Encyclopedia of the Eye PDF
Preview Encyclopedia of the Eye
EDITOR-IN-CHIEF DarleneA.Dartt SchepensEyeResearchInstitute HarvardMedicalSchool Boston,MA,USA EDITORS BIOGRAPHIES DarleneA.Dartt Senior Scientist, the Harold F. Johnson Research Scholar, and Senior Scientist, Schepens Eye Research Institute; AssociateProfessor,DepartmentofOphthalmology,HarvardMedicalSchool,Boston,MA,USA.AttheSchepensEye ResearchInstituteDr.DarttservedastheActingDirectorofResearchandtheDirectorofScientificAffairsfor10years. She received her AB degree from Barnard College (Columbia University) in New York City and her PhD from the Department of Physiology at the University of Pennsylvania in Philadelphia. After postdoctoral fellowships at the UniversityofCopenhagen,Denmark,andTuftsUniversitySchoolofMedicineinBoston,Dr.DarttjoinedtheSchepens EyeResearchInstitutein1985.Herprimaryresearchinterestistheneuralregulationoftearproduction.Herresearch focusesonthesignalingpathwaysusedbynervesandgrowthfactorsinthelacrimalglandandconjunctivalgobletcellsto induce secretion and proliferation and howdysregulation of these pathways can lead to dry-eye syndromes in mouse modelsandhumans,particularlyafterrefractivesurgery.ShehasbeencontinuouslyfundedbyNIHsince1980forthis work.Dr.DarttdirectstheInstitute’sDepartmentofDefenseResearchProgram;shealsochairedfourMilitaryVision Research Symposia. She served on and chaired the ARVO Cornea Program Planning Committee and served on the Members in Training Committee. She was a founding member, a member of the organizing committee, and more recentlytreasurerfortheTearFilmandOcularSurfaceSociety.Shehasbeenontheorganizingcommitteeforthethree Biennial Cornea Conferences in Boston, MA. She served as Vice President for North America for the International SocietyforEyeResearchandaCouncilorfor theInternationalSocietyforContactLensWear.Sheisontheeditorial boardforInvestigativeOphthalmologyandVisualScience,ExperimentalEyeResearch,andTheOcularSurface.Sheservedonthe review panel for Fight For Sight and has been an ad hoc member of numerous NEI and NIAMS study sections. She receivedtheLewR.WassermanmeritawardbyResearchtoPreventBlindnessandisaGoldFellowofARVO. xvii xviii EditorsBiographies JosephC.Besharse Dr.BesharseiscurrentlytheMarvinWagnerProfessorandChairoftheDepartmentofCellBiology,Neurobiology,and Anatomyatthe Medical College of Wisconsin. He received his BA degree from Hendrix College (Conway,Arkansas) and MA and PhD degrees from Southern Illinois University. After a brief period on the faculty at Old Dominion UniversityinVirginia,hemovedtoColumbiaUniversityasaPostdoctoralFellowinRetinalResearch.Hewasappointed tothefacultyofDepartmentofCellBiologyandAnatomyatEmoryUniversitySchoolofMedicinein1977,becoming fullProfessorin1984.HeassumedthepositionofProfessorandChairofAnatomyandCellBiologyattheUniversity KansasSchoolofMedicine(KansasCity)in1989andmovedontohispresentpositionatMedicalCollegeofWisconsin in1997.Dr.Besharse’sresearchhasfocusedonmembraneturnover,circadianclocks,andmicrotubule-basedtransportin photoreceptors, with special attention to fundamental pathways that are disrupted in photoreceptor degenerative diseases.HisresearchhasbeencontinuouslyfundedbytheNationalInstitutesofHealthsince1978.Inadditiontohis roles as Department Chair and a Principal Investigator, he directs an NIH funded training program invision science. Dr.BesharseisapastmemberoftheeditorialboardsofInvestigativeOphthalmologyandVisionScienceandVisionNeuroscience and served as the Section Editor for the ‘Retina’section of Experimental Eye Research. He has served as a trustee of the AssociationforResearchinVisionandOphthalmology,andhasservedtwofulltermsasamemberoftwodifferentNIH studysections.AmonghisawardsaretheAlconResearchAwardforoutstandingresearchinVisionScience(1993),the Alumni Achievement Award from Southern Illinois University (1998) and the Distinguished Service Award from the AssociationforResearchinVisionandOphthalmology(2005).HeisaGoldFellowofARVO. RezaDana