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This fourth volume of Contemporary Topics In Immunobiology treats in vertebrate immunity. Specifically, the results represent several approaches to humoral and cellular immunity. It is evident that invertebrates do have function ing immune systems. For example, cellular immunity is characterized by both specificity and memory, but it is still problematical whether vertebrate immune capacity evolved directly from invertebrates. Most of the manuscripts were formally presented at the International Symposium on Invertebrate Pathology, University of Minnesota, August 1972, held in connection with the 25th anniversary celebration of the American Insti tute of Biological Sciences. I wish to express my appreciation to the contributors and to beg their indulgence in what may have been overzealous editing. This was done, though, in the interest of clarity and to seek uniformity. Because of earlier problems, time limitations did not permit consultations between submission of manuscripts and final editing. For assistance, I extend a special note of gratitude to Mrs. Lois Gehringer who unselfishly retyped many of the manuscripts. The preparation of this volume was aided partially by NSF Grant GB17767, two grants from The California Institute for Cancer Research, and a grant from The Brown-Hazen Corporation. E.L.C. Contents Introduction: General Comments and a Note on Taxonomy ....................... .

Contemporary Topics in Immunobiology: Volume 4 Invertebrate Immunology

Introduction: General Comments and a Note on Taxonomy.- 1. Invertebrate Precursors to Immune Responses.- Self and Not-Self.- "Self-Recognition" in Colonial Marine Forms.- Sponges.- Colonial Ascidians.- General Aspects.- A Note on Self-Incompatibility in Plants.- Coelomate Invertebrates.- Hemagglutinins and Opsonins.- Hemocytes.- Analogies with Vertebrate Immune Responses.- References.- 2. An Electron Microscope Study of the Fate of Bacteria Phagocytized by Granulocytes of Crassostrea virginica.- The Present Problem.- Discussion and Conclusions.- Summary.- Acknowledgment.- References.- 3. On the Immediate Fate of Bacteria in the Land Snail Hel.- Clearance Mechanisms.- Discussion and Conclusions.- Acknowledgments.- References.- 4. Metabolism of Insect Hemocytes during Phagocytosis.- The Present Problem.- Human Phagocytosis.- Phagocytosis by Cockroach Hemocytes.- Summary of the Metabolic Events in Phagocytosis.- Acknowledgments.- References.- 5. The Coelomocytes of Nereid Polychaetes.- Fine Structure of Coelomocytes.- Discussion.- Conclusions.- Acknowledgments.- References.- 6. On the Ultrastructure of Invertebrate Hemocytes: An Interpretation of Their Role in Comparative Hematology.- The Blood.- Molluscan Hemocytes.- Annelid Hemocytes.- Sipunculid Hemocytes.- Arthropod Hemocytes.- Hemocytes of Echinoderms and Urochordates.- Conclusions.- References.- 7. The Octopus White Body: An Ultrastructural Survey.- The Leukopoietic Organs of Cephalopods.- Hemocytoblasts.- Hemocytoblasts to Leukoblasts.- Leukoblasts.- Secondary Leukoblasts to Mature Leukocytes.- Final Comment.- References.- 8. Earthworm Coelomocyte Immunity.- Historical Observations.- The Present Problem.- The Coelomocytes.- Coelomic Cavity.- Phagocytosis.- Interrelationships of Coelomocytes.- Coelomic Cell Responses to Tissue Grafts.- Evolution of Delayed Hypersensitivity and the Anamnestic Response.- Final Comment.- References.- 9. An Approach to Studying Evolution of Cellular Immunity.- Cellular Immunity in an Invertebrate.- Activity of5 1 Cr Labeled Earthworm Coelomocytes.- Behavior of Labeled Normal Coelomocytes in Normal Hosts.- Behavior of Labeled Normal Coelomocytes in Wounded Hosts.- Behavior of Normal and Sensitized Coelomocytes in Grafted Hosts.- Summary of Labeled Coelomocyte Studies.