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RespiratorysystemWikipedia

Respiratorysystem
FromWikipedia,thefreeencyclopedia

Therespiratorysystem(calledalsorespiratoryapparatus,ventilatorysystem)isabiologicalsystem
consistingofspecificorgansandstructuresusedfortheprocessofrespirationinanorganism.Therespiratory
systemisinvolvedintheintakeandexchangeofoxygenandcarbondioxidebetweenanorganismandthe
environment.

Respiratorysystem

Inairbreathingvertebrateslikehumanbeings,respirationtakesplaceintherespiratoryorganscalledlungs.The
passageofairintothelungstosupplythebodywithoxygenisknownasinhalation,andthepassageofairoutof
thelungstoexpelcarbondioxideisknownasexhalationthisprocessiscollectivelycalledbreathingor
ventilation.Inhumansandothermammals,theanatomicalfeaturesoftherespiratorysystemincludetrachea,
bronchi,bronchioles,lungs,anddiaphragm.Moleculesofoxygenandcarbondioxidearepassivelyexchanged,
bydiffusion,betweenthegaseousexternalenvironmentandtheblood.Thisexchangeprocessoccursinthe
alveoli(airsacs)inthelungs.[1]
Infishandmanyinvertebrates,respirationtakesplacethroughthegills.Otheranimals,suchasinsects,have
respiratorysystemswithverysimpleanatomicalfeatures,andinamphibianseventheskinplaysavitalrolein
gasexchange.Plantsalsohaverespiratorysystemsbutthedirectionalityofgasexchangecanbeoppositetothat
inanimals.Therespiratorysysteminplantsalsoincludesanatomicalfeaturessuchasholesontheundersidesof
leavesknownasstomata.[2]

Acomplete,schematicviewofthehuman
respiratorysystemwiththeirpartsandfunctions.
Details

Contents
1 Comparativeanatomyandphysiology
1.1 Horses
1.2 Elephants
1.3 Birds
1.4 Reptiles
1.5 Amphibians
1.6 Fish
2 Anatomyininvertebrates
2.1 Arthropods
2.1.1 Insects
2.2 Molluscs
https://en.wikipedia.org/wiki/Respiratory_system

Identifiers
Latin systemarespiratorium
TA

06.0.00.000(http://www.unifr.ch/ifaa/Publi
c/EntryPage/TA98%20Tree/Entity%20TA9
8%20EN/6.0.00.000%20Entity%20TA98%2
0EN.htm)

FMA 7158(http://xiphoid.biostr.washington.edu/f
ma/fmabrowserhierarchy.html?fmaid=7158)
Anatomicalterminology
[editonWikidata]
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2.2 Molluscs
3 Physiologyinmammals
3.1 Ventilation
3.1.1 Control
3.1.2 MechanicsofInhalationandexhalation
3.2 Gasexchange
3.3 Physiologicalresponsestolowatmosphericpressures
3.4 Immunefunctions
3.5 Metabolicandendocrinefunctionsofthelungs
3.5.1 Vocalization
3.5.2 Temperaturecontrol
3.5.3 Coughingandsneezing
4 Development
4.1 Humansandmammals
5 Disease
6 Plants
7 Footnote
8 References
9 Externallinks

Comparativeanatomyandphysiology
Horses
Horsesareobligatenasalbreatherswhichmeansthattheyaredifferentfrommanyothermammalsbecausetheydonothavetheoptionofbreathingthroughtheir
mouthsandmusttakeinoxygenthroughtheirnoses.

Elephants
Theelephantistheonlyanimalknowntohavenopleuralspace.Rather,theparietalandvisceralpleuraarebothcomposedofdenseconnectivetissueandjoinedto
eachothervialooseconnectivetissue.[3]Thislackofapleuralspace,alongwithanunusuallythickdiaphragm,arethoughttobeevolutionaryadaptations
allowingtheelephanttoremainunderwaterforlongperiodsoftimewhilebreathingthroughitstrunkwhichemergesasasnorkel.[4]

Birds

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Therespiratorysystemofbirdsdifferssignificantlyfromthatfoundinmammals,containinguniqueanatomicalfeaturessuchasairsacs.Thelungsofbirdsalsodo
nothavethecapacitytoinflateasbirdslackadiaphragmandapleuralcavity.Gasexchangeinbirdsoccursbetweenaircapillariesandbloodcapillaries,rather
thaninalveoli.

