Vertebrate:comparative anatomy,function,evolution/
Kenneth Kardong
- New Delhi: Tata mcgrow hill education, 2010.
- 782p.
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