InadditiontohisroleasSeniorScientistandCo-DirectorofResearchattheSchepensEyeResearchInstitute,Dr.Reza Danaholds theClaesH.DohlmanChairinOphthalmologyatHarvardMedicalSchool.HeisDirectoroftheCornea and Refractive Surgery Service at the Massachusetts Eye and Ear, and serves as the Vice Chairman of the Harvard Department of Ophthalmology and Associate Chief of Ophthalmology for Academic Programs at Massachusetts Eye and Ear. After graduating Summa Cum Laude from St. Paul’s School in New Hampshire, he completed his bachelor’s (Phi Beta Kappa), graduate, and medical education at Johns Hopkins University. He performed his residency in EditorsBiographies xix Ophthalmology at the Illinois Eye and Ear Infirmary in Chicago, followed by a clinical fellowship in cornea and externaldiseases attheWillsEyeHospitalinPhiladelphia.Dr.Danathen completed additionalfellowship trainingin immunologyanduveitisattheMassachusettsEyeandEarInfirmary,andinocularandtransplantationimmunologyat theSchepensEyeResearchInstitute.Dr.DanaalsoholdsanMSdegreeinmanagementfromHarvardUniversity.Reza hasbeenamemberofthefull-timeHarvardfacultysince1995.Asaclinician-scientist,hehasaparticularinterestinthe molecular and cellular mechanisms of inflammation as they pertain to the ocular surface and anterior segment pathologies,includingdryeye,allergy,woundhealingresponses,andtransplantrejection. EDITORS AFFILIATIONS EDITOR-IN-CHIEF DeanBok(Retina) DarleneA.Dartt JulesSteinEyeInstitute SchepensEyeResearchInstitute DavidGeffenSchoolofMedicine HarvardMedicalSchool UniversityofCalifornia, Boston,MA LosAngeles,CA USA USA SENIOREDITORS PatriciaD’Amore(VascularBiology) JosephBesharse SchepensEyeResearchInstitute DepartmentofCellBiology, HarvardMedicalSchool NeurobiologyandAnatomy Boston,MA MedicalCollegeofWisconsin USA Milwaukee,WI USA HenryEdelhauser(OcularSurface) RezaDena EmoryUniversityEyeCenter MassachusettsEyeandEarInfirmary Atlanta,GA SchepensEyeResearchInstitute USA HarvardMedicalSchool 20StanifordStreet LindaMcLoon(Orbit) Boston,MA DepartmentsofOphthalmologyandNeuroscience USA UniversityofMinnesota Minneapolis,MN SECTIONEDITORS USA BarbaraBattelle(ComparativeEye) UniversityofFlorida WhitneyLaboratory JerryNiederkorn(ImmuneHomeostasisoftheEye) St.Augustine,FL UniversityofTexasSouthwesternMedicalCenter USA Dallas,TX USA DavidBeebe(Lens) WashingtonUniversitySchoolofMedicine SaintLouis,MO ThomasReh(RetinalDevelopemnt) USA UniversityofWashington SchoolofMedicine PeterBex(VisualOptics) Seattle,WA SchepensEyeResearchInstitute USA HarvardMedicalSchool Boston,MA Ernst Tamm (Aqueous Humor Dynamics and Optic USA Nerve) PaulBishop(Vitreous) InstituteofHumanAnatomyandEmbryology UniversityofManchester UniversityofRegensburg Manchester Regensburg UK Germany xxi PREFACE Eyeresearch hasgrownexplosivelyover theyears,andsimultaneouslywith itindividual scientistshave become more specialized.Coupledwiththisdichotomyisthewiderangeofdisciplinescoveredineyeresearch,fromvisualopticsto butterflyvisionandcornealstructuretobloodvesselgrowth.Thesevariablesnecessitateareferenceworkthatprovides concise,easilyunderstandableinformationonawiderangeofsubjects,andgivesguidanceonthemostimportantarticles toreadfromtheliterature.TheEncyclopediaoftheEyewascreatedtoserveassuchareferencework. Eacharticleintheencyclopediacontainstext(easilyunderstandablefor scientistsoutsidethefieldofthearticle),a numberofmultimediaandcolorfiguresthatillustratethemostimportantpointsofthearticle,andalistofreferencesto provide more in-depth information. Articles are primarilydirected at scientists looking for an entry point into a field tangentialtotheirspecialtyaswellasatgraduatestudentsandpostdoctoralfellowsineyeresearch.Thearticleswillbe especially useful to scientists designing introductory or generalized courses that cover diverse fields of eye research. Scientistswritingreviewarticlesorchapterswillalsofindtheencyclopediaarticlesespeciallyusefulasastartingpoint. Themanyintroductoryarticlesarewrittenatalevelthatuniversityundergraduatescanunderstand,butincludeenough