chromosome biology
TRANSCRIPT
CHROMOSOME BIOLOGY
CHROMOSOME BIOLOGY
Kudi Appels CSIRO, Division of Plant Industry
Canberra, ACT, Australia
Kosalind Morris Deparlment of Agronomy
University of Nebraska, Lincoln, Nebraska
Bikram S. GUI Deparlment of Pathology
Kansas state University, Manhattan, Kansas
Cedric E. May Agricultura) Research Institute Wagga Wagga, NSW, Australia
Illustrations Rudi Appels and Jon Raupp
Springer Science+Business Media, LLC
Library ~ Congress CataIoging-in-PublicatiOll
Cbromosome biology I Rudi Appels ... [et al.]. p.em. lncIudes bibliograpbical references anei index. ISBN 978-1-4613-7470-1 ISBN 978-1-4615-5409-7 (eBook) DOI 10.1007/978-1-4615-5409-7 1. Chromosomes. 2. Cytogenetics. 3. DNA. 1. Appels, Rudi. QH600.C494 1998 572.8'7-«21 97-15812
CIP
Copyright o 1998 by Springer Science+Busine •• Media New York: Second Printing 2003. Originally published by Kluwer Academic Publishers in 1998 1bis printing is a digital duplication of the origioal edition.
AlI rigbts reserved. No part of this publication may be reproduced, stored in a retrieval system or cransmitted in any form or by any meaos, electronic, mecbaoical, photo-c:opyiDg, microfilming, recording, or otherwise, without the prior written permiBsion of the publisher, wiIb the exception of any liIaterial supplied specificaIly for tbe pmpose of being entered anei executed on a computer system, for exclusive use by the purchaser of the work. Permission for books published in tbe USA: [email protected] Permissions for books publisbed in Europe: [email protected] Printtd 0" ocid-free paper.
Contents
PART I INTRODUCTION TO THE STUDY OF CHROMOSOMES
Chapter 1. Introduction 3
1.1 The value of chromosome research 3 1.2 Life-forms and their genetic material 4 1.3 Chromosomes and protosomes 4 1.4 The origin of cellular organelles 5 1.5 Conclusions 6
Bibliography 6
Chapter 2. A Historical Perspective on Chromosome Structure, Function, and Behavior 7
2.1 The pre-l900 period 7 2.2 The 1900-1950 period 8 2.3 The 1950-1996 period 11
Bibliography 17
Chapter 3. Microscopes: Basic Tools for Cytogenetics 22
3.1 Light microscopy 22 3.1.1 The standard light microscope 22 3.1.2 Microscopy using ultraviolet light 23 3.1.3 The phase-contrast microscope 24 3.1.4 The interference-contras\ microscope 24 3.1.5 The confocal microscope 25
3.2 Electron microscopy 26 3.2.1 The transmitting electron microscope 26 3.2.2 The scanning electron microscope 27
3.3 Scanning probe microscopy 27 3.4 Flow cytophotometry 28 3.5 Computer-assisted image analysis 28
Bibliography 29
PART II MITOSIS, MEIOSIS, AND CHROMOSOMAL KARYOTYPES
Chapter 4. Chromosomes in the Mitotic Cell Cycle 33
4.1 The cell cycle consists of two cycles occurring in concert 33 4.1.1 The chromosomes replicate and divide during mitosis 33 4.1.2 The assembly of the spindle apparatus 37 4.1.3 The plane of cell division controls the fate of cells 39
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4.2 Universal mechanisms regulate the cell cycle 39 4.2.1 Kinase and cyclin proteins regulate the eukaryotic cell cycle 40 4.2.2 Cell-fusion experiments indicate that diffusible cellular components control
the cell cycle 41 4.2.3 Overview of the mutations and proteins important in mitosis 42 4.2.4 Oncogenes code for proteins that influence the phosphorylation!
