This book also looks at the molecules and cells that make bones and cartilages and how they differ in various parts of the body and across species. It answers such questions as Is bone always bone? Do bones that develop indirectly by replacing other tissues, such as marrow, tendons or ligaments, differ from one another? Is fish bone the same as human bone? Can sharks even make bone? and many more.
* Complete coverage of every aspect of bone and cartilage
* Full of interesting and unusual facts
* The only book available that integrates development and evolution of the skeleton
* Treats all levels from molecular to clinical, embryos to evolution
* Written in a lively, accessible style
* Extensively illustrated and referenced
* Integrates analysis of differentiation, growth and patterning
* Covers all the vertebrates as well as invertebrate cartilages
* Identifies the stem cells in embryos and adults that can make skeletal tissues
Bones and Cartilage provides the most in-depth review ever assembled on the topic. It examines the function, development and evolution of bone and cartilage as tissues, organs and skeletal systems. It describes how bone and cartilage is developed in embryos and are maintained in adults, how bone reappears when we break a leg, or even regenerates when a newt grows a new limb, or a lizard a tail. This book also looks at the molecules and cells that make bones and cartilages and how they differ in various parts of the body and across species. It answers such questions as "e;Is bone always bone?? "e;Do bones that develop indirectly by replacing other tissues, such as marrow, tendons or ligaments, differ from one another?? "e;Is fish bone the same as human bone?? "e;Can sharks even make bone?? and many more.* Complete coverage of every aspect of bone and cartilage* Full of interesting and unusual facts* The only book available that integrates development and evolution of the skeleton* Treats all levels from molecular to clinical, embryos to evolution* Written in a lively, accessible style* Extensively illustrated and referenced* Integrates analysis of differentiation, growth and patterning* Covers all the vertebrates as well as invertebrate cartilages* Identifies the stem cells in embryos and adults that can make skeletal tissues
Bones and Cartilage: Developmental and Evolutionary Skeletal Biology 2
Epigraph 3
Contents 6
Preface 20
Abbreviations 24
Part I Skeletal Tissues 30
Chapter 1 Types of Skeletal Tissues 32
BONE 33
CARTILAGE 34
DENTINE 34
ENAMEL 36
INTERMEDIATE TISSUES 37
Cementum 37
Enameloid 39
Chondroid and chondroid bone 40
BONE OR CARTILAGE 40
NOTES 41
Chapter 2 Bone 42
DISCOVERY OF THE BASIC STRUCTURE OF BONE 42
CELLULAR BONE 44
OSTEOCYTES 45
INTRAMEMBRANOUS VERSUS ENDOCHONDRAL BONE 46
Embryonic origins 46
Other modes 46
Metabolic differences 47
Morphogenetic differences 47
OSTEONES 48
GROWTH 50
REGIONAL REMODELING 50
AGEING 51
Osteones over time 52
ACELLULAR BONE 53
Caisson disease and abnormal acellular bone in mammals 53
Acellular bone in teleost fishes 53
Development 54
Resorption 54
Repair of fractures 55
Ca++ regulation 55
Aspidine 59
BONE IN CARTILAGINOUS FISHES (SHARKS AND RAYS) 59
NOTES 59
Chapter 3 Cartilage 62
TYPES 62
CHONDRONES 63
CARTILAGE GROWTH 64
CARTILAGE CANALS 65
SECONDARY CENTRES OF OSSIFICATION 65
ELASTIC CARTILAGE 68
Elastic fibres 68
The cells 68
Elastic cartilage intermediates 70
SHARK CARTILAGE 70
Development and mineralization 70
Growth 71
Inhibition of vascular invasion 71
LAMPREYS 71
Mucocartilage 72
Lamprin 72
Mineralization 73
HAGFISH 74
NOTES 74
Part II Natural Experiments 78
Chapter 4 Invertebrate Cartilages 80
CHONDROID, CARTILAGE OR NEITHER 80
ODONTOPHORE CARTILAGE IN THE CHANNELED WHELK, BUSYCON CANALICULATUM 81
BRANCHIAL (GILL BOOK) CARTILAGE IN THE HORSESHOE CRAB, LIMULUS POLYPHEMUS 81
CRANIAL CARTILAGES IN SQUID, CUTTLEFISH AND OCTOPUSES 82
Composition of the extracellular matrix 82
Glycosaminoglycans (GAGs) 82
Collagens 82
TENTACULAR CARTILAGE IN POLYCHAETE ANNELIDS 84
LOPHOPHORE CARTILAGE IN AN ARTICULATE BRACHIOPOD, TEREBRATALIA TRANSVERSA 85
