Advances in Biomembranes and Lipid Self-Assembly, Volume 23 PDF
Preview Advances in Biomembranes and Lipid Self-Assembly, Volume 23
EDITORIAL BOARD Dr.MibelAguilar(MonashUniversity,Australia) Dr.AngelinaAngelova (Universite´deParis-Sud,France) Dr.PaulA.Beales (UniversityofLeeds,UnitedKingdom) Dr.Habil.RumianaDimova (MaxPlanckInstituteofColloidsandInterfaces,Germany) Dr.YuruDeng (ChangzhouUniversity,China) Prof.Dr.NirGov (TheWeizmannInstituteofScience,Israel) Prof.Dr.WojciechGóźdź (InstituteofPhysicalChemistryPolishAcademy ofSciences,Poland) Prof.Dr.ThomasHeimburg (NielsBohrInstitute,UniversityofCopenhagen,Denmark) Prof.Dr.TiborHianik (ComeniusUniversity,Slovakia) Prof.Dr.WolfgangKnoll (Max-Planck-Institutfu¨rPolymerforschung,Mainz,Germany) Prof.Dr.AngelicaLeitmannovaLiu (MichiganStateUniversity,USA) Dr.IlyaLevental(UniversityofTexas,USA) Prof.Dr.ReinhardLipowsky (MPIofColloidsandInterfaces,Potsdam,Germany) Prof.Dr.SylvioMay (NorthDakotaStateUniversity,USA) Prof.Dr.PhilippeMeleard (EcoleNationaleSuperieuredeChimiedeRennes,France) Prof.Dr.YoshinoriMuto (Gifu,Japan) Prof.Dr.V.A.Raghunathan(RamanResearchInstitute,India) Dr.AminSadeghpour (UniversityofLeeds,UnitedKingdom) Prof.KazutamiSakamoto (TokyoUniversityofScience,Japan) Prof.Dr.BernhardSchuster(UniversityofNaturalResourcesandLifeSciences,Vienna) Prof.Dr.P.B.SunilKumar (IndianInstituteofTechnologyMadras,India) Prof.Dr.MathiasWinterhalter (JacobsUniversityBremen,Germany) AcademicPressisanimprintofElsevier 50HampshireStreet,5thFloor,Cambridge,MA02139,USA 525BStreet,Suite1800,SanDiego,CA92101-4495,USA TheBoulevard,LangfordLane,Kidlington,Oxford,OX51GB,UK 125LondonWall,London,EC2Y5AS,UK Firstedition2016 Copyright©2016ElsevierInc.Allrightsreserved. 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ISBN:978-0-12-804715-6 ISSN:2451-9634 ForinformationonallAcademicPresspublications visitourwebsiteathttp://store.elsevier.com/ CONTRIBUTORS Y.Akao UnitedGraduateSchoolofDrugDiscoveryandMedicalInformationSciences, GifuUniversity,Gifu,Japan D.Bochicchio PhysicsDepartment,UniversityofGenoa,Genoa,Italy M.Bryszewska DepartmentofGeneralBiophysics,FacultyofBiologyandEnvironmentalProtection, UniversityofLodz,Poland H.Chamati InstituteofSolidStatePhysics,BulgarianAcademyofSciences,Sofia,Bulgaria A.Ciach InstituteofPhysicalChemistry,PolishAcademyofSciences,Warsaw,Poland W.Go´z´dz´ InstituteofPhysicalChemistry,PolishAcademyofSciences,Warsaw,Poland T.Hianik FacultyofMathematics,PhysicsandInformatics,ComeniusUniversity, Bratislava,Slovakia M.Ionov DepartmentofGeneralBiophysics,FacultyofBiologyandEnvironmentalProtection, UniversityofLodz,Poland I.Junkar JozˇefStefanInstitute,Ljubljana,Slovenia L.Monticelli MolecularMicrobiologyandStructuralBiochemistry(MMSB),UniversityofLyon, CNRSUMR5086,Lyon,France J.I.Pavlicˇ InstituteofSolidStatePhysics,BulgarianAcademyofSciences,Sofia,Bulgaria;InstitutJozˇef Stefan,Ljubljana,Slovenia K.Sakamoto DepartmentofPureandAppliedChemistry,TokyoUniversityofScience,Noda, Chiba,Japan R.Trobec InstitutJozˇefStefan,Ljubljana,Slovenia N.Yamada UnitedGraduateSchoolofDrugDiscoveryandMedicalInformationSciences, GifuUniversity,Gifu,Japan ix PREFACE Today’s scientists are still greatly challenged by countless aspects of biomembranes and by developing novel artificial systems based on lipid self-assembly, mainly because they resemble structurally and dynamically highly complex multicomponent systems. Nevertheless, over the last decade,anewtrendisbecomingmoreandmoreevident:complexsystems are increasingly tackled. From novel asymmetric cell membrane models to theengineeringofhybridnanoparticles,complexityseemsnolongerdeter- rent.Atthesametime,thearisingmultifacetedscientificquestionsdemand increasingly an interdisciplinary approach. Thus in the field of biomembranes and related lipid self-assembly research, mathematicians, physicists, chemists, biologists, engineers, and medics are starting to work closer together than ever. ThenewlytitledElsevierbookseriesAdvancesinBiomembranesandLipid