Tendon - Wikipedia

A tendon or sinew is a tough, high-tensile-strength band of dense fibrous connective tissue that connects muscle to bone. It is able to efficiently transmit ... Tendon FromWikipedia,thefreeencyclopedia Jumptonavigation Jumptosearch Typeoftissuethatconnectsmuscletobone Forotheruses,seeTendon(disambiguation). "Sinew"redirectshere.ForthesongbyPurityRing,seeWomb(album). TendonTheAchillestendon,oneofthetendonsinthehumanbody(fromGray'sAnatomy,1sted.,1858)Micrographofapieceoftendon;H&EstainDetailsIdentifiersLatintendoMeSHD013710THH3.03.00.0.00020FMA9721Anatomicalterminology[editonWikidata] Atendonorsinewisatough,high-tensile-strengthbandofdensefibrousconnectivetissuethatconnectsmuscletobone.Itisabletoefficientlytransmitthemechanicalforcesofmusclecontractiontotheskeletalsystemwithoutsacrificingitsabilitytowithstandsignificantamountsoftension. Tendonsaresimilartoligaments;botharemadeofcollagen.Ligamentsconnectonebonetoanother,whiletendonsconnectmuscletobone. Contents 1Structure 1.1Extracellularmatrix 1.1.1Ultrastructureandcollagensynthesis 1.2Tenocytes 2Functions 2.1Mechanics 2.2Healing 2.2.1Effectsofactivityonhealing 3Societyandculture 3.1Culinaryuses 4Clinicalsignificance 4.1Injury 5Otheranimals 6Seealso 7References Structure[edit] Histologically,tendonsconsistofdenseregularconnectivetissue.Themaincellularcomponentoftendonsarespecializedfibroblastscalledtendoncells(tenocytes).Tenocytessynthesizetheextracellularmatrixoftendons,abundantindenselypackedcollagenfibers.Thecollagenfibersareparalleltoeachotherandorganizedintotendonfascicles.Individualfasciclesareboundbytheendotendineum,whichisadelicatelooseconnectivetissuecontainingthincollagenfibrils[1][2]andelasticfibres.[3]Groupsoffasciclesareboundedbytheepitenon,whichisasheathofdenseirregularconnectivetissue.Thewholetendonisenclosedbyafascia.Thespacebetweenthefasciaandthetendontissueisfilledwiththeparatenon,afattyareolartissue.[4]NormalhealthytendonsareanchoredtobonebySharpey'sfibres. Extracellularmatrix[edit] Thedrymassofnormaltendons,whichmakesup30-45%oftheirtotalmass,iscomposedof: 60-85%collagen 60-80%collagenI 0-10%collagenIII 2%collagenIV smallamountsofcollagensV,VI,andothers 15-40%non-collagenousextracellularmatrixcomponents,including: 3%cartilageoligomericmatrixprotein, 1-2%elastin, 1–5%proteoglycans, 0.2%inorganiccomponentssuchascopper,manganese,andcalcium.[5][6][7][8] WhiletypeIcollagenmakesupmostofthecollagenintendon,manyminorcollagensarepresentthatplayvitalrolesinpropertendondevelopmentandfunction.TheseincludetypeIIcollageninthecartilaginouszones,typeIIIcollageninthereticulinfibresofthevascularwalls,typeIXcollagen,typeIVcollageninthebasementmembranesofthecapillaries,typeVcollageninthevascularwalls,andtypeXcollageninthemineralizedfibrocartilageneartheinterfacewiththebone.[5][9] Ultrastructureandcollagensynthesis[edit] Collagenfibrescoalesceintomacroaggregates.Aftersecretionfromthecell,cleavedbyprocollagenN-andC-proteases,thetropocollagenmoleculesspontaneouslyassembleintoinsolublefibrils.Acollagenmoleculeisabout300 nmlongand1–2 nmwide,andthediameterofthefibrilsthatareformedcanrangefrom50–500 nm.Intendons,thefibrilsthenassemblefurthertoformfascicles,whichareabout10 mminlengthwithadiameterof50–300μm,andfinallyintoatendonfibrewithadiameterof100–500μm.[10] Thecollagenintendonsareheldtogetherwithproteoglycan(acompoundconsistingofaproteinbondedtoglycosaminoglycangroups,presentespeciallyinconnectivetissue)componentsincludingdecorinand,incompressedregionsoftendon,aggrecan,whicharecapableofbindingtothecollagenfibrilsatspecificlocations.