dephosphorylation cycle 42 Bibliography 42
Chapter S. Meiosis and Gamete Formation: A View of Chromosomes in the Germline 44
5.1 The meiotic and postmeiotic cycles 44 5.1.1 Meiosis I 44 5.1.2 Meiosis II 50 5.1.3 Postmeiotic events 51 5.1.4 Achiasmate meiosis 51 5.1.5 Meiosis in organisms with multiple or diffuse centromeres 52 5.1.6 Other variations in the meiotic process 54
5.2 Chromosome synapsis and the synaptonemal complex 54 5.2.1 The synaptonemal complex forms between chromosomes located close to each other 54 5.2.2 The synaptonemal complex is a flexible association between chromosome pairs 57 5.2.3 DNA topoisomerase II unravels interlocked bivalents 57 5.2.4 Presynaptic chromosome alignment at zygotene begins at multiple sites 58 5.2.5 Mechanisms of homologous recognition 59
5.3 The mechanism of crossing over and genetic recombination 60 5.3.1 Crossing over occurs at the four-strand stage during pachynema 60 5.3.2 The recombination process 60 5.3.3 Relationship between genetic recombination and cytological chiasmata 61 5.3.4 The distribution of chiasmata 62 5.3.5 Mechanisms for chiasma maintenance at metaphase I 62
5.4 Distributive system of segregation 63 5.5 Many genes control the meiotic process 63
Bibliography 64
Chapter 6. Chromosome Morphology and Number 67
6.1 Mitotic karyotype analysis 67 6.1.1 Number and morphology of chromosomes are usually constant for an organism 67 6.1.2 Prophase chromosomes reveal additional cytological landmarks 69
6.2 Special staining techniques reveal cytological landmarks in metaphase chromosomes 69 6.2.1 C bands reveal the location of cOIistitutive heterochromatin 71 6.2.2 G, R, and Q bands reveal chromomere structure 71 6.2.3 Combinations of stains can enhance the resolution of bands 71 6.2.4 Many specialized techniques reveal functional domains in metaphase chromosomes 74
6.3 Meiotic karyotype analysis 76 6.3.1 Origins of cytogenetics are based on pachytene and metaphase I chromosome
analyses 76 6.3.2 Giant lampbrush chromosomes and diplotene karyotypes 76
6.4 Polyteny produces giant chromosomes in some differentiated tissues 78 6.5 Sex chromosomes 78
6.5.1 Sex determination in different organisms 79 6.5.2 Structure and behavior of sex chromosomes 80
6.6 B chromosomes 80 6.6.1 Unusual behavior of B chromosomes 81 6.6.2 Genetic functions associated with B chromosomes 81 6.6.3 Origin of B chromosomes may be related to cytoplasmic-specific introgression during
interspecific hybridization 81 Bibliography 82
PART m CHROMOSOME STRUCTURE AND REARRANGEMENTS
Chapter 7. Structural Stability of Chromosomes
7.1 Chromosome structural stability 87 7.2 Chromosome instability and repair 88
7.2.1 Timing of chromosome breakage 88 7.2.2 Major groups of chromosome abnormalities 88 7.2.3 Repair of damaged chromosomes 88
7.3 Sister-chromatid exchange 91 7.4 Agents and conditions inducing chromosome breakage 92
7.4.1 Ionizing radiations 92 7.4.2 Ultraviolet radiation ' 92 7.4.3 Chemically induced damage to chromosomes 93 7.4.4 Chromosomal changes resulting from tissue culture 94 7.4.5 Genetic predisposition to chromosomal breakage 94 7.4.6 Chromosome breakage and the aging process 95
7.5 Transposable elements and chromosome instability 95 7.5.1 The genetics of transposable elements 95 7.5.2 Transposable elements in prokaryotes 98
7.6 Dispersed repeated DNA sequences and chromosome instability 98 7.7 Gametocidal chromosomes 99
Bibliography 100
Chapter 8. Losses and Gains of Chromosome Segments
8.1 Range of structural changes 103 8.2 Deletions and duplications of chromosome segments 103 8.3 Detection of deletions and duplications 105 8.4 Behavior of modified chromosomes during cell division 106 8.5 Effects of deletions and duplications on viability 107