MINERALIZATION OF INVERTEBRATE CARTILAGES 85
CARTILAGE ORIGINS 85
NOTES 91
Chapter 5 Intermediate Tissues 93
CHONDROID AND CHONDROID BONE 95
MODULATION AND INTERMEDIATE TISSUES 95
CARTILAGE FROM FIBROUS TISSUE AND METAPLASIA 97
METAPLASIA OF EPITHELIAL CELLS TO CHONDROBLASTS OR OSTEOBLASTS 97
Chondroid 98
Teleosts 98
Mammals 99
CHONDROID BONE 100
Teleosts 100
Mammals 100
Chondroid bone and pharyngeal jaws 101
TISSUES INTERMEDIATE BETWEEN BONE AND DENTINE 103
Dentine 104
Cementum 105
ENAMELOID: A TISSUE INTERMEDIATE BETWEEN DENTINE AND ENAMEL 106
NOTES 110
Chapter 6 An Evolutionary Perspective 112
FOSSILIZED SKELETAL TISSUES 112
ALL FOUR SKELETAL TISSUES ARE ANCIENT 113
EVOLUTIONARY EXPERIMENTATION 115
Intermediate tissues in fossil agnatha 115
DINOSAUR BONE 116
DEVELOPING FOSSILS 117
PROBLEMATICA 117
PALAEOPATHOLOGY 118
CONODONTS 119
NOTES 120
Part III Unusual Modes of Skeletogenesis 122
Chapter 7 Horns and Ossicones 124
HORNS 124
DISTRIBUTION OF HORNS AS ORGANS 125
Bovidae 125
Rhinos 127
Titanotheres 128
Pronghorn antelopes 128
Giraffes 129
HORN AS A TISSUE 130
DEVELOPMENT AND GROWTH OF HORNS 130
NOTES 131
Chapter 8 Antlers 132
ANTLERS 132
Size and absence 132
INITIATION OF ANTLER FORMATION 133
Pedicle formation 133
The antler bud and dermal–epidermal interactions 134
HORMONAL CONTROL OF PEDICLE DEVELOPMENT AND GROWTH 135
ANTLER REGENERATION 135
The shedding cycle 135
HISTOGENESIS OF ANTLERS 136
White-tailed deer, American elk, European fallow and roe deer 137
Rocky Mountain mule deer 139
Sika deer 139
HORMONES, PHOTOPERIOD AND ANTLER GROWTH 139
Sika deer Photoperiod and testosterone 139
Parathyroid hormone and calcitonin 142
NOTES 142
Chapter 9 Tendons and Sesamoids 144
TENDONS AND SKELETOGENESIS 144
Fibrocartilage in tendons 145
Rodent Achilles tendons 146
Ossification of avian tendons 146
Formation and composition of tendon fibrocartilages 146
Condensation 146
Scleraxis 147
Composition 148
SESAMOIDS 148
Amphibians 149
Reptiles 150
Birds 150
Teleosts 151
NOTES 151
Part IV Stem Cells 154
Chapter 10 Embryonic Stem Cells 156
STEM CELLS 156
SET-ASIDE CELLS 158
STEM CELLS FOR PERIOSTEAL OSTEOGENESIS IN LONG BONES 160
MODULATION OF SYNTHETIC ACTIVITY AND DIFFERENTIATIVE PATHWAYS OF CELL POPULATIONS 162
Fibroblast–chondroblast modulation 162
Modulation of glycosaminoglycan synthesis 162
MODULATION OF SYNTHETIC ACTIVITY AND DIFFERENTIATIVE PATHWAYS IN SINGLE CELLS 163
Degradative activity 163
NOTES 165
Chapter 11 Stem Cells in Adults 167
FIBROBLAST COLONY-FORMING CELLS 167
OSTEOGENIC PRECURSOR CELLS 168
Clonal analysis 169
Lineages of cells 169
Dexamethasone 169
EPITHELIAL INDUCTION OF ECTOPIC BONE 170
Epithelial cell lines 171
NOTES 173
Part V Skeletogenic Cells 176
Chapter 12 Osteo- and Chondroprogenitor Cells 178
IDENTIFYING OSTEO- AND CHONDROPROGENITOR CELLS 179
Execrable terminology 179
Features 179
Cell cycle dynamics 179
BIPOTENTIAL PROGENITOR CELLS FOR OSTEOGENESIS AND CHONDROGENESIS 180
Bipotential cell populations or bipotential cells? 180
Uncovering bipotentiality 180
Discovering bipotentiality 181
Biochemical and metabolic markers 181
Collagen types 182
The tumor suppressor gene p53 182
CONDYLAR CARTILAGE ON THE CONDYLAR PROCESS OF THE MAMMALIAN DENTARY 184
Histodifferentiation and scurvy 184
One or two cell populations 185
Evidence against bipotentiality 185
Evidence supporting bipotentiality 186
All or some? 188
SECONDARY CARTILAGE ON AVIAN MEMBRANE BONES 188
NOTES 193
Chapter 13 Dedifferentiation Provides Progenitor Cells for Jaws and Long Bones 195
CONDYLAR CARTILAGE OF THE MAMMALIAN TEMPOROMANDIBULAR JOINT 195
The temporomandibular joint 195
Hypertrophic chondrocytes survive 195
Hypertrophic chondrocytes transform to osteoprogenitor cells 196
MECKEL’S CARTILAGE 197
Mammalian Meckel’s 197
Prx-1, Prx-2 202
Alx-3 205
Ptx-1 205
DEDIFFERENTIATION DURING ENDOCHONDRAL BONE FORMATION 205
Rodent ribs 206
Mice 206
Rats 206
Appendicular long bones 207
Enzyme activity 207
Evidence from 3 H-thymidine-labeling and other approaches 208
Murine interpubic joints 210
NOTES 210
