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A MARYLAND SEA GRANT BOOK

The Eastern Oyster:

Crassostrea virginica

Victor S. Kennedy, Roger I.E. Newell and Albert F. Eble, Editors

Book, 772 pages, hard cover, 8-1/2" x 11", 400 photographs, drawings and diagrams.

UM-SG-TS-96-01. ISBN 0-943-676-61-4.

Second printing, a high-quality on-demand reproduction of the original casebound edition. Allow one month for delivery.

Cost: New, lower price, $75.00 plus $4.50 state tax (Maryland residents only) and shipping and handling (rates)

Acknowledgments
Foreword
Authors

Chapter 1: A Catalogue of Selected Species of Living Oysters (Ostreacea) of the World
Melbourne R. Carriker and Patrick M. Gaffney

Introduction, p. 1
Catalogue, p. 3
References, p. 15

Chapter 2: General Anatomy
Albert F. Eble and Robert Scro

Introduction, p. 19
The Shell, p. 19
The Mantle, p. 19: Anatomy, p. 19; Histology, p. 21
Adductor Muscle, p. 23: Anatomy, p. 23; Histology, p. 24
Heart and Circulation, p. 25
Gills, p. 25: Anatomy, p. 25; Histology and Fine Structure, p. 27
Labial Palps, p. 37: Anatomy, p. 37; Histology, p. 38
Digestive System, p. 39: Mouth, p. 39; Esophagus, p. 41; Stomach, p. 41; Style Sac-Midgut, p. 44; Intestine -- Ascending, Median, and Descending Portions, p. 47; Rectum, p. 49; Digestive Gland, p. 49
Excretory System, p. 52: Renopericardial Canal, p. 54; Kidney, p. 55
Reproductive System, p. 59: Anatomy, p. 59; Histology p. 59; Oocytes, p. 66
Larval Stages, p. 66: Development, p. 66; Anatomy and Histology, p. 66; Visceral Cavity, p. 67; Velum, p. 68; Mantle, p. 68; Foot, p. 68; Digestive System, p. 68
References, p. 71

Chapter 3: The Shell and Ligament
Melbourne R. Carriker


Introduction, p. 75
Larval Shell, p. 75: Terminology, p. 75; Dimensions, p. 77; Identification, p. 78; Stages, p. 93; Mantle, p. 93; Attachment, p. 96; Shell Microstructure, p. 97
Adult Shell, p. 100: Shell Macroform, p. 103; Ligament, p. 110; Periostracum, p. 116; Valves, p. 119; Evolution of Mechanical Properties, p. 154; Environmental Microstructural Effects, p. 154; Genetic Modification, p. 155; Breakage and Regeneration, p. 156; Shell Anomalies, p. 156
In Prospect, p. 157
References, p. 159

Chapter 4: Adductor and Mantle Musculature
Carol M. Morrison

Structure and Function of the Adductor Muscle, p. 169: Translucent Adductor Muscle, p. 170; Opaque Adductor Muscle, p. 175
Musculature of Mantle and Mantle Lobes, p. 178
References, p. 181

Chapter 5: Mechanisms and Physiology of Larval and Adult Feeding
Roger I.E. Newell and Christopher J. Langdon

Introduction, p. 185
Larval Oysters, p. 185: Development and Structure of the Velum, p. 185; Function of Velar Ciliary Bands, p. 188; Mechanisms of Particle Capture, p. 189; Food Ingestion and Selection, p. 190; Effects of Food Size and Concentration on Larval Feeding, p. 192; Change in Feeding Associated with Metamorphosis, p. 194
Adult Oysters, p. 194: Gill Anatomy, p. 195; Ciliation of the Gill Filaments, p. 199; Mechanisms of Particle Capture, p. 202; Ciliated Tracts on the Gill Surface, p. 204; Particle Retention on the Gill, p. 204; Post-Capture Sorting of Particles, p. 208; Pseudofeces Production, p. 216; Measurement of Clearance Rates, p. 217; Endogenous Regulation of Ventilation and Clearance Rates, p. 218; Exogenous Factors Affecting Feeding Activity, p. 219
Epilogue, p. 222
References, p. 223