[11]Theproteoglycansareinterwovenwiththecollagenfibrils –theirglycosaminoglycan(GAG)sidechainshavemultipleinteractionswiththesurfaceofthefibrils –showingthattheproteoglycansareimportantstructurallyintheinterconnectionofthefibrils.[12]ThemajorGAGcomponentsofthetendonaredermatansulfateandchondroitinsulfate,whichassociatewithcollagenandareinvolvedinthefibrilassemblyprocessduringtendondevelopment.Dermatansulfateisthoughttoberesponsibleforformingassociationsbetweenfibrils,whilechondroitinsulfateisthoughttobemoreinvolvedwithoccupyingvolumebetweenthefibrilstokeepthemseparatedandhelpwithstanddeformation.[13]Thedermatansulfatesidechainsofdecorinaggregateinsolution,andthisbehaviorcanassistwiththeassemblyofthecollagenfibrils.Whendecorinmoleculesareboundtoacollagenfibril,theirdermatansulfatechainsmayextendandassociatewithotherdermatansulfatechainsondecorinthatisboundtoseparatefibrils,thereforecreatinginterfibrillarbridgesandeventuallycausingparallelalignmentofthefibrils.[14] Tenocytes[edit] Thetenocytesproducethecollagenmolecules,whichaggregateend-to-endandside-to-sidetoproducecollagenfibrils.Fibrilbundlesareorganizedtoformfibreswiththeelongatedtenocytescloselypackedbetweenthem.Thereisathree-dimensionalnetworkofcellprocessesassociatedwithcollageninthetendon.Thecellscommunicatewitheachotherthroughgapjunctions,andthissignallinggivesthemtheabilitytodetectandrespondtomechanicalloading.[15]Thesecommunicationshappenbytwoproteinsessentialy :connexin43,presentwherethecellsprocessesmeetandincellbodiesconnexin32,presentonlywheretheprocessesmeet.[16] Bloodvesselsmaybevisualizedwithintheendotendonrunningparalleltocollagenfibres,withoccasionalbranchingtransverseanastomoses. Theinternaltendonbulkisthoughttocontainnonervefibres,buttheepitenonandparatenoncontainnerveendings,whileGolgitendonorgansarepresentatthemyotendinousjunctionbetweentendonandmuscle. Tendonlengthvariesinallmajorgroupsandfrompersontoperson.Tendonlengthis,inpractice,thedecidingfactorregardingactualandpotentialmusclesize.Forexample,allotherrelevantbiologicalfactorsbeingequal,amanwithashortertendonsandalongerbicepsmusclewillhavegreaterpotentialformusclemassthanamanwithalongertendonandashortermuscle.Successfulbodybuilderswillgenerallyhaveshortertendons.Conversely,insportsrequiringathletestoexcelinactionssuchasrunningorjumping,itisbeneficialtohavelongerthanaverageAchillestendonandashortercalfmuscle.[17] Tendonlengthisdeterminedbygeneticpredisposition,andhasnotbeenshowntoeitherincreaseordecreaseinresponsetoenvironment,unlikemuscles,whichcanbeshortenedbytrauma,useimbalancesandalackofrecoveryandstretching.[18]Inaddictiontendonsallowmusclestobeatanoptimaldistancefromthesitewheretheyactivelyengageinmovement,passingthroughregionswherespaceispremium,likethecarpaltunnel.[16] Functions[edit] Magnifiedviewofatendon Traditionally,tendonshavebeenconsideredtobeamechanismbywhichmusclesconnecttoboneaswellasmusclesitself,functioningtotransmitforces.Thisconnectionallowstendonstopassivelymodulateforcesduringlocomotion,providingadditionalstabilitywithnoactivework.However,overthepasttwodecades,muchresearchhasfocusedontheelasticpropertiesofsometendonsandtheirabilitytofunctionassprings.Notalltendonsarerequiredtoperformthesamefunctionalrole,withsomepredominantlypositioninglimbs,suchasthefingerswhenwriting(positionaltendons)andothersactingasspringstomakelocomotionmoreefficient(energystoringtendons).