8.5.1 Effects on viability 107 8.5.2 Effects on fertility 108
8.6 Phenotypic effects of deletions and duplications 109 8.6.1 Deletions 109 8.6.2 Duplications III
8.7 Evolutionary significance of duplications III Bibliography 113
Chapter 9. Rearrangements of Chromosome Segments
9.1 9.2
9.3
9.4
9.5 9.6 9.7
How chromosome rearrangements are detected 115 Inversions 115
9.2.1 Behavior of inversions during mitosis and meiosis 115 9.2.2 Effects of heterozygous inversions on fertility 118
Reciprocal translocations 119 9.3.1 Behavior of translocations during mitosis and meiosis 122 9.3.2 Effects of heterozygous translocations on fertility 125
Phenotypic effects of chromosome rearrangements 126 9.4.1 Chromosomal rearrangements and position-effect variegation
Relationships between chromosomal aberrations and cancer in humans Chromosome rearrangements in natural populations 132 Some specific uses of reciprocal translocations 133
9.7.1 Source of stable duplications 133 9.7.2 Use of translocations to control insect pests 134
Bibliography 135
127 129
PART IV CHROMOSOME POLYPLOIDY, ANEUPLOIDY, AND HAPLOIDY
Chapter 10. Multiples of Basic Chromosome Numbers-Polyploidy
10.1 Designating ploidy levels and somatic versus gametic chromosome numbers 139
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103
115
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10.2 10.3 10.4 10.5 10.6
10.7
10.8
10.9 10.10
10.1.1 Diploidy 139 10.1.2 Polyploid types in relation to genome composition
Polyploidy occurs in whole organisms or part of organisms Whole-organism polyploidy 141 Tissue or organ polyploidy 145 Effects of polyploidy on phenotypes and internal traits 146 Meiotic behavior and fertility in polyploids 146
10.6.1 Meiotic behavior of autopolyploids 147 10.6.2 Fertility of autopolyploids 148 10.6.3 Meiotic behavior of allopolyploids 149 10.6.4 Fertility of allopolyploids 150
Gene segregations in polyploids 150 10.7.1 Allopolyploids 150 10.7.2 Autopolyploids 150
Genome analyses in polyploids 152
139 140
1O~8.1 Identifying diploid ancestors of allopolyploids 154 10.8.2 Quantitative tests of genome relationships in polyploid hybrids 10.8.3 Assigning chromosomes to genomes 155
154
10.8.4 Homoeologous relationships between chromosomes of different genomes 10.8.5 Species-specific translocations 157
Adaptability of natural polyploids 157 Production and uses of induced polyploids Bibliography 158
157
Chapter 11. Deviations from Basic Chromosome Numbers-Aneuploidy
11.1 Types of aneuploids 160 11.2 Occurrence and viability of aneuploidy 160 11.3 Modes of origin of aneuploidy 162
11.3.1 Nondisjunction of the centromeres 162 11.3.2 Univalent state of chromosomes 162
11.4 Conditions that cause aneuploidy 163 11.4.1 Mutations in genes controlling meiotic pairing and separation 163 11.4.2 Occurrence of haploid states 164 11.4.3 Crosses involving autotriploids 164 11.4.4 Crosses between related species 164
155
11.4.5 Reciprocal translocations with irregular chromosome distributions during meiosis 164
11.5 11.6
11.4.6 Robertsonian translocations 164 11.4.7 Aging of cells or organisms 11.4.8 Environmental conditions 11.4.9 Viral infections 166 11.4.10 Physiological stress 166
164 165
Effects of aneuploidy on phenotypes and internal traits Meiotic behavior, fertility, and transmission of aneuploids
11.6.1 Trisomies and related types 169 11.6.2 Monosomics and related types 171
Bibliography 173
Chapter 12. The Cytogenetic Analysis of Haploids
12.1 The origins of haploids 175 12.1.1 Natural occurrences of haploids 175 12.1.2 The induction of haploids 176
