VOLUMENINETYSIX A DVANCES IN PARASITOLOGY Echinococcus and Echinococcosis, Part B SERIES EDITOR D. ROLLINSON J. R. STOTHARD Life Sciences Department Department of Parasitology The Natural History Museum, Liverpool School of Tropical London, UK Medicine Liverpool, UK [email protected] [email protected] EDITORIAL BOARD T. J. C. ANDERSON R. C. OLIVEIRA Department of Genetics, Texas Centro de Pesquisas Rene Rachou/ Biomedical Research Institute, CPqRR - A FIOCRUZ em Minas San Antonio, TX, USA Gerais, Rene Rachou Research Center/CPqRR - The Oswaldo Cruz M. G. BASA(cid:1)N~EZ Foundation in the State of Minas Professor of Neglected Tropical Gerais-Brazil, Brazil Diseases, Department of Infectious R. E. SINDEN Disease Epidemiology, Faculty of Medicine (St Mary’s Campus), Immunology and Infection Imperial College London, Section, Department of Biological London, UK Sciences, Sir Alexander Fleming Building, Imperial College of S. BROOKER Science, Technology and Medicine, London, UK Wellcome Trust Research Fellow and Professor, London School of D. L. SMITH Hygiene and Tropical Medicine, Johns Hopkins Malaria Research Faculty of Infectious and Tropical, Institute & Department of Diseases, London, UK Epidemiology, Johns Hopkins BloombergSchoolofPublicHealth, R. B. GASSER Baltimore, MD, USA Faculty of Veterinary and Agricultural Sciences, The R. C. A. THOMPSON University of Melbourne, Parkville, Head, WHO Collaborating Centre Victoria, Australia for the Molecular Epidemiology of Parasitic Infections, Principal N. HALL Investigator, Environmental School of Biological Sciences, Biotechnology CRC (EBCRC), School Biosciences Building, University of of Veterinary and Biomedical Liverpool, Liverpool, UK Sciences, Murdoch University, Murdoch, WA, Australia J. KEISER X.-N. ZHOU Head, Helminth Drug Development Unit, Department Professor, Director, National of Medical Parasitology and Institute of Parasitic Diseases, Infection Biology, Swiss Tropical Chinese Center for Disease Control and Public Health Institute, Basel, andPrevention,Shanghai,People’s Switzerland Republic of China VOLUME NINETY SIX A DVANCES IN PARASITOLOGY Echinococcus and Echinococcosis, Part B Edited by R.C.A. THOMPSON School of Veterinary and Life Sciences, Murdoch University, Murdoch Australia P. DEPLAZES Institute of Parasitology, University of Z€urich, Z€urich, Switzerland A.J. LYMBERY Freshwater Fish Group and Fish Health Unit, School of Veterinary and Life Sciences Murdoch University, Murdoch, Australia AcademicPressisanimprintofElsevier 125LondonWall,LondonEC2Y5AS,UnitedKingdom TheBoulevard,LangfordLane,Kidlington,OxfordOX51GB,UnitedKingdom 50HampshireStreet,5thFloor,Cambridge,MA02139,UnitedStates 525BStreet,Suite1800,SanDiego,CA92101-4495,UnitedStates Firstedition2017 Copyright©2017ElsevierLtd.Allrightsreserved. Nopartofthispublicationmaybereproducedortransmittedinanyformorbyanymeans,electronicor mechanical,includingphotocopying,recording,oranyinformationstorageandretrievalsystem,without permissioninwritingfromthepublisher.Detailsonhowtoseekpermission,furtherinformationabout thePublisher’spermissionspoliciesandourarrangementswithorganizationssuchastheCopyright ClearanceCenterandtheCopyrightLicensingAgency,canbefoundatourwebsite:www.elsevier.com/ permissions. 