[19]Energystoringtendonscanstoreandrecoverenergyathighefficiency.Forexample,duringahumanstride,theAchillestendonstretchesastheanklejointdorsiflexes.Duringthelastportionofthestride,asthefootplantar-flexes(pointingthetoesdown),thestoredelasticenergyisreleased.Furthermore,becausethetendonstretches,themuscleisabletofunctionwithlessorevennochangeinlength,allowingthemuscletogeneratemoreforce. Themechanicalpropertiesofthetendonaredependentonthecollagenfiberdiameterandorientation.Thecollagenfibrilsareparalleltoeachotherandcloselypacked,butshowawave-likeappearanceduetoplanarundulations,orcrimps,onascaleofseveralmicrometers.[20]Intendons,thecollagenfibreshavesomeflexibilityduetotheabsenceofhydroxyprolineandprolineresiduesatspecificlocationsintheaminoacidsequence,whichallowstheformationofotherconformationssuchasbendsorinternalloopsinthetriplehelixandresultsinthedevelopmentofcrimps.[21]Thecrimpsinthecollagenfibrilsallowthetendonstohavesomeflexibilityaswellasalowcompressivestiffness.Inaddition,becausethetendonisamulti-strandedstructuremadeupofmanypartiallyindependentfibrilsandfascicles,itdoesnotbehaveasasinglerod,andthispropertyalsocontributestoitsflexibility.[22] Theproteoglycancomponentsoftendonsalsoareimportanttothemechanicalproperties.Whilethecollagenfibrilsallowtendonstoresisttensilestress,theproteoglycansallowthemtoresistcompressivestress.Thesemoleculesareveryhydrophilic,meaningthattheycanabsorbalargeamountofwaterandthereforehaveahighswellingratio.Sincetheyarenoncovalentlyboundtothefibrils,theymayreversiblyassociateanddisassociatesothatthebridgesbetweenfibrilscanbebrokenandreformed.Thisprocessmaybeinvolvedinallowingthefibriltoelongateanddecreaseindiameterundertension.[23]However,theproteoglycansmayalsohavearoleinthetensilepropertiesoftendon.Thestructureoftendoniseffectivelyafibrecompositematerial,builtasaseriesofhierarchicallevels.Ateachlevelofthehierarchy,thecollagenunitsareboundtogetherbyeithercollagencrosslinks,ortheproteoglycans,tocreateastructurehighlyresistanttotensileload.[24]Theelongationandthestrainofthecollagenfibrilsalonehavebeenshowntobemuchlowerthanthetotalelongationandstrainoftheentiretendonunderthesameamountofstress,demonstratingthattheproteoglycan-richmatrixmustalsoundergodeformation,andstiffeningofthematrixoccursathighstrainrates.[25]Thisdeformationofthenon-collagenousmatrixoccursatalllevelsofthetendonhierarchy,andbymodulatingtheorganisationandstructureofthismatrix,thedifferentmechanicalpropertiesrequiredbydifferenttendonscanbeachieved.[26]Energystoringtendonshavebeenshowntoutilisesignificantamountsofslidingbetweenfasciclestoenablethehighstraincharacteristicstheyrequire,whilstpositionaltendonsrelymoreheavilyonslidingbetweencollagenfibresandfibrils.[27]However,recentdatasuggeststhatenergystoringtendonsmayalsocontainfascicleswhicharetwisted,orhelical,innature-anarrangementthatwouldbehighlybeneficialforprovidingthespring-likebehaviourrequiredinthesetendons.[28] Mechanics[edit] Mainarticle:Softtissue Tendonsareviscoelasticstructures,whichmeanstheyexhibitbothelasticandviscousbehaviour.Whenstretched,tendonsexhibittypical"softtissue"behavior.Theforce-extension,orstress-straincurvestartswithaverylowstiffnessregion,asthecrimpstructurestraightensandthecollagenfibresalignsuggestingnegativePoisson'sratiointhefibresofthetendon.Morerecently,testscarriedoutinvivo(throughMRI)andexvivo(throughmechanicaltestingofvariouscadaverictendontissue)haveshownthathealthytendonsarehighlyanisotropicandexhibitanegativePoisson'sratio(auxetic)insomeplaneswhenstretchedupto2%alongtheirlength,i.e.withintheirnormalrangeofmotion.