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LungWikipedia

Lung
FromWikipedia,thefreeencyclopedia

Thelungsaretheprimaryorgansofrespirationinhumansandmanyotheranimalsincludingafewfishand
somesnails.Inmammalsandmostothervertebrates,twolungsarelocatednearthebackboneoneithersideof
theheart.Theirfunctionintherespiratorysystemistoextractoxygenfromtheatmosphereandtransferitinto
thebloodstream,andtoreleasecarbondioxidefromthebloodstreamintotheatmosphere,inaprocessofgas
exchange.Respirationisdrivenbydifferentmuscularsystemsindifferentspecies.Mammals,reptilesandbirds
usetheirmusculoskeletalsystemstosupportandfosterbreathing.Inearlytetrapods,airwasdrivenintothe
lungsbythepharyngealmusclesviabuccalpumping,amechanismstillseeninamphibians.Inhumans,the
primarymusclethatdrivesbreathingisthediaphragm.Thelungsalsoprovideairflowthatmakesvocalsounds
includinghumanspeechpossible.
Humanshavetwolungs,arightlungandaleftlung.Theyaresituatedwithinthethoraciccavityofthechest.
Therightlungisbiggerthantheleft,whichsharesspaceinthechestwiththeheart.Thelungstogetherweigh
approximately1.3kilograms(2.9lb),andtherightisheavier.Thelungsarepartofthelowerrespiratorytract
thatbeginsatthetracheaandbranchesintothebronchiandbronchiolesandwhichreceiveairbreathedinviathe
conductingzone.Thesedivideuntilairreachesmicroscopicalveoli,wheretheprocessofgasexchangetakes
place.Together,thelungscontainapproximately2,400kilometres(1,500mi)ofairwaysand300to500million
alveoli.Thelungsareenclosedwithinasaccalledthepleuralsacwhichallowstheinnerandouterwallstoslide
overeachotherwhilstbreathingtakesplace,withoutmuchfriction.Thissacencloseseachlungandalsodivides
eachlungintosectionscalledlobes.Therightlunghasthreelobesandthelefthastwo.Thelobesarefurther
dividedintobronchopulmonarysegmentsandlobules.Thelungshaveauniquebloodsupply,receiving
deoxygenatedbloodsentfromtheheartforthepurposesofreceivingoxygen(thepulmonarycirculation)anda
separatesupplyofoxygenatedblood(thebronchialcirculation).
Thetissueofthelungscanbeaffectedbyanumberofdiseases,includingpneumoniaandlungcancer.Chronic
diseasessuchaschronicobstructivepulmonarydiseaseandemphysemacanberelatedtosmokingorexposure
toharmfulsubstances.Diseasessuchasbronchitiscanalsoaffecttherespiratorytract.

Lung

Diagramofthelungswiththerespiratorytract
visible,anddifferentcoloursforeachlobe
Details
System Respiratorysystem
Identifiers
Latin

Inembryonicdevelopment,thelungsbegintodevelopasanoutpouchingoftheforegut,atubewhichgoesonto
formtheupperpartofthedigestivesystem.Whenthelungsareformedthefetusisheldinthefluidfilled
amnioticsacandsotheydonotfunctiontobreathe.Bloodisalsodivertedfromthelungsthroughtheductus
arteriosus.Atbirthhowever,airbeginstopassthroughthelungs,andthediversionaryductcloses,sothatthe
lungscanbegintorespire.Thelungsonlyfullydevelopinearlychildhood.
https://en.wikipedia.org/wiki/Lung

pulmo

Greek (pneumon)
MeSH A04.411(https://www.nlm.nih.gov/cgi/mes
h/2011/MB_cgi?mode=&term=Lung)
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Medicaltermsrelatedtothelungoftenbeginwithpulmo,fromtheLatinpulmonarius(ofthelungs)asin
pulmonology,orwithpneumo(fromGreek"lung")asinpneumonia.

Contents
1 Structure
1.1 Grossanatomy
1.2 Rightlung
1.3 Leftlung
1.4 Respiratorysystem
1.5 Bloodsupply
1.6 Nervesupply
1.7 Microanatomy
2 Development
2.1 Firstbreath
3 Function
3.1 Breathing
3.2 Respiration
3.3 Protection
3.4 Other
4 Clinicalsignificance
4.1 Lungfunctiontesting
5 Otheranimals
5.1 Birds
5.2 Reptiles
5.3 Amphibians
5.4 Lungfish
5.5 Invertebrates
6 Evolutionaryorigins
7 Seealso
8 Furtherreading
9 References

TA

A06.5.01.001(http://www.unifr.ch/ifaa/Pu
blic/EntryPage/TA98%20Tree/Entity%20T
A98%20EN/06.5.01.001%20Entity%20T
A98%20EN.htm)

FMA

68877(http://xiphoid.biostr.washington.ed
u/fma/fmabrowserhierarchy.html?fmaid=6
8877)
Anatomicalterminology
[editonWikidata]