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ISBN:978-0-12-812399-7 ISSN:0065-308X ForinformationonallAcademicPresspublicationsvisitour websiteathttps://www.elsevier.com/books-and-journals Publisher:ZoeKruze AcquisitionEditor:AlexWhite EditorialProjectManager:HeleneKabes ProductionProjectManager:VigneshTamil Designer:AlanStudholme TypesetbyTNQBooksandJournals CONTRIBUTORS VOLUME 95 PART A C.A.AlvarezRojas TheUniversityofMelbourne,Parkville,VIC,Australia D.Antolova InstituteofParasitologySAS,Kosice,SlovakRepublic K.Brehm UniversityofWu€rzburg,Wu€rzburg,Germany P.S.Craig UniversityofSalford,GreaterManchester,UnitedKingdom G.Cringoli UniversityofNaplesFedericoII,Napoli,Italy M.delaRue UniversityofSantaMaria,SantaMariaRS,Brazil P.Deplazes UniversityofZu€rich,Zurich,Switzerland J.Eckert UniversityofZurich,Zurich,Switzerland P.Giraudoux UniversityofFranche-ComtéandInstitutUniversitairedeFrance,Besancon,France M.F.Harandi ResearchcentreofHydatidDiseaseinIran,KermanUniversityofMedicalSciences, Kerman,Iran D.Jenkins CharlesSturtUniversity,WaggaWagga,NSW,Australia E.J.Jenkins UniversityofSaskatchewan,Saskatoon,SK,Canada U.Koziol UniversityofWu€rzburg,Wu€rzburg,Germany;UniversidaddelaRepu(cid:2)blica,Montevideo, Uruguay S.Lahmar NationalSchoolofVeterinaryMedicine,SidiThabet,Tunisia j ix x Contributors A.J.Lymbery MurdochUniversity,Murdoch,WA,Australia J.Magambo MeruUniversityofScienceandTechnology,Meru,Kenya A.Massolo UniversityofCalgary,Calgary,Alberta,Canada L.Rinaldi UniversityofNaplesFedericoII,Napoli,Italy T.Romig UniversityofHohenheim,Stuttgart,Germany J.M.Schurer UniversityofSaskatchewan,Saskatoon,SK,Canada;UniversityofWashington,Seattle,WA, UnitedStates K.Takahashi RakunoGakuenUniversity,Ebetsu,Japan R.C.A.Thompson MurdochUniversity,Murdoch,WA,Australia P.R.Torgerson UniversityofZu€rich,Zurich,Switzerland M.Wassermann UniversityofHohenheim,Stuttgart,Germany VOLUME 96 PART B O.Akhan HacettepeUniversity,Ankara,Turkey C.Budke TexasA&MUniversity,CollegeStation,TX,UnitedStates A.Casulli WorldHealthOrganizationCollaboratingCentrefortheepidemiology,detectionand controlofcysticandalveolarechinococcosis,IstitutoSuperiorediSanit(cid:3)a,Rome,Italy F.J.Conraths Friedrich-Loeffler-Institut,FederalResearchInstituteforAnimalHealth,Instituteof Epidemiology,Greifswald-InselRiems,Germany P.S.Craig UniversityofSalford,GreaterManchester,UnitedKingdom B.Gottstein UniversityofBern,Bern,Switzerland Contributors xi D.Hegglin UniversityofZurich,Zurich,Switzerland P.Kern UniversityHospitalofUlm,Ulm,Germany M.W.Lightowlers UniversityofMelbourne,Werribee,VIC,Australia A.MenezesdaSilva EstradadaLuz,Lisboa,Portugal N.Mu€ller InstituteofParasitology,VetsuisseFaculty,UniversityofBern,Bern,Switzerland B.Mu€llhaupt UniversityHospitalofZurich,Zu€rich,Switzerland E.Ortona IstitutoSuperiorediSanit(cid:3)a,Rome,Italy M.Siles-Lucas IRNASA-CSIC,Salamanca,Spain A.Siracusano IstitutoSuperiorediSanit(cid:3)a,Rome,Italy P.Soboslay UniversityofTu€bingen,Tu€bingen,Germany P.R.Torgerson UniversityofZurich,Zurich,Switzerland K.A.Vizcaychipi NationalInstituteofInfectiousDiseases,BuenosAires,Argentina D.A.Vuitton WHOCollaboratingCentre,UniversitédeFranche-Comté,Besançon,France J.Wang UniversityofBern,Bern,Switzerland Q.Wang SichuanProvincialCenterforDiseaseControlandPrevention,Chengdu,China PREFACE Our aim in this thematic volume of Advances in Parasitology is to provide a completesynthesisofwhatisknownaboutthecestodeparasiteEchinococcus andthediseasesitcauses,echinococcosis(hydatiddisease).Itbuildsonthe successoftwopreviousvolumes‘TheBiologyofEchinococcusandHydatid disease’ and ‘Echinococcus and Hydatid Disease’ published by Allen and UnwinandCABInternational,respectively,anddetailsthemajoradvances thathavetakenplacesince.The10chaptersdemonstratethatinadditionto its medical, veterinary and economic significance, Echinococcus is an intriguing biological