Encyclopedia of Biophysics PDF
Preview Encyclopedia of Biophysics
Encyclopedia of Biophysics Gordon C. K. Roberts Editor Encyclopedia of Biophysics With1597Figuresand131Tables Editor GordonC.K.Roberts HonoraryProfessorofBiochemistry DepartmentofBiochemistry UniversityofLeicester Leicester,UK ISBN978-3-642-16711-9 ISBN978-3-642-16712-6(eBook) ISBN978-3-642-16713-3(printandelectronicbundle) DOI10.1007/978-3-642-16712-6 SpringerHeidelbergNewYorkDordrechtLondon LibraryofCongressControlNumber:2012949366 FirstEdition:CopyrightEuropeanBiophysicalSocieties’Association(EBSA).Englisheditionpublished bySpringer-VerlagBerlinHeidelberg2013.Allrightsreserved. #EuropeanBiophysicalSocieties’Association(EBSA)2013 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartofthe materialisconcerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,recitation, broadcasting,reproductiononmicrofilmsorinanyotherphysicalway,andtransmissionorinformation storageandretrieval,electronicadaptation,computersoftware,orbysimilarordissimilarmethodology nowknownorhereafterdeveloped.Exemptedfromthislegalreservationarebriefexcerptsinconnection withreviewsorscholarlyanalysisormaterialsuppliedspecificallyforthepurposeofbeingenteredand executed on a computer system, for exclusive use by the purchaser of the work. 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Printedonacid-freepaper SpringerispartofSpringerScienceþBusinessMedia(www.springer.com) To Hilary Preface PerhapsthefirstissueinintroducinganEncyclopaediaofBiophysicsisthequestion “What is Biophysics?”. The answer to this has varied considerably over time and betweendifferent‘biophysicists’,particularlydependingonwhethertheyhavecome tobiophysicsfromphysicsorphysiology.Forthepresentpurposes,wehaveadopted thedefinitiongivenbytheNobelPrize-winningphysiologistA.V.Hill(Science124, 1233, 1956): Biophysics is “the study of biological function, organization, and structure by physical and physicochemical ideas and methods.” – to which one would now add methods of mathematical analysis and computer modelling. An important feature of Hill’s definition is that it includes both physical methods and physical ideas. The astonishing developments in physical methods over the last hundredyearshavemadethemubiquitousinbiologicallaboratories–andindeedin hospitals.However,toquoteHillagain“theemploymentofphysicalinstrumentsin a biological laboratory does not make one a biophysicist - otherwise any user of amicroscope,abalance.....orapHmeterwoulddropautomaticallyintothatclass.” Beyondsimplytheuseofphysicalmethods,itisthecombinationofbothphysicaland biologicalideas,intuitionsandexperiencethatmakesaBiophysicist. The application of physical methods in biology has a very long history. For example,AntonievanLeeuwenhoek(1632–1723)developedimprovedmicroscope lenses which allowed him to be the first to observe and describe single-celled organisms–thebeginningofmicrobiology.The18thcenturysawmuchspeculation about ‘animal electricity’, culminating in the 1780s in Luigi Galvani’s famous experiments on the relation of electricity to muscle contraction. The middle of the 19th century saw what was probably the first explicitly biophysical programme of research,inwhichagroupofphysiologists(duBois-Reymond,Ludwig,vonBr€ucke, von Helmholtz) proclaimed their intention to ‘reduce physiology to physics and chemistry’.OfcoursethestateofphysicsatthetimewassuchastomakeBiophysics averyprematureventure,butthisgroupdidmakeimportantdiscoveries,notablyin physiologicalopticsandelectrophysiology,usingphysicalmethods. Anassociateof this group, Adolf Fick – well-known for his law of diffusion – published what is probablythefirstbiophysicstext,DieMedizinischePhysik(1856). Inthe20thand21stcenturiesthere hasbeenadramatic floweringofBiophysics. Theearlypartof20thcenturysawthedevelopmentofphysicaltoolswhicharenow familiarinbiology–fromX-raydiffraction(vonLaue,Bragg)totheultracentrifuge (Svedberg)andtheelectronmicroscope(Knoll&Ruska)–andthismethodological development continues apace, notably with single-molecule techniques. The 1920s and1930ssawthebeginningsofphysicochemical(Cohn,Edsall,Linderstrøm-Lang) and structural (Astbury, Bernal, Hodgkin, Perutz) studies of proteins. At the same vii viii Preface time, the first Departments or Institutes of Biophysics began to be established. In Germany these were commonly focussed on the study of radiation effects on organisms, while in the USA they concentrated largely on physiology. Many of theselaterexpandedintootherareasofBiophysics,andmanymoreDepartmentsof Biophysics – and Biophysical Societies – were established in the USA, Europe and Israel in the 1940s and 1950s. The power of the Biophysical approach was demonstrated unequivocally in the 1950s with the determination of the first three- dimensional structure of a protein, the structure of DNA and the Hodgkin-Huxley modeloftheactionpotentialinnerves. Overthelast50years,Biophysicshascontinuedtodevelopatanastonishingpace. Biophysicistsstudylifeateverylevel,fromatomsandmoleculestocells,organisms, andenvironments.MolecularandCellularBiophysics:Thisisperhapsthepredom- inantstrandofmodernbiophysics.Itincludes,forexample,structural,functionaland simulation studies of macromolecules and macromolecular assemblies of ever- increasing complexity and imaging of cells at ever-increasing resolution. Applied Biophysics:Theapplicationsofbiophysicalmethodsareperhapsmostnotableinthe areaofmedicalimaging-includingX-rayCATscans,magneticresonanceimaging, positron emission tomography and ultrasound scans. Therapeutic applications include radiation therapy of increasing sophistication and cardiac defibrillators. Inadditiontomedicalapplications,therearenowincreasingandexcitingapplications innanotechnology.EnvironmentalBiophysics:Formanyyearsanimportantareaof biophysics has been the study of the effects of ionising - and indeed non-ionising - radiation onorganisms. Another keyarea ofconsiderable current importance is the developmentofmathematicalmodelsof,e.g.,heatandmasstransferatthelevelof organismsandecosystems. ThisEncyclopediaisintendedtoprovidearesourcebothforbiophysicistsinter- estedinapproachesoutsidetheirimmediatesub-disciplineandforpeoplecomingto biophysics from either the physical or biological direction. The emphasis is very much onmolecular andcellular biophysics,butsome discussion ofimaging andof nanotechnologyisincluded.Justasthereisoverlapbetweenchemistryandphysics, sothereisoverlapbetweenbiophysicsandbiochemistry;ourfocusisofcourseonthe techniques and uses of biophysics, but biochemical context is included where appropriate. TheEncyclopediaconsistsoftwokindsofentries,SystemsandTechniques. • IntheSystemssections,biophysicalapproachestoparticularbiologicalsystemsor problems–fromproteinstructuretomembranes,ionchannelsandreceptors–are described. These sections, which have an emphasis on the integration of the differenttechniques,thereforeprovideanentryintoBiophysicsfromthebiolog- icalmorethanfromthetechnique-orientedphysicsdirection. • IntheTechniquessections,eachofthewiderangeofmethodswhichfallunderthe headingofBiophysicsareexplainedindetail,togetherwiththeirstrengthsandthe limitations of the information each provides. Experimental techniques covered range from diffraction, through a wide range of spectroscopic methods (X-ray, optical, EPR, NMR), kinetics, thermodynamics and hydrodynamics, to imaging (from electron microscopy to live cell imaging and MRI). The important and increasingly powerful computational, modelling and simulation approaches are alsoincluded. Each of the Sections includes concise introductions to the major concepts and methods, and outlines of more specific topics, in each case with links to a limited Preface ix number of carefully selected key reviews and/or papers in the scientific literature. Extensive cross-referencing (hyperlinks in the online version) between different articlesallowsaccesstorelatedtopicsinauser-friendlymanner. TheonlineversionoftheEncyclopediaisintendedtodevelopcontinuously,both bytheupdatingofexistingentriesandbytheadditionofnewarticles.Inthiswayit willprovidetheresourcestohelpbothpractisingbiophysicistsandnewcomerstothe fieldtokeeppacewiththecontinuingandexcitingdevelopmentsinBiophysics. Leicester GordonC.K.Roberts UK May2012
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