Sex chromosomes and sex determination in vertebrates:
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| Main Author: | |
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| Format: | Book |
| Language: | English |
| Published: |
Boca Raton u.a.
CRC Press
1994
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| Subjects: | |
| Online Access: | Inhaltsverzeichnis |
| Physical Description: | 308 S. Ill., graph. Darst. |
| ISBN: | 0849345715 |
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| 100 | 1 | |a Solari, Alberto J. |e Verfasser |4 aut | |
| 245 | 1 | 0 | |a Sex chromosomes and sex determination in vertebrates |c Alberto J. Solari |
| 264 | 1 | |a Boca Raton u.a. |b CRC Press |c 1994 | |
| 300 | |a 308 S. |b Ill., graph. Darst. | ||
| 336 | |b txt |2 rdacontent | ||
| 337 | |b n |2 rdamedia | ||
| 338 | |b nc |2 rdacarrier | ||
| 650 | 4 | |a Chromosomes sexuels | |
| 650 | 4 | |a Sexe - Détermination génétique | |
| 650 | 4 | |a Vertébrés - Génétique | |
| 650 | 4 | |a Sex Chromosomes | |
| 650 | 4 | |a Sex Determination (Genetics) | |
| 650 | 4 | |a Sex chromosomes | |
| 650 | 4 | |a Sex determination, Genetic | |
| 650 | 4 | |a Vertebrates |x Genetics | |
| 650 | 4 | |a Vertebrates |x genetics | |
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Record in the Search Index
| _version_ | 1804124134402162688 |
|---|---|
| adam_text | SEX CHROMOSOMES
and
SEX DETERMINATION
in
VERTEBRATES
Alberto J Solari
CRC Press
Boca Raton Ann Arbor London Tokyo
TABLE OF CONTENTS
Chapter 1
INTRODUCTION: THE ROLE OF SEX CHROMOSOMES 1
I Peculiar Behavior and Morphology of Sex
Chromosomes 1
II The Chromosomal Basis of Sex Determination 2
III The Fundamental Components of Chromosomes 3
IV The Centromere 4
V Telomeres 6
VI Differences Between Autosomes and Sex
Chromosomes 6
VII Sex Chromosome Differentiation and the Regulation
of Recombination 7
VIII Meiotic Chromosomes and Synaptonemal Complexes 7
IX General Properties of Sex Chromosomes and
Relationships with Sex Determination 8
Chapter 2
SEX CHROMOSOMES AND SEX DETERMINATION IN
AMPHIBIA 9
I Sex Chromosomes 9
A Introduction 9
B Occurrence of Sex Chromosomes Among
Amphibians 10
C Amphibian Sex-Chromosome Types 10
D Description of Sex-Chromosome Types 11
1 Homomorphic Sex Chromosomes 11
2 Heteromorphisms for Banding Patterns 12
a Heteromorphism in the Replication
Pattern 12
b C-Banding Heteromorphisms in
Isomorphic Gonosomes 13
3 Heteromorphisms Originated in a
Pericentric Inversion 14
4 Heteromorphisms in Size 15
5 Supernumerary Sex Chromosomes 17
6 Intraspecific Variable Sex Chromosomes 18
7 Multiple Sex Chromosomes 18
II Gonadal Inversion in Amphibia 19
A Introduction 19
B Temperature-Induced Gonadal Inversion 20
C Lampbrush Chromosomes and Sex Reversal in
the Urodela 21
D Hormone-Induced Gonadal Inversion 22
E Sex Reversal by Grafting or Chirurgical
Resection of the Gonads 23
III Sex-Linked Genes and Gene Dosage Effects in
Amphibia 23
IV Genetic Design of Sex Determination in Amphibia 24
Chapter 3
SEX CHROMOSOMES AND SEX DETERMINATION IN
REPTILES 27
I Introduction and General Perspective 27
n Sex Chromosomes in Reptiles 27
A Chelonia 27
B Sauria 28
1 Simple Male Heteromorphism (XY) 29
2 Simple Female Heteromorphism (ZW) 30
3 Multiple Sex Chromosomes 30
4 General Comment 30
C Ophidia 31
1 General • 31
2 Differentiation of the W Chromosome 33
3 Multiple Sex Chromosomes in Snakes 33
D Meiotic Behavior of Sex Chromosomes in
Reptiles 34
E Studies on Synaptonemal Complexes 35
