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Invertebrate Immunity

ADVAVV NCES IN EXPERIMENTATT L MEDICINE AND BIOLOGY

Editorial Board:

NATHANAA BACK, State University of New York at BuffaloIRUNUU R. COHEN, The Weizmann Institute of ScienceABEL LAJTHA, N.S. Kline Institute for Psychiatric ResearchJOHN D. LAMBRIS, University of PennsylvaniaRODOLFO PAOLETTI, University of Milan

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Volume 703INFLAMMATION AND RETINAL DISEASE John D. Lambris and Anthony P. Adamis

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Volume 707HORMONAL AND GENETIC BASIS OF SEXUAL DIFFERENTIATION DISORDERS

AND HOT TOPICS IN ENDOCRINOLOGY Maria I. New and Joe Leigh Simpson

Volume 708INVERTEBRATE IMMUNITY Kenneth Söderhäll

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Invertebrate ImmunityEdited by

Kenneth Söderhäll

Department of Comparative Physiology, Uppsala University, Uppsala, Sweden

Springer Science+Business Media, LLC

Landes Bioscience

Invertebrate immunity / edited by Kenneth Söderhäll. p. ; cm. -- (Advances in experimental medicine and biology ; v. 708) Includes bibliographical references and index. ISBN 978-1-4419-8058-8 1. Invertebrates--Immunology. I. Söderhäll, Kenneth. II. Series: Advances in experimental medicine and biology ; v. 708. 0065-2598 [DNLM: 1. Invertebrates--immunology. W1 AD559 v.708 2010 / QL 362.85]L QL362.85.I58 2010 571.9’612--dc22 2010042470

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Invertebrate Immunity, edited by Kenneth Söderhäll. Landes Bioscience / Springer Science+Business Media, LLC dual imprint / Springer series: Advances in Experimental Medicine and Biology.

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v

DEDICATAA ION

This volume is dedicated to all my present and former PhD students, post-docs and fellow scientists who worked in my laboratory and who have contributed toour research on invertebrate immunity.

vii

PREFAFF CE

The total number of animal species range up to around 1,000,000 species, of which invertebrates constitute more than 95%, including a vast diversity of organisms from unicellular protozoans to the much more complex echinoderms and protochordates.Insects are by far the largest group of animals within the invertebrates and have received a large amount of research interest due to their importance as vectors for human and animal diseases, such as malaria, and the serious harm done by insects to crops and food.

Thus a long-standing research interest in diseases, pathogens and immune responses of insects there has been. The finding by the late Professor Hans G. Boman and colleaguesthat Drosophila responded to a challenge with dead or live bacteria by the synthesis of antibacterial peptides initiated an intense research interest on the mechanism of thisinduction. In 1996 Jules Hoffmann, Bruno Lemaitre and colleagues published a paper in which they for the first time showed that Toll was involved in the production of the TTantifungal peptide drosomycin. Subsequently, this led to the discovery of TollTT -like receptors (TLRs) in vertebrates and their importance in immune responses, particularly in mammals. Research on the role of TLRs in immune responses is very intense and theimportance of these receptors is probably greater in mammals than in most invertebrates.The finding that Toll was involved in induction of an antimicrobial peptide opened the TTpossibility of performing detailed genetic studies of the signaling pathways involved in theproduction of antimicrobial peptides. The completion of the Drosophila genome in 2000 made it possible to carry out powerful molecular genetic analysis of the immune system of this insect. For the past 10 years several genomes of insects and other invertebrates havebeen sequenced, making comparisons between invertebrates possible, not only betweenDrosophila and mammals. Several chapters in this book deal with immune responses in different groups of insects. It is evident that the immune responses are very similar between insects, but that there are also differences (Chapters 8-12). Interesting research is now being performed on mosquitoes because they are vectors for many human diseases. One important aspect, for example, is how malaria avoids any immune responses whileentering into and being inside the vector mosquitoes (Chapter 12). Interesting studies are now carried out on developmental biology and innate immunity in Hydra (Chapter 1), and there is now interest in studying immune reactions in other invertebrate groups,

viii PREFAFF CE

especially in the model organism Caenorhabiditis elegans (Chapter 6), but also in leechesand earthworms (Chapters 4 and 5).