12.2 Detection and phenotypes of haploids 179 12.3 Meiotic behavior and fertility in haploids 179
12.3.1 Monohaploids 180 12.3.2 Polyhaploids 180 12.3.3 Secondary associations 181
12.4 Uses of haploids in plants 181 12.4.1 Screening for mutations using haploids 181
166 169
12.4.2 Uses of haploids in breeding and genetic research 182
160
175
12.5 Production and uses of doubled haploids in plants 182 12.5.1 Methods for doubling haploids 182 12.5.2 Genetic composition of doubled haploids 182 12.5.3 Uses of doubled haploids 183
Bibliography 184
PART V GENE LOCATIONS, GENETIC MANIPULATIONS, AND REARRANGEMENTS
Chapter 13. Locating Genes in Diploids Using Chromosome Aberrations
13.1 Diploid analysis using trisomies 189 13.2 Use of primary trisomies 189 13.3 Use of telotrisomies, aerotrisomies, and tertiary trisomies 192
13.3.1 Telotrisomies 192 13.3.2 Aerotrisomies and tertiary trisomies 193
13.4 Diploid analysis using monosomies 194 13.5 Diploid analysis using translocations and inversions 195
13.5.1 The inverted or translocated chromosome must be identified 195 13.5.2 The breakpoints that produce the inversion or translocation must be determined
precisely in order to map the genes accurately 195 13.6 General procedure using inversions or translocations to locate genes 196
13.6.1 Use of A-A reciprocal translocations 196 13.6.2 Use of B-A translocations 197 13.6.3 Use of Inversions 199
13.7 Diploid analysis using deletions 199 Bibliography 200
Chapter 14. Locating Genes in Polyploids Using Chromosome Aberrations
14.1 Polyploid analysis using monosomics 202 14.2 Polyploid analysis using nullisomics 204 14.3 Polyploid analysis using telochromosomes and isochromosomes 204
14.3.1 Chromosome-arm locations of genes based on dosage effects 204 14.3.2 Mapping gene-centromere distances using telochromosomes 205
14.4 Polyploid analysis using intervarietal chromosome substitutions 206 14.4.1 Development of intervarietal chromosome substitution lines 206 14.4.2 Genetic parameters for chromosome substitution lines and their hybrids 208 14.4.3 Cytogenetic studies using intervarietal chromosome substitution lines 208 14.4.4 Crosses involving substitution lines 209
Contents ix
189
201
14.5 Early detection of genetic differences between donor and recipient homologous chromosomes 210 14.6 Chromosome irregularities associated with monosomics and substitution lines 211
14.6.1 Monosomic shift 211 14.6.2 Misdivision of monosomes 211 14.6.3 Reciprocal translocations 211
14.7 Genetic heterogeneity 212 Bibliography 212
Chapter 15. Gene Transfers by Chromosome Manipulations 214
15.1 The reason for gene transfers 214 15.2 Interspecific and intergeneric crosses 215 15.3 Transfers at the genome level (amphiploids) 216 15.4 Transfers of individual chromosomes (addition and substitution lines) 217
15.4.1 Alien-addition lines 217 15.4.2 Alien-substitution lines 218
15.5 Transfers of alien chromosome arms 220 15.6 Transfers of alien-arm segments 220
15.6.1 Homoeologous pairing 220 15.6.2 Translocations 221
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15.7 Methods for detecting alien chromatin 223 Bibliography 224
PART VI DNA ORGANIZATION, STRUCTURE, AND REPLICATION
Chapter 16. Organization of DNA Sequences in Protosomes and Chromosomes
16.1 The variable amount of DNA and the arrangement of sequences in organisms 229 16.2 Coding and noncoding DNA sequences 229 16.3 DNA of the virus SV40 encodes for overlapping genetic information and contains repetitive
sequences 230 16.4 Bacterial DNA is characterized by the occurrence of groups of genes under a common control 232
16.4.1 The operon organization of genes 232 16.4.2 Transcription attenuation 233