Chapter 14 Dedifferentiation and Urodele Amphibian Limb Regeneration 212
DEDIFFERENTIATION 212
Morphological dedifferentiation 213
Functional dedifferentiation 213
Hyaluronan 213
BLASTEMA FORMATION 215
Aneurogenic limbs 216
More than one cell fate 216
MYOBLAST AND CHONDROBLAST FATES 217
FACTORS CONTROLLING DEDIFFERENTIATION 218
Innervation 218
Aneurogenic limbs 218
Proliferation 218
Not the stump 219
Electrical signals? 220
Hox genes 220
FgfR-1 and FgfR-2 220
Radical fringe 220
WHY CAN’T FROGS REGENERATE? 221
Augmenting regeneration 223
FINGERTIPS OF MICE, MONKEYS AND MEN 224
Comparison with urodele limb regeneration 224
NOTES 225
Chapter 15 Cells to Make and Cells to Break 226
CLASTS AND BLASTS 226
RESORPTION 226
COUPLING BONE RESORPTION TO BONE FORMATION 227
COUPLING OSTEOBLASTS AND OSTEOCLASTS 227
SOME MOLECULAR PLAYERS 229
WHEN COUPLING GOES AWRY 230
TRAP-STAINING FOR OSTEOCLASTS 231
Mammalian osteoclasts 231
Teleost osteoclasts 232
NITRIC OXIDE – IT’S A GAS 232
PROGENITOR CELLS FOR OSTEOBLASTS AND OSTEOCLASTS 232
Japanese quail–domestic fowl chimaeras 234
OSTEOPETROSIS AND OSTEOCLAST ORIGINS 234
OSTEOCLAST–PHAGOCYTE–MACROPHAGE OR OSTEOCLAST–MONOCYTE LINEAGES? 237
Phagocyte/macrophage origin 237
Interleukins 238
IL-1 238
Osteogenesis 238
Chondrogenesis 238
IL-6 238
IL-10 238
Evidence against monocytes 239
Evidence for monocytes 239
CHONDROCLASTS AND OSTEOCLASTS 240
SYNOVIAL CELLS 240
NOTES 240
Part VI Embryonic Origins 244
Chapter 16 Skeletal Origins: Somitic Mesoderm 246
SOMITIC MESODERM AND THE ORIGIN OF THE VERTEBRAE 246
PARAXIAL MESODERM . SOMITES 247
SCLEROTOME FORMATION AND MIGRATION 247
RESEGMENTATION 249
SOMITIC CONTRIBUTION TO LIMB BUDS 251
Formation of muscle 251
Innervation and myogenesis 251
Signals to initiate a limb bud 252
A COMMENT ON PECTORAL GIRDLES 252
THE CLAVICLE: EVEN MORE SURPRISING 253
Humans 254
Other mammals 254
Mammals that lack clavicles 255
Birds 256
Wishbone or clavicles 256
NOTES 256
Chapter 17 Skeletal Origins: Neural Crest 259
DIFFERENT MESENCHYMES, SAME TISSUES 259
NEURAL CREST AS A SOURCE OF SKELETAL CELLS 260
EVIDENCE OF SKELETOGENIC POTENTIAL 260
Ablation and transplantation experiments 261
Marker experiments 262
3H-thymidine 262
Xenopus laevis–Xenopus borealis chimaeras 262
Quail–chick chimaeras 262
Genetic markers for murine neural crest 263
Information from mutants 265
REGIONALIZATION OF THE CRANIAL NEURAL CREST 268
THE VENTRAL NEURAL TUBE 268
MIGRATION OF NCC: THE ROLE OF THE ECM 269
NOTES 270
Chapter 18 Epithelial–Mesenchymal Interactions 272
URODELE AMPHIBIANS: CHONDROGENESIS 272
AVIAN MANDIBULAR SKELETON: CHONDROGENESIS AND OSTEOGENESIS 273
Isolated mesenchyme – chondrogenesis 276
Isolated mesenchyme – osteogenesis 276
Ruling out any role for Meckel’s cartilage 276
Molecular mechanisms 277
OSTEOGENESIS IN AVIAN MAXILLARY ARCH SKELETON 278
MAMMALIAN MANDIBULAR SKELETON 278
Endothelin-1 (Et-1) 279
The Dlx gene family and craniofacial development 279
TELEOST MANDIBULAR ARCH SKELETON 281
Fgf 281
Hoxd-4 and retinoic acid 281
Limb development 281
Craniofacial development 281
Fish 282
Endothelin-1 (Et-1) 282
Mutants 282
LATERAL LINE, NEUROMASTS AND DERMAL BONE 282
Hope from a single trout 282
TERATOMAS 283
Germ-layer combinations 283
MESENCHYME SIGNALS TO EPITHELIUM 284
SPECIFICITY OF EPITHELIAL–MESENCHYMAL INTERACTIONS 284
NOTES 285
Part VII Getting Started 288
Chapter 19 The Membranous Skeleton: Condensations 290
THE MEMBRANOUS SKELETON 290
CONGENITAL HYDROCEPHALUS (ch) 292
CHARACTERIZING CONDENSATIONS 293
HOW CONDENSATIONS ARISE 295
Altered mitotic activity 295
Changing cell density 295
Aggregation and/or failure to disperse 296
Limb buds and limb regeneration 296
Molecular control 297
ESTABLISHING BOUNDARIES 298
Syndecan and tenascin 298
Fgfs 299
Wnt-7a 299
NOTES 299
Chapter 20 From Condensation to Differentiation 301
CONDENSATION GROWTH 301
Lessons from mutants 302
talpid3 302
bpH 303
ADHERE, PROLIFERATE AND GROW 303
Gap junctions 303
Limb-bud mesenchyme 303
Craniofacial mesenchyme 303
Transcription Factors and Hox genes 303
POSITION AND SHAPE 305