phenomenon. They detail the major advances that havetakenplaceduringthelast20years,particularlyinourunderstanding of taxonomy, genetic variation, developmental biology, host–parasite relationships, geographic distribution and host range, diagnosis, control andclinicalmanagement.Inaddition,wehaveincludedachaptercovering historical aspects of echinococcosis that highlights major contributions to knowledge about the parasite and the diseases it causes. In this respect, four doyens of the field and contributors to the previous volume have passed away in recent years: Desmond Smyth, Michael Gemmell, Robert Rausch and Rudolf Ammann. We are therefore pleased to have the opportunity that this volume provides to detail and highlight the major contributions they have made to the field. AlthoughmajoradvanceshavebeenmadeinresearchonEchinococcusand echinococcosis, many questions remain, particularly in the areas of developmental biology and host–parasite relationships. Control efforts have had limited impact globally, and Echinococcus is an emerging problem in some parts of the world. As with many zoonoses, control is hampered by anthropogenic factors that influence both domestic and wild cycles of transmission. The overriding theme of the book is that a comprehensive understand- ing of the biology of Echinococcus is essential for the effective treatment and control of echinococcosis. The links between laboratory knowledge and field applications are emphasized throughout the book. Consequently, we hope that research workers, teachers and students of parasitology, clinicians j xiii xiv Preface andfieldworkerswillfindthisworkanindispensablesourceofinformation, butthatitwillalso provideamodel fortheintegrationof basic andapplied research in parasitology. Andrew Thompson, Peter Deplazes and Alan Lymbery CHAPTERONE Immunology of Alveolar and Cystic Echinococcosis (AE and CE) B. Gottstein*,1, P. Soboslayx, E. Ortona{, J. Wang*, A. Siracusano{, jj D.A. Vuitton *UniversityofBern,Bern,Switzerland xUniversityofTu€bingen,Tu€bingen,Germany {IstitutoSuperiorediSanit(cid:2)a,Rome,Italy jjWHOCollaboratingCentre,UniversitédeFranche-Comté,Besançon,France 1Correspondingauthor:E-mail:[email protected] Contents 1. BriefIntroductiontoAlveolarEchinococcosisandCysticEchinococcosis 2 1.1 Theparasites,theirbasicbiology 2 1.2 Biologyofinfectioninintermediatehosts 3 1.3 Alveolarechinococcosisandcysticechinococcosisrepresentverydifferent 5 diseases 2. BackgroundImmunology-BasedonMurineStudies 7 2.1 Innateimmunity,macrophages,dendriticcellsandsusceptibilitytoinfection 7 2.2 TheroleofdifferentThcelltypes(focusingonTregsandTh17)andrespective 10 cytokines/chemokines 3. OverviewonImmunologicalAspectsonHumanAlveolarEchinococcosisand 14 CysticEchinococcosis 3.1 Innateimmunity 14 3.2 Theroleofmacrophages/Kupffercells 15 3.3 Theroleofdendriticcells 16 3.4 Acquiredimmunity:theroleofdifferentThcelltypes(focusingonTregsand 17 Th17),includingcytokines/chemokines 4. ImmunologyandImmunoregulationofSusceptibility/MorbidityinAlveolar 19 EchinococcosisandCysticEchinococcosis 4.1 Immunomodulationthatleadstoalveolarechinococcosis/disease 19 4.2 ImmunomodulationthatleadstoCE/disease 22 5. ImmunologyandImmunoregulationofResistancetoAlveolarEchinococcosisand 23 CysticEchinococcosis 5.1 Immunomodulationthatleadstoinactivation/dyingout/abortionofalveolar 23 echinococcosis 5.2 Immunomodulationthatleadstoinactivation/dyingout/abortionofcystic 24 echinococcosis AdvancesinParasitology,Volume96 ISSN0065-308X ©2017ElsevierLtd. j http://dx.doi.org/10.1016/bs.apar.2016.09.005 Allrightsreserved. 1