MitochondriaStructureandFunction-TheoreticalDetails Inanattempttobeconciseandunderstandable,introductorylevelcoursesandtextbooksfrequentlypresentconceptsthataretechnicallycorrect,butleadtomisconceptionsonthepartofthestudentbecausetheyomittoomuch.IndiscussionsaboutmitochondriastudentsfrequentlycomeawaywithasuperficialunderstandingofthetruenatureofKrebs"cycle,electrontransport,respiratorycontrol,andoxidativephosphorylation. Thesepagesweredesignedtosupplementlaboratoryworkwithmitochondriabyprovidingbackgroundinasmuchdetailasthestudentmightwish.Studentsatanylevelarelikelytoencountertermswithwhichtheyareunfamiliar.Aglossaryoftermsisincludedinthisprojectforyourconvenience.Abriefoverviewofmitochondriastructureandfunctionispresentedjusttogetyoustarted.Youcanthenwindyourwaythroughthemainpathofdetailedinformation. Mitochondriaarebacteria-sizedorganelles,foundinthecytoplasmofvirtuallyalleukaryoticcells.Theyareespeciallyabundantincellsandpartsofcellsthatareassociatedwithactiveprocesses.Forexample,inflagellatedprotozoaorinmammaliansperm,mitochondriaareconcentratedaroundthebaseoftheflagellumorflagella.Incardiacmuscle,mitochondriasurroundthecontractileelements.Hummingbirdflightmuscleisoneoftherichestsourcesofmitochondriaknown.Thus,fromtheirdistributionaloneonesUSPectsthattheyareinvolvedinenergyproduction. Weknownowthatmulticellularorganismsprobablycouldnotexistwithoutmitochondria.Mitochondriamakeefficientuseofnutrientmolecules,requiringoxygenintheprocess.Theyare,infact,whyweneedoxygenatall. Thedouble-membranedmitochondrioncanbelooselydescribedasalargewrinkledbagpackedinsideofasmaller,unwrinkledbag.Thetwomembranescreatedistinctcompartmentswithintheorganelle,andarethemselvesverydifferentinstructureandinfunction. Theoutermembraneisarelativelysimplephospholipidbilayer,containingproteinstructurescalledporinswhichrenderitpermeabletomoleculesofabout10kilodaltonsorless(thesizeofthesmallestproteins).Ions,nutrientmolecules,ATP,ADP,etc.canpassthroughtheoutermembranewithease. Theinnermembraneisfreelypermeableonlytooxygen,carbondioxide,andwater.Itsstructureishighlycomplex,includingallofthecomplexesoftheelectrontransportsystem,theATPsynthetasecomplex,andtransportproteins.Thewrinkles,orfolds,areorganizedintolamillae(layers),calledthecristae(singlular:crista).Thecristaegreatlyincreasethetotalsurfaceareaoftheinnermembrane.Thelargersurfaceareamakesroomformanymoreoftheabove-namedstructuresthaniftheinnermembranewereshapedliketheoutermembrane. Themembranescreatetwocompartments.Theintermembranespace,asimplied,istheregionbetweentheinnerandoutermembranes.Ithasanimportantroleintheprimaryfunctionofmitochondria,whichisoxidativephosphorylation. ThematrixcontainstheenzymesthatareresponsIBLeforthecitricacidcyclereactions.Thematrixalsocontainsdissolvedoxygen,water,carbondioxide,therecyclableintermediatesthatserveasenergyshuttles,andmuchmore.Becauseofthefoldsofthecristae,nopartofthematrixisfarfromtheinnermembrane.Thereforematrixcomponentscanquicklyreachinnermembranecomplexesandtransportproteins. Electronmicrographshaverevealedthethreedimensionalstructureofmitochondria.However,sincemicrographsarethemselvestwodimensional,theirinterpretationcanbemisleADIng. Textsfrequentlyshowapictureofa"typical"mitochondrionasabacteria-sizedellipsoid(perhaps0.5by1micrometer).However,theyvarywidelyinshapeandsize.Electronmicrographsseldomshowsuchvariation,becausetheyaretwo-dimensionalimages. Isolatedmitochondria,suchasfromhomogenizedmuscletissue,showaroundedappearanceinelectronmicrographs,implyingthatmitochondriaaresphericalorganelles. Mitochondriainsitucanbefreeinthecytoplasmorpackedinamongmorerigidstructures,suchasamongthemyofibrilsofcardiacmuscletissue.Incellssuchasmuscle,itisclearthatmitochondriaarenotspherical,andoftenarenotevenellipsoid.Insometissues,themitochondriaarealmostfilamentous,acharacteristicthattwodimensionalmicrographsmayfailtoreveal. Aplanarsectioncutsthroughoneorseveralpartsoftheorganelle,makingitappearthatthereismorethanone.Theimageweseeofacircularorellipsoidalorganellemaydisguisethetruenatureofthemitochondrion.Structure