ESTABLISHING CONDENSATION SIZE 306
Bmps 306
Fibronectin 306
Hyaluronan 306
Extrinsic control 307
FROM CONDENSATION TO OVERT DIFFERENTIATION 307
The molecular cascades 309
Bmps 309
Tenascin and N-CAM 309
Runx-2 310
NOTES 311
Chapter 21 Skulls, Eyes and Ears: Condensations and Tissue Interactions 313
THE BONY SKULL 313
Avian skull development 314
Mammalian skull development 316
THE CARTILAGINOUS SKULL 317
Type II collagen 317
Otic, optic and nasal capsules 317
The otic vesicle 317
Morphogenesis 318
TYMPANIC CARTILAGES 319
SCLERAL CARTILAGE 320
Heterogeneity 320
Chondrogenic mesenchyme 320
Pigmented retinal epithelium (PRE) 320
Morphogenesis 321
SCLERAL OSSICLES 322
Ossicle number 322
Scleral papillae 323
An epithelial–mesenchymal interaction 323
Scaleless mutant fowl 324
A role for tenascin? 324
NOTES 326
Part VIII Similarity and Diversity 328
Chapter 22 Chondrocyte Diversity 330
SEGREGATION FROM PRECURSORS 330
PERICHONDRIA 331
MORPHOGENETIC SPECIFICITY 332
CARTILAGES OF DIFFERENT EMBRYOLOGICAL ORIGINS 333
CHONDROCYTE HYPERTROPHY 334
TYPE X COLLAGEN 334
Discovery and regulation of synthesis 334
Syndromes and mutations 335
Type X does not always indicate hypertrophy 336
Regulation of chondrocyte hypertrophy 336
Tgf 337
Bmps 337
Type X and mineralization 338
Birds 338
Frogs 338
Rickets 338
MATRIX VESICLES 338
HYPERTROPHIC CHONDROCYTES AND SUBPERIOSTEAL OSSIFICATION 340
Brachypod (bpH ) in mice 340
Early changes 341
Fibulae 341
A role for Wnts 341
NOTES 343
Chapter 23 Cartilage Diversity 345
STERNAL CHONDROCYTES 345
Synthesis of collagen and glycosaminoglycan (GAG) 345
Differential expression of type II collagen 345
Differential synthesis and organization of collagen types 345
Type X collagen and hypertrophy 347
Fibronectin 347
Nanomelia 347
TUMOUR INVASION 347
VASCULARITY 348
RESISTING VASCULAR INVASION 349
INHIBITORS OF ANGIOGENESIS AND VASCULAR INVASION 350
Vascular endothelial growth factor (Vegf) 350
PTH-PTHrP 351
INTERPUBIC JOINTS AND THE TRANSFORMATION OF CARTILAGE TO LIGAMENT 351
Cartilage . ligament 353
Mediation by oestrogen and relaxin 354
NOTES 355
Chapter 24 Osteoblast and Osteocyte Diversity 357
OSTEOCYTIC OSTEOLYSIS 357
INITIATING OSTEOGENESIS IN VITRO FROM EMBRYONIC MESENCHYME 359
OSTEOGENIC CELLS IN VITRO 359
Folded periostea 361
Establishing isolated osteoblasts and initiating osteogenesis in vitro 362
Calvarial osteoblasts in vitro 362
Isolating subpopulations of calvarial osteogenic cells 363
Chondrogenesis from rodent and avian osteogenic cells 364
Clonal cultures 365
NOTES 365
Chapter 25 Bone Diversity 367
HETEROGENEITY OF RESPONSE TO SODIUM FLUORIDE 367
Enhanced proliferation and osteogenesis 367
Interaction with hormonal action 368
Osteoporosis 369
Chondrogenesis 369
Mineralization 369
Mechanical properties of bone 369
ALVEOLAR BONE OF MAMMALIAN TEETH 369
Origin 369
Physiology and circadian rhythms 369
PENILE AND CLITORAL CARTILAGES AND BONES 371
Os penis 373
Os clitoridis 373
Hormonal control 373
Digits and penile bones 374
Hoxd-12, Hoxd-13 AND POLYPHALANGY 374
OESTROGEN-STIMULATED DEPOSITION OF MEDULLARY BONE IN LAYING HENS 374
OESTROGEN-STIMULATED RESORPTION OF PELVIC BONES IN MICE 375
NOTES 376
Part IX Maintaining Cartilage in Good Times and Bad 378
Chapter 26 Maintaining Differentiated Chondrocytes 380
DIFFERENTIATED CHONDROCYTES 380
SYNTHESIS AND DEPOSITION OF CARTILAGINOUS EXTRACELLULAR MATRIX 381
Synthesis of chondroitin sulphate 381
Synthesis of type II collagen 382
SYNTHESIS OF COLLAGEN AND CHONDROITIN SULPHATE BY THE SAME CHONDROCYTE 382
Collagen gel culture 382
FEEDBACK CONTROL OF THE SYNTHESIS OF GLYCOSAMINOGLYCANS 382
Evidence from organ culture 382
Evidence from chondrocyte cell cultures 383
INTERACTIONS BETWEEN GLYCOSAMINOGLYCANS AND COLLAGENS WITHIN THE EXTRACELLULAR MATRIX 383
Synthesis of collagen and chondroitin sulphate are regulated independently 383
Hypertrophy 384
THE INTERACTIVE EXTRACELLULAR MATRIX 384
NOTES 385
Chapter 27 Maintenance Awry – Achondroplasia 387
GENETIC DISORDERS OF COLLAGEN METABOLISM 387
CARTILAGE ANOMALY (Can) IN MICE 388
ACHONDROPLASIA (Ac) IN RABBITS 389
