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Alois Lametschwandtner Bernd Minnich Color Atlas of Adult Xenopus laevis Microvasculature of Tissues and Organs Xenopus laevis Color Atlas of Adult (cid:129) Alois Lametschwandtner Bernd Minnich Color Atlas of Adult Xenopus laevis Microvasculature of Tissues and Organs AloisLametschwandtner BerndMinnich DepartmentofEnvironmentandBiodiversity DepartmentofEnvironmentandBiodiversity UniversityofSalzburg UniversityofSalzburg Salzburg,Austria Salzburg,Austria ISBN978-3-031-05109-8 ISBN978-3-031-05110-4 (eBook) https://doi.org/10.1007/978-3-031-05110-4 #TheEditor(s)(ifapplicable)andTheAuthor(s),underexclusivelicensetoSpringerNatureSwitzerlandAG2022 Thisworkissubjecttocopyright.AllrightsaresolelyandexclusivelylicensedbythePublisher,whetherthewholeor part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting,reproductiononmicrofilmsorinanyotherphysicalway,andtransmissionorinformationstorageand retrieval,electronicadaptation,computersoftware,orbysimilarordissimilarmethodologynowknownorhereafter developed. Theuseofgeneraldescriptivenames,registerednames,trademarks,servicemarks,etc.inthispublicationdoesnot imply,evenintheabsenceofaspecificstatement,thatsuchnamesareexemptfromtherelevantprotectivelawsand regulationsandthereforefreeforgeneraluse. Thepublisher,theauthorsandtheeditorsaresafetoassumethattheadviceandinformationinthisbookarebelievedto betrueandaccurateatthedateofpublication.Neitherthepublishernortheauthorsortheeditorsgiveawarranty, expressedorimplied,withrespecttothematerialcontainedhereinorforanyerrorsoromissionsthatmayhavebeen made.Thepublisherremainsneutralwithregardtojurisdictionalclaimsinpublishedmapsandinstitutionalaffiliations. ThisSpringerimprintispublishedbytheregisteredcompanySpringerNatureSwitzerlandAG Theregisteredcompanyaddressis:Gewerbestrasse11,6330Cham,Switzerland Preface The cardiovascular system of animals and men is the first system to develop and to function. Failureofthesystemresultsinseverefunctionaldisturbancesanddependingupontheseverity itleadstodeath.Atpresent,failureofthissystemhasbecomethemaincauseforhumandeath inWesterncountries. Historically, there is a long-lasting interest in the anatomy of the cardiovascular system. Initially,bloodvesselswereexposedincorpsesofmanandanimalsbymacroscopicdissections and inspection with bare eyes or magnifying glasses. To locate distinct blood vessels, air, liquids,orsolidifyingmaterialswereinjected,wherebytheinventionoftheanatomicalsyringe allowed higher pressures resultinginbetterfillings oflargervascularterritories. Preparations, however,werestilldelicateandcouldbedemonstratedtoasmallaudienceonly.Tremendous progresswasmadewhenpolymerizing(hardening)resinsbecameavailable,whichwithstood thecorrosiveactionsofalkaliandacidsneededtomacerateandtoexposeinjectedindividual bloodvesselsforadetailedmacroscopicandmicroscopicinspection.Thelimitedresolutionof available light microscopes still did not allow to analyze the delicate microvasculature. A hallmark was set when Takuro Murakami in 1971 used the scanning electron microscope to studyresinvascularcorrosioncasts.Sincehispioneeringstudy,morethan1500articleswere publishedonthemicrovascularanatomyoftissuesandorgansinhealthanddiseaseusingthe techniqueofscanningelectronmicroscopyofvascularcorrosioncasts. ThisbookColorAtlasofAdultXenopuslaevis:MicrovasculatureofTissueandOrgansis theresultofworkdoneoverthepast45yearsbytheauthorsandtheirbachelors,masters,and/or PhDstudentsattheUniversityofSalzburg,Austria.Wepublishedouroriginalfindingsinpeer- reviewedjournalsofanatomy,embryology,histology,andmorphology,andareverythankful tocopyrightownersforgrantingcopyrightsoffigures,legends,andtextextracts,whichwere modifiedinsomedetailsandthusformthebackboneofthisatlas. To allow the readers, who are not familiar with vascular corrosion casts, a quick and easy identification of cast vascular structures as arterial, venous, or capillary vessels, we colored arterial vesselsredandvenousvesselsblueusingPhotoshop7.0(AdobeInc,Redwood,CA). Capillariesidentifythemselvesasthethinnestvesselswithdiametersnotlargerthan15μm. Wehopethatreadersenjoythewealthofcolorizedfiguresdisplayingthetissueandorgan- specific microvascular anatomy in adult Xenopus laevis, a longtime model organism in biologicalandbiomedicalresearch. Salzburg,Austria AloisLametschwandtner BerndMinnich v Acknowledgments The authors are very thankful to Synnöve Tholo for her long-lasting valuable technical assistanceovermanyyearsandtoORDrWolf-DietrichKrautgartnerforhiscontinuousexpert helpinscanningelectronmicroscopy.WethankDrHeidiBartel;ClaudiaDeutschmann,MSc; Dr Victoria Engelhardt; Nicole Fagerer, MSc; Nikolas Gerlach, MSc; Monika Höll, MSc; Mathias Hopfinger, MSc; Jan Philip Höpner, MSc; Christine Radner, BSc; Johannes Rattey, MSc;andDrWasanTangphokhanon,whobytheirbachelors,masters,and/ordoctoraltheses onthemicrovascularanatomyofselectedlarvaloradultorgansofXenopuslaevis,substantially contributedtothisatlas. AgrantbytheFondszurFörderungderwissenschaftlichenForschung,ProjektP-19050-B 17(toA.L.)isthankfullyacknowledged. Allstudies,thepresentatlasisbasedupon,wereapprovedbytheEthicsCommitteeofthe University of Salzburg, Austria, and by the Federal Government (BMBWK-66.012/0018- BrGT/2006). vii Contents 1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1 References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 2 MaterialsandMethods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5 2.1 Animals. . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . .. 5 2.2 Histomorphology(LightMicroscopy). . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5 2.3 VascularCorrosionCasting. . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . .. . . .. 5 2.4 ScanningElectronMicroscopy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6 2.5 Color-CodingofArterialandVenousVesselsinSEMMicrographs. . . . . . . . 6 2.6 IdentificationofCast(Replicated)VascularStructures. . . . . . . . . . . . . . . .. 7 2.6.1 VesselNature(Arterial,Venous,andCapillary/Sinusoid). . . . . . . . .. 7 2.6.2 FlowGoverningStructures. . .. . . .. . . . .. . . .. . . . .. . . .. . . . ... 7 2.6.3 SignsofSproutingandNon-sprouting(Intussusceptive MicrovascularGrowth)Angiogenesis. . . . . . . . . . . . . . . . . . . . . . . . 9 2.6.4 Artifacts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 9 2.7 AnalysesofVascularNetworks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 11 2.7.1 VesselSegments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 11 2.7.2 BranchingsandMergings(Confluences). . . . . . . . . . . . . . . . . . . . . . 12 2.7.3 Anastomoses. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15 References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 3 MicrovasculatureofXenopusTissuesandOrgans. . . . . . . . . . . . . . . . . . . . .. 21 3.1 Integument.. . . .. . . .. . . .. . . .. . . . .. . . .. . . .. . . .. . . .. . . .. . . . ... 21 3.1.1 Histomorphology. .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . . ... 21 3.1.2 VascularAnatomy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 21 3.2 CentralNervousSystemandSensoryOrgans. . . . . . . . . . . . . . . . . . . . . . .. 24 3.2.1 BrainandSpinalCord. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 24 3.2.2 Eyes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 60 3.2.3 OlfactoryandAccessoryOlfactory(Vomeronasal)Organs. . . . . . . .. 69 3.2.4 LateralLineOrgans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 81 3.2.5 CarotidLabyrinths. . . . .. . . . . .. . . . . .. . . . . .. . . . . .. . . . . .. .. 86 3.3 RespiratoryOrgans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . .. 90 3.3.1 Trachea. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 90 3.3.2 Bronchi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 96 3.3.3 Lungs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 105 3.3.4 Integument(SeeSect.3.1). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 112 3.4 DigestiveOrgans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 112 3.4.1 MaxillaryTeeth. . . . .. . . . . . . . . .. . . . . . . . . . .. . . . . . . . . .. . .. 112 3.4.2 Esophagus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 115 3.4.3 Stomach. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 121 3.4.4 SmallIntestine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 132 3.4.5 LargeIntestine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 143 ix x Contents 3.4.6 Liver. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153 3.4.7 Gallbladder. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170 3.4.8 Pancreas. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177 3.4.9 BileDuctsandPancreaticDucts. . . . . . . . . . . . . . . . . . . . . . . . . . . . 183 3.5 ExcretoryOrgans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192 3.5.1 Kidneys. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192 3.5.2 UrinaryBladder. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 205 3.6 ReproductiveOrgans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218 3.6.1 Ovaries. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 218 3.6.2 Oviducts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 222 3.6.3 Testes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 235 3.7 LymphaticOrgans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 244 3.7.1 Spleen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244 3.8 EndocrineOrgans. . . . . . . . .. . . . . . . . . . . . . .. . . . . . . . . . . . . . .. . . . .. 252 3.8.1 Hypophysis. . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . .. 252 3.8.2 Epiphysis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 260 3.8.3 ThyroidGlands. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261 3.8.4 IsletsofLangerhans. . . . . . .. . . . . . . .. . . . . . . . .. . . . . . . . .. . .. 267 3.8.5 Gonads(SeeSects.3.6.1and3.6.3). . . . . . . . . . . . . . . . . . . . . . . .. 273 References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 273 About the Authors AloisLametschwandtnerPhDisaretiredProfessoroftheUniversityofSalzburg,Austria.He is an expert in scanning electron microscopy of vascular casts and animal histology and physiology. Bernd Minnich PhD is a Professor at the University of Salzburg, Austria. Bernd Minnich’s researchfocusesonvascularbiology,electronmicroscopy,andbiostatistics. xi Abbreviations a Artery aa Arteriole aaa Arterio-arterialanastomosis ab Anteriorbranchofcerebralcarotidartery abra Anteriorbronchialartery abrv Anteriorbronchialvein ac Acinus acmc Anteriorcavityofmiddlechamber acn Accumbensnucleus acv Anteriorcavalvein aduv Anteriorduodenalvein aec Anterioreyechamber afaa afferentarteriole agaa Afferentglomerulararteriole aima Anteriorinferiormesencephalicartery aina Anteriorintestinalartery al Alveolus aob Accessoryolfactorybulb aov Anteriororbitalvein ap Alarplate apana Anteriorpancreaticartery asma Anteriorsuperiormesencephalicartery asp Adultspleen aspa Accessorysplenicartery asv Ascendingvein at Arterialtrunk atrv Anteriortrachealvein aua Auricularartery av Abdominalvein b Basket ba Basilarartery baina Branchofanteriorintestinalartery bc Basalcell Bc Bowmancapsule bcia Branchofciliaryartery bciv Branchofciliaryvein bd Bileduct bo Body bot Bottom bp Basalplate bpv Branchofportalvein br Bronchus xiii