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Subcellular Biochemistry 99 J. Robin Harris Jon Marles-Wright Editors Macromolecular Protein Complexes IV Structure and Function Subcellular Biochemistry Volume 99 SeriesEditor J. Robin Harris, Institute of Molecular Physiology, University of Mainz, Mainz, Germany AdvisoryEditors TapasK.Kundu,TranscriptionandDiseaseLaboratory,JNCASR,Bangalore,India ViktorKorolchuk,InstituteforCellandMolecularBiosciences,Newcastle University,NewcastleuponTyne,UK VictorBolanos-Garcia,DepartmentofBiologicalandMedicalSciences,Oxford BrookesUniversity,Oxford,UK JonMarles-Wright,SchoolofNaturalandEnvironmentalSciences,Newcastle University,NewcastleuponTyne,UK The book series SUBCELLULAR BIOCHEMISTRY is a renowned and well recognized forum for disseminating advances of emerging topics in Cell Biology and related subjects. All volumes are edited by established scientists and the individual chapters are written by experts on the relevant topic. The individual chaptersofeachvolumearefullycitableandindexedinMedline/Pubmedtoensure maximumvisibilityofthework. (cid:129) J. Robin Harris Jon Marles-Wright Editors Macromolecular Protein Complexes IV Structure and Function Editors J.RobinHarris JonMarles-Wright InstituteofMolecularPhysiology BiosciencesInstitute UniversityofMainz NewcastleUniversity Mainz,Germany NewcastleuponTyne,UK ISSN0306-0225 ISSN2542-8810 (electronic) SubcellularBiochemistry ISBN978-3-031-00792-7 ISBN978-3-031-00793-4 (eBook) https://doi.org/10.1007/978-3-031-00793-4 ©TheEditor(s)(ifapplicable)andTheAuthor(s),underexclusivelicensetoSpringerNatureSwitzerland AG2022 Thisworkissubjecttocopyright.AllrightsaresolelyandexclusivelylicensedbythePublisher,whether thewholeorpartofthematerialisconcerned,specificallytherightsoftranslation,reprinting,reuseof illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similarordissimilarmethodologynowknownorhereafterdeveloped. Theuseofgeneraldescriptivenames,registerednames,trademarks,servicemarks,etc.inthispublication doesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexemptfromtherelevant protectivelawsandregulationsandthereforefreeforgeneraluse. The publisher, the authors and the editors are safe to assume that the advice and information in this bookarebelievedtobetrueandaccurateatthedateofpublication.Neitherthepublishernortheauthorsor theeditorsgiveawarranty,expressedorimplied,withrespecttothematerialcontainedhereinorforany errorsoromissionsthatmayhavebeenmade.Thepublisherremainsneutralwithregardtojurisdictional claimsinpublishedmapsandinstitutionalaffiliations. ThisSpringerimprintispublishedbytheregisteredcompanySpringerNatureSwitzerlandAG Theregisteredcompanyaddressis:Gewerbestrasse11,6330Cham,Switzerland Preface TheSubcellularBiochemistryseriesismovingtowardVolume100!Havingstarted as a quarterly journal in 1973, under the editorship of the late Donald Roodyn, it evolvedin1978toabookseries.Thediversityofcontentoverthepast~50yearshas been considerable, whilst retaining the underlying broad aim of presenting review chaptersonthediversesubcellularcomponents,fromtheorganelletothemolecular level.Recentyearshaveseenastrongemphasisuponthestructureandfunctionof macromolecularcomplexes,resultinginseveralthematicallyrelatedvolumes. Volume 82 (2017) was devoted to Fibrous Proteins, followed by Volume 83, Macromolecular Protein Complexes. Because of the large amount of data available at this time, and the willingness of scientists to contribute to the series, Volume87andVolume88(2018)werepublished,onMembraneandVirusProteins Complexes, respectively. Subsequently, a further book (Volume 93, 2019) entitled MacromolecularProteinComplexesIIpulledtogetherallaspectsofthebroadfield andwasfollowedbyMacromolecularProteinComplexesIII(2021).Severalearlier related books have been included in the series, generating an almost encyclopedic coverage,withsomemacromoleculartopicsalreadybeingupdated. Thisvolumehascontinuedthisapproachandnowcontainsdefinedsub-sections: Soluble Protein Complexes, Membrane Protein Complexes, Fibrous Protein Com- plexes,andVirusProteinComplexes.OwingtotheexceptionallydifficultCOVID- 19 times faced by scientists in academia and in research institutes, a number of initially agreed interesting chapters have been lost during production, but the remaining seventeen chapters presented here have enabled us to compile a strong andinterestingbook. As in previous volumes, x-ray crystallography and cryo-electron microscopy of thestructuralbiologyofproteincomplexespredominates,withfunctionalconsider- ationsalsoapparentinmostchapters.Indeed,theimpactofmoleculargeneticsand the study of dynamic/flexible states and assembly/disassembly of macromolecular structures have become increasingly important, along with benefits for the under- standingofdiseasestates. v vi Preface Rather than expanding here upon some individual chapters, we simply wish to emphasizetheon-goingstrengthofthisgroupofchaptersandreferthereadertothe chapter list of the book. We have been impressed by the quality and outstanding interest ofthematerialinthesechaptersandtheenthusiasm shownbyourauthors. We hope that this book, as print version and available online as the e-book and individuale-chapters,willbeofuseandinteresttothebiosciencecommunity. Mainz,Germany J.RobinHarris NewcastleuponTyne,UK JonMarles-Wright Contents 1 FattyAcidSynthase:Structure,Function,andRegulation. . . . . . . 