ACHONDROPLASIA (Cn) IN MICE 389
FgfR-3 389
CHONDRODYSPLASIA (Cho) IN MICE 391
Sprouty 391
BRACHYMORPHIC (Bm) MICE 392
STUMPY (Stm) MICE 392
NANOMELIA (nm) IN DOMESTIC FOWL 392
INDUCED MICROMELIA 393
METABOLIC REGULATION AND STABILITY OF DIFFERENTIATION 393
NOTES 394
Chapter 28 Restarting Mammalian Articular Chondrocytes 396
MAMMALIAN ARTICULAR CHONDROCYTES IN VITRO 396
A role for oxygen 397
Responsiveness to environmental signals 397
MECHANISMS OF ARTICULAR CARTILAGE REPAIR 398
Dividing again in vitro 398
Dividing again in vivo 401
DNA synthesis vs. division 401
Osteotomy and trauma 402
NOTES 402
Chapter 29 Repair of Fractures and Regeneration of Growth Plates 404
A BRIEF HISTORY OF FRACTURE REPAIR 404
Standardizing the fracture 405
Motion 405
Non-unions and persistent non-unions 406
Growth factors and fracture repair 408
Bmps 409
Jump-starting repair 409
REGENERATION OF GROWTH PLATES IN RATS, OPOSSUMS AND MEN 409
NOTES 410
Part X Growing Together 412
Chapter 30 Initiating Skeletal Growth 414
WHAT IS GROWTH? 414
NUMBERS OF STEM CELLS 414
CELL MOVEMENT AND CELL VIABILITY 415
Epithelia and Fgf/FgfR-2 415
METABOLIC REGULATION 415
Creeper (cp) fowl 416
Tibia/fibula 416
Growth retardation 416
A growth inhibitor 417
MECHANICAL STIMULATION AND CHONDROBLAST DIFFERENTIATION/GROWTH 417
MECHANICAL STIMULI AND METABOLIC ACTIVITY 418
Transduction 418
Membrane potential 419
SKELETAL RESPONSES MEDIATED BY cAMP 419
Matrix synthesis and condensation 419
Hormones 419
Teeth and alveolar bone 420
Electrical stimulation 420
cAMP AND PRECHONDROBLAST PROLIFERATION 420
Long bones 420
Limb regeneration 421
Condylar cartilage 421
NOTES 421
Chapter 31 Form, Polarity and Long-Bone Growth 424
FUNDAMENTAL FORM 424
POLARITY 425
Polarized cells 425
LONG-BONE GROWTH 426
Growth plates 427
Growth-plate dynamics 428
New cells, bigger cells and matrix 428
Cell proliferation 429
Birds and mammals 431
Clones and timing 431
Hormonal involvement 432
Growth at opposite ends 432
Diurnal and circadian rhythms 432
Rhythms are under hormonal control 433
A role for the periosteum in regulation of the growth plate? 433
Periosteal sectioning 435
Feedback control 435
NOTES 436
Chapter 32 Long Bone Growth: A Case of Crying Wolff? 438
WOLFF, VON MEYER OR ROUX 438
RESPONSE TO PRESSURE 439
CONTINUOUS OR INTERMITTENT MECHANICAL STIMULI 440
SCALING AND VARIATION: WHEN WOLFF MEETS THE DWARFS 441
GRAVITY 441
TRANSDUCTION OF MECHANICAL STIMULI 443
NOTES 443
Part XI Staying Apart 446
Chapter 33 The Temporomandibular Joint and Synchondroses 448
THE MAMMALIAN TEMPOROMANDIBULAR JOINT (TMJ) 448
Mechanical factors 449
The condylar process 449
The angular process 450
Diet 450
Other functional approaches 451
CRANIAL SYNCHONDROSES 452
As pacemakers 453
Limited growth potential 454
As adaptive 455
NOTES 456
Chapter 34 Sutures and Craniosynostosis 458
SUTURAL GROWTH AS SECONDARY AND ADAPTIVE 458
Alizarin 460
Working with the functional matrix 462
SUTURAL CARTILAGE 463
THE DURA 463
CRANIOSYNOSTOSIS 464
Msx-2 465
Fgf receptors 465
Sutural growth 465
Sutural fusion 466
NOTES 466
Part XII Limb Buds 470
Chapter 35 The Limb Field and the AER 472
THE MESODERMAL LIMB FIELD 472
ECTODERMAL RESPONSIVENESS 473
MESODERM SPECIFIES FORE- VS. HIND LIMB 474
ROLES FOR THE ECTODERM ASSOCIATED WITH THE LIMB FIELD 476
Limb-bud growth 479
Cell proliferation 479
Suppressing the flank 479
Mitotic rate in limb mesenchyme 480
Proximo-distal patterning of the limb skeleton 480
MESENCHYMAL FACTORS MAINTAIN THE AER 481
AEMF 481
The PNZ 481
SPECIFICITY OF LIMB-BUD EPITHELIUM 482
SPECIFICITY OF DISTAL LIMB MESENCHYME 484
THE TEMPORAL COMPONENT 485
A MECHANICAL ROLE FOR THE EPITHELIUM? 485
NOTES 486
Chapter 36 Adding or Deleting an AER 487
AER REGENERATION 487
EXPERIMENTAL REMOVAL OF THE AER 488
FAILURE TO MAINTAIN AN AER: WINGLESS (wl) MUTANTS 489
Mutual interaction 490
EXPERIMENTAL ADDITION OF AN AER 491
MUTANTS WITH DUPLICATED LIMBS 491
An enlarged AER 491
Duplicating the AER 493
Narrow or subdivided AERs 496
NOTES 496
Chapter 37 AERs in Limbed and Limbless Tetrapods 498