Thehumanlungsflanktheheartand
greatvesselsinthechestcavity

Structure
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Grossanatomy
Thelungsarelocatedinthechestoneithersideoftheheartintherib
cage.Theyareconicalinshapewithanarrowroundedapexatthetop
andabroadbasethatrestsonthediaphragm.[1]Theapexofthelung
extendsintotherootoftheneck,reachingshortlyabovethelevelofthe
sternalendofthefirstrib.Thelungsstretchfromclosetothebackbone
"Meetthelungs"fromtheKhan
intheribcagetothefrontofthechestanddownwardsfromthelower
academy
partofthetracheatothediaphragm.[1]Theleftlungsharesspacewith
theheart,withanimpressioninitsmedialsurfacecalledthecardiac
impression.[2]Thefrontandoutersidesofthelungfacetheribs,whichmakelightindendationsontheir
surfaces.Thebottomofthelungsissmoothandrestsonthediaphragm,matchingitsconcavity.Themedial
surfaceofthelungsfacestowardsthecentreofthechest,andliesagainsttheheart,greatvessels,andthe
carinawherethetwomainbronchibranchofffromthebaseofthetrachea.[2]
Bothlungshaveacentralrecessioncalledthehilumattherootofthelung,wherethebloodvesselsandairwayspassintothelungs.[1]Therearealso
bronchopulmonarylymphnodesonthehilum.[2]
Thelungsaresurroundedbythepulmonarypleurae.Thepleuraearetwoserousmembranestheouterparietalpleuralinestheinnerwalloftheribcageandthe
innervisceralpleuradirectlylinesthesurfaceofthelungs.Betweenthepleuraeisapotentialspacecalledthepleuralcavitycontainingpleuralfluid.Eachlungis
dividedintolobesbytheinvaginationsofthepleuraasfissures.Thefissuresaredoublefoldsofpleurathatsectionthelungsandhelpintheirexpansion.[3]
Thelobesofthelungsarefurtherdividedintobronchopulmonarysegmentsbasedonthelocationsofbronchioles.Segmentsfortheleftandrightlungareshown
inthetable.[4]Thesegmentalanatomyisusefulclinicallyforlocalisingdiseaseprocessesinthelungs.[4]

Rightlung
Therightlunghasbothmorelobesandsegmentsthantheleft.Itisdividedintothreelobes,anupper,middle,andalower,bytwofissures,oneobliqueandone
horizontal.Theupper,horizontalfissure,separatestheupperfromthemiddlelobe.Itbeginsinthelowerobliquefissureneartheposteriorborderofthelung,and,
runninghorizontallyforward,cutstheanteriorborderonalevelwiththesternalendofthefourthcostalcartilageonthemediastinalsurfaceitmaybetraced
backwardtothehilum.[1]
Thelower,obliquefissure,separatesthelowerfromthemiddleandupperlobes,andiscloselyalignedwiththeobliquefissureintheleftlung.[1][3]

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Themediastinalsurfaceoftherightlungisindentedbyanumberofnearbystructures.Theheartsitsinanimpression
calledthecardiacimpression.Abovethehilumofthelungisanarchedgroovefortheazygosvein,andabovethisisa
widegrooveforthesuperiorvenacavaandrightinnominateveinbehindthis,andclosetothetopofthelungisagroove
fortheinnominateartery.Thereisagroovefortheesophagusbehindthehilumandthepulmonaryligament,andnearthe
lowerpartoftheesophagealgrooveisadeepergroovefortheinferiorvenacavabeforeitenterstheheart.[2]

Leftlung

Lobesandbronchopulmonary
segments[4]
Rightlung
Leftlung
Upper
Apical
Anterior
Posterior

Theleftlungisdividedintotwolobes,anupperandalower,bytheobliquefissure,whichextendsfromthecostaltothe
mediastinalsurfaceofthelungbothaboveandbelowthehilum.[1]Theleftlung,unliketheright,doesnothavemiddle
Middle
lobe,thoughitdoeshaveahomologousfeature,aprojectionoftheupperlobetermedthelingula.Itsnamemeanslittle
tongue.Thelingulaontheleftservesasananatomicparalleltotherightmiddlelobe,withbothareasbeingpredisposed
Medial
[5][6]
Lateral
tosimilarinfectionsandanatomiccomplications.
Therearetwobronchopulmonarysegmentsofthelingula:superior
andinferior.[1]
Lower
Themediastinalsurfaceoftheleftlunghasalargecardiacimpressionwheretheheartsits.Thisisdeeperandlargerthan
thatontherightlung,atwhichleveltheheartprojectstotheleft.[2]
Onthesamesurface,immediatelyabovethehilum,isawellmarkedcurvedgroovefortheaorticarch,andagroove
belowitforthedescendingaorta.Theleftsubclavianartery,abranchofftheaorticarch,sitsinagroovefromthearchto
neartheapexofthelung.Ashallowergrooveinfrontofthearteryandneartheedgeofthelung,lodgestheleft
innominatevein.Theesophagusmaysitinawidershallowimpressionatthebaseofthelung.[2]