For the past 10 years research has increasingly been focused on crustaceans (shrimp,crabs and crayfish) (see Chapter 13) and mollusks (scallop, oysters and mussels) (see Chapters 2-3), mainly because of their importance as farmed species for consumption. In these aquatic animals antibacterial peptides have a greater variation and each peptide appears to be produced in several isoforms induced by different bacterial species,indicating that there seems to be some sort of specific response to different bacterial species. The ways in which hemocytes are synthetized have also been studied in detailin crayfish, and the finding of a family of astakines similar to prokineticins, which are involved in hematopoiesis, show that these animals may be well suited for studies on hemocytes and their synthesis (Chapter 13).

Recent research has shown that there are great differences in immune responses between different invertebrate groups, but of course also that there are many similarities. Toll receptors are present in some invertebrate groups but so far the importance of their TTfunction has been mainly studied in insects, while sea urchins, which have more than200 Toll receptors still await further studies in toll receptor function (TT Chapter 14). The significance of another group of molecules, the so-called Dscams, which seem to berestricted to insects and crustaceans and are also present in vertebrates, is difficult to define. Their exact role in immunity is unkown at this moment.

To further emphasize the diversity between different invertebrate animal groups, TTthe clotting reaction can serve as a good example and is described in two chapters (Chapters 7 and 13). In crustaceans it comprises a transglutaminase and a clotting protein, whereas in horse-shoe crabs clotting it is induced by microbial polysaccharides(LPS and beta-1,3-glucans) so that a proteolytic cascade is activated and terminates withcoagulogen being cleaved by the proclotting enzyme to form the clot (Chapter 7). There is no similarity between the clotting proteins of crustaceans and horse-shoe crabs, theprocess is totally different. The only similarity is that transglutaminase is involved in both groups of animals.

The melanization reaction is an important innate immune response and which is present in most invertebrates. It was first noted by Söderhäll and Unestam in 1977that this process, i.e., activation of prophenolocxidase, was induced in the presence of beta-1,3-glucans. Subsequently, this was shown in insects and several other invertebrates.Recently Lee and colleagues managed to show that the prophenoloxidase activatingsystem and the induction of the Toll pathway in an insect share the same proteolyticTTcascade (Chapter 9). Interestingly, melanization is a highly conserved immune response which is present in nearly all vertebrates where it provides protection against UV-VV light and other stressors. Melanization in vertebrates is catalysed by tyrosinase, which has no homology with prophenoloxidase except for the copper binding sites. However, the reactions tyrosinase and phenoloxidase catalyze are exactly the same, converting phenolsto quinones and subsequently melanin. Melanization is responsible for skin color in humans, and in other animals, and it is also an important component in the brain and eyes.

Another innate immune system which is present in both vertebrates and invertebratesis the complement system, and to date a complement-like system seems to be operable in echinoderms (Chapter 14), horseshoe crabs (Chapter 7) and tunicates (Chapter 15). In insects the so-called thiolester-containing proteins are proposed to be complement-like

ixPREFAFF CE

and have been shown to function as opsonic proteins in mosquitoes (Chapter 12). It remains to be shown whether true complement proteins are present in other protostome groups and not only in horse-shoe crabs.

A growing interest in invertebrate immunity is the study of the link betweenAphysiology and immunity. Surely diet, ageing, reproduction, reproductive behavior, time of day, and use of pathogens are all likely to have an effect on immunity and immune studies. The time of day for immune and challenge studies are important, and both in Drosophila and crayfish it has been shown that the immune system varies in efficiencyduring a day. This means that scientists should be aware of these facts when planning or making experiments.

In this book I have gathered scientists who are working with different invertebrates, and it can be seen that the insects are the still attracting most research and researchers. However, an increasing interest is emerging to study new invertebrate groups, especially those where the genome is known, as seen in Chapters 1, 5, 14 and 15. Even though Drosophila has been and still is an excellent model for immune studies, it is now clear that there are great differences between immune responses in Drosophila and that of several other invertebrates, which indeed calls for more research on other invertebrates.

