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Life - as a matter of fat : the emerging science of lipidomics PDF

273 Pages·2005·5.166 MB·English
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the frontiers collection the frontiers collection SeriesEditors: D.Dragoman M.Dragoman A.C.Elitzur M.P.Silverman J.Tuszynski H.D.Zeh Thebooksinthiscollectionaredevotedtochallengingandopenproblemsattheforefront of modern physics and related disciplines, including philosophical debates. In contrast totypicalresearchmonographs,however,theystrivetopresenttheirtopicsinamanner accessiblealsotoscientificallyliteratenon-specialistswishingtogaininsightintothedeeper implicationsandfascinatingquestionsinvolved.Takenasawhole,theseriesreflectsthe needforafundamentalandinterdisciplinaryapproachtomodernscience.Itisintendedto encouragescientistsinallareastoponderoverimportantandperhapscontroversialissues beyond their own speciality. Extending from quantum physics and relativity to entropy, time and consciousness – the Frontiers Collection will inspire readers to push back the frontiersoftheirownknowledge. QuantumMechanicsandGravity ByM.Sachs Mind,MatterandQuantumMechanics ByH.Stapp Quantum–ClassicalCorrespondence ByA.O.Bolivar Quantum–ClassicalAnalogies ByD.DragomanandM.Dragoman KnowledgeandtheWorld: ChallengesBeyondtheScienceWars EditedbyM.Carrier,J.Roggenhofer, G.Ku¨ppers,andPh.Blanchard Life–AsaMatterofFat ByO.G.Mouritsen Serieshomepage–springeronline.com Ole G. Mouritsen LIFE – AS A MATTER OF FAT The Emerging Science of Lipidomics 123 Prof.OleG.Mouritsen UniversityofSouthernDenmark,MEMPHYS–CenterforBiomembranePhysics, PhysicsDepartment,Campusvej55,5230OdenseM,Denmark email:[email protected] SeriesEditors: Prof.DanielaDragoman UniversityofBucharest,PhysicsFaculty,SolidStateChair,POBoxMG-11, 76900Bucharest,Romania email:[email protected] Prof.MirceaDragoman NationalResearchandDevelopmentInstituteinMicrotechnology,POBox38-160, 023573Bucharest,Romania email:[email protected] Prof.AvshalomC.Elitzur Bar-IlanUniversity,UnitofInterdisciplinaryStudies, 52900Ramat-Gan,Israel email:[email protected] Prof.MarkP.Silverman DepartmentofPhysics,TrinityCollege, Hartford,CT06106,USA email:[email protected] Prof.JackTuszynski UniversityofAlberta,DepartmentofPhysics,Edmonton,AB, T6G2J1,Canada email:[email protected] Prof.H.DieterZeh UniversityofHeidelberg,InstituteofTheoreticalPhysics,Philosophenweg19, 69120Heidelberg,Germany email:[email protected] ISSN1612-3018 ISBN3-540-23248-6SpringerBerlinHeidelbergNewYork LibraryofCongressControlNumber:2004115728 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,recitation,broadcasting, reproductiononmicrofilmorinanyotherway,andstorageindatabanks.Duplicationofthispublicationor partsthereofispermittedonlyundertheprovisionsoftheGermanCopyrightLawofSeptember9,1965,in itscurrentversion,andpermissionforusemustalwaysbeobtainedfromSpringer.Violationsareliableto prosecutionundertheGermanCopyrightLaw. SpringerisapartofSpringerScience+BusinessMedia springeronline.com ©Springer-VerlagBerlinHeidelberg2005 PrintedinGermany Theuseofgeneraldescriptivenames,registerednames,trademarks,etc.inthispublicationdoesnotimply, evenintheabsenceofaspecificstatement,thatsuchnamesareexemptfromtherelevantprotectivelawsand regulationsandthereforefreeforgeneraluse. Typesettingbytheauthor DataconversionbyTechBooks CoverdesignbyKünkelLopka,WerbeagenturGmbH,Heidelberg Printedonacid-freepaper 54/3141/jl-543210 To Myer Bloom, mentor in science and life Acknowledgments Theresearchworkunderlyingthepictureoffats,lipids,andmembranesadvo- catedinthepresentbookderivesfrommanydifferentscientistsandstudents in several laboratories across the world. The specific examples and data pre- sentedarehoweverbiasedtowardtheworkcarriedoutbytheauthorandhis collaborators over the last almost twenty years. Therefore, the book should not be considered an authoritative monograph but more a personal perspec- tive on a diverse and rapidly expanding field of science. The work has been supported by a number of public and private funding agencies, notably the Danish Natural Science Research Council, the Danish Technical Research Council, the Danish Medical Research Council via the Center for Drug De- sign and Transport, the Villum Kann Rasmussen Foundation, the Carlsberg Foundation,theVeluxFoundation,theHasselbladFoundation,andtheDan- ish National Research Foundation. The author has over the years benefited from stimulating interaction and fruitful collaboration with a large number of colleagues and students, in particular from the Center for Biomembrane Physics (MEMPHYS) and from the Canadian Institute for Advanced Re- search’s program on the Science of Soft Surfaces and Interfaces under the directorship of Professor Myer Bloom. The author is greatly indebted to the following: Thomas Andresen, Luis Bagatolli, Rogert Bauer, Thomas Baek- mark, Gerhand Besold, Rodney Biltonen, Thomas Bjørnholm, Myer Bloom, DavidBoal,ThomasHøngerCallisen,RobertCantor,BerndDammann,Jes- perDavidsen,LarsDuelund,EvanEvans,SvenFrøkjær,TamirGil,Henriette Gilhøj, Per Lyngs Hansen, Jonas Henriksen, Pernille Høyrup, John Hjort Ipsen, Ask Jacobsen, Morten Ø. Jensen, Claus Jeppesen, Kent Jørgensen, Thomas Kaasgaard, Danielle Keller, Lars Kildemark, Dennis Kim, Paavo Kinnunen,BeateKlo¨sgen,PerKnudsen,MohamedLaradji,ChadLeidy,Jes- per Lemmich, Ling Miao, Kell Mortensen, Mohan Narla, David Needham, Morten Nielsen, Jaan Noolandi, Adrian Parsegian, Tina