[29]Afterthis'toe'region,thestructurebecomessignificantlystiffer,andhasalinearstress-straincurveuntilitbeginstofail.Themechanicalpropertiesoftendonsvarywidely,astheyarematchedtothefunctionalrequirementsofthetendon.Theenergystoringtendonstendtobemoreelastic,orlessstiff,sotheycanmoreeasilystoreenergy,whilstthestifferpositionaltendonstendtobealittlemoreviscoelastic,andlesselastic,sotheycanprovidefinercontrolofmovement.Atypicalenergystoringtendonwillfailataround12-15%strain,andastressintheregionof100-150MPa,althoughsometendonsarenotablymoreextensiblethanthis,forexamplethesuperficialdigitalflexorinthehorse,whichstretchesinexcessof20%whengalloping.[30]Positionaltendonscanfailatstrainsaslowas6-8%,butcanhavemoduliintheregionof700-1000MPa.[31] Severalstudieshavedemonstratedthattendonsrespondtochangesinmechanicalloadingwithgrowthandremodelingprocesses,muchlikebones.Inparticular,astudyshowedthatdisuseoftheAchillestendoninratsresultedinadecreaseintheaveragethicknessofthecollagenfiberbundlescomprisingthetendon.[32]Inhumans,anexperimentinwhichpeopleweresubjectedtoasimulatedmicro-gravityenvironmentfoundthattendonstiffnessdecreasedsignificantly,evenwhensubjectswererequiredtoperformrestivenessexercises.[33]Theseeffectshaveimplicationsinareasrangingfromtreatmentofbedriddenpatientstothedesignofmoreeffectiveexercisesforastronauts. Healing[edit] Thetendonsinthefootarehighlycomplexandintricate.Therefore,thehealingprocessforabrokentendonislongandpainful.Mostpeoplewhodonotreceivemedicalattentionwithinthefirst48hoursoftheinjurywillsufferfromsevereswelling,pain,andaburningsensationwheretheinjuryoccurred. Itwasbelievedthattendonscouldnotundergomatrixturnoverandthattenocyteswerenotcapableofrepair.However,ithassincebeenshownthat,throughoutthelifetimeofaperson,tenocytesinthetendonactivelysynthesizematrixcomponentsaswellasenzymessuchasmatrixmetalloproteinases(MMPs)candegradethematrix.[34]Tendonsarecapableofhealingandrecoveringfrominjuriesinaprocessthatiscontrolledbythetenocytesandtheirsurroundingextracellularmatrix. Thethreemainstagesoftendonhealingareinflammation,repairorproliferation,andremodeling,whichcanbefurtherdividedintoconsolidationandmaturation.Thesestagescanoverlapwitheachother.Inthefirststage,inflammatorycellssuchasneutrophilsarerecruitedtotheinjurysite,alongwitherythrocytes.Monocytesandmacrophagesarerecruitedwithinthefirst24hours,andphagocytosisofnecroticmaterialsattheinjurysiteoccurs.Afterthereleaseofvasoactiveandchemotacticfactors,angiogenesisandtheproliferationoftenocytesareinitiated.TenocytesthenmoveintothesiteandstarttosynthesizecollagenIII.[35][36]Afterafewdays,therepairorproliferationstagebegins.Inthisstage,thetenocytesareinvolvedinthesynthesisoflargeamountsofcollagenandproteoglycansatthesiteofinjury,andthelevelsofGAGandwaterarehigh.[37]Afteraboutsixweeks,theremodelingstagebegins.Thefirstpartofthisstageisconsolidation,whichlastsfromaboutsixtotenweeksaftertheinjury.Duringthistime,thesynthesisofcollagenandGAGsisdecreased,andthecellularityisalsodecreasedasthetissuebecomesmorefibrousasaresultofincreasedproductionofcollagenIandthefibrilsbecomealignedinthedirectionofmechanicalstress.[36]Thefinalmaturationstageoccursaftertenweeks,andduringthistimethereisanincreaseincrosslinkingofthecollagenfibrils,whichcausesthetissuetobecomestiffer.Gradually,overaboutoneyear,thetissuewillturnfromfibroustoscar-like.