Upper
Anterior
Apicoposterior
Lower
Superior
Anterior
Posterior
Medial
Lateral

Superior
Anterior Lingula
Posterior
Superior
Medial
Inferior
Lateral

Theleftlung(left)andrightlung(right).Thelobesofthelungscan
beseen,andthecentralrootofthelungisalsopresent.
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Respiratorysystem
Thelungispartoftherespiratorysystem,andcontainsthemajorityofthelowerrespiratorytractafterthe
trachea.[7]Thetracheareceivesairfromthepharynxandtravelsdowntoaplacewhereitsplits(thecarina)intoa
rightandleftbronchus.Thesesupplyairtotherightandleftlungs,splittingprogressivelyintothesecondaryand
tertiarybronchiforthelobesofthelungs,andintosmallerandsmallerbronchiolesuntiltheybecomethe
respiratorybronchioles.Theseinturnsupplyairthroughalveolarductsintothealveoli,wheretheexchangeof
gasestakeplace.[7]Oxygendiffusesthroughthewallsofthealveoliintotheenvelopingcapillaries(smallblood
vessels).[8]
Estimatesofthetotalsurfaceareaoflungsvaryfrom50to75squaremetres(540to810sqft)[7][9]roughlythe
sameareaasonesideofatenniscourt.[9][10]
Thebronchiintheconductingzonearereinforcedwithhyalinecartilageinordertoholdopentheairways.The
bronchioleshavenocartilageandaresurroundedinsteadbysmoothmuscle.[9]Airiswarmedto37C(99F),
humidifiedandcleansedbytheconductionzoneparticlesfromtheairbeingremovedbytheciliaonthe
respiratoryepitheliumliningthepassageways.[11]

Bloodsupply
Thehumanlunghasadualbloodsupply.[7]
Thetissueofthelungsreceiveoxygenatedbloodviathebronchialcirculation,aseriesofarteriesthatleavethe
aortaandarepartofthesystemiccirculation.Thereareusuallythreearteries,andtheybranchalongsidethe
bronchiandbronchioles.[7]Thebloodvolumeofthelungsisabout450millilitresonaverage,about9percentof
thetotalbloodvolumeoftheentirecirculatorysystem.Thisquantitycaneasilyfluctuatefrombetweenonehalf
andtwicethenormalvolume.[12]

Diagramoftherespiratorysystem,which
includestheupperrespiratorytractandlower
respiratorytract,ofwhichthelungsarethe
majorpart.

Thelungsalsoreceivedeoxygenatedbloodfromtheheartandsupplyitwithoxygen,inaprocessknownasrespiration.Inthisprocess,venousbloodinthebody
collectsintherightatriumandispumpedfromtherightventriclethroughthepulmonarytrunkandthepulmonaryarteriesintotheleftandrightlungs.Blood
passesthroughsmallcapillariesnexttothealveoliinthelung,receivesoxygen,andtravelsbacktotheheart.Thisiscalledthepulmonarycirculation.The
oxygenatedbloodisthenpumpedtotherestofthebody.[7]

Nervesupply
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Thelungsaresuppliedbynervesoftheautonomicnervoussystem.Inputfromtheparasympatheticnervoussystemoccursviathevagusnerve.Whenstimulated
byacetylcholine,thiscausesconstrictionofthesmoothmuscleliningthebronchusandbronchioli,andincreasesthesecretionsfromglands.[13]Thelungsalso
haveasympathetictonefromnorepinephrineactingonthebeta2receptorsintherespiratorytract,whichcausesbronchodilation.[13]
Theactionofbreathingtakesplacebecauseofnervesignalssentbytherespiratorycentresinthebrainstem,alongthephrenicnervetothediaphragm.Thisis
describedinmoredetailbelow.[14]

Microanatomy
Thelungscontaintherespiratorytractanditslining,whichterminateinalveoli,thetissueinbetween(called
interstitiumorparenchyma),andveins,arteries,nervesandlymphaticvessels.[2][15]
Therespiratorytractbeginswiththetracheaandbronchi.Thesestructuresarelinedwithcolumnarepithelialcellsthat
possesscilia,smallfrondlikeprojections.Interspersedwiththeepithelialcellsaregobletcellswhichproducemucous,
andclubcellswithactionssimilartomacrophages.Surroundingtheseinthetracheaandbronchiarecartilagerings,
whichhelptomaintainstability.[9]Bronchiolespossessthesamecolumnarepitheliallining,butarenotsurroundedby
cartilagerings.Instead,theyareencircledbyalayerofsmoothmuscle.[9]Therespiratorytractendsinlobules.These
consistofarespiratorybronchiole,whichbranchesintoalveolarductsandalveolarsacs,whichinturndivideinto
alveoli.[2]
Theepithelialcellsthroughouttherespiratorytractsecreteepithelialliningfluid(ELF),thecompositionofwhichis
tightlyregulatedanddetermineshowwellmucociliaryclearanceworks.[16][17]:Section4pages78(Page47ff)whichinturn
iscoatedwithalayerofsurfactant.[7]

Arespiratorylobule,thefunctionalunit
ofthelung.Thealveolus,respiratory
bronchioleandsurroundingbloodvessels
canbeseen.