Kenneth Söderhäll Department of Comparative Physiology, Uppsala University

Uppsala, Sweden

xi

ABOUT THE EDITOR...

KENNETH SÖDERHÄLL studied at L Uppsala University and obtained his MSc degree in 1972 and PhD in 1978. He was promoted to associate professor (Docent) in 1980. After receiveing his PhD he worked in many different laboratories as a post-doc or as a guest scientist at the University of Montpellier/St.Christol, the Medical CellBiology department at TromsöTT University, the Millport Marine Biological Station in Scotland, University College of Wales in Swansea, the Marine Biological Laboratory at Woods Hole and John Hopkins University. In 1986 he held a Royal Society Fellowship at Swansea. In collaboration with Valerie J. Smith, he developed a new method to isolate and separate blood cells from invertebrates,which was based on using an anticoagulant with a low pH and EDTA, a method now used to isolate most invertebrate blood cells. When he returned to Uppsala he took a position as a researcher at the Swedish Science Research Council in 1987, obtaining a chair and appointment to professor and head of department at Uppsala University in 1989. He continued with research mainly on the proPO-system in arthropods. His group was first to clone proPO from an invertebrate and he has worked with this so-called melanization reaction for several years. His research team has also deciphered the clotting reaction in crustaceans and shown that it consists of a clotting protein present in plasma and a clotting enzyme, a transglutaminase present in the blood cells. This clotting system is distinct from that of a horseshoe crabs’ in which a

proteinase cascade demonstrates the great diversity in immune processes in invertebrates. He has published 230 original papers and 65 reviews in journals and books, and he has edited 4 books. He is presently on the editorial board on Fish and Shellfish Immunology, and is the editor of Journal of Experimental Biology. Dr. Söderhäll previously served onJournal Experimental Zoology, Animal Biology, Journal of Invertebrate Pathology and Developmental and Comparative Immunology. Since 2000 he has coedited the journalDevelopmental and Comparative Immunology and beginning in 2009 he is editor-in-chief for this journal. He was president for the International Society of Developmental and Comparative Immunology from 2006-2009. Currently he heads the Department of Comparative Physiology at Uppsala University (http://www.fu.uu.se/jamfys/is.html).

xiii

PARTICIPAPP NTS

René Augustin Zoological InstituteChristian-Albrechts-UniversityKielGermany

Martin BilejInstitute of MicrobiologyAcademy of Sciences

of the Czech RepublicPragueCzech Republic

Thomas C.G. BoschZoological InstituteChristian-Albrechts-UniversityKielGermany

Katherine M. BuckleyDepartment of Medical

Biophysics and Department of Immunology

Sunnybrook Health Sciences CentreUniversity of TorontoTTToronto,TT OntarioCanada

Veronika BurešováInstitute of ParasitologyBiology CentreAcademy of Sciences

of the Czech RepublicUniversity of South Bohemia��!"��

y

Czech Republic

Lage CereniusDepartment of Comparative

PhysiologyUppsala UniversityUppsalaSweden

Jun-Ho ChaiNational Research Laboratory

of Defense ProteinsCollege of PharmacyPusan National UniversityBusanKorea

Lori A. Clow#$�&��'��*��Alexandria,VirginiaUSA

Sirlei DaffreDepartamento de ParasitologiaInstituto de Ciências BiomédicasUniversidade de São PauloSão PauloBrazil