[37] Matrixmetalloproteinases(MMPs)haveaveryimportantroleinthedegradationandremodelingoftheECMduringthehealingprocessafteratendoninjury.CertainMMPsincludingMMP-1,MMP-2,MMP-8,MMP-13,andMMP-14havecollagenaseactivity,meaningthat,unlikemanyotherenzymes,theyarecapableofdegradingcollagenIfibrils.ThedegradationofthecollagenfibrilsbyMMP-1alongwiththepresenceofdenaturedcollagenarefactorsthatarebelievedtocauseweakeningofthetendonECMandanincreaseinthepotentialforanotherrupturetooccur.[38]Inresponsetorepeatedmechanicalloadingorinjury,cytokinesmaybereleasedbytenocytesandcaninducethereleaseofMMPs,causingdegradationoftheECMandleadingtorecurringinjuryandchronictendinopathies.[36] Avarietyofothermoleculesareinvolvedintendonrepairandregeneration.Therearefivegrowthfactorsthathavebeenshowntobesignificantlyupregulatedandactiveduringtendonhealing:insulin-likegrowthfactor1(IGF-I),platelet-derivedgrowthfactor(PDGF),vascularendothelialgrowthfactor(VEGF),basicfibroblastgrowthfactor(bFGF),andtransforminggrowthfactorbeta(TGF-β).[37]Thesegrowthfactorsallhavedifferentrolesduringthehealingprocess.IGF-1increasescollagenandproteoglycanproductionduringthefirststageofinflammation,andPDGFisalsopresentduringtheearlystagesafterinjuryandpromotesthesynthesisofothergrowthfactorsalongwiththesynthesisofDNAandtheproliferationoftendoncells.[37]ThethreeisoformsofTGF-β(TGF-β1,TGF-β2,TGF-β3)areknowntoplayaroleinwoundhealingandscarformation.[39]VEGFiswellknowntopromoteangiogenesisandtoinduceendothelialcellproliferationandmigration,andVEGFmRNAhasbeenshowntobeexpressedatthesiteoftendoninjuriesalongwithcollagenImRNA.[40]Bonemorphogeneticproteins(BMPs)areasubgroupofTGF-βsuperfamilythatcaninduceboneandcartilageformationaswellastissuedifferentiation,andBMP-12specificallyhasbeenshowntoinfluenceformationanddifferentiationoftendontissueandtopromotefibrogenesis. Effectsofactivityonhealing[edit] Inanimalmodels,extensivestudieshavebeenconductedtoinvestigatetheeffectsofmechanicalstrainintheformofactivitylevelontendoninjuryandhealing.Whilestretchingcandisrupthealingduringtheinitialinflammatoryphase,ithasbeenshownthatcontrolledmovementofthetendonsafteraboutoneweekfollowinganacuteinjurycanhelptopromotethesynthesisofcollagenbythetenocytes,leadingtoincreasedtensilestrengthanddiameterofthehealedtendonsandfeweradhesionsthantendonsthatareimmobilized.Inchronictendoninjuries,mechanicalloadinghasalsobeenshowntostimulatefibroblastproliferationandcollagensynthesisalongwithcollagenrealignment,allofwhichpromoterepairandremodeling.[37]Tofurthersupportthetheorythatmovementandactivityassistintendonhealing,ithasbeenshownthatimmobilizationofthetendonsafterinjuryoftenhasanegativeeffectonhealing.Inrabbits,collagenfasciclesthatareimmobilizedhaveshowndecreasedtensilestrength,andimmobilizationalsoresultsinloweramountsofwater,proteoglycans,andcollagencrosslinksinthetendons.[35] Severalmechanotransductionmechanismshavebeenproposedasreasonsfortheresponseoftenocytestomechanicalforcethatenablethemtoaltertheirgeneexpression,proteinsynthesis,andcellphenotype,andeventuallycausechangesintendonstructure.Amajorfactorismechanicaldeformationoftheextracellularmatrix,whichcanaffecttheactincytoskeletonandthereforeaffectcellshape,motility,andfunction.Mechanicalforcescanbetransmittedbyfocaladhesionsites,integrins,andcell-celljunctions.Changesintheactincytoskeletoncanactivateintegrins,whichmediate“outside-in”and“inside-out”signalingbetweenthecellandthematrix.G-proteins,whichinduceintracellularsignalingcascades,mayalsobeimportant,andionchannelsareactivatedbystretchingtoallowionssuchascalcium,sodium,orpotassiumtoenterthecell.