Hox Genes

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  • Edition: 1st
  • Format: Hardcover
  • Copyright: 2010-06-25
  • Publisher: Landes Bioscience

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The purpose of this book is to argue, and to convince those who may not agree, that the Hox genes are indeed so important that their study will not end with the 20th century. The book is divided into three major sections. The first section covers aspects of the regulation of Hox gene expression and the structure and function of the now justifiably well-known homeobox. The second section offers insights and discussions of the sometimes contentious issues of the origin and evolution of the aforementioned Hox complexes. In the third and last section the role of the resident loci in the specification of body plans and meristic identity of thearthropods is presented.

Table of Contents

Mechanisms of Activity
Regulation of Hox Activity: Insights from Protein Motifsp. 3
Abstractp. 3
Introductionp. 3
The Homeodomainp. 4
The Hexapeptide Motifp. 8
Additional Hox Functional Motifsp. 9
Conclusionp. 10
CIS-Regulation in the Drosophila Bithorax Complexp. 17
Abstractp. 17
Genetics of the Bithorax Complex: The Model of Ed Lewisp. 17
The BX-C Encodes Only Three Genes, Ubx, abd-A and Abd-Bp. 19
The Segment-Specific Functions Act as Segment/Parasegment-Specific Enhancersp. 22
Initiation and Maintenance Phase in BX-C Regulationp. 23
Initiation, Maintenance and Cell Type-Specific Elements within the Cis-Regulatory Domainp. 25
The Cis-Regulatory Regions Are Organized in Segment-Specific Chromosomal Domainsp. 26
Chromatin Boundaries Flank the Parasegment-Specific Domainsp. 28
Elements Mediating Long-Distance Cis- and Trans- Regulatory Interactionsp. 28
Transvection Studiesp. 28
Promoter Targeting Sequencesp. 31
Promoter Tethering Elementp. 32
Intergenic Transcription in the BX-Cp. 32
MicroRNAs in the BX-Cp. 34
Conclusionp. 35
Maintenance of Hox Gene Expression Patternsp. 41
Abstractp. 41
Introductionp. 41
Genetics of PcG and trxG Genesp. 42
PcG Proteins and Their Complexesp. 43
TrxG Proteins and Their Complexesp. 46
ETP Proteinsp. 46
PcG and trxG Response Elementsp. 47
Recruitment of Maintenance Proteins to Maintenance Elementsp. 48
Role of Maintenance Proteins in Regulation of Transcriptionp. 50
Epigenetic Marksp. 51
Release of PcG Silencingp. 53
Role of PcG Proteins in Chromatin Replicationp. 54
Role of PcG Proteins in Stem Cellsp. 54
Conclusionp. 55
Future Research in the Fieldp. 55
Control of Vertebrate Hox Clusters by Remote and Global Cis-Acting Regulatory Sequencesp. 63
Abstractp. 63
Introductionp. 63
Colinearity and Clustering of the Homeotic Genes: An Obligatory Functional Link?p. 64
Vertebrate Hox Clusters Are More Clustered Than Othersp. 65
Global Regulation of the Complex through Shared Mechanisms: The Retinoic Acid Connectionp. 66
High-Order Structures Over the Complex and Colinearityp. 66
Control of Vertebrate Hox Genes by Shared Internal Enhancersp. 67
The Ins and Outs of Hoxd Gene Regulationp. 67
The Role of the Flanking Regions in the Control of Vertebrate Hox Genesp. 68
Control of the HoxD Cluster through Remote Enhancersp. 69
Regulation of the HoxD Cluster and More: Global Control Regions and Regulatory Landscapesp. 70
Remote Enhancers for the Other Vertebrate Hox Clusters?p. 71
An Evolutionary Success Story and an Increasing Need for a Global Regulationp. 73
Conclusion and Outlook for Hox Gene Regulation in the 21st Centuryp. 74
Evolution of Hox Genes and Complexes
The Early Evolution of Hox Genes: A Battle of Belief?p. 81
Abstractp. 81
The Hox Systemp. 81
Phylogenetic Evidencep. 82
Opposing Viewsp. 84
Conclusionp. 87
Evolution of Hox Complexesp. 91
Abstractp. 91
Introductionp. 91
Origin of the ProtoHox Genep. 91
Origin of the Hox Cluster from a ProtoHox Cluster, or Not?p. 92
Expansion and Contraction of the Number of Hox Genes in Evolutionp. 96
Conclusionp. 98
The Nematode Story: Hox Gene Loss and Rapid Evolutionp. 101
Abstractp. 101
Introduction: Hox Gene Loss, the Third Wayp. 101
The Caenorhabditis elegans Hox Cluster, an Extreme Case of Gene Lossp. 102
Tracing Hox Gene Loss through the Nematode Phylum: Mode and Tempop. 104
Sea Squirts and Nematodes: Why Do Both Groups Lose Hox Genesp. 105
Hox Gene Loss in Flagrantep. 106
Nematode Hox Gene Function: A Story of Novelty, Conservation and Redeploymentp. 106
Conclusionp. 108
Are the Deuterostome Posterior Hox Genes A Fast-Evolving Class?p. 111
Abstractp. 111
The Distribution of the Posterior Hox Genes in the Metazoap. 111
Early Duplications of the Posterior Hox Genesp. 113
The 'Deuterostome Posterior Flexibility' Hypothesisp. 114
The Mechanistic Basis of Deuterostome Posterior Flexibilityp. 116
Conclusion and Future Directionsp. 118
Biological Function
Hox Genes and the Body Plans of Chelicerates and Pycnogonidsp. 125
Abstractp. 125
Arthropods, Mandibulates vs Cheliceratesp. 125
Chelicerate Hox Genesp. 126
Chelicerate Hox Genes and the Chelicerate vs Mandibulate Body Planp. 127
Hox Genes and the Enigmatic Sea Spider Body Planp. 130
Conclusionp. 131
Hox3/-zen and the Evolution of Extraembryonic Epithelia in Insectsp. 133
Abstractp. 133
Introductionp. 133
Setting the Stage: Morphological Evolution of Extraembryonic Developmentp. 134
Variants of zen Expression and Function in Insects and Possible Morphological Correlatesp. 135
The Amnioserosa Gene-Network in Evolutionary Perspectivep. 140
Conclusionp. 141
Hox Genes and Brain Development in Drosophilap. 145
Abstractp. 145
Introductionp. 145
Expression and Function of Hox Genes in Embryonic Brain Developmentp. 146
Genetic Interactions between Hox Genes in Embryonic Brain Developmentp. 148
Hox Genes in Postembryonic Brain Developmentp. 149
Evolutionary Conservation of Hox Gene Action in Brain Developmentp. 151
Conclusionp. 152
Homeosis and Beyond. What is the Function of the Hox Genes?p. 155
Abstractp. 155
What Are the Hox Genes?p. 155
The Hox Genes' Explosionp. 156
What Is the Function of a Gene?p. 156
Hox Genes' Function at the Molecular and Cellular Levelsp. 157
Hox Genes and Homeosisp. 157
Homeosis as a Differential Functionp. 157
Hox Genes as 'Meta-Selector' Genesp. 158
The Hox Specificity Paradoxp. 158
Posterior Prevalencep. 159
An Evolutionary Paradox: Morphological Differentiation and the Hox Repertoirep. 159
Hox and Neuronal Homeosisp. 159
Morphological Homeosis as a Derived Propertyp. 160
Why Does the 'Hox System' Make Sense?p. 160
Conclusionp. 161
Indexp. 167
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