Vertebrate:comparative anatomy,function,evolution/ Kenneth Kardong

Comparative Vertebrate Morphology 1
Designs of Students 2
Vertebrate Design—Form and Function 3
Grand Design 3
Historical Predecessors—Evolution 3
The Process behind the Change 4
Linnaeus 4
Naturalists 5
J'B. de Lamarck 5
Acquired Characteristics 6
Upward to Perfection 6
Natural Selection 7
A. R. Wallace 7
Charles Darwin 8
Critics and Controversy 9
Historical Predecessors-
Georges Cuvier 10
Richard Owen 11
Why Are There No Flying Elephants? 13
Morphological Concepts 14
Similarities 14
Symmetry 16
Segmentation 16
Evolutionary Morphology 18
Function and Biological Role 18
Preadaptation 18
Evolution as Remodeling 20
Phylogeny 20
Of Bean Stalks and Bushes 20
Simplification 22
Patterns of Phylogeny 22
Grades and Clades 23
Contents
-Morphology 10
Paleontology 27
Fossilization and Fossils 27
Recovery and Restoration 30
From Animal to Fossil 33
Dating Fossils 35
Stratigraphy 35
Index Fossils 35
Radiometric Dating 35
Geological Ages 35
Tools of the Trade 39
The Question 39
The Function 40
The Biological Role 43
Overview 45
CHAPTER two
Origin of Chordates 47
Chordate Phylogeny 47
Chordate Characteristics
Notochord 49
Pharyngeal Slits 51
Endostyle or Thyroid Gland 52
Dorsal and Tubular Nerve Cord 52
Postanal Tail 52
Chordate Body Plan 52
Protochordates 53
Hemichordata 54
Enteropneusta—"Acorn Warms" 55
Pterobranchia 57
Hemichordate Phylogenetic Affinities 58
Urochordata 58
Ascidiacea—"Sea Squirts" 60
Larvacea (Appendicularia) 61
Thaliacea 64
Cephalochordata 65
Overview of Protochordates 70
Chordate Origins 71
Cephalochordates from Annelids and An! .ropods 71
Cephalochordates from Echinoderms 72
The Evidence 71
Appearance of Chordate Characteristics 73
Selective Advantage 73
Larval Echinoderm to Chordate Tadpole 74
The Critics 75
Phylogeny, Paedomorphism, and Origins 75
Overview 77
CHAPTER three
The Vertebrate Story
Introduction 80
Innovations 81
Vertebral Column 81
Head 82
Origin of Vertebrates 82
Step 1: Pre Vertebrate 82
Step 2: Agnathan 83
Step 3: Gnathostome 83
The Controversial Calci Chordates 84
Vertebrate Classification 84
Agnathans 84
Living Agnathans 86
Myxinoidea 86
Petromyzontida 86
Early Vertebrate Fossils 86
Conodonts 87
Ostracoderms 89
Pteraspidomorphi 89
Other Ostracoderms 89
Overview of Agnathan Evolution 91
Gnathostomes 92
Placodermi 93
Chondrichthyes 93
Elasmobranchii—Sharks andRays 95
Holocephali—Chimaeras 95
Teleostomi 96
Acanthodii 96
Osteichthyes 96
Actinopterygii 98
Sarcopterygii 99
Overview of Fish Phylogeny 101
Tetrapods 101
Primitive Tetrapods 102
Labyrinthodonts 102
Lepospondyls 104
Lissamphibia-Modem Amphibians 105
Urodela (Caudata) 105
Salientia (Anura) 106
Gymnophiona (Apoda) 106
80
Amniotes 106
Stem-Amniotes 108
Sauropsids 108
Mesosdwrs 108
Reptilia 108
Synapsida 116
Pelycosauria 117
Therapsida 117
Mammalia 120
CHAPTER four
Biological Design 125
Introduction: Size and Shape 125
Size 128
Relationships among Length, Area, and Volume 128
Surface Area 130
Volume and Mass 130
Shape 131
Allometry 131
Transformation Grids 132
On THE Consequences of Being the Right Size 134
Biomechanics 134
Fundamental Principles 135
Basic Quantities—Length, Time, and Mass 135
Units 136
Derived Quantities—Velocity, Acceleration, Force,
and Relatives 136
Reference Systems 137
Center of Mass 137
Vectors 137
Basic Force Laws 138
Free Bodies and Forces 138
Torques and Levers 139
Land and Fluid 141
Life on Land: Gravity 141
Life in Fluids 142
Machines 144
Strength of Materials 145
Loads 146
Biological Design and Biological Failure 146
Tissue Response to Mechanical Stress 147
Responsiveness of Bone 148