[37] Societyandculture[edit] Sinewwaswidelyusedthroughoutpre-industrialerasasatough,durablefiber.Somespecificusesincludeusingsinewasthreadforsewing,attachingfeatherstoarrows(seefletch),lashingtoolbladestoshafts,etc.Itisalsorecommendedinsurvivalguidesasamaterialfromwhichstrongcordagecanbemadeforitemsliketrapsorlivingstructures.Tendonmustbetreatedinspecificwaystofunctionusefullyforthesepurposes.Inuitandothercircumpolarpeopleutilizedsinewastheonlycordageforalldomesticpurposesduetothelackofothersuitablefibersourcesintheirecologicalhabitats.TheelasticpropertiesofparticularsinewswerealsousedincompositerecurvedbowsfavouredbythesteppenomadsofEurasia,andNativeAmericans.Thefirststonethrowingartilleryalsousedtheelasticpropertiesofsinew. Sinewmakesforanexcellentcordagematerialforthreereasons:Itisextremelystrong,itcontainsnaturalglues,anditshrinksasitdries,doingawaywiththeneedforknots. Culinaryuses[edit] Mainarticle:Tendon(meal) Tendon(inparticular,beeftendon)isusedasafoodinsomeAsiancuisines(oftenservedatyumchaordimsumrestaurants).Onepopulardishissuanbaoniujin,inwhichthetendonismarinatedingarlic.ItisalsosometimesfoundintheVietnamesenoodledishphở. Clinicalsignificance[edit] Injury[edit] Tendonsaresubjecttomanytypesofinjuries.Therearevariousformsoftendinopathiesortendoninjuriesduetooveruse.Thesetypesofinjuriesgenerallyresultininflammationanddegenerationorweakeningofthetendons,whichmayeventuallyleadtotendonrupture.[35]Tendinopathiescanbecausedbyanumberoffactorsrelatingtothetendonextracellularmatrix(ECM),andtheirclassificationhasbeendifficultbecausetheirsymptomsandhistopathologyoftenaresimilar. Thefirstcategoryoftendinopathyisparatenonitis,whichreferstoinflammationoftheparatenon,orparatendinoussheetlocatedbetweenthetendonanditssheath.Tendinosisreferstonon-inflammatoryinjurytothetendonatthecellularlevel.Thedegradationiscausedbydamagetocollagen,cells,andthevascularcomponentsofthetendon,andisknowntoleadtorupture.[41]Observationsoftendonsthathaveundergonespontaneousrupturehaveshownthepresenceofcollagenfibrilsthatarenotinthecorrectparallelorientationorarenotuniforminlengthordiameter,alongwithroundedtenocytes,othercellabnormalities,andtheingrowthofbloodvessels.[35]Otherformsoftendinosisthathavenotledtorupturehavealsoshownthedegeneration,disorientation,andthinningofthecollagenfibrils,alongwithanincreaseintheamountofglycosaminoglycansbetweenthefibrils.[36]Thethirdisparatenonitiswithtendinosis,inwhichcombinationsofparatenoninflammationandtendondegenerationarebothpresent.Thelastistendinitis,whichreferstodegenerationwithinflammationofthetendonaswellasvasculardisruption.[5] Tendinopathiesmaybecausedbyseveralintrinsicfactorsincludingage,bodyweight,andnutrition.Theextrinsicfactorsareoftenrelatedtosportsandincludeexcessiveforcesorloading,poortrainingtechniques,andenvironmentalconditions.[34] Otheranimals[edit] OssifiedtendonfromanEdmontosaurusbonebedinWyoming(LanceFormation) Insomeorganisms,notablybirds,[42]andornithischiandinosaurs,[43]portionsofthetendoncanbecomeossified.Inthisprocess,osteocytesinfiltratethetendonandlaydownboneastheywouldinsesamoidbonesuchasthepatella.Inbirds,tendonossificationprimarilyoccursinthehindlimb,whileinornithischiandinosaurs,ossifiedaxialmuscletendonsformalatticeworkalongtheneuralandhaemalspinesonthetail,presumablyforsupport. Seealso[edit] Thisarticleusesanatomicalterminology. WikimediaCommonshasmediarelatedtoTendons. 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