1 AybegN.Günenc,BenjaminGraf,HolgerStark,andAshwinChari 2 ATP-BindingCassetteTransporters:Snap-onComplexes?. . . . . . . 35 IqraYounus,SofiaKochkina,CheriC.Choi,WenjuanSun, andRobertC.Ford 3 TheForkProtectionComplex:ARegulatoryHubattheHead oftheReplisome. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83 DanielB.Grabarczyk 4 Ferritin-LikeProteins:AConservedCoreforaMyriadofEnzyme Complexes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109 RahulBanerjee,VivekSrinivas,andHugoLebrette 5 StillnoRestfortheReductases:RibonucleotideReductase(RNR) StructureandFunction:AnUpdate. . . . . . . . . . . . . . . . . . . . . . . . 155 MarcusJ.C.Long,PhillippeLy,andYimonAye 6 StructureandDynamicsoftheHumanMulti-tRNASynthetase Complex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199 MyungHeeKimandBeomSikKang 7 OntheRegulationofMitosisbytheKinetochore,aMacromolecular ComplexandOrganisingHubofEukaryoticOrganisms. . . . . . . . . 235 VictorM.Bolanos-Garcia PartI MembraneProteinComplexes 8 TheConformationalDynamicsofHeterotrimericGProteins DuringGPCR-MediatedActivation.. . . . . .. . . . .. . . . . .. . . . . .. 271 DonghoonAhnandKaYoungChung vii viii Contents 9 RegulationofLyticMachineriesbytheFtsEXComplexinthe BacterialDivisome. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 285 MartínAlcorlo,SisethMartínez-Caballero,RafaelMolina, andJuanA.Hermoso 10 Structure,Function,andRegulationoftheKainateReceptor. . . . . 317 SurbhiDhingra,JuhiYadav,andJaneshKumar 11 Structure,Function,andVariationsofthePhotosystemI-Antenna SupercomplexfromDifferentPhotosyntheticOrganisms. . . . . . . . 351 Jian-RenShen 12 StructureandFunctionofMycobacterial Arabinofuranosyltransferases. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 YongZiTanandFilippoMancia PartII FibrousProteinComplexes 13 StructureandAssemblyoftheBacterialFlagellum. . . . . . . . . . . . . 395 NatalieS.Al-OtaibiandJulienR.C.Bergeron 14 ActomyosinComplex. . .. . . . .. . . .. . . . .. . . .. . . . .. . . .. . . . .. 421 IanPepperandVitoldE.Galkin 15 StructureofMotileCilia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 471 TakashiIshikawa 16 Segment-Long-Spacing(SLS)andthePolymorphicStructures ofFibrillarCollagen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 495 YujiaXuandMicheleKirchner PartIII VirusProteinComplexes 17 ViralCapsidandPolymeraseinReoviridae. . . . . . . . . . . . . . . . . . 525 HongrongLiuandLingpengCheng Chapter 1 Fatty Acid Synthase: Structure, Function, and Regulation AybegN.Günenc,BenjaminGraf,HolgerStark,andAshwinChari Abstract Fatty acid (FA) biosynthesis plays a central role in the metabolism of livingcellsasbuildingblocksofbiologicalmembranes,energyreservesofthecell, andprecursorstosecondmessengermolecules.Inkeepingwithitscentralmetabolic role,FAbiosynthesisimpactsseveralcellularfunctionsanditsmisfunctionislinked todisease,suchascancer,obesity,andnon-alcoholicfattyliverdisease.CellularFA biosynthesisisconductedbyfattyacidsynthases(FAS).AllFASenzymescatalyze similar biosynthetic reactions, but the functional architectures adopted by these cellularcatalystscandiffersubstantially.ThisvariabilityinFASstructureamongst variousorganismsandtheessentialroleplayedbyFAbiosyntheticpathwaysmakes this metabolic route a valuable target for the development of antibiotics. Beyond cellularFAbiosynthesis,thequestforrenewableenergysourceshaspiquedinterest inFAbiosyntheticpathwayengineeringtogeneratebiofuelsandfattyacidderived chemicals. For these applications, based on FA biosynthetic pathways, to succeed, detailed metabolic, functional and structural insights into FAS are required, along withanintimateknowledgeintotheregulationofFAS.Inthisreview,wesummarize our present knowledge about the functional, structural, and regulatory aspects ofFAS. Keywords Fattyacidbiosynthesis·Fattyacidsynthase·ACP·FASstructure·FAS regulation·Biofuels·Oleochemicals A.N.Günenc·B.Graf·A.Chari(*) ResearchGroupforStructuralBiochemistryandMechanisms,MaxPlanckInstitutefor MultidisciplinarySciences,Göttingen,Germany e-mail:[email protected];[email protected]; [email protected] H.Stark DepartmentofStructuralDynamics,MaxPlanckInstituteforMultidisciplinarySciences, Göttingen,Germany e-mail:[email protected] ©TheAuthor(s),underexclusivelicensetoSpringerNatureSwitzerlandAG2022 1 J.R.Harris,J.Marles-Wright(eds.),MacromolecularProteinComplexesIV, SubcellularBiochemistry99,https://doi.org/10.1007/978-3-031-00793-4_1