Reptiles
Theanatomicalstructureofthelungsislesscomplexinreptilesthaninmammals,withreptileslackingtheveryextensive
airwaytreestructurefoundinmammalianlungs.Gasexchangeinreptilesstilloccursinalveolihowever,reptilesdonot
possessadiaphragm.Thus,breathingoccursviaachangeinthevolumeofthebodycavitywhichiscontrolledby
contractionofintercostalmusclesinallreptilesexceptturtles.Inturtles,contractionofspecificpairsofflankmuscles
governsinspirationorexpiration.[5]

Amphibians
Boththelungsandtheskinserveasrespiratoryorgansinamphibians.Theventilationofthelungsinamphibiansuses
XrayvideoofafemaleAmerican
positivepressureventilation.Muscleslowertheflooroftheoralcavity,enlargingitanddrawinginairthroughthenostrils
alligatorwhilebreathing.
(whichusesthesamemechanicspressure,volume,anddiffusionasamammalianlung).Withthenostrilsandmouth
closed,theflooroftheoralcavityisforcedup,whichforcesairdownthetracheaintothelungs.Theskinoftheseanimalsis
highlyvascularizedandmoist,withmoisturemaintainedviasecretionofmucusfromspecializedcells.Whilethelungsareofprimaryimportancetobreathing
control,theskin'suniquepropertiesaidrapidgasexchangewhenamphibiansaresubmergedinoxygenrichwater.[6]

Fish
Inmostfish,respirationtakesplacethroughgills.(Seealsoaquaticrespiration.)Lungfish,however,dopossessoneortwolungs.Thelabyrinthfishhave
developedaspecialorganthatallowsthemtotakeadvantageoftheoxygenoftheair.

Anatomyininvertebrates
Arthropods
Somespeciesofcrabusearespiratoryorgancalledabranchiostegallung.[7]Itsgilltissueisformedsoastoincreasethesurfaceareaandthelungismoresuitedto
takingoxygenfromtheairthanfromwater.Someofthesmallestspidersandmitescanbreathesimplybyexchanginggasthroughthesurfaceofthebody.Larger
spiders,scorpionsandotherarthropodsuseaprimitivebooklung.
Insects
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Mostinsectsbreathpassivelythroughtheirspiracles(specialopeningsintheexoskeleton)andtheairreachesthebodybymeansofaseriesofsmallerandsmaller
pipescalled'trachaea'whentheirdiameterisrelativelylargeand'tracheoles'whentheirdiameterisverysmall.Diffusionofgasesiseffectiveoversmalldistances
butnotoverlargerones,thisisoneofthereasonsinsectsareallrelativelysmall.Insectswhichdonothavespiraclesandtrachaea,suchassomeCollembola,
breathedirectlythroughtheirskins,alsobydiffusionofgases.[8]Thenumberofspiraclesaninsecthasisvariablebetweenspecies,howevertheyalwayscomein
pairs,oneoneachsideofthebody,andusuallyonepersegment.SomeoftheDiplurahaveeleven,withfourpairsonthethorax,butinmostoftheancientforms
ofinsects,suchasDragonfliesandGrasshopperstherearetwothoracicandeightabdominalspiracles.However,inmostoftheremaininginsectsthereareless.It
isatthislevelofthetracheolesthatoxygenisdeliveredtothecellsforrespiration.Thetracheaarewaterfilledduetothepermeablemembraneofthesurrounding
tissues.Duringexercise,thewaterlevelretractsduetotheincreaseinconcentrationoflacticacidinthemusclecells.Thislowersthewaterpotentialandthewater
isdrawnbackintothecellsviaosmosisandairisbroughtclosertothemusclecells.Thediffusionpathwayisthenreducedandgasescanbetransferredmore
easily.
Insectswereoncebelievedtoexchangegaseswiththeenvironmentcontinuouslybythesimplediffusionofgasesintothetrachealsystem.Morerecently,however,
largevariationininsectventilatorypatternshavebeendocumentedandinsectrespirationappearstobehighlyvariable.Somesmallinsectsdodemonstrate
continuousrespirationandmaylackmuscularcontrolofthespiracles.Others,however,utilizemuscularcontractionoftheabdomenalongwithcoordinated
spiraclecontractionandrelaxationtogeneratecyclicalgasexchangepatternsandtoreducewaterlossintotheatmosphere.Themostextremeformofthese
patternsistermeddiscontinuousgasexchangecycles(DGC).[9]

Molluscs
Molluscsgenerallypossessgillsthatallowexchangeofoxygenfromanaqueousenvironmentintothecirculatorysystem.Theseanimalsalsopossessaheartthat
pumpsbloodwhichcontainshemocyaninineasitsoxygencapturingmolecule.Hence,thisrespiratorysystemissimilartothatofvertebratefish.Therespiratory
systemofgastropodscanincludeeithergillsoralung.

Physiologyinmammals
Ventilation
Inrespiratoryphysiology,theventilationrateistherateatwhichgasentersorleavesthelung.Itiscategorizedunderthefollowingdefinitions:

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Measurement

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Equation

Description

Minute
ventilation

tidalvolume*respiratoryrate[1][2] thetotalvolumeofgasenteringthelungsperminute.

Alveolar
ventilation

(tidalvolumedeadspace)*
respiratoryrate[1]

thevolumeofgasperunittimethatreachesthealveoli,therespiratoryportionsofthelungswheregas
exchangeoccurs.

Deadspace
ventilation

deadspace*respiratoryrate[3]

thevolumeofgasperunittimethatdoesnotreachtheserespiratoryportions,butinsteadremainsinthe
airways(trachea,bronchi,etc.).