Biophysics and Other Physical Processes 152
Diffusion and Exchange 152
Pressures and Partial Pressures 152
Countercurrent, Concurrent, and Crosscurrent
Exchange 153
Optics 154
Depth Perception 155
Accommodation 155
Overview 156
CHAPTER five
Life History 158
Introduction 158
Early Embryology 159
Fertilization 159
Cleavage 161
Amphioxus 161
Fishes 162
Amphibians 162
Reptiles and Birds 162
Mammab 163
Overview of Cleavage 164
Gastrulation and Neurulation 164
Amphioxus 166
Fishes 166
Amphibians 167
Reptiles and Birds 168
Mammals 169
Organogenesis 172
Histogenesis 174
Epithelium 174
Covering and Lining Epithelium 176
Glandular
Epithelium 176
Connective Tissues 177
General Connective Tissues 177
Special Connective Tissues 178
Bone Development and Growth 179
Endochondral Bone Devebpment 179
Intramembranous
Bone Development 181
Comparative Bone Histology 181
Bone Remodeling and Repair 183
Joints 184
Neural Crest and Ectodermal Placodes 186
Extraembryonic Membranes 1-86
Reptiles and Birds 187
Mammals 187
Eutherian Placenta 188
Other Placentae 189
Overview of Early Embryonic Development 189
Development of the Coelom and its
Compartments 191
Maturation 194
Metamorphosis 194
Heterochrony 195
Peramorphosis 195
Paedomorphosis 197
OnTOGENY ANDPHYLOGENY 197
Biogenetic Law 197
von Baer's Law 198
Overview of the Biogenetic Law 199
Hox Genes and Their Kingdoms
Egg to Adult 200
Shaping Up: Positions and Parts
Evolutionary Significance 201
Epigenomics 202
Induction 202
Phylogeny 203
Overview 206
CHAPTER SIX
Integument 209
Embryonic Origin 210

General Features OF THE Integument 210
Dermis 210
Epidermis 212
Phylogeny 213
Integument of Fishes 213
Primitive Fishes 214
Chondrichthyes 214
•Bony Fishes 214
Integument of Tetrapods 216
Amphibians 216
Reptiles 217
Birds 218
Mammals 221
Specializations of the Integument
Nails, Claws, Hooves 225.
Horns and Antlers 226
Baleen 227
Scales 228
Dermal Armor 228
Mucus 228
Color 229
Overview 230
CHAPTER seven
Skeletal System:
The Skull 234
Introduction 235
Chondrocranium
Embryology 235
Splanchnocranium
Embryology 238
Origin of Jaws 239
Types of Jaw Attachments
Dermatocranium 242
235
Parts of the Dermatocranium
Dermal Bone Series 242

Overview of Skull Morphology 243
Braincase 243
Jaws 246
Hyoid Apparatus 246
Cranial Kinesis 246
Phylogeny of the Skull 247
Agnathans 247
Ostracoderms 247
Cyclostomes 248
Gnathostomes 249
Fishes 249
Early Tetrapods 256
Primitive Amniotes 258
Modem Reptiles 260
Birds 265
Synapsids 267
Overview of Skull Function and Design 275
Prey Capture 277
Feeding in Water 277
Feeding in Air 280
Swallowing 280
Overview 280
Cranial Neural Crest 280
Emergence of Mammals 281
Evolutionary Modifications of Immature Forms:
Akinesis in Mammals 282
Composite Skull 285
CHAPTER eight
Skeletal System:
The Axial Skeleton 289
Introduction 289
Basic Components 290
Vertebrae 290
Regions of the Vertebral Column 290
Centra 290
Ribs 292
Sternum 294
Gastralia 294
Embryonic Development 296
Fishes 296
Tetrapods 297
Phylogeny 298
Fishes 298
Agnathans 298
Gnathostomes 298
Contents
TetrapaKs 303
Early Tetrapods
Amniotes 308
303
Form and Function 311
Fluid Environment 311
Terrestrial
Environment 312
Design of Vertebrae 313
Direction of the Neural Sprite 313
Height of the Neural Spine 314
Regionalization of the Vertebral Column
Overview 318
CHAPTER nine
Skeletal System: The
Appendicular Skeleton 320
Introduction 320
Basic Components 321
Fins 321
Limbs 321
Origin of Paired Fins 322
Gill-Arch Theory 322
Fin-Fold Theory 323
Embr^'onic Development ofTetrapod Limbs 324
Phylogeny 325
Fishes 325
Agnathans 325
Placoderms 326
Chondrichthyans 326
Acanthodians 328
Bony Fishes 328
