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Abstract
Master the concepts you need to know with Human Embryology and Developmental Biology. Dr. Bruce M. Carlson's clear explanations provide an easy-to-follow "road map" through the most up-to-date scientific knowledge, giving you a deeper understanding of the key information you need to know for your courses, exams, and ultimately clinical practice.
- Consult this title on your favorite e-reader with intuitive search tools and adjustable font sizes. Elsevier eBooks provide instant portable access to your entire library, no matter what device you're using or where you're located.
- Visualize normal and abnormal development with hundreds of superb clinical photos and embryological drawings.
- Access the fully searchable text online, view animations, answer self-assessment questions, and much more at www.studentconsult.com.
- Grasp the molecular basis of embryology , including the processes of branching and folding - essential knowledge for determining the root of many abnormalities.
- Understand the clinical manifestations of developmental abnormalities with clinical vignettes and Clinical Correlations boxes throughout.
Table of Contents
Section Title | Page | Action | Price |
---|---|---|---|
Front cover | cover | ||
Inside front cover | ifc1 | ||
Human Embryology and Developmental Biology, 5/e | i | ||
Copyright page | iv | ||
Dedication | v | ||
Preface to the Fifth Edition | vii | ||
Table of Contents | ix | ||
Developmental Tables | xi | ||
I Early Development and the Fetal-Maternal Relationship | 1 | ||
1 Getting Ready for Pregnancy | 2 | ||
Gametogenesis | 2 | ||
Phase 1: Origin and Migration of Germ Cells | 2 | ||
Phase 2: Increase in the Number of Germ Cells by Mitosis | 2 | ||
Phase 3: Reduction in Chromosomal Number by Meiosis | 4 | ||
Stages of Meiosis | 4 | ||
Meiosis in Females | 4 | ||
Meiosis in Males | 7 | ||
Phase 4: Final Structural and Functional Maturation of Eggs and Sperm | 7 | ||
Oogenesis | 7 | ||
Spermatogenesis | 12 | ||
Abnormal Spermatozoa | 14 | ||
Preparation of the Female Reproductive Tract for Pregnancy | 15 | ||
Structure | 15 | ||
Ovaries and Uterine Tubes | 15 | ||
Uterus | 16 | ||
Vagina | 16 | ||
Hormonal Control of the Female Reproductive Cycle | 16 | ||
Hypothalamic Control | 16 | ||
Pituitary Gland (Hypophysis) | 16 | ||
Ovaries and Placenta | 16 | ||
Reproductive Target Tissues | 17 | ||
Hormonal Interactions with Tissues during Female Reproductive Cycles | 18 | ||
Hormonal Interactions Involved with Reproduction in Males | 20 | ||
Summary | 20 | ||
Review Questions | 22 | ||
References | 23 | ||
2 Transport of Gametes and Fertilization | 24 | ||
Ovulation and Egg and Sperm Transport | 24 | ||
Ovulation | 24 | ||
Egg Transport | 24 | ||
Sperm Transport | 26 | ||
Formation and Function of the Corpus Luteum of Ovulation and Pregnancy | 27 | ||
Fertilization | 28 | ||
Penetration of the Corona Radiata | 28 | ||
Attachment to and Penetration of the Zona Pellucida | 28 | ||
Binding and Fusion of Spermatozoon and Egg | 30 | ||
Prevention of Polyspermy | 31 | ||
Metabolic Activation of the Egg | 31 | ||
Decondensation of the Sperm Nucleus | 31 | ||
Completion of Meiosis and the Development of Pronuclei in the Egg | 31 | ||
What is Accomplished by Fertilization? | 32 | ||
Summary | 32 | ||