Alveoliconsistoftwotypesofalveolarcellandanalveolarmacrophage.ThetwotypesofcellareknownastypeIandtypeIIalveolarcells[7](alsoknownas
pneumocytes).[2]TypesIandIImakeupthewallsandseptaofthealveoli.TypeIcellsprovide95%ofthesurfaceareaofeachalveoliandareflat("squamous"),
andTypeIIcellsgenerallyclusterinthecornersofthealveoliandhaveacuboidalshape.[9]Despitethis,cellsoccurinaroughlyequalratioof1:1or6:4.[7][9]
TypeIaresquamousepithelialcellsthatmakeupthealveolarwallstructure.Theyhaveextremelythinwallsthatenableaneasygasexchange.[7]ThesetypeIcells
alsomakeupthealveolarseptawhichseparateeachalveolus.Theseptaconsistofanepithelialliningandassociatedbasementmembranes.[9]TypeIcellsarenot
abletodivide,andconsequentlyrelyondifferentiationfromTypeIIcells.[9]
TypeIIarelargerandtheylinethealveoliandproduceandsecreteELFandsurfactant.[7]TypeIIcellsareabletodivideanddifferentiatetoType1cells.[9]

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Thealveolarmacrophageshaveanimportantimmunologicalrole.Theyremovesubstanceswhichdepositinthe
alveoliincludinglooseredbloodcellsthathavebeenforcedoutfrombloodvesels.[9]
Thelungissurroundedbyaserousmembraneofvisceralpleura,whichhasanunderlyinglayerofloose
connectivetissueattachedtothesubstanceofthelung.[18]

Development

Crosssectionaldetailofthelung

Thedevelopmentofthehumanlungsarisefromthelaryngotracheal
grooveanddeveloptomaturityoverseveralweeksinthefoetusandfor
severalmonthsfollowingbirth.[19]Thelarynx,trachea,bronchiandlungs
begintoformduringthefourthweekofembryogenesis.[20]fromthe
respiratorybud(ordiverticulum)whichappearsventrallytothecaudal
portionoftheforegut.[21]

Attheendofthefourthweekthelungbuddividesintotwo,therightandleftprimarybronchialbuds.[22]Duringthefifth
weektherightbudbranchesintothreesecondarybronchialbudsandtheleftbranchesintotwosecondarybronchialbuds.
Lungsduringdevelopment,showing
Thesegiverisetothelobesofthelungs,threeontherightandtwoontheleft.Overthefollowingweek,thesecondary
primitivelungbudsthathavenotyet
[22]
fullydividedintomaturelungs.
budsbranchintotertiarybuds,abouttenoneachside. Fromthesixthweektothesixteenthweek,themajorelementsof
[23]
thelungsappearexceptthealveoli. Fromweek16toweek26,thebronchienlargeandlungtissuebecomeshighly
vascularised.Bronchiolesandalveolarductsalsodevelop.Duringtheperiodcoveringthe26thweekuntilbirththeimportantbloodairbarrierisestablished.
SpecialisedtypeIalveolarcellswheregasexchangewilltakeplace,togetherwiththetypeIIalveolarcellsthatsecretepulmonarysurfactant,appear.The
surfactantreducesthesurfacetensionattheairalveolarsurfacewhichallowsexpansionoftheterminalsaccules.Thesesacculesformattheendofthebronchioles
andtheirappearancemarksthepointatwhichlimitedrespirationwouldbepossible.[24]

Firstbreath
Atbirth,thebaby'slungsarefilledwithfluidsecretedbythelungsandarenotinflated.Afterbirthitscentralnervoussystemreactstothesuddenchangein
temperatureandenvironment.Thistriggersthefirstbreath,withinabout10secondsafterdelivery.[25]Beforebirth,thelungsarefilledwithfetallungfluid.[26]
Afterthefirstbreath,thefluidisquicklyabsorbedintothebodyorexhaled.Theresistanceinthelung'sbloodvesselsdecreasesgivinganincreasedsurfacearea
forgasexchange,andthelungbeginstobreathespontaneously.Thisaccompaniesotherchangeswhichresultinanincreasedamountofbloodenteringthelung
tissues.[25]

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Atbirththelungsareveryundevelopedwithonlyafractionofthealveolipresent.Thealveolicontinuetoformuntilthethirdyear.Interalveoliseptahavea
doublecapillarynetworkinsteadofthesinglenetworkofthedevelopedlung.Onlyafterthematurationofthecapillarynetworkcanthelungenteranormalphase
ofgrowth.Followingtheearlygrowthinnumbersofalveolithereisanotherstageofthealveolibeingenlarged.[27]

Function
Breathing
Breathingreferstotheprocessbywhichairentersandleavesthelungs.[28]Itislargelydrivenbythe
musculardiaphragmatthebottomofthethorax.Contractionofthediaphragmpullsthebottomofthe
cavityinwhichthelungisencloseddownward,increasingvolumeandthusdecreasingpressure,causingair
toflowintotheairways.Anumberofmusclesassistinthisaction,includingtheintercostalmusclesand
musclesthatlinetheabdominalwall.Thelungisnotcapableofexpandingitself,andonlyexpandsbecause
ofanegativepressurebetweenthetwopleuralwallsoutsidethelung.Thelungsarepulledbythisnegative
pressureandexpandorcontracttofillitout.[29]Thisdrawsairintothelungs,whichpassesthroughthe
respiratorytractandintothealveoli,whererespirationoccurs.[13]
Duringnormalbreathing,exhalationispassiveandnomusclesarecontracted(thediaphragmrelaxes).[13]
Theprocessofbreathingoccursbecauseofsignalssentalongthephrenicnerveintherespiratorycentres
situatedinthemedullaofthebrainstem.Therespiratorycentresaresituatedbelowthefourthventricleand
containstwomaingroupsofneurons,aventralrespiratorygroupandadorsalrespiratorygroup.Theventral
respiratorygroupisresponsibleformusclemovementsassociatedwithrespiration(suchasofthepharynx,
larynxandintercostalmuscles)andmaycontainthe"pacemaker"cellsresponsiblefordrivingtherateofrespirationinthepreBotzingercomplex.Thedorsal
respiratorygroupseemsprimarilyresponsibleforcontrolofthediaphragmandintegrationofsignalstransmittedbackviathevagusnervethataffecttherateof
respiration.[14]Withinthesetwogroupsaremixedgroupsofnerves,inspiratoryneurons,andexpiratoryneurons.Thesespontaneouslygeneratesignals(action
potentials)responsibleforinspirationandexpirationwhicharethentransmittedtothediaphragmbythephrenicnerve.[14]
Lungsofaraccoonbeingmanuallyinflatedduringa
dissection