Nolwenn M. DheillyDepartment of Biological

SciencesMacquarie UniversitySydneyAustralia

xiv PARTICIPANTSPP

Ilka EngelmannCentre d’Immunologie

de Marseille-LuminyUniversité de la MéditerranéeMarseilleFrance

Julie GhoshDepartment of Biological

SciencesGeorge Washington UniversityWashington, DCUSA

Nam-Chul HaNational Research Laboratory


*��<"�=�"��>�Institute of ParasitologyBiology CentreAcademy of Sciences


y

Czech Republic

Tor HaugNorwegian College

of Fishery ScienceUniversity of TromsoTromsoNorway

John H. HensonDepartment of BiologyBiochemistry and Molecular

Biology ProgramDickinson CollegeCarlisle, PennsylvaniaUSA

Julián F. HillyerDepartment of Biological

SciencesInstitute for Global HealthVanderbilt UniversityNashville, TenesseeeUSA

Hyun-Ok HwangNational Research Laboratory


Haobo JiangDepartment of Entomology

and Plant PathologyOklahoma State UniversityStillwater, OklahomaUSA

Pikul JiravanichpaisalMolecular Aquatic Biology

and Genetic LaboratoryNational Center for Genetic

Engineering and BiotechnologyNational Science and Technology

Development Agency?�"��Thailand

Radka JoskováInstitute of MicrobiologyAcademy of Sciences


Michael R. KanostDepartment of BiochemistryKansas State UniversityManhattan, KansasUSA

Shun-ichiro KawabataDepartment of BiologyKyushu UniversityFukuokaJapan

Chan-Hee KimDivision of Infectious DiseaseDepartment of MedicineUniversity of Massachusetts

Medical SchoolWorcester, MassachusettsUSA

xvPARTICIPAPP NTS

&��@�\^�Institute of ParasitologyBiology CentreAcademy of Sciences


y

Czech Republic

Shoichiro KurataKKGraduate School

of Pharmaceutical SciencesTohokuTT UniversitySendaiJapan

Kenji KurokawaKKNational Research Laboratory


Bok Luel LeeNational Research Laboratory


Chun LiNorwegian College

of Fishery ScienceUniversity of TromsoTTTromsoTTNorway

Hai-peng LiuState Key Laboratory of Marine

Environmental ScienceCollege of Oceanography

and Environmental ScienceXiamen UniversityFujianChina

Eric S. LokerCenter for Evolutionary

and Theoretical ImmunologyDepartment of BiologyUniversity of New MexicoAlbuquerque, New MexicoUSA

Cheng Man LunDepartment of Biological


Audrey J. MajeskeDepartment of Biological


Valeria MatrangaConsiglio Nazionale

delle RichercheInstituto di Biomedicina

e immunologia Molecolare “Albert Monroy”

PalermoItaly

Sham V. NairDepartment of Biological


Masaru NonakaDepartment of Biological

SciencesGraduate School of ScienceThe University of TokyoTTHongo, TokyoTTJapan

Ji-Won ParkNational Research Laboratory


xvi PARTICIPAPP NTS

Jiang RuiRRNational Research Laboratory


Kyung-KK Hwa RyuRRNational Research Laboratory


Sandro SacchiDepartment of Biological


Michel SalzetUniversité Lille Nord De FranceLaboratoire de Neuroimmunologie

et Neurochimie EvolutivesUniversité LilleLilleFrance

Honoo SatakeSuntory Institute for Bioorganic

ResearchWakayamadai, Shimamoto, MishimaOsakaJapan

Catherine S. SchrankelDepartment of Biological


Marcela ŠilerováInstitute of MicrobiologyAcademy of Sciences


Keun-Hwa ParkNational Research Laboratory


Petra ProcházkováInstitute of MicrobiologyAcademy of Sciences


Nathalie PujolCentre d’Immunologie

de Marseille-LuminyUniversité de la MéditerranéeMarseilleFrance

Limei QiuKey Laboratory of Experimental

Marine BiologyInstitute of OceanologyChinese Academy of SciencesQingdaoChina

David A. RaftosDepartment of Biological


Jonathan P. RastDepartment of Medical

BiophysicsDepartment of ImmunologySunnybrook Health Sciences CentreUniversity of TorontoTTToronto, TT OntarioCanada