Self-Assembly (ABLSA) will try to embrace this vivid scientific spirit and therefore will cover a broad range of topics from theoretical membrane models, over novel biomimetic membrane systems to various two- and three-dimensional lipid self-assemblies. Planar lipid bilayers are widely investigated due to their ubiquity in nature and find their application in the design of liposomal dispersions. Increasingly evident though, also non- lamellar membrane phases play an important role in nature, especially in dynamic processes such as vesicle fusion and cell communication. Self- assembled lipid structures have an enormous potential ranging from model systemstocellmembranesandfrombiosensingtocontrolleddrugdelivery. Likewise,thecontributionsofVolume23arespanningawiderangeofnew andexcitingtopicsincludingcoarsegrainmoleculardynamicsonpreformed vesicles (Hassan Chamati), simulation studies on the membrane bending modulus (Luca Monticelli), investigations on the importance of planarity in biomembranes (Kazutami Sakamoto), membranes interaction with den- drimersanddendricomplexes(TiborHianik),interactionsofcellsandplate- letswithbiomaterialsurfaces(ItaJunkar),Casimirpotentialsinnear-critical mixtures (Alina Ciach), bicontinuous cubic phases in the light of latest developments (Wojciech Gozdz), and extracellular vesicles in cancer are discussed (Yukihiro Akao). We wish to express our gratitude to all authors who contributed their chapterstotheVolume23ofABLSA,toMs.ShellieBryantandMs.Poppy xi xii Preface GarrawayfromElsevierOfficeinLondon,Mr.MageshKumarMahalingam fromElsevierdivisioninChennai,andtoallmembersoftheEditorialBoard who helped to prepare this volume of ABLSA. ALESˇ IGLICˇ CHANDRASHEKHAR V. KULKARNI MICHAEL RAPPOLT CHAPTER ONE The Importance of Planarity for Lipid Bilayers as Biomembranes K. Sakamoto1 DepartmentofPureandAppliedChemistry,TokyoUniversityofScience,Noda,Chiba,Japan 1Correspondingauthor:e-mailaddress:[email protected] Contents 1. Introduction 2 2. HowCPPTranslocatesintotheCytosol 4 3. PoreFormation 14 4. HowDoesCurvatureModulationbyChangingVesicleSizeEffecttheCPP Translocation? 18 5. Conclusion 20 Acknowledgments 21 References 21 Abstract Questionslike“Whatisthecellorvesiclesizethatfreeslipidmoleculesinbilayersfrom membranecurvaturestrains?”and“Whyisthesizeofself-reproducibleeukaryoticcell over10μm?” aretopics regarding membraneshape deformationcaused by cationic peptide interacted with anionic head group of membrane lipids. The mechanism of direct translocation (cytolysis) of cell-penetrating peptides (CPPs) and antimicrobial peptides(AMPs)throughbiomembranesisdiscussedinrelationtomembranecurva- ture and lipid mobility. It is confirmed that cytolysis happens when cationic CPP or AMPmoleculesinteractwithanioniclipidheadgroupstogeneratelocalandtransient catenoidporeswithnegativeGaussiancurvature.ThetranslocationofCPPandAMP,or moregenerally,thevesicleshapechangecausedbyphysicalorchemicalmodifiersis possible only under the condition that the lipid is under a lamellar liquid crystal (L ) d phasetobemobile.Thisconditioncouldbegivenprimarilyfortheplanarmembrane withnegligiblecurvaturestraintomakelocalorevenwholetopologicalmodification. Theremustbeathresholdlevelinthevesiclesizethatforceslipidstobecomeimmobile tomaintainvesiclestructures,whichexceedsthebendingenergyprovidedbyCPPor AMPtoreleasecurvaturestrain.Assuch,planarityisimportantforbiomembranestobe flexibleenoughtoaccommodatenecessarylocaltopologychanges,whichisthekeyof controllingthestimuliresponsivebarrierfunctionofcellmembrane. AdvancesinBiomembranesandLipidSelf-Assembly,Volume23 #2016ElsevierInc. 1 ISSN2451-9634 Allrightsreserved. http://dx.doi.org/10.1016/bs.abl.2016.01.001 2 K.Sakamoto 1. INTRODUCTION Inthischapter,thequestion“Whatisthecellorvesiclesizethatfrees lipidmoleculesinbilayersfrommembranecurvaturestrains”isthetopicfor discussion. This question also relates to the question “Why is the size of a self-reproducibleeukaryoticcellover10μm?”