Control
Ventilationoccursviatherespiratorycenterinthemedullaoblongataandtheponsofthebrainstem.Theseareasofthebrainformaseriesofinterconnectedbrain
cellswithinthelowerandmiddlebrainstemwhichreceiveinformationaboutthearterialpartialpressureofoxygen(
)andthearterialpartialpressureof
carbondioxide(
).Thisinformationdeterminestheaveragemediumtermrateofventilationofthealveoliofthelungs,tokeepthearterial
and
constant.Therespiratorycenterdoessoviamotorneuronswhichactivatetheskeletalmusclesofrespiration(inparticularthediaphragm).Sectionsthroughthe
medullaoblongataandponssuggestthattherespiratorycenterconsistsofapneumotaxiccenter,anapneusticcenter,andadorsalandventralrespiratorygroups.
Thislattergroupisespeciallysensitiveduringinfancy,andtheneuronscanbedestroyediftheinfantisdroppedand/orshakenviolently.Theresultcanbedeath
dueto"shakenbabysyndrome".[10]
Thebreathingrateincreaseswhenthe
inthebloodincreases.Thisisdetectedby"peripheral"
chemoreceptorsintheaorticandcarotidbodies,butin
particularbythe"central"
chemoreceptorsontheanteriorsurfaceofthemedullaoblongataofthebrainstem.(The"peripheralchemoreceptors"aremore
sensitivetothearterial
thantheyaretothearterial
.)Exercisealsoincreasestherespiratoryrate,partlyinresponsetothemovementofthelimbs
detectedbyproprioceptorsinthemusclesandjoints,anincreaseinbodytemperature,thereleaseofepinephrinefromtheadrenalglands,andfrommotorimpulses
originatingfromthebrain.[11]Informationreceivedfromstretchreceptorsinthelungslimitstidalvolume(i.e.thedepthofinhalationandexhalation).
MechanicsofInhalationandexhalation
Inmammals,breathingin(inhalation)atrestisprimarilyduetothecontractionandflatteningofthediaphragm,adomedmusclethatseparatesthethoraciccavity
fromtheabdominalcavity.Whenthediaphragmcontractsitpushestheabdominalorgansdownward,butsincethepelvicfloorpreventsthelowermostabdominal
organsfrommovinginthatdirection,theabdomen,infact,bulgesforwards(oroutwards).Intheprocessthesizeofthethoraciccavityhasincreasedinvolume(as
hasthevolumeofthebodyasawhole).Thisincreasedthoracicvolumeresultsinafallinpressureinthethorax,whichcausestheexpansionofthelungs.During
exhalation(breathingout),atrest,thediaphragmrelaxes,returningthechestandabdomentoapositionwhichisdeterminedbytheiranatomicalelasticity(i.e.the
positioninthecadaver,orinananimalthathasbeengivenamusclerelaxantunderanesthesia).Thisisthe"restingmidposition"ofthethoraxwhenthelungs

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containthefunctionalresidualcapacityofair,whichintheadulthuman
hasavolumeofabout2.5liters.[12]Restingexhalationlastsabouttwiceas
longasinhalationbecausethediaphragmrelaxesmoregentlythanit
contractsduringinhalation.Thispreventsunduenarrowingoftheairways,
fromwhichtheairescapesmoreeasilythanfromthealveoli.

Theeffectofthecontractionofthe
accessorymusclesofinhalation,pullingthe
frontoftheribcageupwards.This
increasestheanteroposteriordiameterof
thethorax,contributingtotheexpansionin
thevolumeofthechest.Asimilareffect
causesthetransversediameterofthechest
toincrease,becausenotonlydotheribs
slantdownwardsfromthebacktothe
front,but,inthecaseofthelowerribs,
alsofromthemidlinedownwardstothe
sidesofthechest.

Duringheavybreathing(hyperventilation),as,forinstance,during
exercise,the"accessorymusclesofinhalation"(ofwhichthefirsttobe
recruitedaretheintercostalmuscles,butincludealargenumberofother
musclesseebelow)pulltheribsupwards,bothinthefrontandonthe
sides.Thisincreasesthevolumeoftheribcage,addingtothevolume
increasecausedbythedescendingdiaphragm.Duringtheensuing
exhalationtheribcageisactivelypulleddownwards(frontandsides)by
theabdominalmuscles,whichnotonlydecreasesthesizeoftheribcage,
butalsopushestheabdominalorgansupwardsagainstthediaphragm
whichconsequentlybulgesdeeplyintothethorax.Theendexhalatory
lungvolumeisnowwellbelowtherestingmidpositionandcontainsfar
lessairthantheresting"functionalresidualcapacity".However,ina
normalmammal,thelungscannotbeemptiedcompletely.Inanadult
humanthereisalwaysstillatleast1literofresidualairleftinthelungs
aftermaximumexhalation.