16.5 Repetitive DNA sequences comprise up to 2% of the E. coli proto some 234 16.6 Small, circular DNA molecules (plasmids) in E. coli code for 1-3 genes 235 16.7 Repetitive sequences are a major feature in eukaryotic nuclear DNA 236
16.7.1 Repetitive-sequence families are characterized by episodes of amplification and deletion during evolution 237
16.7.2 Dispersed repetitive sequences fall into two broad classes 237 16.8 Repetitive sequences in and around eukaryotic genes 239
16.8.1 The histone genes 239 16.8.2 The hemoglobin genes 240
16.9 RNA processing is an integral aspect of understanding the way DNA sequences are organized 241 Bibliography 241
Chapter 17. Variable Structure and Folding of DNA
17.1 The structure of DNA 244 17.1.1 Polarity of the sugar-phosphate backbone is a key feature of DNA 244 17.1.2 Variation in the structure of DNA 244 17.1.3 Curvature of DNA can result from a particular sequence of base pairs 246
17.2 Structural modification of DNA by methylation 248 17.3 Interaction of small molecules with DNA 250
17.3.1 Water 250 17.3.2 Spermine, ions, and atoms 251 17.3.3 Dye and antibiotic molecules 251
17.4 Protein molecules interact with DNA to determine its folding and transcription properties 253 17.4.1 Helix-tum-helix modules 253 17.4.2 Zn finger modules 253 17.4.3 Leucine-zipper modules 255 17.4.4 Distortion of the DNA helix by TATA-box binding protein 255
17.5 Folding of DNA into chromatin: the nucleosome 256 17.5.1 Histone proteins 256 17.5.2 Nonhistone proteins are also important in the organization of DNA into
chromosomes 258 17.5.3 Chromatin domains 260
17.6 Structural changes in chromatin during gene expression 262 17.7 Folding of DNA-protein complexes on a broader scale within the interphase nucleus 263
Bibliography 265
Chapter 18. Replication of Protosomes and Chromosomes
18.1
18.2
18.3
Semiconservative DNA replication 18.1.1 Protosomes 270 18.1.2 Chromosomes 270
DNA synthesis in prokaryotes 18.2.1 The proteins involved 18.2.2 Fidelity of replication
DNA replication in chromosomes
270
272 272 274 274
229
244
270
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18.3.1 Origins of replication 274 18.3.2 Proteins involved in chromosomal DNA replication 277 18.3.3 The assembly of newly formed chromosomes 277 18.3.4 The synthesis of histones in the S phase of the cell cycle 278 18.3.5 Synthesis of histones can occur outside the limits of the S phase 278
18.4 DNA synthesis outside the normal cell cycle is limited to specific regions of the genome 279 Bibliography 280
PART VII IN SITU HYBRIDIZATION, CHROMOSOME LANDMARKS, GENETIC MAPPING
Chapter 19. Exploring Chromosomes by In Situ Biochemical Reactions
19.1
19.2
19.3 19.4 19.5 19.6
Principles underlying in situ reactions 285 19.1.1 Preparation of tissues for study 285 19.1.2 Specificity in the primary reaction 286 19.1.3 The secondary or "capture" reaction to detect the results of the primary reaction
Sources of radioactive and nonradioactive DNA and RNA probes 288 19.2.1 In situ hybridization with fluorescently labeled DNA (FISH) 289 19.2.2 In situ hybridization to DNA fibers (Fiber-FISH) 290
In situ reactions to modify accessible regions of DNA 290 In situ biochemical reactions for assaying specific enzymatic activities 291 Traditional stains 292 The concept of reporter molecules and genes 293 Bibliography 294
Chapter 20. Molecular Analysis of Chromosomal Landmarks
20.1 Heterochromatin and euchromatin 296 20.1.1 The nature of euchromatin 296 20.1.2 The highly condensed nature of heterochromatin 298 20.1.3 The DNA sequences of heterochromatin 299 20.1.4 Some chromosomal proteins are more prominent in heterochromatin 300 20.1.5 Heterochromatin contains genes at a lower frequency than euchromatin 301 20.1.6 Unusual biological properties of heterochromatin 302