Review Questions | 35 | ||
References | 36 | ||
3 Cleavage and Implantation | 37 | ||
Cleavage | 37 | ||
Morphology | 37 | ||
Molecular, Genetic, and Developmental Control of Cleavage | 38 | ||
Parental Imprinting | 42 | ||
X-Chromosome Inactivation | 43 | ||
Developmental Properties of Cleaving Embryos | 45 | ||
Experimental Manipulations of Cleaving Embryos | 45 | ||
Stem Cells and Cloning | 47 | ||
Embryo Transport and Implantation | 50 | ||
Transport Mechanisms by the Uterine Tube | 50 | ||
Zona Pellucida | 51 | ||
Implantation into the Uterine Lining | 51 | ||
Embryo Failure and Spontaneous Abortion | 52 | ||
Summary | 52 | ||
Review Questions | 55 | ||
References | 56 | ||
4 Molecular Basis for Embryonic Development | 58 | ||
Fundamental Molecular Processes in Development | 58 | ||
Transcription Factors | 58 | ||
Homeobox-Containing Genes and Homeodomain Proteins | 58 | ||
HOX Genes | 60 | ||
Pax Genes | 61 | ||
Other Homeobox-Containing Gene Families | 62 | ||
T-Box Gene Family | 62 | ||
Helix-Loop-Helix Transcription Factors | 63 | ||
Basic Helix-Loop-Helix Proteins | 63 | ||
Forkhead Gene Family | 63 | ||
Zinc Finger Transcription Factors | 63 | ||
Sox Genes | 63 | ||
WT1 | 63 | ||
Signaling Molecules | 63 | ||
Transforming Growth Factor-β Family | 63 | ||
Fibroblast Growth Factor Family | 66 | ||
Hedgehog Family | 66 | ||
Wnt Family | 66 | ||
Other Actions of Signaling Molecules | 67 | ||
Receptor Molecules | 68 | ||
Signal Transduction | 68 | ||
Small RNAs | 70 | ||
Retinoic Acid | 71 | ||
Developmental Genes and Cancer | 72 | ||
Summary | 73 | ||
Review Questions | 73 | ||
References | 73 | ||
5 Formation of Germ Layers and Early Derivatives | 75 | ||
Two-Germ-Layer Stage | 75 | ||
Gastrulation and the Three Embryonic Germ Layers | 76 | ||
Regression of the Primitive Streak | 79 | ||
Notochord and Prechordal Plate | 80 | ||
Induction of the Nervous System | 80 | ||
Neural Induction | 80 | ||
Early Formation of the Neural Plate | 85 | ||
Cell Adhesion Molecules | 85 | ||
Summary | 89 | ||
Review Questions | 90 | ||
References | 90 | ||
6 Establishment of the Basic Embryonic Body Plan | 92 | ||
Development of the Ectodermal Germ Layer | 92 | ||
Neurulation: Formation of the Neural Tube | 92 | ||
Segmentation in the Neural Tube | 93 | ||
Morphological Manifestations of Segmentation | 93 | ||
Mechanisms of Early Segmentation of the Neural Tube | 95 | ||
Segmentation in the Hindbrain Region | 95 | ||
Formation and Segmentation of the Spinal Cord | 96 | ||
Neural Crest | 97 | ||
Sensory Placodes and Secondary Inductions in the Cranial Region | 97 | ||
Development of the Mesodermal Germ Layer | 97 | ||
Basic Plan of the Mesodermal Layer | 97 | ||
Paraxial Mesoderm | 97 | ||
Formation of Individual Somites | 99 | ||
Organization of the Somite and the Basic Segmental Body Plan | 102 | ||
Intermediate Mesoderm | 103 | ||
Lateral Plate Mesoderm | 104 | ||
Formation of the Coelom | 104 | ||
Extraembryonic Mesoderm and the Body Stalk | 104 | ||
Early Stages in the Formation of the Circulatory System | 104 | ||
Heart and Great Vessels | 104 | ||
Blood and Blood Vessels | 107 | ||
Development of the Endodermal Germ Layer | 107 | ||
Basic Structure of a 4-Week-Old Embryo | 111 | ||
Gross Appearance | 111 | ||
Circulatory System | 111 | ||
Derivatives of the Embryonic Germ Layers | 111 | ||