Mattias RothDepartment of Biological


xviiPARTICIPAPP NTS

Aurélie TasiemskiTTUniversité Lille Nord De FranceLaboratoire de Neuroimmunologie

et Neurochimie EvolutivesUniversité LilleFrance

Andreas VilcinskasInstitut für Phytopathologie

und Angewandte ZoologieJustus-Liebig-Universität GießenHeinrich-GießenGermany

Lingling WangKey Laboratory of Experimental


Huan ZhangKey Laboratory of Experimental


L. Courtney SmithDepartment of Biological


Irene SöderhällDepartment of Comparative


Kenneth SöderhällDepartment of Comparative


Linsheng SongInstitute of OceanologyChinese Academy of SciencesQingdaoChina

Klara StensvågNorwegian College

of Fishery ScienceUniversity of TromsoTTTromsoTTNorway

xix

CONTENTS

1. CNIDARIAN IMMUNITY: A TALE OF TWO BARRIERS ...............................1

René Augustin and Thomas C.G. Bosch

Abstract......................................................................................................................................... 1Introduction: Cnidaria Provide Information about the Evolution of Immunity................... 1Cnidarian Immune Responses Are Performed by TwoTT Epithelial Barriers........................... 3Cnidarians Do Not Live Alone but Are Intimately Associated with Symbionts..................... 5Cnidarians Recognize Microbial Associated Molecular Patterns through Germ

Line Encoded Pattern Recognition Receptors (PRRs) ..................................................... 7Cnidarians Produce Antimicrobial Peptides Which

Are Effective Even Against Human Pathogens ................................................................. 9How Do Cnidaria Distinguish Self from Nonself? .................................................................. 11Concluson and Perspective—Where Will the TaleTT Lead Us? ................................................ 13

2. GASTROPOD IMMUNOBIOLOGY ...................................................................17

Eric S. Loker

Abstract....................................................................................................................................... 17Introduction: An Homage to Gastropod Antiquity and Diversity......................................... 18Infectious Challenges to Gastropods........................................................................................ 18Gastropod Genome Projects—Awaiting the Deluge............................................................... 21The Distinctive Architecture of Gastropod Immune Systems Offers Many

Opportunities for Study .................................................................................................... 21��

Host-Parasite Systems ....................................................................................................... 28Addressing Fundamental Issues in Immunology Using Gastropod Models......................... 30Opportunities to Expand the Relevance of Gastropod Immunity......................................... 33Conclusion .................................................................................................................................. 33

xx CONTENTS

3. BIVAVV LVE IMMUNITY ..........................................................................................44

Linsheng Song, Lingling Wang, Limei Qiu and Huan Zhang

Abstract....................................................................................................................................... 44Introduction................................................................................................................................ 44Hemocytes and Phagocytosis .................................................................................................... 45Immune Recognition.................................................................................................................. 46Immunity Signaling Pathways .................................................................................................. 50Immune Effector ........................................................................................................................ 53Conclusion .................................................................................................................................. 60

4. EARTHWORM IMMUNITY................................................................................66

Martin Bilej, Petra Procházková, Marcela Šilerová and Radka Josková

Abstract....................................................................................................................................... 66Introduction................................................................................................................................ 66Basic Information on Earthworm Anatomy............................................................................ 67Cellular Defense Mechanisms................................................................................................... 68Humoral Defense Mechanisms ................................................................................................. 70Conclusion .................................................................................................................................. 75

5. LEECH IMMUNITY: FROM BRAIRR N TO PERIPHERARR L

RESPONSES ...................................................................................................80

Aurélie Tasiemski and Michel SalzetTT

Abstract....................................................................................................................................... 80Introduction................................................................................................................................ 80The Medicinal Leech as a Model for Studying the Immune Response

of the CNS........................................................................................................................... 81B/Theromyzon Tessulatum as aTT Model for Studying the Peripheral

Immune Response .............................................................................................................. 92Conclusion and Perspectives................................................................................................... 101