AERs ACROSS THE TETRAPODS 498
Amphibians 498
Anurans 498
Urodeles 499
Reptiles 499
Mammals 499
Mice 499
Chimaeras 500
Humans 501
LIMBLESS TETRAPODS 501
Evolutionary patterns 501
Gaining limbs back 501
Ecological correlates of limblessness 502
The developmental basis of limblessness in snakes and legless lizards 503
Inability to maintain an AER 504
Molecular mechanisms 505
NOTES 505
Part XIII Limbs and Limb Skeletons 508
Chapter 38 Axes and Polarity 510
ESTABLISHING AXES AND POLARITY 510
THE A-P AXIS AND THE ZPA 510
A role for Fgf-2 511
dHand and Shh 511
Wnts and Fgf 512
ZPAs abound 513
D-V POLARITY 513
P-D POLARITY AND THE PROGRESS ZONE 513
Extension to amphibian limb regeneration 513
CONNECTING D-V AND P-D POLARITY 514
THALIDOMIDE AND LIMB DEFECTS 514
Time of action 515
Mode of action 515
NOTES 517
Chapter 39 Patterning Limb Skeletons 519
MORPHOGENESIS AND GROWTH 519
PROGRAMMED CELL DEATH (APOPTOSIS) 520
Posterior and anterior necrotic zones (PNZ, ANZ) 520
Interdigital cell death 521
A role for BmpR-1 522
The opaque patch 523
CELL ADHESION AND MORPHOGENESIS: TALPID (ta) MUTANT FOWL 523
Talpid2 524
Talpid3 524
NOTES 526
Chapter 40 Before Limbs There Were Fins 527
DORSAL MEDIAN UNPAIRED FINS 527
Teleost fish 527
Life style 527
Developmental origins 528
Evolutionary origins 528
PAIRED FINS 532
Fin buds and fin folds 532
Fin skeletons 533
Retinoic acid 534
…Regeneration 535
An RA-Shh link 535
FIN REGENERATION 536
FINS . SUCKERS 536
FINS . LIMBS22 536
FROM MANY TO FEWER DIGITS 537
NOTES 538
Part XIV Backbones and Tails 540
Chapter 41 Vertebral Chondrogenesis: Spontaneous or Not? 542
SELF-DIFFERENTIATION OR INDUCTION? 542
MORPHOGENESIS 543
Spinal ganglia and vertebral morphogenesis 544
CHONDROGENESIS IN VITRO 545
SPONTANEOUS CHONDROGENESIS? 545
Environmental influences 546
Cell division and cell death 546
NOTES 547
Chapter 42 The Search for the Magic Bullet 548
INTEGRITY OF NOTOCHORD/SPINAL CORD AND VERTEBRAL MORPHOGENESIS 548
Fish skeletal defects 548
FOR HOW LONG ARE NOTOCHORD AND SPINAL CORD ACTIVE? 549
CAN DERMOMYOTOME OR LATERAL-PLATE MESODERM CHONDRIFY? 549
THE SEARCH FOR THE MAGIC BULLET 550
A role for ectoderm? 550
Cartilage cells as cartilage inducers 551
CHONDROCYTE EXTRACELLULAR MATRIX 552
NOTOCHORD AND SPINAL CORD EXTRACELLULAR MATRICES 552
GLYCOSAMINOGLYCANS 552
Collagens 553
FUNCTION OF NOTOCHORD AND SPINAL CORD MATRIX PRODUCTS 554
KEY ROLES FOR Pax-1 AND Pax-9 554
CONCLUSIONS 556
NOTES 556
Chapter 43 Tail Buds, Tails and Taillessness 558
EMBRYOLOGICAL ORIGIN 558
THE VENTRAL EPITHELIAL RIDGE (VER) 558
Tbx GENES 560
TAIL GROWTH 560
Genes or environment 560
Temperature 560
TEMPERATURE-INDUCED CHANGE IN VERTEBRAL NUMBER: MERISTIC VARIATION 563
Natural variation and adaptive value 563
Studies with teleost fish 564
Studies with avian embryos 564
Studies with mammals 564
Studies with amphibian embryos 565
Temperature plus… 565
TAILLESSNESS 565
AND THEREBY HANGS A TAIL 566
Fish tails 566
LIZARDS’ TAILS: AUTOTOMY 566
NOTES 566
Part XV Evolutionary Skeletal Biology 568
Chapter 44 Evolutionary Experimentation Revisited 570
VARIATION 570
Variation of individual elements 570
Variation that tests a hypothesis 571
Pattern variation 572
ADAPTIVE VALUE 572
METAMORPHOSIS 573
MINIATURIZATION 573
HETEROCHRONY 576
Process heterochrony 576
Coupling and uncoupling dermal and endochondral ossification 576
Primates 577
NEOMORPHS 577
The preglossale of the common pigeon 577
Digits 578
Secondary jaw articulations 578
A Boid intramaxillary joint 579
Regenerated joints 579
Wishbones 579
Limb rudiments in whales 579
Turtle shells 580
Development 580
Evolutionary history 580
ATAVISMS 583
Limb skeletal elements in whales 584
Mammalian teeth 584
Teleosts and taxic atavisms 584
Late-developing bones in anurans 585
NEOMORPH OR ATAVISM? 585
NOTES 586
References 588
Index 766
Preface