Theentirelyunconsciousandautomaticbreathingonwhichthelifeofthe
animaldependscanbetemporarilyoverriddenbyconsciousoremotion
drivenmovementsofairinandoutofthelungs.Speechinhumansis
generatedbyaspecializedformofexhalation,butotherformsofcommunication(e.g.crying,yelping,yawning,
barking,baying,hissing,panting,sighing,shouting,laughingetc.)alsorelyonabalancebetweenbreathingforblood
gashomeostasisandtheemotionalorothermessagesthatneedtobeconveyedtotheanimal'sconspecifics.
Tenmusclescanbeusedforinhalation:[13]
Diaphragm,IntercostalMuscles,Scalenes,PectoralisMinor,SerratusAnterior,Sternocleidomastoid,Levator
Costarum,Upper/SuperiorTrapezius,LatissimusDorsi,andSubclavius.
Eightareusedforforcedexhalation:[14]
Internalintercostal,ObliquusInternus,ObliquusExternus,LevatorAni,TriangularisSterni,Transversalis,
Pyramidalis,andRectusAbdominus.
https://en.wikipedia.org/wiki/Respiratory_system

Acartoonillustratingthemechanismsfor
forcefulinhalation(left),andforceful
exhalation(right).Duringforceful
inhalationthediaphragm(thedomed,
almosthorizontalstructureinred,between
thethoraciccavity(uppercompartment)
andabdominalcavity(lower
compartment),contractsforcingthe
abdominalcontentsdownwards,causing
theabdomentobulgeoutwards.Atthe
sametimetheaccessorymusclesof
inhalationcausethetransversediameterof
thethoraxtoincreaseasdescribedinthe
illustrationofthemovementoftheribson
theleft.Duringforcedexhalationthe
powerfulmusclesoftheabdominalwall
pulltheloweredgesoftheribcage
downwardsdecreasingtheanteroposterior
andtransversediametersofthechest
cavity,whileatthesametimeforcingthe
abdominalorgansupagainstthediaphragm
causingittobulgedeeplyintothe
chest.Theforcewithwhichaircanbe
expelledfromthelungsisconsiderable
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Gasexchange

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greaterthantheforcewithwhichaircan
beinhaled.Thisistheresultofthepower
ofthemusclesoftheabdominalwall
exceedingthatofalltheaccessorymuscles
ofinhalation.Themostvigorous
exhalatoryeffortsoccurduringcoughing,
sneezingandtheblowingoutof,for
instance,candles.

Themajorfunctionoftherespiratorysystemisthe
equilibrationofthebloodgaseswiththoseinthe
alveolarair.Thelungsofanaveragepersonatrest,
andbreathingnormallycontainbetweenabout2.5
and3litersofair.Thissemistagnantvolumeof
air[note1]thatalwaysremainsinthelungalveoli
afteranormalexhalationistermedthefunctionalresidualcapacity.[12]Witheachbreathonly
about350ml(i.e.lessthan15%)ofthisalveolarairisexpelledintotheambientairtobe
replacedwiththesamevolumeoffresh,butmoistened,atmosphericair.Itisthereforeobvious
thatthecompositionofthealveolarair(orfunctionalresidualcapacity)changesverylittle
undernormalcircumstances:thealveolarpartialpressureofoxygen(
)remainsveryclose
to105mmHg(14kPa),andthatofcarbondioxide(
)variesminimallyfrom40mmHg
(5.3kPa)throughouttherespiratorycycle(ofinhalationandexhalation).[12]Thecorresponding
partialpressuresofoxygenandcarbondioxideintheambient(dry)airatsealevelare160
mmHg(21kPa)and0.3mmHg(0.04kPa)respectively.[12]
Thechangesinthecompositionofthealveolarairduringanormal
breathingcycleatrest.Thescaleontheleft,andtheblueline,
indicatethepartialpressuresofcarbondioxideinkPa,whilethaton
therightandtheredline,indicatethepartialpressuresofoxygen,
alsoinkPa(toconvertkPaintommHg,multiplyby7.5).