information to satisfy the more advanced needs of graduate students and postdoctoral fellows. The more in-depth articlesonspecialistresearchareasareidealforpostdoctoralfellowsandexperiencedscientists.Theplentifulmultime- diafigureswillbeespeciallyhelpfulinunderstandingthemorecomplicatedpointsaswellasillustratingbasicprocesses andanatomy. TheformatoftheEncyclopediaoftheEyewasselectedbytheeditor-in-chiefincollaborationwithtwosenioreditors, Drs.RezaDanaandJosephC.Besharse.Wecoveredthefieldofeyeresearchwithfocusonspecialtiesinocularsurface, immunology, clinical research, and retina. To choose the articles to be included in the encyclopedia, we selected 11 majorareasof eyeresearch thatcover theentire field, withtheexception ofvisualprocessing.Theareasincludedthe ocularsurface,ocularimmunology,vascularbiology,orbit,aqueoushumor,lens,vitreous,retina,visualoptics,compara- tiveeyeresearch,andoculardevelopment.Thus,thescopeoftheencyclopediaincludesallfieldsofeyeresearch,except visual processing.Thelatter area has beenthe focus ofseveralrecent major referenceworks.Aprominent scientist in eachareawaschosenasasectioneditor.Eachofthesectioneditorsincollaborationwiththeeditor-in-chiefandsenior editorsdevelopedalistofarticlestocoverallimportanttopicsinthe11majorareas.Theintroductoryarticlesandthose within-depthdiscussionsofmorefocusedspecializedareasbothareincludedintheencyclopedia.Thesectioneditors thenselectedanexpertastheauthorforeacharticle.Finishedarticleswerereviewedbythreeindividuals,thesection editors, senior editors, and the editor-in-chief. Thus, the Encyclopedia of the Eye is a collaborative product of the most eminentscientistsineyeresearch. Eye research has become specialized and compartmentalized over theyears and, to a large extent, has split among anatomicallinesfromtheanteriortotheposterioreye.Forexample,cornearesearchandretinaresearchfollowdifferent pathsandarebasedondifferentunderlyingconceptsofcellularorganization.TheEncyclopediaoftheEyebridgesthisgap by including articles on all facets of eye research. In particular, we have included several topics usually not found in visionresearchreferenceworks.Theseincludeorbit,vitreous,comparativeeyeresearch,andoculardevelopment.These areashaveuniquespecializationsthatarefascinatingbythemselvesandhighlightdifferencesamongwell-knowntissue ororganismfunctions.Twoarticlesinparticular illustrate these points:oneonthemetabolismofintraocular muscles, which is unique to these nonfatiguing muscles; the other on color vision in butterflies, which allows butterflies to recognizeeachother. Importantly,studyoftheeyeanditsvariouscomponentsinvolvespathwaysfoundinothertissuesandprovidesinsight intothefunctionoftheothertissuesandorgans.Inaddition,eyeresearchhasbeenapacesettingdisciplineinmanyareas. Thus,theEncyclopediaoftheEyewillprovideknowledgeandinsighttoscientistsfromwide-rangingdisciplines,inaddition toeyeresearchitself.Forexample,eyeresearchhasprovidedseminaldiscoveriesthatimpactotherfields,suchasgene therapy, genetics of human disease, angiogenesis, and retinal signaling pathways. Several unique areas of eye research xxiii xxiv Preface includetheimmune-privilegedresponseoftheeye,thepreventionofbloodvesselgrowthinthecorneaandlens,andthe roleofaGTP-bindingproteincoupledcascadeinretinalphotoreceptorsignaling.ForallthesereasonstheEncyclopediaof theEyewillbevaluablefornonvisionbiomedicalscientists. Articles are arranged in alphabetical order. The best way to find an article is to look at the subject index, as both introductoryand morespecializedarticles are included inthe encyclopedia. It could be advantageousto startwith an introductory article to obtain knowledge about anatomy and the basic functions and then follow up with a more specialized,in-deptharticle.Furthermore,asubjectcouldbeincludedinanintroductory,broaderarticle andperhaps underanamethatisnotthereader’sfirstguess. An experienced, prompt, reliable,and thorough editorial staff is essential to the