- Discussion.- References.- 10. Cellular Aspects of Graft Rejection in Earthworms and Some Other Metazoa.- Graft Rejection in Lumbricids.- Self-Destruction of Grafts.- Leukocyte Activity.- Second-Set Anamnestic Response.- Comparison between the Graft Reaction in Lumbricids and Some Other Metazoa.- Induced Self-Destruction of the Graft.- Cellular Immune Reactions.- Phylogenetic Implications.- References.- 11. Graft Rejection and the Regulation of Length in Hydra viridis.- Position of Coelenterates in Invertebrate Immunology Research.- Separation.- Intraspecific Grafts.- Multiply Grafted Animals.- Doubling the Gastric Region (2g) or Peduncle (2p).- Higher Degrees of Multiplicity.- The Diffusion Gradient.- Concluding Remarks.- References.- 12. Tissue Transplantation in Diverse Marine Invertebrates.- Corals.- Pearl Oysters (Pinctada margaritifera).- Echinoderms.- Discussion and Conclusions.- Acknowledgments.- References.- 13. Specific Receptors in Relation to the Evolution of Immunity.- Specific Uptake of Protein by Sea Urchin Coelomocytes.- Selective Uptake of Amino Acids by Amoebae.- Evolutionary Implications.- Acknowledgments.- References.- 14. Recognition of Foreignness in the Fresh-Water Crayfish, Parachaeraps bicarinatus.- Preparation of Hemocyte Monolayers.- Recognition of Erythrocytes by Crayfish Hemocytes.- Rate of Elimination of Bacteria from the Circulation of the Crayfish.- Factors Involved in the Removal of Bacteria from the Circulation of the Crayfish.- In Vitro Studies on the Phagocytosis of Bacteria by Crayfish Hemocytes.- References.- 15. Insect Immunity to Parasitic Nematodes.- Simple Encapsulation.- Melanotic Encapsulation.- Humoral Melanization.- Intracellular Melanization.- Tissue Responses.- Conclusion.- Summary.- References.- 16. Control Mechanisms of Leafhopper Endosymbiosis.- Lysozyme Content.- The Process of Endosymbiosis.- Conclusions.- References.- 17. Cellular and Noncellular Recognition of and Reactions to Fungi in Crayfish.- Reactions in the Hemolymph.- Reactions in Inner Organs.- Reactions in the Integument.- Final Remarks.- References.- 18. Insect Hemocytes and the Problem of Host Recognition of Foreignness.- Hemocytes and the Problem of Recognition of Foreignness.- Possible Mechanism of Hemocyte Activation.- Concluding Statement.- References.- 19. Lysozymelike Activities in the Hemolymph of Crassostrea virginica.- Lysozyme Characteristics.- Analytical Disc Electrophoresis.- Preparative Disc Electrophoresis.- Ion Exchange Chromatography.- Final Comment.- References.- 20. Induction of Internal Defense Mechanisms in the Lobster, Homarus americanus.- Disease and Immunity in Crustaceans.- Final Remarks.- References.- 21. Comparison of a Natural Agglutinin in the Hemolymph of the Blue Crab, Callinectes sapidus, with Agglutinins of Other Invertebrates.- Discussion.- General.- Agglutinin Titers.- Physical and Chemical Properties.- Molecular Weight Studies.- Specificity and Function.- Summary.- References.- 22. Characteristics of the Agglutinin in the Scorpion, Androctonus australis.- Elimination of SRBC by Scorpions and Increase in Hemocytes.- Increase in Hemagglutinins after Immunization.- Specificity of Naturally Occurring Agglutinins.- Effect of Temperature and Dialysis.- Immunoelectrophoresis and Paper Electrophoresis of Hemolymph.- The Cellular Machinery.- Summary.- Acknowledgments.- References.- 23. Hemagglutinins: Primitive Receptor Molecules Operative in Invertebrate Defense Mechanisms.- Biological Studies.- Physical and Chemical Studies.- Comments and Conclusions.- Acknowledgments.- References.- 24. Tumors in Drosophila and Antibacterial Immunity.- Tumors in Drosophila.- Bacterial Immunity in Strains of Drosophila with and without Hereditary Melanotic Tumors.- Acknowledgments.- References.- 25. A Final Comment on Invertebrate Immunity.