6. INNATEAA IMMUNITY IY N C. ELEGANS .............................................................105

Ilka Engelmann and Nathalie Pujol

Abstract..................................................................................................................................... 105Introduction.............................................................................................................................. 105Routes of Infection ................................................................................................................... 106Pathogen Recognition .............................................................................................................. 106Signalling Pathways Involved in the Immune Response ...................................................... 109TranscriptionTT Factors Involved in the Immune Response ................................................... 112Effector Molecules Involved in the Immune Response......................................................... 113Modulation of the Immune Response by the Nervous System ............................................ 114Immune Response to Pore-Forming ToxinsTT ........................................................................... 114Epidermal Immune Response to the Fungus Drechmeria Coniospora................................ 115Conclusion ................................................................................................................................ 117

xxiCONTENTS

7. IMMUNOCOMPETENT MOLECULES AND THEIR RESPONSE

NETWORK IN HORSESHOE CRABRR S .....................................................122

Shun-ichiro Kawabata

Abstract..................................................................................................................................... 122Introduction and Historical Background .............................................................................. 122The Molecular Mechanism Underlying LPS-TriggeredTT Hemocyte Exocytosis.................. 123��!�� .............................................. 124Function of a Toll-TT Like Receptor on the Hemocyte .............................................................. 126The Coagulation Cascade Triggered byTT LPS and BDG ....................................................... 126BDG Recognition by Factor G................................................................................................ 127Pathogen Recognition by Hemocyte- and Plasma-Derived Lectins .................................... 128Sterilization by Antimicrobial Peptides ................................................................................. 128Conversion of Hemocyanin to Phenoloxidase by Antimicrobial Peptides

and Coagulation Factors ................................................................................................. 129TGase-Dependent Cross-Linking in the Immune System.................................................... 130Pathogen Recognition by the Complement System .............................................................. 131Conclusion and Future Prospects in the Horseshoe Crab Innate Immune System........... 132

8. TICK INNATEAA IMMUNITY ...............................................................................137

&��@�\^��*��<"�=�"��>��_��>��\ and Sirlei Daffre

Abstract..................................................................................................................................... 137Introduction.............................................................................................................................. 138Defense Mechanisms in the Tick Haemocoel......................................................................... 139Immunity in the Tick Gut ....................................................................................................... 152Immune Molecules in Other Tick Tissues.............................................................................. 156Conclusion ................................................................................................................................ 157

9. BEETLE IMMUNITY..........................................................................................163

Ji-Won Park, Chan-Hee Kim, Jiang Rui, RR Keun-Hwa Park, KyungKK -Hwa Ryu, RRJun-Ho Chai, Hyun-Ok Hwang, Kenji Kurokawa, NamKK -Chul Ha, Irene Söderhäll, Kenneth Söderhäll and Bok Luel Lee

Abstract..................................................................................................................................... 163Introduction.............................................................................................................................. 163Biochemical Characterization of Pattern Recognition Proteins in Beetles ........................ 164Serine Protease Zymogens Regulate the proPo Cascade in Beetles .................................... 166A Modular Serine Protease is the Immediate Downstream Protease

of the PGRP-SA/GNBP1 Complex ................................................................................. 167Three Serine Protease Zymogens Are Involved in Lys-LL TypeTT PG-Dependent

Activation of the TollTT Signaling Cascade ....................................................................... 168��"��#��$��"� �-1,3-Glucan-Dependent

TollTT Signaling Cascade..................................................................................................... 170Three Protease-Serpin Complexes Cooperatively Regulate the Innate Immune

Responses in Beetles......................................................................................................... 173Conclusion ................................................................................................................................ 176

xxii CONTENTS

10. IMMUNITY IN LEPIDOPTERARR N INSECTS ................................................181

Haobo Jiang, Andreas Vilcinskas and Michael R. Kanost

Abstract..................................................................................................................................... 181Introduction.............................................................................................................................. 182Hemocytes................................................................................................................................. 182Recognition of Microorganisms.............................................................................................. 183Antimicrobial Peptides and Proteins...................................................................................... 186Extracelluar and Intracellular Signal TransductionTT Stimulating Antimicrobial

Peptide Synthesis.............................................................................................................. 188Prophenoloxidase Activation System ..................................................................................... 190Inhibitory Regulation of Hemolymph Proteinases by Serpins ............................................ 191Lepidopteran Immune Responses to Different Types of InfectionTT ...................................... 192Conclusion ................................................................................................................................ 194