Summary | 112 | ||
Review Questions | 115 | ||
References | 115 | ||
7 Placenta and Extraembryonic Membranes | 117 | ||
Extraembryonic Tissues | 117 | ||
Amnion | 117 | ||
Yolk Sac | 118 | ||
Allantois | 119 | ||
Chorion and Placenta | 120 | ||
Formation of Chorionic Villi | 120 | ||
Establishing the Uteroplacental Circulation | 122 | ||
Gross Relationships of Chorionic and Decidual Tissues | 122 | ||
Formation and Structure of the Mature Placenta | 124 | ||
Structure of the Mature Placenta | 125 | ||
Umbilical Cord | 126 | ||
Placental Circulation | 126 | ||
Structure of a Mature Chorionic Villus | 126 | ||
Placental Physiology | 126 | ||
Placental Hormone Synthesis and Secretion | 129 | ||
Placental Immunology | 129 | ||
Placenta After Birth | 130 | ||
Placenta and Membranes in Multiple Pregnancies | 130 | ||
Summary | 133 | ||
Review Questions | 134 | ||
References | 135 | ||
8 Developmental Disorders: | 136 | ||
General Principles | 136 | ||
Periods of Susceptibility to Abnormal Development | 139 | ||
Patterns of Abnormal Development | 141 | ||
Causes of Malformations | 141 | ||
Genetic Factors | 141 | ||
Abnormal Chromosome Numbers | 141 | ||
Polyploidy | 141 | ||
Monosomy and Trisomy | 142 | ||
Abnormal Chromosome Structure | 143 | ||
Genetic Mutations | 143 | ||
Environmental Factors | 144 | ||
Maternal Infections | 144 | ||
Chemical Teratogens | 145 | ||
Folic Acid Antagonists | 145 | ||
Androgenic Hormones | 145 | ||
Anticonvulsants | 145 | ||
Sedatives and Tranquilizers | 145 | ||
Antineoplastic Agents | 146 | ||
Alcohol | 146 | ||
Retinoic Acid (Vitamin A) | 146 | ||
Antibiotics | 147 | ||
Other Drugs | 147 | ||
Physical Factors | 148 | ||
Ionizing Radiation | 148 | ||
Other Physical Factors | 148 | ||
Maternal Factors | 148 | ||
Mechanical Factors | 148 | ||
Developmental Disturbances Resulting in Malformations | 149 | ||
Duplications and Reversal of Asymmetry | 149 | ||
Faulty Inductive Tissue Interactions | 149 | ||
Absence of Normal Cell Death | 149 | ||
Failure of Tube Formation | 149 | ||
Disturbances in Tissue Resorption | 149 | ||
Failure of Migration | 149 | ||
Developmental Arrest | 149 | ||
Destruction of Formed Structures | 149 | ||
Failure to Fuse or Merge | 149 | ||
Hypoplasia and Hyperplasia | 149 | ||
Receptor Defects | 150 | ||
Defective Fields | 150 | ||
Effects Secondary to Other Developmental Disturbances | 150 | ||
Germ Layer Defects | 150 | ||
Summary | 151 | ||
Review Questions | 152 | ||
References | 152 | ||
II Development of the Body Systems | 155 | ||
9 Integumentary, Skeletal, and Muscular Systems | 156 | ||
Integumentary System | 156 | ||
Epidermis | 156 | ||
Structural Development | 156 | ||
Immigrant Cells in the Epidermis | 156 | ||
Epidermal Differentiation | 158 | ||
Dermis | 159 | ||
Dermal-Epidermal Interactions | 160 | ||
Epidermal Appendages | 161 | ||
Hair | 161 | ||
Nails | 163 | ||
Mammary Glands | 164 | ||
Skeleton | 165 | ||
Axial Skeleton | 168 | ||
Vertebral Column and Ribs | 168 | ||
Skull | 172 | ||
Appendicular Skeleton | 177 | ||
Muscular System | 178 | ||
Skeletal Muscle | 181 | ||
Determination and Differentiation of Skeletal Muscle | 181 | ||
Muscle Transcription Factors | 183 | ||
Histogenesis of Muscle | 185 | ||
Morphogenesis of Muscle | 186 | ||
Muscles of the Trunk and Limbs | 186 | ||
Muscles of the Head and Cervical Region | 187 | ||