Tetrapods 330
Pectoral Girdle 330
Pelvic Girdle 333
Manus and Pes 333
Evolution of the Appendicular System
Dual Origin of the Pectoral Girdle 336
Adaptive Advantage of Lobe Fins 337
Onto the Land 341
Form and Function
Swimming 342
Terrestrial Locomotion
Early Gaits 342
Early Modes of Locomotion
Cursorial Locomotion 347
Aerial Locomotion 350
Origin of Bird Flight 360
Fossorial Locomotion 360
Overview 362
342
342
343
315
336
CHAPTER ten
The Muscular System 365
Introduction 365
Organization of Muscles 366
Classification of Muscles 366
Skeletal Muscle 367
Cardiac Muscle 368
Smooth Muscle 368
Structure of Skeletal Muscles 368
Tendons 370
Basis of Muscle Contraction 370
Resting andActive Muscle 370
Molecular Mechanisms of Contraction 370
Muscle Function 371
Muscle Fibers 371
Tension-Length Curves for a Single
Muscle Fiber 371
Properties of Muscle Fibers 371
Muscle Organs and Fibers 373
Whole Muscle Force Generation 373
Tension-Length Curves fora Whole Muscle
Graded Force 376
Cross-Sectional Area 376
Fiber Orientation 377
Velocity of Shortening 377
Distance of Shortening 378
Bone-Muscle Lever Systems 378
Sequencing of Muscle Actions 380
Overview of Muscle Mechanics 381
Muscle Actions 381
Muscle Homologies 383
Embryonic Origin of Muscles 384
Postcranial Musculature 384
Appendicular Musculature 384
Axial Musculature 385
Cranial Musculature 386
Jaw and Pharyngeal Musculature 386
Extrinsic EyeMuscles 386
Comparative Anatomy 387
Postcranial Musculature 387
Axial Musculature 387
Appendicular Musculature 390
Cranial Musculature 396
Branchiomeric Musculature 396
Hypobranchial Musculature 400
Overview 401
374
chaptek eleven
The Respiratory System 406
Introduction 406
Respiratory Organs
Gills 409
Lungs 409
Gas Bladders 409
Cutaneous Respiratory Organs 410
Accessory Air-breathing Organs 410
Breathing and Embryos 411
Ventilatory Mechanisms 414
Cilia 414
Muscular Mechanisms 414
Water Ventilation: Dual Pump 414
Air Ventilation: BuccalPump 415
Air Ventilation: Aspiration Pump 416
Phylogeny 417
Agnathans 417
Elasmobranchs 419
BonyFishes 420
Overview of Fish Respiration 420
Gills 420
Lungs and Gas Bladders 421
Amphibians 423
Amphibian Larvae 423
Amphibian Adults 425
Reptiles 426
Mammals 427
Ventilation 428
Gas Exchange 429
Birds 430
Form and Function 432
Patterns of Gas Transfer 432
Rates of Gas Transfer 433
Breathing in Water 437
Breathing in Air 437
Evolution of Respiratory Organs 437
Acid-Base Regulation 437
Ventilation 438
Ciliary Pumps 438
Muscular Pumps 439
Water-to-Land Transition 439
Air-breathing Organs 439
Advantages of Movement to Land
Air-breathing
Mechanisms 440
Bird Lungs and Air Sacs 441
Overview 442
409
439
Contents
XI
CHAPTER twelve
The Circulatory System 445
Introduction 445
Cardiovascular System 446
Blood 446
Arteries, Veins, and Capillaries 446
Arteries 447
Hemodynamics of Circulation 447
Veins 448
Microcirculation 448
Single and Double Circulation 449
Embryonic Development of the Cardiovascular
System 450
Phylogeny of the CardiovascularSystem 451
Arterial Vessels 455
Venous Vessels 460
Hearts 467
Basic Vertebrate Heart 467
Fishes 469
Amphibians 472
Reptiles 474
Birds and Mammals 481
Cardiovascular System: Matching Design
to Environmental Demands 481
Accessory Air-breathing Organs 484
Diving Birds andMammab 484
Heart Flow 484
Ontogeny ofCardiovascular Function 485
Fetal Circulation in PlacentalMammab 485
Changes at Birth 485
Heat Transfer 487
Lymphatic System 488
Lymphatic Vessels 489
Lymphatic Tissue 491
Form and Function 491
Overview 492
CHAPTER thirteen
The Digestive System 496
Introduction 496
Preview 497 j
Components of the Digestive System 497 j
Xll
Buccal Cavity 497 ?