III Environmental Sex Determination (ESD) in Reptiles 35
A Temperature Effects in Laboratory Conditions 36
B Temperature Effects in Natural Environments 36
IV Experimental Sex Reversal By Hormones in Reptiles 37
V A Unified Hypothesis on Temperature and Hormonal
Effects 38
VI DNA Sequences Specific of Reptilian Gonosomes The
Bkm DNA 39
VII The Design of Sex Determination in Reptiles 40
Chapter 4
SEX CHROMOSOMES, SEX DETERMINATION, AND
MEIOSIS IN BIRDS 43
I Introduction 43
II General Comments on Avian Karyotypes 43
III Mitotic Sex Chromosomes in Birds 44
A General 44
B Sex Chromosomes in the Ratitae 45
C Sex Chromosomes in Carinate Birds 46
D The Z and W Chromosomes of the Chicken
(Gallus domesticus) 47
IV Gonosome-Autosome Translocations in the Chicken 49
V The Meiotic Behavior of Gonosomes in the Chicken 51
A Meiosis in the Male (Cock) 51
B Meiosis in the Female (Fowl) 52
C Microspreading of Oocytes Partial Synapsis
Between the W and Z Chromosomes 52
D Recombination Between the Z and W
Chromosomes 55
E Consequences of Regular Recombination
Between the Z and W Chromosomes 56
F The Presumed Homology Between the Z and W
Chromosomes: Results of the Study of Z-
Autosome Translocations 58
G The ZW Pair After Pachytene: Lampbrush
Organization 59
VI Gene Contents of the Z and W Chromosomes in the
Chicken 60
A Linkage Map in the Chicken 60
B Specific DNA of the W Chromosome in the
Chicken 60
C The Absence of Dosage Compensation of Genes
in the Avian Gonosomes 61
VII Sex Determination in Birds 62
A Aneuploidy and Gonosomal Behavior in Birds 62
B Gonadal Asymmetry in Birds 63
C Gonadal Morphogenesis in the Chicken 64
D Spontaneous Sex Reversal and Experimental
Gonadal Inversion in Birds 66
E Gonadectomy and Gonadal Inversion 66
F Gonadal Inversion By Estradiol Administration
to Male Bird Embryos 68
G Gonadal Inversion in ZZW Triploid Chickens 69
VIII Design of Sex Determination in Birds 71
Chapter 5
SEX CHROMOSOMES AND SEX DETERMINATION IN
MAMMALS: THE MONOTREMES 75
I Sex Chromosomes 75
A Introduction 75
B Sex Chromosomes in Monotremes 75
C Gonosomes of the Platypus 76
D Male Meiosis in the Platypus 77
E Sex Chromosomes in the Echidna 79
n Sex Determination in Monotremes 79
Chapter 6
SEX CHROMOSOMES IN METATHERIANS
(MARSUPIALS) 81
I Introduction 81
n General Remarks on Marsupial Karyotypes 81
m Sex Chromosomes of Marsupials 82
A Morphology of Sex Chromosomes in Marsupials 82
1 Simple XY-XX Systems 82
a Group With a Small-Sized
X-Chromosome 82
b Group With Mid- and Large-Sized
X-Chromosomes 84
2 Multiple Gonosome Systems 84
B Gonosome Elimination From Somatic Tissues 85
IV Meiosis in Male Marsupials: A Basic Difference With
Eutherian Mammals , 86
A X-Y Behavior in Monodelphis dimidiata 86
B Meiosis in Other American Marsupials 88
C Male Meiosis in Australian Marsupials 89
D Meiotic Behavior of Complex Gonosome
Systems 90
V Sex Determination in Marsupials 90
A Gene Contents of the Marsupial X and Y
Chromosomes 90
B X-Chromosome Inactivation in Marsupials 91
C Intersexual States in Marsupials 92
D Design of Sex Determination in Marsupials 93
Chapter 7
SEX CHROMOSOMES IN EUTHERIAN MAMMALS: GENE
DOSAGE COMPENSATION 95
I Introduction 95
II Gene Dosage Compensation Between Males and
Females in Eutherian Mammals 95
m Cytological Aspects of the X-Chromosome Inactivation 96
A Sex (X) Chromatin (Barr Body) 96
B Origin of the X-Chromatin Body and Regularity
of Its Presence 99
C Late Replication 101
IV Molecular Differences Between Active and Inactive
X-Chromosomes 101
V Lyon s Hypothesis and the Process of