Anomalies of Skeletal Muscles | 189 | ||
Cardiac Muscle | 189 | ||
Smooth Muscle | 190 | ||
Summary | 190 | ||
Review Questions | 191 | ||
References | 191 | ||
Skin | 191 | ||
Skeleton | 192 | ||
Muscle | 192 | ||
10 Limb Development | 193 | ||
Initiation of Limb Development | 193 | ||
Regulative Properties and Axial Determination | 193 | ||
Outgrowth of Limb Bud | 194 | ||
Apical Ectodermal Ridge | 195 | ||
Mesoderm of Early Limb Bud | 196 | ||
Structure and Composition | 196 | ||
Mesodermal-Ectodermal Interactions and the Role of Mesoderm in Limb Morphogenesis | 197 | ||
Zone of Polarizing Activity and Morphogenetic Signaling | 198 | ||
Morphogenetic Control of Early Limb Development | 199 | ||
Control of Proximodistal Segmentation | 199 | ||
Molecular Signals in Limb Development | 200 | ||
Cell Death and Development of Digits | 202 | ||
Development of Limb Tissues | 205 | ||
Skeleton | 205 | ||
Musculature | 206 | ||
Tendons | 208 | ||
Innervation | 208 | ||
Vasculature | 208 | ||
Summary | 211 | ||
Review Questions | 214 | ||
References | 214 | ||
11 Nervous System | 216 | ||
Establishment of the Nervous System | 216 | ||
Early Shaping of the Nervous System | 216 | ||
Histogenesis Within the Central Nervous System | 218 | ||
Proliferation Within the Neural Tube | 218 | ||
Cell Lineages in Histogenesis of the Central Nervous System | 219 | ||
Fundamental Cross-Sectional Organization of the Developing Neural Tube | 220 | ||
Craniocaudal Pattern Formation and Segmentation | 222 | ||
Patterning in the Hindbrain and Spinal Cord Regions | 222 | ||
Patterning in the Midbrain Region | 224 | ||
Patterning in the Forebrain Region | 226 | ||
Peripheral Nervous System | 226 | ||
Structural Organization of a Peripheral Nerve | 226 | ||
Patterns and Mechanisms of Neurite Outgrowth | 228 | ||
Neurite-Target Relations During Development of a Peripheral Nerve | 229 | ||
Factors Controlling Numbers and Kinds of Connections Between Neurites and End Organs in the Peripheral Nervous System | 231 | ||
Autonomic Nervous System | 231 | ||
Sympathetic Nervous System | 231 | ||
Parasympathetic Nervous System | 232 | ||
Differentiation of Autonomic Neurons | 232 | ||
Congenital Aganglionic Megacolon (Hirschsprung’s Disease) | 233 | ||
Later Structural Changes in the Central Nervous System* | 233 | ||
Histogenesis Within the Central Nervous System | 233 | ||
Spinal Cord | 235 | ||
Myelencephalon | 236 | ||
Metencephalon | 237 | ||
Mesencephalon | 238 | ||
Diencephalon | 240 | ||
Telencephalon | 240 | ||
Ventricles, Meninges, and Cerebrospinal Fluid Formation | 244 | ||
Cranial Nerves | 245 | ||
Development of Neural Function | 245 | ||
Summary | 251 | ||
Review Questions | 252 | ||
References | 252 | ||
12 Neural Crest | 254 | ||
Developmental History of the Neural Crest | 254 | ||
Origin, Induction, and Specification | 254 | ||
Epitheliomesenchymal Transformation and Emigration from the Neural Tube | 254 | ||
Neural Crest Cell Migration | 255 | ||
Differentiation of Neural Crest Cells | 255 | ||
Major Divisions of the Neural Crest | 258 | ||
Trunk Neural Crest | 258 | ||
Sympathoadrenal Lineage | 259 | ||
Sensory Lineage | 259 | ||
Melanocyte Lineage | 259 | ||
Cranial Neural Crest | 259 | ||
Circumpharyngeal Neural Crest | 263 | ||
Cardiac Crest | 263 | ||
Vagal Crest | 263 | ||
Summary | 266 | ||