Chapter 6: Digestion and Nutrition in Larvae and Adults
Christopher J. Langdon and Roger I.E. Newell

Introduction, p. 231
Adult Oysters, p. 231: Stomach, p. 231; Crystalline Style and Style Sac, p. 233; Digestive Diverticula, p. 238; Mid-Gut and Rectum, p. 243; Hemocytes, p. 244; Gut Microflora, p. 245; Digestion and Absorption Efficiencies, p. 246
Larval Oysters, p. 247
Dissolved Organic Matter as a Source of Nutrition, p. 249
Natural Particulate Sources of Nutrition, p. 252
Artificial Diets, p. 258
References, p. 260

Chapter 7: The Circulatory System
Albert F. Eble

Introduction, p. 271
Hemolymph Vessels -- Nomenclature and General Structure, p. 271
Pericardium, Heart, and Aortae, p. 273: Pericardial Coelom, p. 273; Systemic Heart, p. 273; Physiology, p. 277; The Aortae, p. 281
Circulatory System, p. 282: Visceral Mass, p. 282; The Gills, p. 284; The Adductor Muscle, p. 288; The Kidney, p. 290; Accessory Hearts, p. 292; The Pallial Circulation, p. 293; Venous Return to the Heart, p. 294
Summary, p.296
References, p. 296

Chapter 8: Hemocytes: Forms and Functions
Thomas C. Cheng

Introduction, p. 299
Formed Hemolymph Elements, p. 299
Origin and Development of Hemocytes, p. 305
Subcategories of Granulocytes, p. 308
Hemocyte Behavior, p. 310
Number of Hemocytes, p. 312
Functions of Hemocytes, p. 312
Hemocytes and Humoral Factors, p. 323
References, p. 326

Chapter 9: Reproductive Processes and Early Development
Raymond J. Thompson, Roger I.E. Newell,
Victor S. Kennedy, and Roger Mann

Introduction, p. 335
Sex Ratios and Changes with Size, p. 335
Factors Influencing Gametogenesis, p. 338: Endogenous and Exogenous Control of Reproduction, p. 338
Biochemical Energy Storage Cycles, p. 342: Glycogen, p. 343; Intraspecific Variation, p. 348; Glycogen Concentrations in Triploid Oysters, p. 349; Effects of Environmental Stress Factors, p. 350
Fecundity and Reproductive Effort, p. 351
Factors Influencing Spawning, p. 357
Success of Reproduction, p. 358: Fertilization Success, p. 358
Egg and Larval Development, p. 361
Future Research Directions, p. 363
References, p. 364

Chapter 10: Biology of Larvae and Spat
Victor S. Kennedy

Introduction, p. 371
The Estuarine Habitat, p. 373: Classes of Estuarine Structure, p. 373; Physical Factors Influencing Larvae, p. 375
Larval Movement, p. 379: Larval Swimming, p. 379
Larval Dispersal, p. 382: Tracking Methods, p. 382; Studies of Oyster Larval Dispersal, p. 383; Overview of Dispersion Studies, p. 392; Patchiness of Larval Distributions, p. 393
Experimental Laboratory Studies on Settlement, p. 394: Settlement, p. 394; Cementation, p. 398; Metamorphic Changes, p. 399
Settlement and Recruitment in Nature, p. 399: Settlement and Recruitment Variability, p. 399
Larval and Spat Survival and Growth, p. 403: Food and Feeding, p. 405; Depletion of Biochemical Reserves, p. 406; Predation, Competition, and Recruitment, p. 406; Disease, p. 407; Larval and Spat Growth, p. 407
Future Research Directions, p. 410
References, p. 411