X-Chromosome Inactivation 102
A Sex-Linked Variegation and Sex-Linked Cell
Mosaicism 102
B Time of Inactivation and Appearance of X
Chromatin 103
C Nonrandom Inactivation in Extraembryonic
Tissues 104
D X-Chromosome Reactivation and Differentiation
in the Germ Cells 105
E The Inactivation Center and the Spreading of
Inactivation 105
F The XIST Gene is Located at the XIC Region
and is Transcribed 106
VI Dosage Compensation By X-Chromosome Inactivation
in Eutherian Mammals: General Comments 107
Chapter 8
THE XY BODY AND PARTIAL SYNAPSIS
IN EUTHERIAN MAMMALS 109
I The XY Body 109
A Introduction 109
B Nomenclature of the XY Body 109
C Determination of the Temporal Sequence of
Events During Meiosis 110
D General Morphology of the XY Body 112
1 Variations 112
2 Occurrence 114
E The XY Body in Species With Composite
Sex Chromosomes 114
F Other Exceptions 116
G Relationships with Nucleoli 116
1 Nucleolar Association With the XY Body
in the Mouse 117
2 Nucleolus-Gonosome Relationships in
Man 120
II Partial Synapsis 121
A The Development of Partial Synapsis Between
the X and Y Chromosomes 121
B The Sequence of X-Y Synapsis in the Mouse
(Mus musculus) 121
C Variations in the Synaptic Pattern of the X and
Y Chromosomes 122
1 Axial Differentiations 122
2 Variations Related to Heterochromatin 124
3 Variations in Compound
Sex-Chromosomes 124
D Exceptional XY Bodies Lacking Partial Synapsis
III Functional Aspects of the XY Body 129
A Transcriptional Inactivity of the XY Body 129
B Spreading of Inactivation Towards Autosomal
Chromatin in Searle s Translocation 130
C DNase Sensitivity in the XY Body 131
D Theoretical Aspects of Gonosomal Inactivation in
the XY Body 132
E Time and Mechanism of XY Body Inactivation 134
F Exceptional Gonosomal Bodies 135
Chapter 9
RECOMBINATION IN THE XY PAIR OF EUTHERIAN
MAMMALS 137
I Introduction 137
II Chiasmata and the XY Pair 137
A The Behavior of the XY Pair at Metaphase:
Terminal Vs Chiasmatic Association 137
B Species With Chiasmatic XY Pairs and
Gonosome-Autosome Multivalents 138
G Specificity of Arm Association in XY Pairs 140
D Chromatin Labelling With BrdU and XY
Recombination 140
III Recombination Nodules and the XY Pair 141
A Fine Structure and Recombination Nodules
(RNs) in the XY Pair 141
B Fine Structure of Chiasmata in the XY Pair 142
IV Genetic Recombination in the XY Pair 145
A Regular Meiotic Recombination in the XYSxr
Mouse 145
B Restatement of the Hypothesis of Partial Sex-
Linkage: the Recombining Region 146
C Telomeric Sequences 147
D The Paratelomeric, Pseudoautosomal DXYS14
Sequence in Human Gonosomes 148
E X-Y Recombination at the DXYS14 Sequence 149
F Additional Recombining, Noncoding Sequences
in the Human Gonosomes 150
G Linkage and the Construction of Genetic Maps
of the Human Recombining Region 150
H Genes Located in the Human Recombining
(Pseudoautosomal) Region 152
I Physical Mapping of the Recombining Region in
the Human Species 153
V The Boundary Between the Recombining and the
Differential Regions in the Human XY Pair 153
A The Boundary of the Recombining Region 153
B Comparative Sequences of the Boundary Region
Among the Great Apes 155
C Recombining Regions in Other Species 156
VI General Comments on Recombination in the XY Pair 157
A Reappraisal of Homology in Pairing of
Meiotic Gonosomes 157
B The Problen of Exceptional, Asynaptic
Gonosomes 158
VII Early Dissociation of the X and Y Gonosomes at
Metaphase-I 159
Chapter 10
SEX CHROMOSOMES AND SEX DETERMINATION IN
THE MOUSE 161
I Introduction 161