Review Questions | 266 | ||
References | 267 | ||
13 Sense Organs | 269 | ||
Eye | 269 | ||
Early Events in the Establishment of the Eye | 270 | ||
Formation of the Lens | 274 | ||
Formation of the Cornea | 276 | ||
Retina and Other Derivatives of the Optic Cup | 279 | ||
Neural Retina | 279 | ||
Iris and Ciliary Body | 281 | ||
Vitreous Body and Hyaloid Artery System | 282 | ||
Choroid Coat and Sclera | 282 | ||
Eyelids and Lacrimal Glands | 283 | ||
Ear | 285 | ||
Development of the Inner Ear | 286 | ||
Development of the Middle Ear | 289 | ||
Development of the External Ear | 290 | ||
Summary | 291 | ||
Review Questions | 292 | ||
References | 292 | ||
14 Head and Neck | 294 | ||
Early Development of the Head and Neck | 294 | ||
Tissue Components and Segmentation of Early Craniofacial Region | 294 | ||
Fundamental Organization of the Pharyngeal Region | 295 | ||
Establishing the Pattern of the Craniofacial Region | 297 | ||
Cellular Migrations and Tissue Displacements in the Craniofacial Region | 298 | ||
Development of the Facial Region | 299 | ||
Formation of the Face and Jaws | 299 | ||
Temporomandibular Joint and Its Relationship with the Jaw Joint of Lower Vertebrates | 302 | ||
Formation of the Palate | 303 | ||
Formation of the Nose and Olfactory Apparatus | 305 | ||
Formation of the Salivary Glands | 307 | ||
Formation of the Teeth | 307 | ||
Patterning of the Dentition | 310 | ||
Stages of Tooth Development | 311 | ||
Tissue Interactions in Tooth Development | 311 | ||
Formation of Dentin and Enamel | 315 | ||
Tooth Eruption and Replacement | 315 | ||
Development of the Pharynx and Its Derivatives | 315 | ||
External Development of the Pharyngeal Region | 315 | ||
Pharyngeal Arches | 315 | ||
Pharyngeal Grooves | 323 | ||
Internal Development of the Stomodeal and Pharyngeal Regions | 324 | ||
Pharynx and Pharyngeal Pouches | 324 | ||
Thyroid | 324 | ||
Hypophysis | 324 | ||
Thymus and Lymphoid Organs | 326 | ||
Formation of the Tongue | 326 | ||
Summary | 331 | ||
Review Questions | 332 | ||
References | 333 | ||
15 Digestive and Respiratory Systems and Body Cavities | 335 | ||
Digestive System | 335 | ||
Patterning the Gut | 335 | ||
Formation of the Esophagus | 337 | ||
Formation of the Stomach | 340 | ||
Development of the Spleen | 343 | ||
Formation of the Intestines | 344 | ||
Partitioning of the Cloaca | 346 | ||
Histogenesis of the Intestinal Tract | 347 | ||
Formation of Enteric Ganglia | 349 | ||
Glands of the Digestive System | 352 | ||
Formation of the Liver | 352 | ||
Development of Hepatic Function | 354 | ||
Formation of the Pancreas | 355 | ||
Respiratory System | 359 | ||
Formation of the Larynx | 359 | ||
Formation of the Trachea and Bronchial Tree | 359 | ||
Stages in Lung Development | 362 | ||
Embryonic Stage (Weeks 4 to 7) | 362 | ||
Pseudoglandular Stage (Weeks 8 to 16) | 362 | ||
Canalicular Stage (Weeks 17 to 26) | 362 | ||
Terminal Sac Stage (Weeks 26 to Birth) | 362 | ||
Postnatal Stage | 362 | ||
Body Cavities | 362 | ||
Formation of the Common Coelom and Mesentery | 362 | ||
Formation of the Septum Transversum and Pleural Canals | 365 | ||
Formation of the Diaphragm | 367 | ||
Summary | 372 | ||
Review Questions | 373 | ||
References | 374 | ||
16 Urogenital System | 376 | ||
Urinary System | 376 | ||
Early Forms of the Kidney | 376 | ||