Anumberoffactorsinfluencetherateofbreathing.Carbondioxidelevels,particularlythearterialandcerebrospinalfluidlevels,detectedbymedullarycentral
chemoreceptors,causeanincreaseintherateofbreathingwhenhighandadecreasewhenlow.Theacidityofblood,detectedbyperipheralchemoreceptors,
causesanincreaseintherateofbreathingwhenlowandadecreasewhenhigh.Significanthypoxia,primarilydetectedbyperipheralchemoreceptorsintheaortic
body,cancauseanincreaseintherateofbreathing.Factorsinternaltothelung,includingpulmonarystretchordetectionofirritants,canalsostimulate
breathing.[14]

Respiration
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Respirationistheprocessbywhichoxygenistakenintothebodyandcarbondioxideisexpelled.[30]Respirationis
dividedinto'inspiration,inwhichairistakenintothelungs,andexpiration,inwhichitisexpelledfromthem.Thisgas
exchangetakesplaceinthealveoliduringbreathing.[7]
Inhumans,thetracheadividesintothetwomainbronchithatentertherootsofthelungs.Thebronchicontinuetodivide
withinthelung,andaftermultipledivisions,giverisetobronchioles.Thebronchialtreecontinuesbranchinguntilit
reachesthelevelofterminalbronchioles,whichleadtorespiratorybronchiolesandalveolarsacs.Alveolarsacs,are
madeupofclustersofalveoli,likeindividualgrapeswithinabunch.Theindividualalveoliaretightlywrappedin
capillariesanditisherethatgasexchangeactuallyoccurs.Deoxygenatedbloodfromtheheartispumpedthroughthe
pulmonaryarterytothelungs,whereoxygendiffusesintobloodandisexchangedforcarbondioxideinthehemoglobin
oftheerythrocytes.Theoxygenrichbloodreturnstotheheartviathepulmonaryveinstobepumpedbackintosystemic
circulation.[7]

Gasexchangeoccursinthealveolus,
wherevenousbloodexchangescarbon
dioxideforoxygen.

Protection
Thelungspossessseveralcharacteristicswhichprotectagainstinfection.Thelungtractislinedbyepitheliawithhairlikeprojectionscalledciliathatbeat
rhythmicallyandcarrymucous.Thismucociliaryclearanceisanimportantdefencesystemagainstairborneinfection.Thedustparticlesandbacteriainthe
inhaledairarecaughtinthemucouslayerpresentatthemucosalsurfaceofrespiratorypassagesandaremoveduptowardsthepharynxbytherhythmicupward
beatingactionofthecilia.[9][31][32]TheliningofthelungalsosecretesimmunoglobulinAwhichprotectsagainstrespiratoryinfections[31]gobletcellssecrete
mucous[9]whichalsocontainscontainsseveralantimicrobialcompoundssuchasdefensins,antiproteases,andantioxidates.[31]Inaddition,theliningofthelung
alsocontainsmacrophages,immunecellswhichengulfanddestroydebrisandmicrobesthatenterthelunginaprocessknownasphagocytosisanddendriticcells
whichpresentantigenstoactivatecomponentsoftheadaptiveimmunesystemsuchasTcellsandBcells.[31]
Thesizeoftherespiratorytractandtheflowofairalsoprotectthelungsfromlargerparticles.Smallerparticlesdepositinandbehindthemouth,andlarger
particlesaretrappedafterinhalationbynosehairs.[31]

Other
Inadditiontotheirfunctioninrespiration,thelungsalsohaveanumberofotherfunctions.Theyareinvolvedinthemaintenanceofhomeostasis.Theyhelpinthe
regulationofbloodpressureaspartofthereninangiotensinsystem.Theinnerliningofthebloodvesselssecretesangiotensinconvertingenzyme(ACE)an
enzymethatcatalysestheconversionofangiotensinItoangiotensinII.[33]Thelungsareinvolvedintheblood'sacidbasehomeostasisbyexpellingcarbondioxide
whenbreathing.[29][34]

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Thelungsalsoserveaprotectiverole.Severalbloodbornesubstances,suchasafewtypesofprostaglandins,leukotrienes,serotoninandbradykinin,areexcreted
throughthelungs.[33]Drugsandothersubstancescanbeabsorbed,modifiedorexcretedinthelungs.[29][35]Thelungsfilteroutsmallbloodclotsfromveinsand
preventthemfromenteringarteriesandcausingstrokes.[34]
Thelungsalsoplayapivotalroleinspeechbyprovidingairandairflowforthecreationofvocalsounds.[29][36]