(Fig.1)[1,2].Takinganalog- icalcomparison,theratioofbilayerthickness (ca.5nm)tothediameterof erythrocyte (ca. 10μm) is 1/2000, while those of large unilamellar vesicles (LUVs)with200nmdiameteris1/40.Assmallerparticleshavelargercur- vatures,thestabilityofbilayerswouldbedependentonthecellsize.Kodama etal.reportedthatvesiclesmadeofdimyristoylphosphocholinearethermo- dynamicallystabilizedbytheenthalpyeffect,resultingfromacloselypacked aggregationofthelipidmolecules[2].Theyrevealedthattherelativether- modynamicstabilityofvesiclesattheliquidcrystal(LC)phaseandgelphase are both at the order of SUV ((cid:1)40nm), LUV ((cid:1)200nm), and large mul- tilamellarvesicle(>1μm),respectively.Regardingthetopologicaleffectof the vesicle, the surface energy depends on the curvature as expressed by Laplace equation; P ¼P +2 γ/r, where P is the inside pressure, P in out in out is the outside pressure, γ is the flat surface tension, and r is the radius of the curvature. As shown in Fig. 2, P for LUV (400nm) and SUV in (40nm) are 25 and 250 times larger than giant unilamellar vesicle (GUV; 10μm),respectively;inotherwords,thebendingenergyofthemembrane islargerforthesmallervesicles.Inordertokeepanenclosedsphericalshape to overcome curvature stress, lipids are less mobile and more ordered for ca. 5 nm Thickness of cell membranes (h) 25 nm Microtubule [1] d=x nm 30 nm d Small virus (picornaviruses) [1] h=5 nm ~ 40 nm SUV [2] 150–250 nm Small bacteria such as mycoplasma [1] 200–500 nm Lysosomes [1] ~ 200 nm LUV [2] (1–10 µm) The general sizes for prokaryotes [1] 2 µm E. coli —a bacterium [1] 9 µm Human red blood cell [1] Self-reproducible cells (10–30 µm) Most eukaryotic animal cells [1] 10 µm GUV [2] 100 µm Human egg [1] Fig.1 Sizeofcellandvesicle[1,2].Relativesizeofcellsandvesiclestofindtheimportance oftotalplanarityandlocalcurvaturemodulationforthefunctionofbiomembrane. TheImportanceofPlanarityforLipidBilayers 3 Laplace equation Pin = Pout +2 γ/r Pinr Pout r (nm) Relative 2 γ/r GUV 10,000 1 LUV 400 25 200 50 SUV 40 250 Fig.2 RelativepressuredifferencebyLaplaceequation. smaller vesicles [2]. All these data suggest that the bilayer curvature has a significant effect for the mobility and function of biomembranes. There couldbeathresholddiameterwhichcurvatureoverrulesagainstthemobil- ityoflipidmoleculesinthemembrane,whichisthekeyofcontrollingthe stimuli responsive barrier function of cell membrane. When I was invited to join the Editorial Board of Advances in Planar Lipid Bilayers and Liposomes (APLBL), I was asked about my opinion on changing the name of the journal to Advances in Biomembranes and Lipid Self-Assemblies(ABLSA),withtheexplanationthatAPLBLwasoriginally chosen by the founding editor who started the book series in 2004. He selected this name based on the nature of the research at the time, but thesituationhasseenasignificantchangesince.Thecurrenteditorswere concerned that it is no longer as relevant due to the changing nature of this research area and decided to change the name of the book series to ABLSA. Ihadnoobjection for thisdecisionbutchosethetitleofthischapteras “The Importance of Planarity for Lipid Bilayers as Biomembranes.” The titleexplainsitself,butIwouldliketoemphasizetheimportanceofplanarity in regards to the functionality of lipid bilayers for material trafficking through biomembranes. The reason this title came to me is that we have recentlyfacedcontradictoryresultsfortheeffectofcurvatureofunilamellar vesiclesbychangingtheirsizesagainstthetransmembranetraffickingofcell- penetratingpeptides(CPPs).ThisisthereasonIstartedthischaptertowon- der“Whatisthecellorvesiclesizethatfreeslipidmoleculesinbilayersfrom membrane curvature strains?” which would be closely related to the ques- tion “Why is the size of self-reproducible eukaryotic cells over 10μm?.” I will first review the mechanism of CPP’s translocation through biomembranesintermsoftopologyandfunctions inrelation tothecurva- ture, then continue further to the topic we have faced during the CPP translocation study that relates to the question above. 