II The Normal Sex Chromosomes 161
A The X Chromosome of Mus musculus 161
B The Y Chromosome of Mus musculus 162
C Gene Contents in the X Chromosome 163
D Gene Contents in the Y Chromosome 163
III Structural Variants: The Sxr Region 165
A The Sex-Reversed (Sxr) Condition in the Mouse 165
B Origin of the Sxr Chromosomal Region 165
C Transfer of the Sxr Region to the X
Chromosome 167
D Cytology of Sxr 168
E Sxr Region, Histocompatibility Antigen-Factor
(Hya) and the Tdy Gene 168
F Gene Contents and Variants of the Sxr Region 169
IV Functional Aspects of the Sxr Condition 169
A Phenotype of XYSxr Males 169
B XX amp;crMice 169
C XSxrO Males 170
D XSxr O Males 170
E Meiotic Behavior of Sxr and Its Variants 170
F Sxr Region and Gametogenesis:
XSxrO Diploid Spermatids 174
G Sxr Region and X-Chromosome Activity
During Development 174
H Sex-Reversal, Sex Reversed (T16/HXSxr) Mice
and X-Chromosome Activity 175
I Sxr Region and Formation of the XY Body 176
V Other Gonosomal Variants 176
A The Y* (Rearranged) Chromosome 176
B The XXY Chromosome Constitution and Partial
Y-Disomy in the Mouse 177
Chapter 11
DEVELOPMENTAL AND GENETIC ASPECTS OF SEX
DETERMINATION IN THE MOUSE 179
I Gonadal Morphogenesis in the Mouse 179
A Origin of Germ Cells 179
B Migration of Primordial Germ Cells (PGCs) and
Colonization of the Gonad 179
C The Sexually Undifferentiated Gonad 180
D Sexual Differentiation of the Male Gonad 182
E Sertoli Cells and Activity of Sex-Determining
Genes 183
II Sex in Chimeric Mice 185
A The Death of XX Germ Cells in a Testis 185
B Sex in Female XX *» XY Chimeras and the
Exceptional XY Oocytes 187
C Mixed Cultures of Somatic and Germ Cells 188
III Developmental Potentialities of Germ Cells 189
A Behavior of Ectopic Germ Cells of Either XX Or
XY Constitution 189
B Reappraisal of the Developmental Potentiality of
XX and XY Germ Cells 190
C Gonadal Transplantation 191
IV Genetics of Sex Determination in the Mouse 192
A Autosomal Genes Involved in Sex
Determination 192
B The T-Associated Sex Reversal (Tas) Gene in
Chromosome 17 193
C The Major Sex-Determining Gene: Tdy (Sry) 194
D Induction of Sex Reversal of XX Mice
Transgenic for Sry 195
V General Comments on Sex Determination in the Mouse 197
Chapter 12
UNUSUAL GONOSOMAL SYSTEMS: LEMMINGS AND
AKODON RODENTS 199
I Introduction 199
II Sex Chromosomes and Sex Determination in the Wood
Lemming (Myopus Schisticolor) 199
A Morphology of the Sex Chromosomes 199
B The Two X-Chromosome Types (X and X*) of
the Wood Lemming 200
C Sex Ratio in the Wood Lemming 200
D Sex Ratio and Karyotype in the Offspring of
Different Matings 201
E Meiosis in the Wood Lemming 202
F Occurrence of Gonosomal Trisomics and
Monosomics in the Wood Lemming 203
G Lyonization of X Chromosomes in the X*XY
Wood Lemming 204
H Comments on Sex Determination in the Wood
Lemming 205
III Sex Chromosomes in Lemmings From the Genus
Dicrostonyx 206
IV Sex Chromosomes and Sex Determination in Field Mice
of the Genus Akodon 206
A Sex Chromosomes 206
B Sex Ratio in A azarae 208
C Genotypes in the Offspring of Xx Females 209
D Litter Size and Other Reproductive Parameters in
A azarae 209
E Meiosis in Males of A azarae 209
F Meiosis in XX Females of A azarae 210
G Meiosis in the Variant Xx Females of A azarae 210
H Gonosomal DNA in A azarae 211
I Hypotheses on the Mechanism of Sex
Determination in A azarae 212
J Sex Chromosomes in Other Species of the Genus
Akodon 213
V Other Gonosomal Variants 214
A Sex Chromosomes in Ellobius lutescens 214
B Sex Chromosomes in Microtus oregoni 217