11. FLY IY MMUNITY: RECOGNITION OF PATHOGENS AA

AND INDUCTION OF IMMUNE RESPONSES ......................................205

Shoichiro KurataKK

Abstract..................................................................................................................................... 205Introduction.............................................................................................................................. 205Primary Humoral Responses: Activation of Constitutive Protein Cascades

in the Hemolymph............................................................................................................ 206Secondary Humoral Responses: Induction of Self-Defense Molecules............................... 207Recognition of VariousVV Pathogens in the Hemolymph ......................................................... 210Recognition and Elimination of Intracellular Bacteria in the Cytoplasm.......................... 211�!��%��%��$�� ............................................. 212Conclusion ................................................................................................................................ 213

12. MOSQUITO IMMUNITY .................................................................................218

Julián F. Hillyer

Abstract..................................................................................................................................... 218Introduction.............................................................................................................................. 218Biology of Pathogens inside Mosquitoes ................................................................................ 221Mosquito Compartments and Barriers to Infection ............................................................. 221Molecular Basis of Mosquito Immunity................................................................................. 225Conclusion ................................................................................................................................ 232

13. CRUSTATT CEAN IMMUNITY ............................................................................239

Lage Cerenius, Pikul Jiravanichpaisal, Hai-peng Liu and Irene Söderhäll

Abstract..................................................................................................................................... 239Introduction.............................................................................................................................. 239Pattern Recognition ................................................................................................................. 240Lectins ....................................................................................................................................... 241Hemocytes and Hematopoiesis................................................................................................ 246Antiviral Reactions .................................................................................................................. 249Clotting, Synthesis of Antimicrobial Proteins and Melanisation......................................... 251Conclusion ................................................................................................................................ 253

xxiiiCONTENTS

14. ECHINODERM IMMUNITY ...........................................................................260

L. Courtney Smith, Julie Ghosh, Katherine M. Buckley, Lori A. Clow, Nolwenn M. Dheilly, Tor TT Haug, John H. Henson, Chun Li, Cheng Man Lun, Audrey J. Majeske, Valeria Matranga, Sham V. Nair, Jonathan P. Rast, David A. Raftos, Mattias Roth, Sandro Sacchi, Catherine S. Schrankeland Klara Stensvåg

Abstract..................................................................................................................................... 260Introduction.............................................................................................................................. 261Coelomocytes, the Immune Mediators in Echinoderms....................................................... 261Clot Formation ......................................................................................................................... 268Key Molecules in Echinoderm Immunity .............................................................................. 270Sp185/333—A Diverse Family of Genes and Proteins Expressed in Response

to Immune Challenge....................................................................................................... 272Complement in Echinoderms.................................................................................................. 277Lectins ....................................................................................................................................... 279Scavenger Receptors ................................................................................................................ 281Antimicrobial Peptides ............................................................................................................ 284Gene Expression in Sea Urchin Coelomocytes ...................................................................... 288&��!��&��'��"��#�� ...................................................... 289Regeneration............................................................................................................................. 290Ecotoxicology and the Echinoderm Immune System ........................................................... 290Conclusion ................................................................................................................................ 291

15. UROCHORDATEAA IMMUNITY........................................................................302

Masaru Nonaka and Honoo Satake

Abstract..................................................................................................................................... 302Introduction.............................................................................................................................. 302Toll-TT Like Receptors (TLR)...................................................................................................... 303Complement System ................................................................................................................ 305Allorecognition ......................................................................................................................... 307Conclusion ................................................................................................................................ 309

INDEX........................................................................................................................311

xxiv

ACKNOWLEDGEMENTS

I wish to thank all who reviewed the chapters of this book and the Swedish Science Research Council for continous support of my research.

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Invertebrate Immunity - Home - Springer978-1-4419-8059-5/1.pdf · No part of t his book may be repro duced or transm itted in any form or by ... now carried out on developmental biology