate of Protein
Synthesis at Fertilization
Phosphorylation of Initiation Factors and Associated Proteins
Controls Translation
Regulatory Proteins or RNAs “Mask” Critical Sequences of
Specific mRNAs
Polyadenylation and Deadenylation Control the Translation of
Specific mRNAs
Translational Control in Oocytes, Eggs, and Embryos
Early Embryos Use mRNA Synthesized During Oogenesis
Specific mRNAs Shift from Subribosomal mRNP Particles to
Polysomes during Oocyte Maturation
Translation of Some mRNAs Depends on Their Cytoplasmic
Localization
Translational Control during Spermiogenesis
Protamine mRNA Is Stored in Subribosomal RNP Particles
before Translation
Messenger RNA May Be Earmarked for Storage by Its 5’
UTR or 3’ URT Sequence
Posttranslational Polypeptide Modifications
Polypeptides Are Directed to Different Cellular Destinations
Polypeptides May Undergo Several Posttranslational
Modifications
Protein Degradation Is Differentially Controlled
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CHAPTER 19 / GENETIC AND PARAGENETIC INFORMATION
19.1
19.2
19.3
19.4
19.5
19.6
The Principle of Genetic and Paragenetic Information
Self-Assembly
Self-Assembly Is Under Tight Genetic Control
The Initiation of Self-Assembly Is Accelerated by Seed
Structures
The Conformation of Proteins May Change During SelfAssembly
Aided Assembly
Bacteriophage Assembly Requires Accessory Proteins and
Occurs in a Strict Sequence Order
Aided Assembly Is Common in Prokaryotic and Eukaryotic
Cells
Directed Assembly
One Bacterial Protein Can Assembly into Two Types of
Flagella
Prion Proteins Occur in Normal and Pathogenic Conformations
Tubulin Dimers Assemble into Different Arrays of Microtubules
Centrioles and Basal Bodies Multiply Locally and by Directed
Assembly
Ciliated Protozoa Inherit Accidental Cortical Rearrangements
Global Patterning in Ciliates
The Global Pattern in Ciliates Is Inherited During Fission
The Global Cell Pattern Is Maintained During Encystment
Global Patterning Is Independent of Local Assembly
Paragenetic Information in Metazoan Cells and Organisms
CHAPTER 20 / CELL DIFFERENTIATION
20.1
20.2
20.3
20.4
The Principle of Cell Differentiation
Each Organism Has a Limited Number of Cell Types
The Differentiated State Is Generally Stable
The Same Cell Type May Be Formed Through Different
Developmental Pathways
Cell Differentiation and Cell Division
Some Cells Divide in the Differentiated States
Other Cell Populations Are Renewed from Stem Cells
Stem Cells May Be Unipotent or Pluripotent
Stem Cell Development in Hydra
The Organization of Hydra is Relatively Simple
Interstitial Cells Contain Pluripotent and Unipotent Stem Cells
Growth and Differentiation of Blood Cells
The Same Universal Stem Cell Forms All Types of Blood Cells
as well as Endothelial Cells
Progenitor Cell Commitment May Depend on Stable Control
Circuits of Genes and Transcriptional Factors
Blood Cell Development Depends on Colony-Stimulating
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20.5
20.6
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Factors (CSFs)
Erythrocyte Development Depends on Successive Exposure to
Different CSFs
The Abundance of Blood Cells Is Controlled by Cell
Proliferation and Cell Death
Genetic Control of Muscle Cell Differentiation
Myogenesis is Controlled by a Family of Myogenic bHLH
Proteins
Myogenic bHLH Proteins Have Partially Overlapping Function
in Vivo
Inductive Signals Regulate Myogenic bHLH Gene Activity
Myogenic bHLH Proteins Interact with Cell Cycle Regulators
Myogenic bHLH Proteins Cooperate with MEF2 Factors in
Activating Muscle-Specific Target Genes
Unexpected Potency of Stem Cells and Prospects for Medical Uses
CHAPTER 21 / PATTERN FORMATION AND EMBRYONIC FIELDS
21.1
21.2
21.3
21.4
21.5
Regulation and the Field Concept
Characteristics of Pattern Formation
Pattern Formation Depends Upon Cellular Interactions
The Response to Patterning Signals Depends on Available
Genes
The Response to Patterning Signals Depends on Developmental
History
Patterning Signals Have a High Degree of Generality
The Concept of Receiving and Interpreting Positional Value
Limb Regeneration and the Polar Coordinate Model