The skeleton has fascinated humankind ever since it was realized that, aside from one or several sets of genes, bare bones are our only bequest to posterity. But the skeleton is more than an articulated set of bones: its three-dimensional conformation establishes the basis of our physical appearance; its formation and rate of differentiation determine our shape and size at birth; its postnatal growth orders us among our contemporaries and sets our final stature; while its decline in later life is among the primary causes of loss of the swiftness and agility of youth. Not surprisingly, the skeleton is a central focus of many scientific and biomedical disciplines and investigations.
For the developmental or cell biologist, the skeleton provides an excellent model for studies of gene action, cell differentiation, morphogenesis, polarized growth, epithelial–mesenchymal interactions, programmed cell death, and the role of the extracellular matrix. The skeleton supplies the geneticist with a permanent record of the vicissitudes of its growth, whereby the phenotypic expression of genetic abnormalities can be studied. The orthopaedic surgeon earns a livelihood from correcting abnormalities and breaks, while the orthodontist corrects the position of teeth displaced consequent to alveolar bone dysfunction. Physiologists, biochemists and nutritionists are concerned with the skeleton’s store of calcium and phosphorus and its response to vitamins and hormones. Haematologists, on the other hand, find that the skeleton houses the progenitors of the blood cells. Pathologists endeavour to understand the disease states that result from abnormalities in skeletal cellular differentiation or function; surgeons want to prevent formation of skeletal tissues in the wounds that bear witness to their work. Vertebrate palaeontologists make their living from the analysis of the skeletons of extinct taxa. Veterinarians, physical anthropologists, radiographers, forensic scientists – the list goes on.
Bones come in all shapes and sizes. There are long bones, flat bones, curved bones, bones of irregular and geometrically indefinable shapes, large bones and small. Bones exhibit bumps, ridges, grooves, holes and depressions where they articulate with other bones, attach to tendons and ligaments, and where nerves and blood vessels course through them. Some bones and cartilages arise within the skeleton and are integral parts of it. Others arise outside the skeleton, some as sesamoids or ossifications within tendons or ligaments, others as pathological ossifications in what otherwise would be benign soft tissues. Bones and cartilages may develop during embryonic or foetal life, in larval stages or in adulthood – often late in adulthood – during normal ontogeny, wound repair, or regeneration. Bones modify themselves in response to injury, disease or parasitic infection, in the aftermath of surgery, as a defensive response to predators, as a consequence of domestication or hibernation, and through evolutionary adaptations.
My previous book on the skeleton – Developmental and Cellular Skeletal Biology – was published in 1978. That book concerned itself with how bones and cartilages are made and how these tissues, organs and systems evolved. So too does the present book, which includes and updates the earlier treatment. With respect to skeletal development, I address such questions as the following.
• Is bone always bone, no matter where and under what conditions it forms?
• Do bones that develop indirectly by replacing another tissue – be it cartilage, marrow, connective tissue, fat, tendon or ligament – differ from one another, and/or from bone that develops directly (intramembranously)?
• Is fast-growing the same as slow-growing bone?
• Is fish bone the same as human bone?
• Does bone form continuously or in cycles?
• Do bears make new bone during hibernation?
• Can sharks make bone?
• If cartilage does not contain type II (cartilage-type) collagen, is it still cartilage?
• Does the body contain cells that can differentiate as chondrocytes or as osteocytes and, if so, what factors allow cells to choose their fate?
• Are progenitor (stem) cells for bone and cartilage only found within the skeleton? If not, how do we recognize such cells and activate them for skeletogenesis?