Pedersen, Gu¨nther Peters, Amy Rowat, Jens Risbo, Mads C. Sabra, Erich Sackmann, Adam C. Simonsen, Maria M. Sperotto, Jenifer Thewalt, Christa Trandum, Ilpo Vat- tulainen, Peter Westh, Matthias Weiss, Michael Wortis, and Martin Zucker- mann. Olaf Sparre Andersen is thanked for collaboration concerning a con- ference book on which part of Chap. 7 is based. Michael Crawford shared with me his insight into the relationships between lipids, nutrition, brain VIII Acknowledgments evolutionandhumanhealth.IamindebtedtoRodneyCotterillforusefulad- viseconcerningthebooktitle.FabiennePiegay,LaurenceSideau,andMartin Bennetzen are thanked for thorough and critical reading of the manuscript. I have drawn on the help of a number of colleagues and the permission to use their graphic material for preparing many of the figures of this book. A list of the sources for the figures is provided at the end. Tove Nyberg has kindly drawn many of the figures. Jonas Drotner Mouritsen is thanked for creative suggestions and helpful advice regarding the preparation and design of many of the illustrations. The Villum Kann Rasmussen Foundation is gratefully acknowledged for a generous grant that made it financially possible to arrange for the leave of absence that was required to get free time to start working on the book. Institutionen San Cataldo, where the first version of the book manuscript emerged in the tranquillity of Cell No. 6 of the old convent San Cataldo overlooking the beautiful Amalfi Bay in Southern Italy, is thanked for its hospitality during the month of April 2001. Contents Prologue: Lipidomics – A Science Beyond Stamp Collection .. 1 Part I The Overlooked Molecules 1 Life from Molecules....................................... 9 1.1 The Three Kingdoms of Life ............................. 9 1.2 The Molecules of Life ................................... 10 1.3 The Post-Genomic Era.................................. 16 1.4 A Call for Physics ...................................... 19 2 Head and Tail ............................................ 23 2.1 Fat Family: Fats and Fatty Acids......................... 23 2.2 The Polar Lipids – Both Head and Tail ................... 26 2.3 Cholesterol – A Lipid of Its Own ......................... 28 2.4 Strange Lipids ......................................... 29 2.5 Lipid Composition of Membranes......................... 31 3 Oil and Water ............................................ 33 3.1 Water – The Biological Solvent........................... 33 3.2 The Hydrophobic Effect ................................. 34 3.3 Mediating Oil and Water ................................ 35 3.4 Self-Assembly and the Lipid Aggregate Family ............. 37 3.5 Plucking Lipids ........................................ 40 4 Lipids Speak the Language of Curvature.................. 43 4.1 How Large Is a Lipid Molecule? .......................... 43 4.2 Lipid Molecules Have Shape ............................. 46 4.3 Lipid Structures with Curvature.......................... 46 4.4 Microorganisms’ Sense of Curvature ...................... 51 5 A Matter of Softness ..................................... 53 5.1 Soft Matter............................................ 53 5.2 Soft Interfaces ......................................... 53 5.3 Forces Between Soft Interfaces ........................... 58 X Contents 5.4 Lipid Membranes are Really Soft ......................... 59 6 Soft Shells Shape Up ..................................... 63 6.1 Bending Interfaces...................................... 63 6.2 Spontaneous Curvature ................................. 66 6.3 Shaping Membranes .................................... 67 6.4 Red Blood Cells Shape Up............................... 68 7 Biological Membranes – Models and Fashion ............. 73 7.1 What Is a Model? ...................................... 73 7.2 Brief History of Membrane Models........................ 74 7.3 Do We Need a New Membrane Model?.................... 76 7.4 Theoretical and Experimental Model Systems .............. 78 Part II Lipids Make Sense 8 Lipids in Bilayers – A Stressful and Busy Life ............................................ 81 8.1 Trans-bilayer Structure.................................. 81 8.2 The Lateral Pressure Profile ............................. 83 8.3 How Thick Are Membranes? ............................. 85 8.4 Lively Lipids on the Move ............................... 86 9 The More We Are Together .............................. 91 9.1 Phase Transitions Between Order and Disorder............. 91 9.2 Lipids Have Phase Transitions ........................... 94 9.3 Mixing Different Lipids.................................. 98 9.4 Cholesterol Brings Lipids to Order........................ 100 10 Lipids in Flatland......................................... 105 10.1 Gases, Liquids, and Solids in Two Dimensions.............. 105 10.2 Langmuir and Langmuir-Blodgett Films................... 107 10.3 Pattern Formation in Lipid Monolayers ................... 110 10.4 Lipids Make the Lung Work ............................. 112 11 Social Lipids.............................................. 117 11.1 Lateral Membrane Structure ............................. 117 11.2 Imaging Lipid Domains ................................. 121 11.3 Lipid Rafting .......................................... 124 11.4 Domains and Rafts Carry Function ....................... 127

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