Metanephros | 377 | ||
Later Changes in Kidney Development | 381 | ||
Formation of the Urinary Bladder | 383 | ||
Genital System | 383 | ||
Genetic Determination of Gender | 389 | ||
Specification of Germ Cells, Migration into the Gonads, and Entry into Meiosis | 390 | ||
Establishment of Gonadal Gender | 391 | ||
Origin of the Gonads and Adrenal Cortex | 391 | ||
Differentiation of the Testes | 391 | ||
Differentiation of the Ovaries | 393 | ||
Sexual Duct System | 394 | ||
Indifferent Sexual Duct System | 394 | ||
Sexual Duct System of Males | 394 | ||
Sexual Duct System of Females | 396 | ||
Descent of the Gonads | 398 | ||
Descent of the Testes | 398 | ||
Descent of the Ovaries | 399 | ||
External Genitalia | 399 | ||
Indifferent Stage | 399 | ||
External Genitalia of Males | 401 | ||
External Genitalia of Females | 401 | ||
Summary | 405 | ||
Review Questions | 406 | ||
References | 406 | ||
17 Cardiovascular System | 408 | ||
Development of Blood and the Vascular System | 408 | ||
Embryonic Hematopoiesis | 408 | ||
Cellular Aspects of Hematopoiesis | 410 | ||
Erythropoiesis | 410 | ||
Hemoglobin Synthesis and its Control | 411 | ||
Formation of Embryonic Blood Vessels | 413 | ||
Development of the Arteries | 415 | ||
Aorta, Aortic Arches, and Their Derivatives | 415 | ||
Major Branches of the Aorta | 417 | ||
Arteries of the Head | 418 | ||
Coronary Arteries | 419 | ||
Development of Veins | 420 | ||
Cardinal Veins | 420 | ||
Vitelline and Umbilical Veins | 422 | ||
Pulmonary Veins | 422 | ||
Development of Lymphatic Channels | 423 | ||
Development and Partitioning of the Heart | 425 | ||
Early Development of the Heart | 425 | ||
Cellular Origins | 425 | ||
Looping of the Heart | 426 | ||
Early Atrioventricular Partitioning of the Heart | 426 | ||
Later Partitioning of the Heart | 429 | ||
Separation of the Atria from the Ventricles | 429 | ||
Partitioning of the Atria | 430 | ||
Repositioning of the Sinus Venosus and the Venous Inflow into the Right Atrium | 431 | ||
Partitioning of the Ventricles | 431 | ||
Partitioning of the Outflow Tract of the Heart | 431 | ||
Innervation of the Heart | 433 | ||
Initiation of Cardiac Function and the Conducting System of the Heart | 433 | ||
Fetal Circulation | 434 | ||
Summary | 449 | ||
Review Questions | 451 | ||
References | 451 | ||
18 Fetal Period and Birth | 453 | ||
Growth and Form of the Fetus | 453 | ||
Fetal Physiology | 453 | ||
Circulation | 453 | ||
Fetal Lungs and Respiratory System | 456 | ||
Fetal Movements and Sensations | 458 | ||
Fetal Digestive Tract | 459 | ||
Fetal Kidney Function | 461 | ||
Endocrine Function in the Fetus | 461 | ||
Parturition | 462 | ||
Adaptations to Postnatal Life | 467 | ||
Circulatory Changes at Birth | 467 | ||
Lung Breathing in the Perinatal Period | 469 | ||
Overview | 470 | ||
Summary | 471 | ||
Review Questions | 471 | ||
References | 472 | ||
Answers to Clinical Vignettes and Review Questions | 473 | ||
Chapter 1 | 473 | ||
Index | 479 | ||
A | 479 | ||
B | 480 | ||
C | 481 | ||
D | 484 | ||
E | 485 | ||
F | 487 | ||
G | 489 | ||
H | 490 | ||
I | 491 | ||
J | 492 | ||
K | 492 | ||
L | 493 | ||
M | 493 | ||
N | 496 | ||
O | 497 | ||
P | 498 | ||
Q | 501 | ||
R | 501 | ||
S | 501 | ||
T | 503 | ||
U | 505 | ||
V | 505 | ||
W | 506 | ||
X | 506 | ||
Y | 506 | ||
Z | 506 | ||
Inside back cover | ibc1 |