Clinicalsignificance
Humanlungscanbeaffectedbyavarietyofdiseases.Pulmonologyisthemedicalspecialitythatdealswithdiseasesinvolvingtherespiratorytract,[37]and
cardiothoracicsurgeryisthesurgicalfieldthatdealswithsurgeryofthelungs.[38]
Infectionsofthelungtissuearecalledpneumonia,oftherespiratorytractarecalledbronchitisorbronchiolitis,andofthepleuraesurroundingthelungspleurisy.
Manyinfectionsareduetobacteriaorviruses,howevernotallcausesofpneumoniaorbronchialtreeinflammationarecausedbyinfection.Oneimportantcause
ofbacterialpneumoniaistuberculosis.[31]ChronicinfectionsoftenoccurinthosewithimmunodeficiencyandcanincludeafungalinfectionbyAspergillus
fumigatusthatcanleadtoanaspergillomaforminginthelung.[31][39]
Apulmonaryembolismisabloodclotthatbecomeslodgedinthepulmonaryarteries.Themajorityofemboliarise
becauseofdeepveinthrombosisinthelegs.Pulmonaryembolimaybeinvestigatedusingaventilation/perfusionscan,a
CTscanofthearteriesofthelung,orbloodtestssuchastheDdimer.[31]Pulmonaryhypertensiondescribesan
increasedpressureatthebeginningofthepulmonaryarterythathasalargenumberofdifferingcauses.[31]Otherrarer
conditionsmayalsoaffectthebloodsupplyofthelung,suchasgranulomatosiswithpolyangiitis,whichcauses
inflammationofthesmallbloodvesselsofthelungsandkidneys.[31]
Alungcontusionisabruisecausedbychesttrauma.Itresultsinhemorrhageofthealveolicausingabuildupoffluid
whichcanimpairbreathing,andthiscanbeeithermildorsevere.Thefunctionofthelungscanalsobeaffectedby
compressionfromfluidinthepleuralcavitypleuraleffusion,orothersubstancessuchasair(pneumothorax),blood
(hemothorax),orrarercauses.ThesemaybeinvestigatedusingachestXrayorCTscan,andmayrequiretheinsertion
ofasurgicaldrainuntiltheunderlyingcauseisidentifiedandtreated.[31]

Infarctionofthelungduetoapulmonary
embolism

Asthma,chronicbronchitis,bronchiectasisandchronicobstructivepulmonarydisease(COPD)areallobstructivelung
diseasescharacterisedbyairwayobstruction.Thislimitstheamountofairthatisabletoenteralveolibecauseofconstrictionofthebronchialtree,dueto
inflammation.Obstructivelungdiseasesareoftenidentifiedbecauseofsymptomsanddiagnosedwithpulmonaryfunctiontestssuchasspirometry.Many
obstructivelungdiseasesaremanagedbyavoidingtriggers(suchasdustmitesorsmoking),withsymptomcontrolsuchasbronchodilators,andwithsuppression
ofinflammation(suchasthroughcorticosteroids)inseverecases.OnecommoncauseofCOPDandemphysemaissmoking,andcommoncausesofbronchiectasis
includesevereinfectionsandcysticfibrosis.Thedefinitivecauseofasthmaisnotyetknown.[31]
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Sometypesofchroniclungdiseasesareclassifiedasrestrictivelungdisease,becauseofarestrictionintheamountof
lungtissueinvolvedinrespiration.Theseincludepulmonaryfibrosiswhichcanoccurwhenthelungisinflamedfora
longperiodoftime.Fibrosisinthelungreplacesfunctioninglungtissuewithfibrousconnectivetissue.Thiscanbedue
toalargevarietyofoccupationaldiseasessuchasCoalworker'spneumoconiosis,autoimmunediseasesormorerarelyto
areactiontomedication.[31]
Lungcancercancaneitherarisedirectlyfromlungtissueorasaresultofmetastasisfromanotherpartofthebody.
Therearetwomaintypesofprimarytumourdescribedaseithersmallcellornonsmallcelllungcarcinomas.Themajor
riskfactorforcancerissmoking.OnceacancerisidentifieditisstagedusingscanssuchasaCTscanandasampleof
tissue(abiopsy)istaken.Cancersmaybetreatedbysurgicallyremovingthetumour,radiotherapy,chemotherapyor
combinationsthereof,orwiththeaimofsymptomcontrol.[31]Lungcancerscreeningisbeingrecommendedinthe
UnitedStatesforhighriskpopulations.[40]

Lungtissueaffectedbyemphysemausing
H&Estain

Congenitaldisordersincludecysticfibrosis,pulmonaryhypoplasia(anincompletedevelopmentofthe
lungs)[41]congenitaldiaphragmatichernia,andinfantrespiratorydistresssyndromecausedbyadeficiencyinlungsurfactant.Anazygoslobeisacongenital
anatomicalvariationwhichthoughusuallywithouteffectcancauseproblemsinthoracoscopicprocedures.[42]

Lungfunctiontesting
Lungfunctiontestingiscarriedoutbyevaluatingaperson's
capacitytoinhaleandexhaleindifferentcircumstances.[43]The
inhaledandexhaledbyapersonatrestisthetidalvolume
(normally500750mL)theinspiratoryreservevolumeand
expiratoryreservevolumearetheadditionalamountsapersonis
abletoforciblyinhaleandexhalerespectively.Thesummedtotal
offorcedinspirationandexpirationisaperson'svitalcapacity.
Notallairisexpelledfromthelungsevenafteraforcedbreath
outtheremainderoftheairiscalledtheresidualvolume.
Togetherthesetermsarereferredtoaslungvolumes.[43]

Lungvolumesasdescribedinthetext.