20.2 The nucleolus and ribosomal-RNA genes 302 20.2.1 Identification of the genes involved in forming the nucleolus 302 20.2.2 The nature of ribosomal-RNA genes 303 20.2.3 Ribosomal-RNA synthesis 305 20.2.4 The structure of the nucleolus 306 20.2.5 Nucleolus-organizer-region suppression 307 20.2.6 Ribosomal-RNA gene loci and meiotic-chromosome pairing sites 307 20.2.7 Ribosomal-RNA gene loci and fragile sites 308 20.2.8 Amplification of rDNA gene sequences 308
20.3 The centromere 309 20.3.1 The kinetochore 309 20.3.2 DNA sequences at or near the centromere of mammalian chromosomes 309 20.3.3 DNA sequences from the centromeres of yeast chromosomes 310
20.4 The telomeres 311 20.4.1 The requirement of primers for DNA synthesis creates problems in the
replication of linear molecules 311 20.4.2 The DNA of telomeres in the ciliated protozoan Tetrahymena thermophila 311 20.4.3 Telomeres in other organisms 312 20.4.5 The attachment of telomeres to nuclear membranes and proteins 313
Bibliography 313
Chapter 21. Genetic and Molecular Mapping of Chromosomes
21.1 An overview of mapping in plants, animals, and fungi 318 21.1.1 Studies in animals 318
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285
296
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21.1.2 Studies in plants 319 21.1.3 Studies in yeast 319
21.2 Genetic markers 319 21.2.1 Morphological markers 319 21.2.2 Chromosomal landmarks 320 21.2.3 Disease resistance and susceptibility 320 21.2.4 Biochemical defects 320 21.2.5 Enzymes and other protein markers 321 21.2.6 DNA markers 322
21.3 Chromosome aberrations provide a first step for gene mapping 324 21.4 Data collection and linkage analysis 325
21.4.1 The principles underlying linkage analysis 325 21.4.2 The analysis of family and F2 progenies 326 21.4.3 The LOD score 327 21.4.4 Bayes' theorem 327 21.4.5 The influence of linkage phase on gene mapping 327 21.4.6 Mapping functions 328 21.4.7 Multipoint mapping 328 21.4.8 Interval mapping 328 21.4.9 Quantitative trait loci (QTL) 329
21.5 The practical aspects of gene mapping 330 21.5.1 Polymorphism analysis and selection of parents 330 21.5.2 Backcross and F2 progenies 331 21.5.3 Recombinant-inbred lines, single-seed descent, and doubled haploids 331 21.5.4 Recombinant disomic substitution lines 331 21.5.5 Genetic maps for forest trees 331 21.5.6 Somatic-cell hybrids 332 21.5.7 Extended human families 332 21.5.8 Homozygosity mapping 332
21.6 Construction of chromosome maps 332 21.6.1 Computer analysis to produce a linkage map 332 21.6.2 The concept of a consensus map 332
21.7 Correlation of linkage maps to physical maps 333 21.7.1 Procedures for placing the genetic map into the context of the physical description of a
chromosome 333 21.7.2 The relationship between cytogenetic and genetic maps 333 21.7.3 The length of genetic maps 333
21.8 Wider applications for genetic maps 334 21.8.1 Genetic counseling 334 21.8.2 Animal and plant breeding 335
Bibliography 336
PART vm CYTOPLASMIC DNA, GENETIC ENGINEERING, ORGANISMS OF IMPORTANCE
Chapter 22. Cytoplasmic DNA and Maternally Inherited Traits
22.1 The DNA in mitochondria 341 22.1.1 Vertebrate mtDNA 341 22.1.2 Invertebrate mtDNA 342 22.1.3 Fungal and plant mtDNAs 342
22.2 The DNA of chloroplasts and plastids 342 22.3 The analysis of cytoplasmic traits 343
22.3.1 Reciprocal interspecific hybrids 343 22.3.2 Nuclear substitutions and the genes involved in nucleo-cytoplasmic interactions 343 22.3.3 Nucleo-cytoplasmic interactions and genome evolution 345