success of a major referencework. Theoutstandingcontributionoftheseniorandsectioneditorsareappreciativelyacknowledgedbytheeditor-in-chief. Theeditor-in-chiefwouldliketogratefullyacknowledgethescientificandeditorialhelpofMs.RobinHodges.Ms. Hodges provided invaluable help in keeping track of all the articles in their various stages of the editorial process, contactingauthorsto keepthem onschedule,and assisting the senior editorsDrs.RezaDanaand Joseph C.Besharse. WithouttheexperthelpofMs.Hodges,theEncyclopediaoftheEyewouldnotbetheoutstandingnewmajorreferencework thatitis.Theeditor-in-chiefwouldalsoliketoacknowledgetheextremelyhelpfuladviceofDr.RichardMasland,whose experiencedsuggestionswereinvaluableinsettinguptheeditorialteamaswellasDr.JoeG.Hollyfieldforhishelpful suggestionsfor sectioneditors. DARLENEA.DARTT Editor-in-Chief FOREWORD This is avery long book – if indeed it should be denominated as a book – but it does not need a long Foreword. The editors,whoarewellchosenscholarsoftheeye,haveassembledaremarkableseriesofover200articles,thatcoveravast landscapeonthebiologyoftheeye,anddosowithclarityanddepth.Theycover theeyefromthefronttotheback– fromthecornealtearfilmtotheaxonsoftheopticnerve. Thecoverageisjudicious.Eachsubjecthasanarticlethatreviewsitsbasicbiology,theseareaccompaniedbymore specializedtopics,commonlythoserelatingtodiseaseprocesses.Becausediseaseprocessesaresociallyimportant,they deserve this emphasis – and it should be noted that theyalso have things to teach about the basic biology. The basic biologyispresentedinsubstantialdepth,notonlyasatokengestureinthedirectionofthefundamentals.Thiskindof balance between basic biologyand clinical subjects is rarelyachieved, and the effect is that this is not onlya Google- readyreferenceworkbutonethatcanbereadserially,likeatraditionalexpositoryvolume. TheIntroductiontoanotherencyclopediainthisseriesmakesreferencetoWikipedia,thefreeform‘‘Encyclopediaof Everything’’ created bya communityof volunteer authors. The present encyclopedia has in common with Wikipedia greatbreadth,accessibilitytorollingrevision,andthefactthatitsauthorsarepaidalmostnothing.Itisacredittothe editors,totheauthors,andtoourscientificcommunitythatsomanytalentedpeoplewoulddevotetheirskillandeffort primarilyprobono–forthecommongood. And it is averygreat ‘‘good.’’ I hope and believe thatthiswill become astandard, universal source for information abouttheeye.InthisitalsohasacommunalitywithGoogleandtheInternet,butalsoamajordifference:theinformation that appears in the Encyclopedia of the Eye is curated, by the editors and by the authentic experts whom they have chosentowritethearticles.SoIseethisasanongoingventure,acoreresourcethatwillrollforwardwithnewauthors andneweditorsasthesciencemovesforward.ItisanambitiousventureandIwishitallsuccess. RichardH.Masland CharlesA.PappasProfessorofNeuroscience HarvardMedicalSchool xxv PERMISSION ACKNOWLEDGMENTS ThefollowingmaterialisreproducedwithkindpermissionofNaturePublishingGroup Fig1of‘‘DynamicImmunoregulatoryProcessesthatSustainImmunePrivilegeintheEye’’ Fig1of‘‘MolecularCompositionoftheVitreousandAgingChanges’’ Fig2of‘‘MolecularCompositionoftheVitreousandAgingChanges’’ Fig3of‘‘MolecularCompositionoftheVitreousandAgingChanges’’ Fig6of‘‘MolecularCompositionoftheVitreousandAgingChanges’’ Fig5of‘‘PathologicalRetinalAngiogenesis’’ http://www.nature.com/nature ThefollowingmaterialisreproducedwithkindpermissionofAmericanAssociationfortheAdvancement ofScience Fig2of‘‘ImmunopathogenesisofOnchocerciasis(RiverBlindness)’’ Fig6of‘‘InformationProcessing:HorizontalCells’’ Fig1of‘‘RodandConePhotoreceptorCells:Outer-SegmentMembraneRenewal’’ http://www.sciencemag.org ThefollowingmaterialisreproducedwithkindpermissionofTaylor&FrancisLtd Table2of‘‘DrugDeliverytoCorneaandConjunctiva:Esterase-andProtease-DirectedProdrugDesign’’ Fig6of‘‘DrugDeliverytoCorneaandConjunctiva:Esterase-andProtease-DirectedProdrugDesign’’ http://www.tandf.co.uk/journals
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