4 K.Sakamoto 2. HOW CPP TRANSLOCATES INTO THE CYTOSOL Discreetnessofcellisthekeyoflifewithcellmembranesthatseparate theirinteriorcytosol(orcytoplasm)fromtheirsurroundingenvironment.In this regard, cytosol is never connected to outside media, even during the course of cell fission or fusion. However, a water-soluble peptide called CPP can spontaneously translocate through cell membranes without any particular transporters or receptors [3]. There are two pathways of CPP translocationthroughbiomembranesasshowninFig.3,whichare(i)endo- cytosis followed by endosomal release into cytosol and (ii) direct trans- location through the membrane (cytolysis) [3]. Although well-established mechanisms are reported for the endocytosis, no conclusive mechanism has been found for the cytolysis, which is a more direct and efficient internalizationofwater-solubleCPPthroughhydrophobicbiomembranes. Ingeneral,CPPisconsideredtobesafeagainstmembraneintegrityandcell viability,soCPPcouldbeapotentialDDSvectorwithvarietiesofthebio- active cargosattached to it. There is another type of peptidecalled antimi- crobial peptide (AMP) that also spontaneously translocate into cytosol by pore formation [4]. Typical AMP called cationic amphipathic polypeptide hasstructuralresemblancetoCPPwithcationicaminoacidresidues,which causeAMPtoabsorbontotheanionicheadgroupofmembranelipids.AMP also has substantive hydrophobic moieties, which is different from CPP. Although there are no clear distinctions between CPP and AMP, we take apositiontodefineeachotherasCPPisatypeofpeptidethatspontaneously translocatethroughabiomembranewithoutdamagingthemembraneinteg- rity and cell viability. On the other hand,AMP kills microorganisms, as its Direct internalization Endocytosis (cytolysis) CPP Outer media Inner media Inner media (cytosol) (cytosol) Direct internalization Formation of endosome Fig.3 TranslocationofCPPthroughbiomembrane. TheImportanceofPlanarityforLipidBilayers 5 name tells, primarily by causing an efflux of cytoplasmic constituents through the transient or permanent pore. As the translocation processes, both CPP and AMP first absorb onto the biomembrane, then cause mem- branecurvaturedeformationfromflatbilayertothelocalnegativeGaussian curvature,andleadtoporeformation.Moredetailsofthistopologychange will be discussed later. Despite of plentiful reports on the CPP’s translocation through bio- membranes, a conclusive mechanism is yet to be established as Brock explained in his recent review [3]. We have taken CPP’s cytolysis as local andtemporalphasetransitionsofamphiphilicself-assemblyatitsliquidcrys- talstate(LC)bymodulatingthemoleculargeometryoflipidwithCPP[5,6] as shown in Fig. 4. In general, LC as highly ordered self-organization of amphiphiles has series of structures such as hexagonal, cubic, or lamellar. ThetopologyofeachLCdeterminedbythemoleculargeometryparameter oftheamphiphileiscalledsurfactantparameter(SP)definedbyEq.(1),and SP correspondsto the proportion of the hydrophilic part and hydrophobic part in the amphiphile as shown in Fig. 5 [7,8] SP¼v=ða(cid:3)lÞ (1) where a is the cross-sectional area per molecule at the hydrophilic– hydrophobic interface, l is the length of hydrophobic chains, and v is the volume of hydrophobic parts. SP correlates to the surface curvature of molecular assembly as a LC and can be changed by modulating the value ofa,l,orvthroughchangingthelevelofionization,hydration,solute,tem- perature,ormanyotherchemicalandphysicalmediators.SPisalsorelatedto the curvature of self-assembly by Eq. (2) for surfactant monolayer SP¼v=ða(cid:3)lÞ¼1+Hl+Kl2=3 (2) Local positive curvature change CPP Cytosol Fig. 4 Hypothesized mechanism for the cytosolic translocation of CPP through biomembrane.Alocalandtemporalphasetransitionofmembranelipidsatliquidcrys- talstate(LC)bymodulatingthemoleculargeometryoflipidbyCPPabsorption.
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