22.4 Protoplast fusion and cytoplasmic manipulation 345 22.4.1 Cybrids versus somatic hybrids 346 22.4.2 Genetic incompatibility in somatic hybrids and cybrids 346
341
Contents xiii
22.5 Mitochondrial genes associated with cytoplasmic male sterility 347 22.5.1 RNA editing 347 22.5.2 New proteins in cytoplasmic-male-sterile plants 348
22.6 Mitochondrial-DNA mutations and human diseases 349 Bibliography 349
Chapter 23. Engineering the Genome 351
23.1 Genetic engineering 351 23.2 Agrobacterium transformation of plants 352
23.2.1 T-DNA 352 23.2.2 The Ti plasmid 353 23.2.3 Genes in the Agrobacterium protosome 355 23.2.4 Modifications of the Agrobacterium delivery system 355 23.2.5 Selectable and scorable markers 356
23.3 The production of chimeric genes 356 23.3.1 Regulator sequences 357 23.3.2 The SEKDEL modification 358 23.3.3 Tailoring gene DNA sequences 358 23.3.4 Binary gene constructs 358
23.4 Genetic transformation in dicotyledons 359 23.4.1 Introduction of herbicide resistance 359 23.4.2 New resistance to viruses 359 23.4.3 Introduction of insecticidal genes 361 23.4.4 Introduction of resistance to bacterial and fungal pathogens 361 23.4.5 Antisense genes 362 23.4.6 Plants as factories 362
23.5 Transformation in cereals and other crops 363 23.5.1 Electroporation 363 23.5.2 High-velocity microprojectile bombardment or biolistics 363 23.5.3 Other means of plant transformation 364
23.6 Transgenic animals 365 23.6.1 Transformation in animal-cell cultures 365 23.6.2 Mammalian transfection 365 23.6.3 Retroviral vectors 367 23.6.4 Microinjection in animals 367 23.6.5 Homologous recombination for gene introgression 368 23.6.6 Antisense oligonucleotides 369 23.6.7 Transfer of mice technology to larger animals and humans 369
23.7 Deletion of unnecessary genes 370 23.7.1 Removal of genes using CRFJLOX recombination 370 23.7.2 Removal of genes using maize AciDs transposable elements 370
23.8 Post-transformation control of transgenes and genetic silencing 370 23.9 Mendelian inheritance of transformed DNA 371
Bibliography 371
Chapter 24. Organisms of Importance to Genetics and Cytogenetics 376
24.1 Bacteriophage A (a bacterial virus) 376 24.1.1 Description 376 24.1.2 Genome size 376 24.1.3 Features of interest to genetics and molecular biology 376
24.2 Escherichia coli (coliform bacterium) 377 24.2.1 Description 377 24.2.2 Genome size 377 24.2.3 Features of interest to genetics and molecular biology 377
24.3 Saccharomyces cerevisiae (brewer's yeast) 378 24.3.1 Description 378 24.3.2 Genome size and karyotype 378 24.3.3 Features of interest to genetics and molecular biology 378
24.4 Arabidopsis thaliana (thale weed or wall cress) 379
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24.4.1 Description 379 24.4.2 Genome size and karyotype 379 24.4.3 Features of interest to genetics, cytogenetics, and molecular biology 379
24.5 Lycopersicon esculentum (tomato) 380 24.5.1 Description 380 24.5.2 Genome size and karyotype 380 24.5.3 Features of interest to genetics, cytogenetics, and molecular biology 380
24.6 Zea mays (maize) 380 24.6.1 Description 380 24.6.2 Genome size and karyotype 380 24.6.3 Features of interest to genetics, cytogenetics, and molecular biology 381
24.7 Triticum aestivum (bread wheat) 24.7.1 Description 381 24.7.2 Genome size and karyotype 381 24.7.3 Features of interest to genetics, cytogenetics, and molecular biology 381
24.8 Oryza sativa (rice) 382 24.8.1 Description 382 24.8.2 Genome size and karyotype 382 24.8.3 Features of interest to genetics, cytogenetics, and molecular biology 382