Boundaries 497
Palate 498
Teeth 499
Tongue 509
Pharynx 510
Contents
Alimentary Can?l 513
Esophagus 515
Stomach 515
Intestines 516
Cloaca 518
Specializations of the Alimentary Canal 518
Vascularization of the Gastrointestinal Tract 520
Fishes 520
Tetrapods 521
Associated Glands of Digestion 524
Oral Glaruis 524
Liver 526
Pancreas 527
Function and Evolution of the
Digestive System 528
Absorption 528
Feces 528
Mechanical Breakdown of Food 529
Mastication 529
Gizzards 529
Chemical Breakdown of Food 529
Gastric Fermentation 530
Intestinal Fermentation 533
Gastric versus Intestinal Fermenters 533
Size and Fermentation 535
DigestingToxins 535
Overview 535
CHAPTER fourteen
The Urogenital System 538
Introduction 538
Urinary System 538
Structure of the Mammalian Kidney 538
Embryonic Development 540
Nephrotome to Nephric Tubules 540
Tripartite Concept of Kidney Organization 541
Kidney Phylogeny 544
Fishes 544
Tetrapods 545
Kidney Function and Structure 546
Excretion: Removing the Products of Nitrogen
Metabolism 546
Osmoregulation: Regulating Water and Salt
Balance 548
Evolution 555
Preadaptation 555
Origin of Vertebrates 555
Reproductive System 556
Structure of the Mammalian Reproductive
System 556
Embryonic Development 556
Gonads and Gametes 556
Reproductive Tracts 558
Overview 558
Female Reproductive System 560
Ovary 560
Genital Ducts 560
Oviduct 562
Uterus 563
Male Reproductive System 565
Testis 565
Genital Ducts 565
Copulatory Organs 568
Cloaca 572
Urinary Bladder 577
Function and Evolution 578
Potency and Fertility 578
External and Internal Fertilization 579
Delays in Gestation 581
Overview 583
CHAPTER fifteen
The Endocrine System 585
Survey of Endocrine Organs 585
Thyroid Gland 585
Structure and Phylogeny 585
Function 587
Ultimobranchial Body and Parathyroid Gland 589
Ultimobranchial Body 589
Parathyroid Gland 589
Form and Function 591
Adrenal Gland 591
Structure and Phylogeny 591
Function 594
Pancreatic Islets 595
Structure and Phylogeny 595
Function 595
Pituitary Gland 597
Structure 597
Phylogeny 597
Function 600
Gonads 603
Pineal Gland 603
SecondaryEndocrine Organs 603
Gastrointestinal Tract 604
Kidneys 604
Endocrine Coordination 605
MammalianReproduction 605
Male 605
Female 606
Metamorphosis in Frogs 612
Fundamentals of Hormonal Control 613
Functional andStructural Linkage 613
Target Tissue Responses 614
The Endocrine System and the Environment 614
Evolution 615
Overview 616
CHAPTER sixteen
The Nervous System 618
L-.troduction 618
Types of Cells within the Nervous System 618
Neuroglia 618
Neurons 619
Transmission of Information 619
Neurosecrctory Cells 621
Peripheral Nervous System 621
Spinal Nerves 622
Cranial Nerves 623
Evolution 630
Functions of the Peripheral Nervous System 632
Spinal Reflexes 632
The Autonomic Nervous System 633
Central Nervous System 638
Embryology 639
Spinal Cord 639
Spinal Reflexes 642
SpinalTracts 642
Brain 643
Phylogeny 643
Form and Function 644
Functional Associations ofParts ofthe Central
Nervous System 658
Overview 660
CHAPTER seventeen
Sensory Organs 663
Introduction 663
Components of a Sensory Organ 664
General Sensory Organs 664
FreeSensory Receptors 664
Encapsulated Sensory Receptors 665
Associated Sensory Receiptors 665
Proprioception 665
Mechanisms of Perceiving Stimuli from General
Sensory Receptors 666
Contents
xui
Special Sensory Organs 666
Chemoreceptors 666
Nasal Passages 667
Vomeronasal Area 669
Mouth 670
Radiation Receptors 671
Photoreceptors 67i
Infrared Receptors 684
Mechanoreceptors , 684
Lateral Line System 685
Vestibular Apparatus 686
Auditory System 688
Functions of the Ear 689
Electroreceptors 698
Structure and Phylogeny 698
Form and Function 699
Additional SpecialSensory Organs 701
Overview 701
CHAPTER eighteen
Conclusions 706
Introduction 706
Structural Analysis 709
Functional Analysis 710
XIV
How Does It Work? 710
Functional Coupling, Functional Compromise 711
Contents
Multiple Functions 712
Performance 713 -
Ecolcx31cal Analysis 714
Evolutionary Analysis 714
Historical Constraints 714
Primitive and Advanced 714
Diversity of Type/Unity of Pattern 715
Mosaic Evolution 717
Morphology and Modules 717
Mode and Tempo of Evolution 719
Remodeling 720
Embryonic Changes 721
Hox Genes 721
Evolutionary Significance 722
The Promise of Vertebrate Morphology 722