• Why is aggregation (condensation) of cells so important for the initiation of the skeleton?
• Does the skeleton display daily or circadian rhythms?
• Do similar genes/growth factors regulate the differentiation of osteoblasts and chondroblasts?
• Can mononucleated cells resorb bone?
• How do joints form and remain patent?
• How does activating FGF receptors cause cranial sutures to fuse?
• What can mutants tell us about normal skeletogenesis?
• Does Wolff’s law really govern the structure of bone?
• How do chondroid, chondroid bone, osteoid and bone differ from one another?
• How do antlers, horns and knobs (ossicones) differ one from the other?
• Can we restart cell division in articular cartilage to effect repair?
With respect to the evolution of skeletal tissues, organs and systems, I ask such questions as the following.
• What are the evolutionary relationships between cartilage and bone and between acellular and cellular bone?
• How did novel features such as tetrapod limbs arise from fish fins?
• Can fossilized bone reveal patterns of growth, metabolism or physiology?
• Why are so few aware of the extensive cartilaginous skeletons found in many invertebrates?
• Is five the canonical number of tetrapod digits?
• If tetrapods are vertebrates with limbs, then how can limbless snakes be tetrapods?
• How did snakes lose their limbs?
• How did whales lose their hind limbs and transform their forelimbs into flippers?
• How do we recognize the diverse range of tissues in fossilized skeletons that are intermediate between connective tissues and cartilage, cartilage and bone, bone and dentine, or dentine and enamel?
• Why can some vertebrates regenerate their limbs or tails and others not?
• How does reduction in body size (miniaturization) affect the skeleton?
The answers to the above and many other questions may be found in this book. Sometimes the ‘answers’ are limited to descriptions. In other cases we have an extensive knowledge of the molecular, cellular, developmental and evolutionary processes involved. Some transitions (fins → limbs, for example) are understood in considerable detail, with paleontology, paleobiology, paleohistology, paleopathology, and the study of extant forms through molecular, cell and developmental biology contributing to our understanding. Other transitions – the origin of the turtle shell, for example – are much less well understood, with fossils contributing little and developmental information only beginning to appear.
Discussion of the mechanisms of skeletal development and evolution is organized into 15 parts to enable you to select with ease a topic of special interest. The range of skeletal tissues covered by the book is outlined in Part I. Although primarily devoted to bone and cartilage, Part I introduces dentine and enamel and four skeletal tissues that I call ‘intermediate’ because they display features of two or more of cartilage, bone, dentine and enamel. The four are chondroid, chondroid bone, cementum and enameloid. Discussion of these intermediate tissues is expanded in Part II in the context of what I refer to as ‘natural experiments,’ a category that includes invertebrate cartilages and an examination of the evolution of skeletal tissues.
Unusual tissues are followed in Part III by unusual modes of skeletogenesis, namely, horns, antlers, intratendinous ossifications and sesamoids, and the ossicones (knobs) of giraffes. Parts IV and V deal with the origin of skeletogenic cells, either as stem cells in embryos or adults (Chapters 10 and 11) or as more definitive skeletogenic cells (Part V). Here the emphasis is on those cells that can differentiate either as chondro- or osteoblasts (Chapter 12), on dedifferentiation as a source of skeletogenic cells in normally developing long bones and jaws and in regenerating urodele limbs (Chapters 13 and 14), and on the relationship(s) between the cells that make and the cells that break bone – osteoblasts and osteoclasts (Chapter 15).
I move explicitly into embryonic development in the three chapters in Part VI through examination of the embryonic origins of skeletogenic cells in somitic mesoderm and the neural crest, and an evaluation of the roles of epithelial–mesenchymal interactions in initiating skeletogenesis. The developmental processes that underpin skeletal formation – differentiation, morphogenesis and growth – are mediated through modification of cell division, movement, death (apoptosis) and/or specialization. To our amazement, similar genes and gene networks or pathways may be...
| Erscheint lt. Verlag | 20.6.2005 |
|---|---|
| Sprache | englisch |
| Themenwelt | Sachbuch/Ratgeber |
| Medizinische Fachgebiete ► Chirurgie ► Unfallchirurgie / Orthopädie | |
| Medizinische Fachgebiete ► Innere Medizin ► Endokrinologie | |
| Medizin / Pharmazie ► Medizinische Fachgebiete ► Orthopädie | |
| Studium ► 1. Studienabschnitt (Vorklinik) ► Anatomie / Neuroanatomie | |
| Studium ► 1. Studienabschnitt (Vorklinik) ► Biochemie / Molekularbiologie | |
| Studium ► 1. Studienabschnitt (Vorklinik) ► Histologie / Embryologie | |
| Studium ► 1. Studienabschnitt (Vorklinik) ► Physiologie | |
| Naturwissenschaften ► Biologie ► Evolution | |
| Naturwissenschaften ► Biologie ► Genetik / Molekularbiologie | |
| Naturwissenschaften ► Biologie ► Ökologie / Naturschutz | |
| Naturwissenschaften ► Biologie ► Zoologie | |
| Technik | |
| ISBN-10 | 0-08-045415-1 / 0080454151 |
| ISBN-13 | 978-0-08-045415-3 / 9780080454153 |
| Haben Sie eine Frage zum Produkt? |
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