Regeneration Restores the Elements Distal to the Cut Surface
Intercalary Regeneration Restores Missing Segments Between
Unlike Parts
Limb Regeneration Resembles Embryonic Limb Bud
Development
The Polar Coordinate Model Is Based on Two Empirical Rules
The Polar Coordinate Model Explains Supernumerary Limb
Formation from Misaligned Regenerates
Morphogen Gradients as Mediators of Positional Value
A Morphogen Gradient Can Specify a Range of Positional
Values and Polarity to a Field
Positional Values Specified by Morphogen Gradients May
Elicit Differential Gene Activities and Cell Behaviors
Morphogen Gradients Confer Size Invariance on Embryonic
Fields
Activin Can Form a Morphogen Gradient in Xenopus Embryos
Insect Development Has Been Model by Morphogen Gradients
and by Local Interactions
CHAPTER 22 / GENETIC AND MOLECULAR ANALYSIS OF PATTERN FORMATION IN THE DROSOPHILA
EMBRYO
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22.1
22.2
22.3
22.4
22.5
22.6
22.7
22.8
22.9
Review of Drosophila Oogenesis and Embryogenesis
Cascade of Developmental Gene Regulation
Maternal Genes Affecting the Anteroposterior Body Pattern
The Anteroposterior and Dorsoventral Axes Originate From
the Same Signal
The Anterior Group Generates and Autonomous Signal
The Posterior Group Acts by Depression
Many Posterior-Group Genes Are Also Required for Germ
Line Development
The Terminal Group Relies on the Local Activation of a
Receptor
Segmentation Genes
Gap Genes Are Controlled by Maternal Products and by
Interactions Among Themselves
Pair-Rule Genes Are Controlled by Gap Genes and by Other
Pair-Rule Genes
Segment Polarity Genes Define the Boundaries and Polarity of
Parasegments
Midway Through Embryogenesis, Parasegmental Boundaries
Are Replaced with Segmental Boundaries
Homeotic Genes
Homeotic Genes Are Expressed Regionally and Specify
Segmental Characteristics
The Homeotic Genes of Drosophila Are Clustered in Two
Chromosomal Regions
Homeotic Genes Are Part of a Complex Regulatory Network
The Dorsoventral Body Pattern
The Compartment Hypothesis
Compartments Are Controlled by the Activities of Specific
Selector Gene Activities
The Ultrabithorax+ Expression Domain Coincides with a
Compartment
Engrailed+ Controls the Anteroposterior Compartment
Boundary
Apterous+ Controls the Dorsoventral Compartment Boundary
Appendage Formation and Patterning
Compartment Interactions across Boundaries Organize Pattern
Formation within Compartments
Decapentaplegic Protein Forms a Morphogen Gradient in the
Wing Imaginal Disc
Patterning Genes in Other Insects
CHAPTER 23 / THE ROLE OF HOX GENES IN VERTEBRATE DEVELOPMENT
23.1
Strategies for Identifying Patterning Genes in Vertebrates
Genes Are Cloned on the Basis of Their Chromosomal Map
Position
Promoter Trapping Identifies Patterning Genes
Vertebrate Genes Can Be Isolated with Molecular Probes from
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23.2
23.3
23.4
23.5
23.6
Kalthoff / Second Edition Contents
Drosophila Genes
Homeotic Genes and Hox Genes
The Homeodomain is a Cooperative DNA-Binding Region
Hox Genes Can Be Divided in Paralogy and Orthology Groups
The Homeodomain is Highly Conserved in Evolution
The Role of Hox Genes in the Anteroposterior Body Pattern
The Order of Hox Genes on the Chromosome Is Colinear with
Their Expression in the Embryo
Boundaries of Hox Genes May Cause Transformations
Towards More Anterior Fates
Overexpression of Hox Genes May Cause Transformations
Toward More Posterior Fates
Only Some of the Control Systems of Hox Gene Activity Are
Evolutionarily Conserved
The Dorsoventral Body Pattern
A Lobster May Be Viewed as an Upside-Down Rabbit
Dorsoventral Patterning Is Controlled by the Same Genes in
Flies and Frogs
Pattern Formation and Hox Gene Expression in Limb Buds
Limb Field Initiation Is Affected by FGF Signaling and Hox
Gene Expression
T-box Genes Control the Difference Between Hindlimb and
Forelimb
The Proximodistal Pattern Is Formed in the Progress Zone
Signals From Somites and Lateral Plate Specify the
Dorsoventral Limb Bud Axis
A Zone of Polarizing Activity Determines the Anteroposterior
Pattern
Sonic Hedgehog Protein Has Polarizing Activity