Chapter 13
HUMAN SEX CHROMOSOMES 219
I Introduction 219
II Morphology of the Sex Chromosomes 219
A The Human X Chromosome 219
B The Human Y Chromosome 220
III Gene Contents of the Gonosomes 221
A Genetic Contents of the Human X Chromosome 221
B Deletion Map of the Human Y Chromosome 221
C Gene Contents of the Human Y Chromosome 221
D Physical Maps 223
IV The SRY Gene 224
A Location of the SRY Gene in the Human Y
Chromosome 224
B Some Characteristics of SRY 225
C Function of Y-Linked Genes in Human Sex
Determination 225
V Abnormal Sex Determination 226
A XXMales 226
B Cytogenetic and Molecular Studies on
XX Males 227
C Origin of XX, Y( + ) Males 228
D Origin of XX, Y( - ) Males * 229
E XY Females (Yp-Deleted Variety) 229
F Other Types of XY Females 229
G XY Females With a Mutant SRY Gene 230
H True Hermaphrodites 230
Chapter 14
SEX CHROMOSOMES AND SEX DETERMINATION IN
FISHES 233
I Introduction 233
II Sex Chromosomes of Fishes 233
A Karyology of Fishes 233
B XY-XX Gonosomal Systems in Teleosts 234
C ZZ-ZW Gonosomal Systems in Teleosts 235
D Multiple Gonosome Systems in Teleosts 237
E Other Gonosomal Types 238
F Sex-Specific DNA Sequences 238
III Sex Determination in Fishes 239
A Sex-Linked Pigment Genes and the
Demonstration of the Heterogametic Sex 239
B Matings of Sex-Reversed Males With Sex-
Reversed Females: the Control of Crossing-Over 239
C The Polyfactorial Hypothesis of Sex
Determination in Gonochoristic Teleosts 241
D Hermaphroditism Among Fishes Sequential and
Simultaneous Hermaphrodites 242
E The Genetic Basis of Gonadal Morphogenesis 243
IV Gonadal Morphogenesis 245
A Gonadal Morphogenesis in Gonochorist Species 245
B Sexual Bipotentiality of Germ Cells in
Teleostean Fishes 246
C Experimental Sex Reversal in Teleostean Fishes 246
Chapter 15
SEX CHROMOSOMES AND SPECIALIZED
RECOMBINATION PATTERNS 249
I Introduction 249
II Differences in Gene Recombination Between the Sexes 249
A General 249
B Gonosome-Specific Recombination Differences 250
III Effects of Restricted Recombination 252
A General 252
B Muller s Ratchet 253
C Genetic Hitchhiking and the Eyolution of
Heteromorphic Gonosomes 255
D Operation of Mechanisms For Gonosome
Degeneration 256
IV Origins of Recombination Restriction 257
A Promotion of Recombination Restriction 258
1 The Case of Sexually Antagonistic Genes
Facilitating Recombination Restriction 258
2 Multiplicity of Sex-Determining Loci
Favoring Recombination Restriction 259
B Establishment of Recombination Modifiers 260
V Behavior of Gonosomes With Recombination Restriction
A Differences Between Partial Synapsis and
Recombination Restriction 260
1 Synapsis Exceeds the Homologous and
the Recombining Segments in
Mammalian XY Pairs 261
2 Synapsis Exceeds the Homologous and
the Recombining Regions in Avian ZW
Pairs 262
B Regional Specificity of Recombination
Restriction 263
C Disruption of Recombination Restriction in
Different Genetic Backgrounds 264
VI Gonosomal Condensation During Meiosis in Mammals 265
Appendix
THE DISCOVERY OF PARTIAL SYNAPSIS 267
References 271
Index 303
|
| any_adam_object | 1 |
| author | Solari, Alberto J. |
| author_facet | Solari, Alberto J. |
| author_role | aut |
| author_sort | Solari, Alberto J. |
| author_variant | a j s aj ajs |
| building | Verbundindex |
| bvnumber | BV009786375 |
| callnumber-first | Q - Science |
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| classification_rvk | WG 3540 WG 6710 |
| ctrlnum | (OCoLC)27934923 (DE-599)BVBBV009786375 |
| dewey-full | 596/.01/5 |