Apersondoingaspirometry
test.

Pulmonaryplethysmographsareusedtomeasurefunctionalresidualcapacity.[44]Functionalresidualcapacitycannotbemeasuredbyteststhatrelyonbreathing
out,asapersonisonlyabletobreatheamaximumof80%oftheirtotalfunctionalcapacity.[45]Thetotallungcapacitydependsontheperson'sage,height,weight,
andsex,andnormallyrangesbetween4and6litres.[43]Femalestendtohavea2025%lowercapacitythanmales.Tallpeopletendtohavealargertotallung
capacitythanshorterpeople.Smokershavealowercapacitythannonsmokers.Thinnerpersonstendtohavealargercapacity,andcapacitycanbeincreasedby
physicaltrainingasmuchas40%.[45]
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Otherlungfunctiontestsincludespirometry,measuringtheamount(volume)andflowofairthatcanbeinhaledandexhaled.Themaximumvolumeofbreaththat
canbeexhalediscalledthevitalcapacity.Inparticular,howmuchapersonisabletoexhaleinoneminute(calledFEV1)asaproportionofhowmuchtheyare
abletoexhaleintotal(FEV).Thisratio,theFEV1/FEVratio,isimportanttodistinguishwhetheradiseaseisrestrictiveorobstructive.[31][43]

Otheranimals
Birds
Thelungsofbirdsarerelativelysmall,butareconnectedto
89airsacsthatextendthroughmuchofthebody,andare
inturnconnectedtoairspaceswithinthebones.On
inspiration,airtravelsthroughthetracheaofabirdintothe
89airsacs.Airthentravelscontinuouslyfromtheairsacs
attheback,throughthelungs,andtotheairsacsatthe
front.Fromhere,airisexpelled.Thistypeoflung
constructioniscalledacirculatorylung,asdistinctfrom
thebellowslungpossessedbyotheranimals.[46]Lungsare
relativelyfixedinsize,unlikethefluctuatingsizeofa
bellowstypelung.[47]Thismeansthatbirdsareableto
extractagreaterconcentrationofoxygenfrominhaledair.
Birdsarethusequippedtoflyataltitudesatwhich
mammalswouldsuccumbtohypoxia.Thisalsoallows
themtosustainahighermetabolicratethanmost
equivalentweightmammals.[46]

Oninspiration,airtravelstoairsacsneartheback
ofabird.Thesethenpassthroughthelungstoair
sacsnearthefrontofthebird,whereairisexpelled.

Therespiatorysysteminabird.

Thelungsofbirdsarehoneycomblikeandcontainmillionsoftinypassagescalledparabronchi.Smallsacs
calledcalledatriaradiatefromthewallsofthetinypassages,andarethesiteofgasexchange.Gasexchange
occursbydiffusioninthesewalls,asgastravelsbetweenthebloodvesselsandthelumenofeach
parabronchus.[47]

Theairsacs,whichholdair,donotcontributemuchtogasexchange,despitebeingthinwalled,astheyare
poorlyvascularised.Theairsacsexpandandcontractduetochangesinthevolumeinthethoraxandabdomen.
Thisvolumechangeiscausedbythemovementofthesternumandribsandthismovementisoftensynchronisedwithmovementoftheflightmuscles.[48]
Thistypicalsystemisoneoftwotypesofparabronchifoundinbirds,calledpaleopulmonicparabronchiandisfoundinallbirds.Somebirdspeciesalsohavea
lungstructurewheretheairflowisbidirectional,calledneopulmonicparabronchi.[47]
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Reptiles
Thelungofmostreptileshasasinglebronchusrunningdownthecentre,fromwhichnumerousbranchesreachouttoindividualpocketsthroughoutthelungs.
Thesepocketsaresimilartoalveoliinmammals,butmuchlargerandfewerinnumber.Thesegivethelungaspongeliketexture.Intuataras,snakes,andsome
lizards,thelungsaresimplerinstructure,similartothatoftypicalamphibians.[48]
Snakesandlimblesslizardstypicallypossessonlytherightlungasamajorrespiratoryorgantheleftlungisgreatlyreduced,orevenabsent.Amphisbaenians,
however,havetheoppositearrangement,withamajorleftlung,andareducedorabsentrightlung.[48]
Bothcrocodiliansandmonitorlizardshavedevelopedlungssimilartothoseofbirds,providinganunidirectionalairflowandevenpossessingairsacs.[49]Thenow
extinctpterosaurshaveseeminglyevenfurtherrefinedthistypeoflung,extendingtheairsacsintothewingmembranesand,inthecaseoflonchodectids,
tupuxuara,andazhdarchoids,thehindlimbs.[50]
Reptilianlungstypicallyreceiveairviaexpansionandcontractionoftheribsdrivenbyaxialmusclesandbuccalpumping.Crocodiliansalsorelyonthehepatic
pistonmethod,inwhichtheliverispulledbackbyamuscleanchoredtothepubicbone(partofthepelvis),whichinturnpullsthebottomofthelungsbackward,
expandingthem.Turtles,whichareunabletomovetheirribs,insteadusetheirforelimbsandpectoralgirdletoforceairinandoutofthelungs.[48]