Hox Gene Expression Mediates Anteroposterior Limb
Patterning
The Principle of Reciprocal Interaction
CHAPTER 24 / GENETIC AND MOLECULAR ANALYSIS OF PATTERN FORMATION IN PLANTS
24.1
24.2
24.3
Reproduction and Growth of Flowering Plants
Spore-Forming Generations Alternate with Gamete-Forming
Generations
Plant Embryos Develop Inside the Flower and Fruit
Meristems at the Tips of Root and Shoot Allow Plants to Grow
Continuously
Genetic Analysis of Pattern Formation in Plant Embryos
Similar Methods Are Used for the Genetic Analysis of Plant
and Animal Development
Pattern Formation in Plant Embryos Involves 25 to 50 Specific
Gene Functions
Groups of Genes Control Distinct Patterning Events
Genetic Control of Flower Development
Floral Induction Genes Control the Formation of an
Inflorescence
Kalthoff / Second Edition Contents
24.4
24.5
Meristem Identity Genes Promote the Transition from Floral
Meristems to Inflorescence Meristems
The Role of Homeotic Genes in Flower Patterning
Arabidopsis Has Homeotic Genes
Three Classes of Homeotic Genes Determine the
Morphological Characters of Four Flower Whorls
Double and Triple Homeotic Phenotypes Confirm the Genetic
ABC Model
Homeotic Genes are Controlled by Regulator Genes, Mutual
Interactions, and Feedback Loops
Other Plants Have Patterning Genes Similar to Those of
Arabidopsis
Antirrhinum Has Genes Controlling Organ Variations Within
the Same Whorl
Molecular Analysis of Homeotic Plant Genes
The agamous+ Gene Encodes a Transcription Factor
Genetic and Molecular Studies Reveal Orthologous Genes
between Arabidopsis and Antirrhinum
The Known Plant Homeotic Genes Have a MADS Box
Instead of a Homeobox
CHAPTER 25 / EXPERIMENTAL AND GENETIC ANALYSIS OF CAENORHABIDITIS ELEGANS
DEVELOPMENT
25.1
25.2
25.3
25.4
Normal Development
Hermaphrodites and Males
Fertilization, Cleavage, and Axis Formation
Founder Cells
Gastrulation, Organogenesis, and Histogenesis
Larval Development
Localization and Induction During Early Cleavage
The First Cleavage Generates Blastomeres with Different
Potentials
P Granules Are Segregated into Germ Line Cells
Par Genes Affect Cytoplasmic Localization
Three Maternal Effect Genes Generate a Localized Activity that
Promotes EMS Identity
Determinant Activity for the EMS Blastomere
Determination of Anterior Pharyngeal Muscle Cells Requires
Multiple Inductive Interactions
P2 Polarizes EMS to Form Unequal Daughter Cells
Heterochronic Genes
Mutations in the lin-14+ Gene are Heterochronic
The lin-14+ Gene Encodes a Nuclear Protein That Forms a
Temporal Concentration Gradient
The lin-4+ Gene Encodes Small Regulatory RNAs with
Antisense Complementarity to lin-14 mRNA
Down-Regulation of lin-14+ Expression Is Initiated by a
Developmental Cue
Programmed Cell Death
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Programmed Cell Death in C. elegans Is Controlled by a
Genetic Pathway
 The Strategy of Genetic Mosaic Analysis Shows That ced-3+
and ced-4+ Act Cell-Autonomously
The egl-1, ced-9, and ced-4 Gene Products Control the
Initiation of Apoptosis
The Genes ces-1+ and ces-2+ Control the Programmed Death
of Specific Cells in the Pharynx
The Product of a Human Gene, bcl-2+, Prevents Programmed
Cell Death in C. elegans
Vulva Development
The Vulval Precursor Cells Form an Equivalence Group
The Gonadal Anchor Cell Induces The Primary VPC Fate
A Hypodermal Signal Inhibits Vulva Formation
CHAPTER 26 / SEX DETERMINATION
26.1
26.2
26.3
26.4
26.5
Enviromental Sex Determination
Genotypic Sex Determination
Sex Determination in Mammals
Maleness in Mammals Depends on the Y Chromosome
Dosage Compensation in Mammals Occurs by X Chromosome
Inactivation
The First Morphological Sex Difference Appears in the Gonad
The Testis-Determining Factor Acts Primarily in Prospective
Sertoli Cells
Mapping and Cloning of the Testis-Determining Factor
Translocated Y Chromosome Fragments Causing Sex Reversal
Are Used to Map TDF
The SRY/ Sry Function Is Necessary for Testis Determination
The Mouse Sry+ Gene Can Be Sufficient for Testis
Determination
The SRY+ Gene Controls Primary Sex Differentiation
Sex Determination and Sex Differentiation in Drosophila, C. elegans and Mammals