Thismarkeddifferencebetweenthecompositionofthealveolarairandthatoftheambientair
canbemaintainedbecausethefunctionalresidualcapacityiscontainedinadeadendsac
connectedtotheoutsideairbyafairlynarrowandrelativelylongtube(theairways:nose,
pharynx,larynx,trachea,bronchiandtheirbranchesdowntothebronchioles).Thisanatomy
andthefactthatthelungsarenotemptiedandreinflatedwitheachbreath,providestheanimal
withaveryspecialportableatmosphere,whosecompositiondifferssignificantlyfromthe
[15]
presentdayambientair. Itisthisportableatmosphere(thefunctionalresidualcapacity)towhichthebloodandthereforethebodytissuesareexposednotto
theoutsideair.
Allthebloodreturningfromthebodytissuestotherightsideoftheheartflowsthroughthepulmonarycapillariesbeforebeingpumpedaroundthebodyagain.On
itspassagethroughthelungsthebloodcomesintoclosecontactwiththealveolarair,separatedfromitbyaverythindiffusionmembranewhichisonlyabout0.5
mthick.[12]Thegaspressuresinthebloodwillthereforerapidlyequilibratewiththoseinthealveoli,ensuringthatthearterialbloodthatcirculatestoallthe
tissuesthroughoutthebodyhasanoxygentensionof105mmHg(14kPa),andacarbondioxidetensionof40mmHg(5.3kPa).Thesearterialpartialpressuresof
oxygenandcarbondioxidearehomeostaticallycontrolled.Ariseinthearterial
,and,toalesserextent,afallinthearterial
,willreflexlycausedeeper
andfasterbreathingtillthebloodgastensionreturntonormal.Theconversehappenswhenthecarbondioxidetensionfalls,or,againtoalesserextent,theoxygen
tensionrises:therateanddepthofbreathingarereducedtillbloodgasnormalityisrestored.
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Sincethebloodarrivinginthepulmonarycapillarieshasa
of,onaverage,45mmHg(6kPa),whilethepressure
inthealveolarairis105mmHg,therewillbeanetdiffusionofoxygenintothecapillaryblood,changingthe
compositionofthe3litersofalveolarairslightly.Similarly,sincethebloodarrivinginthepulmonarycapillarieshasa
ofalsoabout45mmHg,whereasthatofthealveolarairis40mmHg(5.3kPa),thereisanetmovementof
carbondioxideoutofthecapillariesintothealveoli.Thechangesbroughtaboutbytheseflowsofindividualgasesinto
andoutofthefunctionalresidualcapacitynecessitatethereplacementofabout15%ofthealveolarairwithambient
airevery5secondsorso.Thisisverytightlycontrollednotonlybythemonitoringofthearterialbloodgases(which
accuratelyreflectcompositionofthealveolarair)bytheaorticandcarotidbodies,butalsobyoxygenandcarbon
dioxidesensorsinthelungs,thoughthelatterprimarilydeterminethediametersofthebronchiolesandpulmonary
capillaries,andarethereforeresponsiblefordirectingtheflowofairandbloodtodifferentpartsofthelungs.
Itisonlyasaresultofmaintainingthecompositionofthe3litersalveolarairthatwitheachbreathsomecarbon
dioxideisdischargedintotheatmosphereandsomeoxygenistakenupfromtheoutsideair.Ifmorecarbondioxide
thanusualhasbeenlostbyashortperiodofhyperventilation,respirationwillbesloweddownorhalteduntilthe
alveolar
hasreturnedto40mmHg(5.3kPa).Itisthereforestrictlyspeakinguntruethattheprimaryfunctionof
therespiratorysystemistoridthebodyofcarbondioxidewaste.Thecarbondioxidethatisbreathedoutwitheach
breathcouldprobablybemorecorrectlybeseenasabyproductofthebodysextracellularfluidcarbondioxideandpH
homeostats
Ifthesehomeostatsarecompromised,thenarespiratoryacidosis,orarespiratoryalkalosiswilloccur.Inthelongrun

thesecanbecompensatedbyrenaladjustmentstotheH+andHCO3 concentrationsintheplasmabutsincethistakes
time,thehyperventilationsyndromecan,forinstance,occurwhenagitationoranxietycauseapersontobreathefast
anddeeplythuscausingadistressingrespiratoryalkalosisthroughtheblowingoffoftoomuchCO2fromtheblood
intotheoutsideair.

Physiologicalresponsestolowatmosphericpressures

Ahighlydiagrammaticillustrationofthe
processofgasexchangeinthemammalian
lungs,emphasizingthedifferencesbetween
thegascompositionsoftheambientair,
thealveolarair(lightblue)withwhichthe
pulmonarycapillarybloodequilibrates,
andthebloodgastensionsinthe
pulmonaryarterial(bluebloodenteringthe
lungontheleft)andvenousblood(red
bloodleavingthelungontheright).All
thegastensionsareinkPa.Toconvertto
mmHg,multiplyby7.5.

Thealveoliareopen(viatheairways)totheatmosphere,withtheresultthatalveolarairpressureisexactlythesame
astheambientairpressureatsealevel,ataltitude,orinanyartificialatmosphere(e.g.adivingchamber,or
decompressionchamber)inwhichtheindividualisbreathingfreely.Withexpansionofthelungs(throughloweringof
thediaphragmandexpansionofthethoraciccage)thealveolarairnowoccupiesalargervolume,anditspressurefallsproportionately,causingairtoflowinfrom
thesurroundings,throughtheairways,tillthepressureinthealveoliisonceagainattheambientairpressure.Thereverseobviouslyhappensduringexhalation.
Thisprocess(ofinhalationandexhalation)isexactlythesameatsealevel,asontopofMt.Everest,orinadivingchamberordecompressionchamber.