Amphibians
Thelungsofmostfrogsandotheramphibiansaresimpleandballoonlike,withgasexchangelimitedtotheoutersurfaceof
thelung.Thisisnotveryefficient,butamphibianshavelowmetabolicdemandsandcanalsoquicklydisposeofcarbon
dioxidebydiffusionacrosstheirskininwater,andsupplementtheiroxygensupplybythesamemethod.Amphibians
employapositivepressuresystemtogetairtotheirlungs,forcingairdownintothelungsbybuccalpumping.Thisis
distinctfrommosthighervertebrates,whouseabreathingsystemdrivenbynegativepressurewherethelungsareinflated
byexpandingtheribcage.[51]Inbuccalpumping,thefloorofthemouthislowered,fillingthemouthcavitywithair.The
throatmusclesthenpressesthethroatagainsttheundersideoftheskull,forcingtheairintothelungs.[52]
Duetothepossibilityofrespirationacrosstheskincombinedwithsmallsize,allknownlunglesstetrapodsareamphibians.
Themajorityofsalamanderspeciesarelunglesssalamanders,whichrespiratethroughtheirskinandtissuesliningtheir
mouth.Thisnecessarilyrestricttheirsize:allaresmallandratherthreadlikeinappearance,maximisingskinsurface
relativetobodyvolume.[53]OtherknownlunglesstetrapodsaretheBorneanflatheadedfrog[54]andAtretochoanaeiselti,a
caecilian.[55]

Theaxolotl(Ambystomamexicanum)
retainsitslarvalformwithgillsinto
adulthood

Thelungsofamphibianstypicallyhaveafewnarrowinternalwalls(septa)ofsofttissuearoundtheouterwalls,increasingtherespiratorysurfaceareaandgiving
thelungahoneycombappearance.Insomesalamanderseventhesearelacking,andthelunghasasmoothwall.Incaecilians,asinsnakes,onlytherightlung
attainsanysizeordevelopment.[48]
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Lungfish
Thelungsoflungfisharesimilartothoseofamphibians,withfew,ifany,internalsepta.IntheAustralianlungfish,thereisonlyasinglelung,albeitdividedinto
twolobes.OtherlungfishandPolypterus,however,havetwolungs,whicharelocatedintheupperpartofthebody,withtheconnectingductcurvingaroundand
abovetheesophagus.Thebloodsupplyalsotwistsaroundtheesophagus,suggestingthatthelungsoriginallyevolvedintheventralpartofthebody,asinother
vertebrates.[48]

Invertebrates
Someinvertebrateshave"lungs"thatserveasimilarrespiratorypurposeas,butarenotevolutionarilyrelatedto,
vertebratelungs.Somearachnids,spidersandscorpions,havestructurescalled"booklungs"usedforatmosphericgas
exchange.Somespeciesofspiderhavefourpairsofbooklungsbutmosthavetwopairs.[56]Scorpionshavespiracleson
theirbodyfortheentranceofairtothebooklungs.[57]
Thecoconutcrabisterrestrialandusesstructurescalledbranchiostegallungstobreatheair.[58]Theycannotswimand
woulddrowninwater,yettheypossessarudimentarysetofgills.Theycanbreatheonlandandholdtheirbreath
underwater.[59]Thebranchiostegallungsareseenasadevelopmentaladaptivestagefromwaterlivingtoenableland
living,orfromfishtoamphibian.[60]

Booklungsofspider(showninpink)

Pulmonatesaremostlylandsnailsandslugsthathavedevelopedasimplelungfromthemantlecavity.Anexternallylocatedopeningcalledthepneumostome
allowsairtobetakenintothemantlecavitylung.[61][62]

Evolutionaryorigins
Thelungsoftoday'sterrestrialvertebratesandthegasbladdersoftoday'sfisharebelievedtohaveevolvedfromsimplesacs,asoutpocketingsoftheesophagus,
thatallowedearlyfishtogulpairunderoxygenpoorconditions.[63]Theseoutpocketingsfirstaroseinthebonyfish.Inmostoftherayfinnedfishthesacsevolved
intoclosedoffgasbladders,whileanumberofcarps,trouts,herrings,catfish,andeelshaveretainedthephysostomeconditionwiththesackbeingopentothe
esophagus.Inmorebasalbonyfish,suchasthegar,bichir,bowfinandthelobefinnedfish,thebladdershaveevolvedtoprimarilyfunctionaslungs.[63]Thelobe
finnedfishgaverisetothelandbasedtetrapods.Thus,thelungsofvertebratesarehomologoustothegasbladdersoffish(butnottotheirgills).[64]

Seealso
Drowning
Interstitiallungdisease
https://en.wikipedia.org/wiki/Lung

WikimediaCommonshas
mediarelatedtolungs.
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Liquidbreathing
Lungmicrobiome
Mechanicalventilation

Furtherreading
DrD.R.Johnson:Introductoryanatomy,respiratorysystem(http://www.leeds.ac.uk/chb/lectures/anatomy7.html),leeds.ac.uk
FranlinkInstituteOnline:TheRespiratorySystem(http://sln.fi.edu/biosci/systems/respiration.html),sln.fi.edu
Avianlungsandrespiration(http://www.people.eku.edu/ritchisong/birdrespiration.html),people.eku.edu

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