One Primary Signal Controls Multiple Aspects of Sex
Differentiation
The X:A Ratio Is Measured by Numerator and Denominator
Elements
Drosophila and C. elegans Have Master Regulatory Genes for
Sex Differentiation
Dosage Compensation Mechanisms Vary Widely
The Somatic Sexual Differentiation Occurs with Different
Degrees of Cell Autonomy
Germ Line Sex Differentiation Involves Interactions with
Somatic Gonadal Cells
CHAPTER 27 / HORMONAL CONTROL OF DEVELOPMENT
27.1
General Aspects of Hormone Action
Kalthoff / Second Edition Contents
27.2
27.3
27.4
27.5
Hormonal Control of Sex Differentiation in Mammals
The Synthetic Pathway for Male and Female Sex Hormones
Are Interconnected
The Genital Ducts Develop from Parallel Precursors, the
Wolffian and Mllerian Ducts
Male and Female External Genitalia Develop from the Same
Embryonic Primordia
Hormonal Control of Brain Development and Behavior in Vertebrates
Androgen and Estrogen Receptors Are Distributed Differently
in the Brain
Androgens Cause Seasonal and Sexually Dimorphic Changes in
the Brains of Birds
Prenatal Exposure to Sex Hormones Affects Adult
Reproductive Behavior and Brain Anatomy in Mammals
Hormonal Control of Insect Metamorphosis
Insect Molting, Pupation, and Metamorphosis Are Controlled
by Hormones
Puffs in Polytene Chromosomes Reveal Genes Responsive to
Ecdysone
Multiple Ecdysone Receptors Have Tissue-Specific Activities
Early Regulatory Genes Diversify and Coordinate the
Responses of Target Cells to Ecdysone
Hormonal Control of Amphibian Metamorphosis
The Hormonal Response in Amphibian Metamorphosis Is
Organ-Specific
Amphibian Metamorphosis Is Controlled by Thyroid Hormone
The Production of Thyroid Hormone Is Controlled by the Brain
Various Defects in Metamorphosis Cause Paedomorphic
Development in Salamanders
The Receptors for Ecdysone and Thyroid Hormone Share
Many Properties
CHAPTER 28 / ORGANISMIC GROWTH AND ONCOGENES
28.1
28.2
28.3
28.4
Measurement and Mechanisms of Growth
Growth is Defined as Change in Mass
Growth May Be Isometric or Allometric
Growth Occurs by Different Mechanisms
Growth Analysis by Heterospecific Transplantation
The Growth Potential of a Limb Is a Property of the Limb’s
Mesoderm
The Optic Cup and the Lens of the Eye Adjust Their Growth
Rates to Each Other
Growth Hormones
Growth Factors
Nerve Growth Factor Promotes the Growth and Differentiation
of Certain Neurons
Other Growth Factors Affect Multiple Target Cells in a Variety
of Ways
The Effects of a Growth Factor May Depend on the Presence
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28.6
28.7
28.8
Kalthoff / Second Edition Contents
of Other Growth Factors
Mechanical Control of Cell Survival and Division
Cell Cycle Control
Tumor-Related Genes
Oncogenes Are Deregulated or Mutated Proto-oncogenes
Proto-oncogenes Encode Growth Factors, Growth Factor
Receptors, Signal Proteins, Transcription Factors, and Other Proteins
Oncogenes Arise from Proto-Oncogenes Through Various
Genetic Events
Tumor Suppressor Genes Limit the Frequency of Cell Divisions
Growth and Pattern Formation
Patterning Signal Control Cell Cycling and Growth
Local Cell Interactions Affect Cell Division and Growth
Organisms Measure Growth by Mass or Size Rather Than by
Cell Number
CHAPTER 29 / SENESCENCE
29.1
29.2
29.3
29.4
29.5
Statistical Definition of Senescence
Evolutionary Theory of Senescence
The Mutation Accumulation Hypothesis Focuses on the Age
Distribution in Natural Populations
Senescence Can Be Ascribed to Antagonistic Pleiotropy
According to the Disposable Soma Hypothesis, Every
Individual is the Temporary Carrier of His/Her Germ Line
Various Physiological Causes of Senescence
Cellular Senescence and Telomerase
Somatic Mammalian Cells Show a Limited Capacity for
Proliferation
Chromosomal Ends Are Protected by Telomeres
Telomerase Activity Counteracts Cellular Senescence
Loss of Telomerase Activity is a Safeguard Against Cancer
Damage from Oxidants and Organismic Senescence
Oxidative Phosphorylation Generates Aggressive Oxidants as
Intermediates
Oxidants Damage Cellular DNA, Lipids and Proteins
Superoxide Radical Stimulates Cell Division and Growth
Antioxidative Agents Delay Senescence