| dewey-hundreds | 500 - Natural sciences and mathematics |
| dewey-ones | 596 - Chordata |
| dewey-raw | 596/.01/5 |
| dewey-search | 596/.01/5 |
| dewey-sort | 3596 11 15 |
| dewey-tens | 590 - Animals |
| discipline | Biologie |
| format | Book |
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| id | DE-604.BV009786375 |
| illustrated | Illustrated |
| indexdate | 2024-07-09T17:40:50Z |
| institution | BVB |
| isbn | 0849345715 |
| language | English |
| oai_aleph_id | oai:aleph.bib-bvb.de:BVB01-006474220 |
| oclc_num | 27934923 |
| open_access_boolean | |
| owner | DE-703 DE-20 DE-11 |
| owner_facet | DE-703 DE-20 DE-11 |
| physical | 308 S. Ill., graph. Darst. |
| publishDate | 1994 |
| publishDateSearch | 1994 |
| publishDateSort | 1994 |
| publisher | CRC Press |
| record_format | marc |
| spelling | Solari, Alberto J. Verfasser aut Sex chromosomes and sex determination in vertebrates Alberto J. Solari Boca Raton u.a. CRC Press 1994 308 S. Ill., graph. Darst. txt rdacontent n rdamedia nc rdacarrier Chromosomes sexuels Sexe - Détermination génétique Vertébrés - Génétique Sex Chromosomes Sex Determination (Genetics) Sex chromosomes Sex determination, Genetic Vertebrates Genetics Vertebrates genetics Wirbeltiere (DE-588)4066376-0 gnd rswk-swf Geschlechtschromosom (DE-588)4157012-1 gnd rswk-swf Geschlechtschromosom (DE-588)4157012-1 s Wirbeltiere (DE-588)4066376-0 s DE-604 HEBIS Datenaustausch application/pdf http://bvbr.bib-bvb.de:8991/F?func=service&doc_library=BVB01&local_base=BVB01&doc_number=006474220&sequence=000002&line_number=0001&func_code=DB_RECORDS&service_type=MEDIA Inhaltsverzeichnis |
| spellingShingle | Solari, Alberto J. Sex chromosomes and sex determination in vertebrates Chromosomes sexuels Sexe - Détermination génétique Vertébrés - Génétique Sex Chromosomes Sex Determination (Genetics) Sex chromosomes Sex determination, Genetic Vertebrates Genetics Vertebrates genetics Wirbeltiere (DE-588)4066376-0 gnd Geschlechtschromosom (DE-588)4157012-1 gnd |
| subject_GND | (DE-588)4066376-0 (DE-588)4157012-1 |
| title | Sex chromosomes and sex determination in vertebrates |
| title_auth | Sex chromosomes and sex determination in vertebrates |
| title_exact_search | Sex chromosomes and sex determination in vertebrates |
| title_full | Sex chromosomes and sex determination in vertebrates Alberto J. Solari |
| title_fullStr | Sex chromosomes and sex determination in vertebrates Alberto J. Solari |
| title_full_unstemmed | Sex chromosomes and sex determination in vertebrates Alberto J. Solari |
| title_short | Sex chromosomes and sex determination in vertebrates |
| title_sort | sex chromosomes and sex determination in vertebrates |
| topic | Chromosomes sexuels Sexe - Détermination génétique Vertébrés - Génétique Sex Chromosomes Sex Determination (Genetics) Sex chromosomes Sex determination, Genetic Vertebrates Genetics Vertebrates genetics Wirbeltiere (DE-588)4066376-0 gnd Geschlechtschromosom (DE-588)4157012-1 gnd |
| topic_facet | Chromosomes sexuels Sexe - Détermination génétique Vertébrés - Génétique Sex Chromosomes Sex Determination (Genetics) Sex chromosomes Sex determination, Genetic Vertebrates Genetics Vertebrates genetics Wirbeltiere Geschlechtschromosom |
| url | http://bvbr.bib-bvb.de:8991/F?func=service&doc_library=BVB01&local_base=BVB01&doc_number=006474220&sequence=000002&line_number=0001&func_code=DB_RECORDS&service_type=MEDIA |
| work_keys_str_mv | AT solarialbertoj sexchromosomesandsexdeterminationinvertebrates |