Chapter 11: Biochemical and Population Genetics
Patrick M. Gaffney

Introduction, p. 423
Biochemical Genetics, p. 423: Enzyme-coding Loci, p. 423; Non-enzyme Proteins, p. 425; Effects of Biochemical Genetic Variation on Physiology, p. 425; Correlates of Heterozygosity, p. 426
Population Genetics, p. 431: Geographic Variation, p. 431; Conclusions Regarding Population Structure, p. 436
Summary, p. 437
References, p. 437

Chapter 12: Chromosomes, Biology, and Breeding
A. Crosby Longwell and S.S. Stiles

Introduction, p. 443
Meiosis, p. 443: Introductory Review, p. 443; Normal Meiosis, p. 444; Fertilization, Abnormal Meiosis, and Cleavage, p. 446
Chromosome Number and Morphology, p. 448: Chromosome Number, p. 448; Chromosome Morphology, p. 449
DNA Content, p. 451
Intraspecific Chromosome Polymorphism, p. 452
Polyploidy and Chromosome Engineering, p. 453: Polyploidy in Nature, p. 453; Absence of Natural Polyploidy, p. 454; Induced Triploids, p. 454; Triploid Sterility, p. 456; Tetraploidy, p. 456; Parthenogenesis, p. 457
Conclusions, p. 458
References, p. 459

Chapter 13: Natural Environmental Factors
Sandra E. Shumway

Introduction, p. 467
Distribution, Tolerance, and Survival, p. 468: Larval Distribution, p. 470
Reproduction Patterns, p. 474
Spawning, p. 478
Larval Settlement, p. 480: Development and Larval Growth, p. 480; Adult Growth, p. 484; Activity, p. 486
Heart Rate, p. 488
Respiration, p. 489
Osmotic Regulation, p. 491
Genetic Adaptations to Environmental Stressors, p. 496
Parasites, Predators, and Diseases, p. 497
Light, pH, and Turbidity, p. 500
Summary, p. 503
References, p. 503

Chapter 14: Environmental Factors: Response to Metals
G. Roesijadi

Introduction, p. 515
Variability in Metal Bioaccumulation and Implications for Biomonitoring, p. 516
Metal Turnover in Eastern Oysters, p. 520
Cellular Mechanisms of Metal Sequestration, p. 521: Granular Deposits, p. 521; Metallothioneins and Other Metal-binding Proteins, p. 521
Toxic Effects of Metals on Oysters, p. 528
Tributyltin and Oysters: An Ongoing Case History, p. 530
Public Health Implications of Metal Bioaccumulation in Oysters, p. 531
Summary, p. 532
References, p. 532

Chapter 15: The Bioaccumulation and Biological Effects of Lipophilic Organic Contaminants
Judith McDowell Capuzzo

Introduction, p. 539
Bioaccumulation, p. 540: General Considerations, p. 540; Petroleum Hydrocarbons, p. 540; Polychlorinated Biphenyls and Other Chlorinated Compounds, p. 543; Organotin Compounds, p. 545
Biological Effects, p. 546: General Considerations, p. 546; Biotransformation and Disease Responses, p. 547; Effects of Petroleum Hydrocarbons and PAHs, p. 549; Effects of Organotin Compounds, p. 550; Exposure to Multiple Contaminants, p. 551
Summary, p. 552
References, p. 553

Chapter 16: Predators, Pests and Competitors
Marie E. White and Elizabeth A. Wilson

Introduction, p. 559
Predators, p. 559: Predatory Gastropods, p. 559; Predation by Crabs, p. 563; Polyclad Flatworms, p. 565; Starfish, p. 567; Fish, p. 567; Predation on Oyster Larvae, p. 568
Pests, p. 568: Cliona spp, p. 568; Polydora spp. and Boccardia spp, p. 569; Pea Crabs, p. 570; Boonea impressa, p. 571; Clams and Mussels, p. 572
Competitors, p. 572
Summary, p. 574
References, p. 575