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However,asonerisesabovesealevelthedensityoftheairdecreasesexponentially,halving
approximatelywithevery5500mriseinaltitude.[16]Sincethecompositionoftheatmospheric
airisalmostconstantbelow80km,asaresultofthecontinuousmixingeffectoftheweather,
theconcentrationofoxygenintheair(mmolsO2perliterofambientair)decreasesatthesame
rateasthefallinairpressurewithaltitude.[17]Therefore,inordertobreatheinthesame
amountofoxygenperminute,thepersonhastoinhaleaproportionatelygreatervolumeofair
perminuteataltitudethanatsealevel.Thisisachievedbybreathingdeeperandfaster(i.e.
hyperventilating)thanatsealevel(seebelow).
Thereis,however,a
complicationthatincreases
thevolumeofairthatneeds
tobeinhaledperminute
(respiratoryminutevolume)
toprovidethesameamount
ofoxygentothelungsat
altitudeasatsealevel.
Agraphshowingtherelationshipbetweentotalatmosphericpressure
Duringinhalationtheairis
andaltitudeabovesealevel.
warmedandsaturatedwith
watervaporduringits
passagethroughthenoseandpharynx.Saturatedwatervaporpressureisdependentonlyon
temperature.Atabodycoretemperatureof37Citis6.3kPa(47.0mmHg),irrespectiveofany
AerialphotoofMountEverestfromthesouth,behindNuptseand
otherinfluences,includingaltitude.[18]Thusatsealevel,wheretheambientatmospheric
Lhotse.
pressureisabout100kPa,themoistenedairthatflowsintothelungsfromthetracheaconsists
ofwatervapor(6.3kPa),nitrogen(74.0kPa),oxygen(19.7kPa)andtraceamountsofcarbon
dioxideandothergases(atotalof100kPa).Indryairthepartialpressureofoxygenatsealevelis21.0kPa,comparedtothe19.7kPaofoxygenenteringthe
alveolarair.(Thetrachealpartialpressureofoxygen(19.7kPa)is21%of[100kPa6.3kPa]).AtthesummitofMt.Everest(atanaltitudeof8,848mor
29,029ft)thetotalatmosphericpressureis33.7kPa,ofwhich7.1kPa,or21%,isoxygen.[16]Theairenteringthelungsalsohasatotalpressureof33.7kPa,of
which6.3kPais,unavoidably,watervapor(asitisatsealevel).Thisreducesthepartialpressureofoxygenenteringthealveolito5.8kPa,or21%of[33.7kPa
6.3kPa=27.4kPa].Thereductioninthepartialpressureofoxygenintheinhaledairisthereforesubstantiallygreaterthanthereductionofthetotalatmospheric
pressureataltitudewouldsuggest(onMtEverest:5.8kPavs.7.1kPa).
Afurtherminorcomplicationexistsataltitude.Considerthefollowingtheoreticalscenario.Ifthevolumeofthelungsweretobeinstantaneouslydoubledatthe
beginningofinhalation,theairpressureinsidethelungswouldbehalved.Thishappensregardlessofaltitude.Thus,halvingofthesealevelairpressure(100kPa)
resultsinanintrapulmonaryairpressureof50kPa.Doingthesameat5500m,wheretheatmosphericpressureisonly50kPa,theintrapulmonaryairpressurefalls
to25kPa.Therefore,thesamechangeinlungvolumeatsealevelresultsina50kPadifferenceinpressurebetweentheambientairandtheintrapulmonaryair,
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whereasitresultinadifferenceofonly25kPaat5500m.Thedrivingpressureforcingairintothelungsduringinhalationisthereforehalvedatthisaltitude.The
rateofinflowofairintothelungsduringinhalationatsealevelisthereforetwicethatwhichoccursat5500m.However,inreality,inhalationandexhalation
occurfarmoregentlyandlessabruptlythanintheexamplegiven.Thedifferencesbetweentheatmosphericandintrapulmonarypressures,drivingairinandoutof
thelungsduringthebreathingcycle,areintheregionofonly23kPa.[19][20]Adoublingormoreofthesesmallpressuredifferencescouldbeachievedbyonly
veryminoradjustmentstothebreathingeffortathighaltitudes.
Alloftheaboveinfluencesoflowatmosphericpressuresonbreathingareaccommodatedprimarilybyhyperventilating(orhyperpnea).Theexactdegreeof
hyperventilationisdeterminedbythebloodgashomeostat,whichregulatesthepartialpressuresofoxygen(
)andcarbondioxide(
)inthearterialblood.
Thishomeostatprioritizestheregulationofthearterialpartialpressureofcarbondioxideoverthatofoxygenatsealevel.Thatistosay,atsealevelthearterial
ismaintainedatverycloseto5.3kPa(or40mmHg)underawiderangeofcircumstances,attheexpenseofthearterial
,whichisallowedtovarywithin
averywiderangeofvalues,beforeelicitingacorrectiveventilatoryresponse.However,whentheatmosphericpressure(andthereforethepartialpressureof
oxygenintheambientair)fallstobelow75%ofitsvalueatsealevel,oxygenhomeostasisisgivenpriorityovercarbondioxidehomeostasis.Thisswitchover
occursatanelevationofabout2500m(orabout8000ft).Ifthisswitchoccursrelativelyabruptly,thehyperventilationathighaltitudewillcauseaseverefallin
thearterialpartialpressureofcarbondioxide,withaconsequentriseinthepHofthearterialplasma.Thisisonecontributortohighaltitudesickness.Ontheother
hand,iftheswitchtooxygenhomeostasisisincomplete,thenhypoxiamaycomplicatetheclinicalpicturewithpotentiallyfatalresults.
Thereareoxygensensorsinthesmallerbronchiandbronchioles.Inresponsetolowpartialpressuresofoxygenintheinhaledairthesesensorsreflexlycausethe
pulmonaryarteriolestoconstrict.[21](Thisistheexactoppositeofthecorrespondingreflexinthetissues,wherelowarterial
valuescausearteriolar
vasodilation.)Ataltitudethiscausesthepulmonaryarterialpressuretoriseresultinginamuchmoreevendistributionofbloodflowtothelungsthanoccursatsea
level.Atsealevelthepulmonaryarterialpressureisverylow,withtheresultthattheapices(tops)ofthelungsreceivefarlessbloodthanthebasis,whichare
relativelyoverperfusedwithblood.Itisonlyinmiddleofthelungs(halfwayfromthebasestotheapices)thatthebloodandairflowtothealveoliareideally
matched.Ataltitudethisvariationintheventilation/perfusionratioofalveolifromthetopsofthelungstothebottomsiseliminated,withallthealveoliperfused
andventilatedinmoreorlessthephysiologicallyidealmanner.Thisisafurtherimportantcontributortotheacclimatatizationtohighaltitudesandlowoxygen
pressures.
Thekidneysmeasuretheoxygencontent(ratherthanthe