24.9 Tetrahymena thermophila (ciliated protozoan) 383 24.9.1 Description 383 24.9.2 Genome size and karyotype 384 24.9.3 Features of interest to genetics, cytogenetics, and molecular biology 384
24.10 Caenorhabditis elegans (nematode eelworm) 24.10.1 Description 384 24.10.2 Genome size and karyotype 385 24.10.3 Features of interest to genetics, developmental genetics, and molecular biology 385
24.11 Drosophilia melanogaster (fruit fly) 385 24.11.1 Description 385 24.11.2 Genome size and karyotype 386 24.11.3 Features of interest to genetics, cytogenetics, and molecular biology 386
24.12 Strongylocentrotus purpuratus (sea urchin) 386 24.12.1 Description 386 24.12.2 Genome size and karyotype 386 24.12.3 Features of interest to genetics, cytogenetics, and molecular biology 386
24.13 Xenopus laevis (clawed toad) 386 24.13.1 Description 386 24.13.2 Genome size and karyotype 387 24.13.3 Features of interest to genetics, cytogenetics, and molecular biology 387
24.14 Gallus domesticus (chicken) 387 24.14.1 Description 387 24.14.2 Genome size and karyotype 387 24.14.3 Features of interest to genetics, cytogenetics, and molecular biology 388
24.15 Mus musculus (common mouse) 388 24.15.1 Description 388 24.15.2 Genome size and karyotype 388 24.15.3 Features of interest to genetics, cytogenetics, and molecular biology 388
24.16 Homo sapiens (humans) 389 24.16.1 Description 389 24.16.2 Genome size and karyotype 389 24.16.3 Features of interest to genetics, cytogenetics, and molecular biology 389
Bibliography 390
Index 391
Preface
The close of the twentieth century is an exciting and stimulating period of discovery in the area of chromosome structure and function. In • 'Chromosome Biology" we target the many different disciplines that are being applied in the analysis of chromosomes in order to provide an admixture of cytogenetics, genetics and molecular biology for application to courses taught singly or cooperatively by cytogeneticists and molecular biologists. The book provides background reading for courses involving anyone discipline. Our intent is to convey to students an appreciation of plant and animal improvement programs, the importance of mapping the human genome, particularly in relation to diseases, and the advantages of modem technology to analyzing the development of different life-forms.
At the start of each Chapter the main concepts discussed are
Rudi Appels
listed so that the subject matter has a clear focus and, where possible, figures illustrate the key concepts throughout the book. Specific references are cited in the figure legends, or in the text, to justify the information provided. General books and reviews are listed at the end of each Chapter.
We are grateful to numerous colleagues for providing helpful critisisms and information; we particularly want to thank Drs G. Hart, D. Smyth, C. Gillies, N. Darvey and D. Hayman; external reviewers provided extensive comments on all or part of the manuscript and made major contributions towards the development of the book. Colleagues and reviewers cannot, however, be held responsible for the final content of the book, since this ultimately reflects the individual interests and biases of the authors.
Chief Research Scientist, CSIRO, Division of Plant Industry, Canberra, ACT, Australia Rosalind Morris Emeritus Professor, Department of Agronomy, University of Nebraska, Lincoln, NE, USA Bikram S. Gill Professor, Department of Plant Pathology, Kansas State University, Manhattan, KS, USA Cedric E. May Principal Research Scientist, NSW Agriculture, Wagga Wagga, NSW Australia W. John Raupp Associate Scientist, Department of Plant Pathology, Kansas State University, Manhattan, KS, USA
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