Chapter 17: Diseases and Defense Mechanisms
Susan E. Ford and M.R. Tripp

Introduction, p. 581
Principal Infectious Diseases, p. 583: Overview, p. 583; Larval Vibriosis, p. 585; Juvenile Oyster Disease, p. 587; Dermo Disease, p. 592; MSX Disease, p. 612; SSO Disease, p. 626
Defense Mechanisms, p. 632: Hemocytes and Their Capabilities, p. 632; Inflammation, p. 632; Response to Mechanical Injury, p. 633; Response to Chemical Injury, p. 633; Response to Foreign Proteins, p. 634; Response to Transplanted Tissue, p. 634; Responses to Various Biological Agents, p. 634; Response to Tumors, p. 638; Humoral Factors, p. 639
References, p. 642

Chapter 18: Culture: Genetic Improvement
Gary Newkirk

Introduction, p. 661
Breeding Programs, p. 662
Approaches to Breeding, p. 664: Inbreeding, p. 664; Alternative Selection Approaches, p. 665; Experimental Selection of Bivalves, p. 667; Hybridization, p. 668; Applied Chromosome Manipulation, p. 669; Biotechnology, p. 670; Summary, p. 670
References, p. 671

Chapter 19: Culture: Application
Michael Castagna, Mary C. Gibbons and Kenneth Kurkowski

Introduction, p. 675
History of Hatchery Development, p. 676
Culture Methods, p. 676: Seawater System, p. 677; Intakes, Pumps, and Lines, p. 677; Filters and Water Sterilization, p. 678; Culture of Unicellular Algae, p. 679
Brood Stock Conditioning and Holding, p. 680: Spawning or Stripping Gametes, p. 681; Rearing Larvae, p. 682
Treatment of Disease, p. 683
Setting and Remote Setting, p. 684
Nursery, p. 685
Competitors, Predators, and Parasites, p. 686
Grow Out, p. 686
References, p. 687

Chapter 20: Transfers and World-wide Introductions
James T. Carlton and Roger Mann

Introductions to Europe, p. 691: France: 1860s to 1870s, p. 691; England: 1870s and Later, p. 691; The "European Export Trade," p. 693; Recent Trends, p. 694
Introductions to the Pacific Coast of North
America, p. 694: British Columbia, p. 694; Washington, p. 695; Oregon, p. 695; California, p. 696; Baja California, p. 697; Establishment of Reproducing Populations on the Pacific Coast, p. 697
Introductions to Hawaii, p. 697
Transfers of Eastern Oysters on the North
American East Coast, p. 697: Phase I: The "Southern Trade," p. 697; Phase II: The "South Atlantic Trade" and the "Seaside Trade," p. 698; Phase III: The "Gulf Trade," p. 698
Failures and Successes of Introduced Populations of Crassostrea virginica, p. 700
Population Genetics and Movements of Crassostrea virginica, p. 700: Mixing of Populations, p. 700; Genetic Analysis of Long-Isolated Populations, p. 700; Accidental Transport of Associated Organisms, p. 701
Epilogue, p. 702
References, p. 702

Chapter 21: Management of Natural Populations
Clyde L. MacKenzie, Jr.

Introduction, p. 707
Elements of Oyster Management, p. 708: Adequate Size of Spawning Stock, p. 708; A Suitable Aquatic Environment for Larval Development, p. 710; A Suitable Substrate Environment for Larval Setting, p. 710; A Suitable Environment for Spat and Seed, p. 711; Growing Seed to Market Size, p. 713
Recent Successes in Increasing Oyster Production, p. 713: Prince Edward Island, p. 713; Long Island Sound, p. 716; Chesapeake Bay, Maryland and Virginia, p. 716; Appalachicola Bay, Florida, p. 716; Louisiana, p. 717; Texas, p. 717
Developing a Management Program, p. 717: Techniques for Examining Beds, p. 718; Environmental Concerns about Cultivating Oyster Beds, p. 718; Formulating a Program, p. 718
References, p. 719