)ofthearterialblood.Whentheoxygencontentofthebloodischronicallylow,asathighaltitude,

theoxygensensitivekidneycellssecreteerythropoietin(EPO)intotheblood.[22]Thishormonestimulatestheredbonemarrowtoincreaseitsrateofredcell
production,whichleadstoanincreaseinthehematocritoftheblood,andaconsequentincreaseinitsoxygencarryingcapacity(duetothenowhighhemoglobin
contentoftheblood).Inotherwords,atthesamearterial
,apersonwithahighhematocritcarriesmoreoxygenperliterofbloodthanapersonwithalower
hematocritdoes.Highaltitudedwellersthereforehavehigherhematocritsthansealevelresidents.[22][23]

Immunefunctions
Airwayepithelialcellscansecreteavarietyofmoleculesthataidinthedefenseoflungs.Secretoryimmunoglobulins(IgA),collectins(includingSurfactantAand
D),defensinsandotherpeptidesandproteases,reactiveoxygenspecies,andreactivenitrogenspeciesareallgeneratedbyairwayepithelialcells.Thesesecretions
canactdirectlyasantimicrobialstohelpkeeptheairwayfreeofinfection.Airwayepithelialcellsalsosecreteavarietyofchemokinesandcytokinesthatrecruit
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thetraditionalimmunecellsandotherstositeofinfections.
Mostoftherespiratorysystemislinedwithmucousmembranesthatcontainmucosalassociatedlymphoidtissue,whichproduceswhitebloodcellssuchas
lymphocytes.

Metabolicandendocrinefunctionsofthelungs
Inadditiontotheirfunctionsingasexchange,thelungshaveanumberofmetabolicfunctions.Theymanufacturesurfactantforlocaluse,asnotedabove.They
alsocontainafibrinolyticsystemthatlysesclotsinthepulmonaryvessels.Theyreleaseavarietyofsubstancesthatenterthesystemicarterialbloodandthey
removeothersubstancesfromthesystemicvenousbloodthatreachthemviathepulmonaryartery.Prostaglandinsareremovedfromthecirculation,buttheyare
alsosynthesizedinthelungsandreleasedintothebloodwhenlungtissueisstretched.Thelungsalsoactivateonehormonethephysiologicallyinactive
decapeptideangiotensinIisconvertedtothepressor,aldosteronestimulatingoctapeptideangiotensinIIinthepulmonarycirculation.Thereactionoccursinother
tissuesaswell,butitisparticularlyprominentinthelungs.Largeamountsoftheangiotensinconvertingenzymeresponsibleforthisactivationarelocatedonthe
surfaceoftheendothelialcellsofthepulmonarycapillaries.Theconvertingenzymealsoinactivatesbradykinin.Circulationtimethroughthepulmonary
capillariesislessthanonesecond,yet70%oftheangiotensinIreachingthelungsisconvertedtoangiotensinIIinasingletripthroughthecapillaries.Fourother
peptidaseshavebeenidentifiedonthesurfaceofthepulmonaryendothelialcells.
Vocalization
Themovementofgasthroughthelarynx,pharynxandmouthallowshumanstospeak,orphonate.Vocalization,orsinging,inbirdsoccursviathesyrinx,anorgan
locatedatthebaseofthetrachea.Thevibrationofairflowingacrossthelarynx(vocalcords),inhumans,andthesyrinx,inbirds,resultsinsound.Becauseofthis,
gasmovementisextremelyvitalforcommunicationpurposes.
Temperaturecontrol
Pantingindogs,catsandsomeotheranimalsprovidesameansofcontrollingbodytemperature.Thisphysiologicalresponseisusedasacoolingmechanism.
Coughingandsneezing
Irritationofnerveswithinthenasalpassagesorairways,caninduceacoughreflexandsneezing.Theseresponsescauseairtobeexpelledforcefullyfromthe
tracheaornose,respectively.Inthismanner,irritantscaughtinthemucuswhichlinestherespiratorytractareexpelledormovedtothemouthwheretheycanbe
swallowed.Duringcoughing,contractionofthesmoothmusclenarrowsthetracheabypullingtheendsofthecartilageplatestogetherandbypushingsofttissue
outintothelumen.Thisincreasestheexpiredairflowratetodislodgeandremoveanyirritantparticleormucus.

