Science Serving Maryland's Coasts



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.

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The Eastern Oyster is the most comprehensive synthesis about the biology of Crassotrea virginica since Paul Galtsoff published his landmark work, The American Oyster, in 1964. The Eastern Oyster extends Galtsoff's findings and adds considerably to details on such topics as anatomy, shell structure, and predators and pests. Its major thrust, however, is holistic, reflecting current approaches to biological processes and covering topics that could not have been studied in such detail thirty years ago. Thus, there are intensive treatments of recent work in the physiology of feeding, population genetics, reproduction and larval biology, pollution and environmental effects, and diseases and mechanisms of defense. Concluding with discussions of the eastern oyster as a valuable ecological and commercial resource, several chapters address public management issues, aquaculture, and the effects of transferring oysters among ecosystems.

Check out the selected contents from Chapter 2 and the table of contents below.

Selected Chapter Summaries and Illustrations

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

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[oyster anatomy]
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Albert F. Eble and Robert Scro synthesize the scientific literature on the eastern oyster's general anatomy (Chapter 2), covering structures such as the mantle, gills, and labial palps as well as the functioning of the digestive, excretory, and reproductive systems. Eble draws extensively on his own unpublished studies. Subsequent chapters cover specific features of oyster anatomy in more detail, for example, Carol Morrison summarizes details on the structure and function of the adductor and mantle musculature (Chapter 4), and Albert Eble surveys the research (Chapter 7) on the heart and circulatory system.

Shell Structure

[oyster anatomy]
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Melbourne Carriker's comprehensive synthesis of the shell structure of the larval, juvenile, and adult eastern oyster begins with Galtsoff's findings in The American Oyster ; he then covers the last three decades of scientific literature on the ontogeny of form, structure, microstructure, biomineralogy, biochemistry, and physiology of shell and shell. The chapter includes 153 micrographs that derive from the author's many years of malacological research.

Diseases and Mechanisms of Defense

[sketch of oyster diseases]
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Few fields in oyster biology have progressed as rapidly in the last decade as studies in oyster diseases and disease mechanisms. Since the early twentieth century, when an outbreak of an unidentified disease led to massive mortalities of eastern oysters in Prince Edward Island, Canada, disease has been a major concern for the oyster industry and resource management. Since the late 1950s, diseases have been responsible for devastating oyster mortalities , particularly in Delaware Bay and Chesapeake Bay. Susan Ford and M.R. Tripp (Chapter 17) describe the major infectious diseases of the eastern oysters — Juvenile Oyster Disease and Dermo, MSX, and SSO — and the oyster's responses to them. They review the subject of defense mechanisms by describing interactions between cellular and humoral elements in oysters and a variety of biotic and abiotic agents.

The oyster's primary means of defense are hemocytes that are found in hemolymph, the oyster's circulatory fluid. Thomas Cheng (Chapter 8) surveys our understanding of hemocytic processes.

Table of Contents


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

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The Eastern Oyster: Crassostrea virginica.
Kennedy, Victor S., Roger I.E. Newell, and Albert F. Eble, eds.. 1996. UM-SG-TS-96-01. 75.00. 772 pp.

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