Development
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Humansandmammals
Therespiratorysystemliesdormantinthehumanfetusduringpregnancy.Atbirth,therespiratorysystembecomesfullyfunctionaluponexposuretoair,although
somelungdevelopmentandgrowthcontinuesthroughoutchildhood.[24]Pretermbirthcanleadtoinfantswithunderdevelopedlungs.Theselungsshow
incompletedevelopmentofthealveolartypeIIcells,cellsthatproducesurfactant.Thelungsofpreterminfantsmaynotfunctionwellbecausethelackof
surfactantleadstoincreasedsurfacetensionwithinthealveoli.Thus,manyalveolicollapsesuchthatnogasexchangecanoccurwithinsomeormostregionsofan
infant'slungs,aconditiontermedrespiratorydistresssyndrome.Basicscientificexperiments,carriedoutusingcellsfromchickenlungs,supportthepotentialfor
usingsteroidsasameansoffurtheringdevelopmentoftypeIIalveolarcells.[25]Infact,onceaprematurebirthisthreatened,everyeffortismadetodelaythe
birth,andaseriesofsteroidshotsisfrequentlyadministeredtothemotherduringthisdelayinanefforttopromotelunggrowth.[26]

Disease
Disordersoftherespiratorysystemcanbeclassifiedintofourgeneralareas:
Obstructiveconditions(e.g.,emphysema,bronchitis,asthma)
Restrictiveconditions(e.g.,fibrosis,sarcoidosis,alveolardamage,pleuraleffusion)
Vasculardiseases(e.g.,pulmonaryedema,pulmonaryembolism,pulmonaryhypertension)
Infectious,environmentalandother"diseases"(e.g.,pneumonia,tuberculosis,asbestosis,particulatepollutants):
Coughingisofmajorimportance,asitisthebody'smainmethodtoremovedust,mucus,saliva,andotherdebrisfromthelungs.Inabilitytocoughcanleadto
infection.Deepbreathingexercisesmayhelpkeepfinerstructuresofthelungsclearfromparticulatematter,etc.
Therespiratorytractisconstantlyexposedtomicrobesduetotheextensivesurfacearea,whichiswhytherespiratorysystemincludesmanymechanismstodefend
itselfandpreventpathogensfromenteringthebody.
DisordersoftherespiratorysystemareusuallytreatedinternallybyapulmonologistandRespiratoryTherapist.

Plants
Plantsusecarbondioxidegasintheprocessofphotosynthesis,andexhaleoxygengasaswaste.Thechemicalequationofphotosynthesisis6CO2(carbon
dioxide)and6H2O(water)andthatmakes6O2(oxygen)andC6H12O6(glucose).Whatisnotexpressedinthechemicalequationisthecaptureofenergyfrom
sunlightwhichoccurs.Photosynthesisuseselectronsonthecarbonatomsastherepositoryforthatenergy.Respirationistheoppositeofphotosynthesis.It
reclaimstheenergytopowerchemicalreactionsincells.Insodoingthecarbonatomsandtheirelectronsarecombinedwithoxygenformingagaswhichiseasily
removedfromboththecellsandtheorganism.Plantsusebothprocesses,photosynthesistocapturetheenergyandrespirationtouseit.

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Plantrespirationislimitedbytheprocessofdiffusion.Plantstakeincarbondioxidethroughholesontheundersidesoftheirleavesknownasstomaorpores.
However,mostplantsrequirelittleair.Mostplantshaverelativelyfewlivingcellsoutsideoftheirsurfacebecauseair(whichisrequiredformetaboliccontent)can
penetrateonlyskindeep.However,mostplantsarenotinvolvedinhighlyaerobicactivities,andthushavenoneedoftheselivingcells.

Footnote
1.Althoughthefunctionalresidualcapacityisdescribedhereasasemistagnantvolumeofair,thisisonlytrueinthesensethatalakeofwaterwithasmallinletandoutlet
seemsstagnantcomparedwiththerestofriver.Inthecaseofthefunctionalresidualcapacitytheentirevolumeoftrappedair(orthelakeintheriveranalogy)isalways
thoroughlymixedwiththeincominginhaledair.Thisisbroughtaboutbythemicroscopicsubdivisionsofthefunctionalresidualcapacityintomanybillionsofsmaller,
minuteairsacs,thealveoli,intowhichtheinhaledairentersturbulently.Undernormalcircumstancesthefunctionalresidualcapacityisfarfromstagnantbutalways
representsathoroughlystirredlargevolumeofthestoredairwithasmallamountofdiluted(withwatervapor)freshinhaledair,aftereachinbreath.

References
1.Maton,AntheaHopkins,JeanSusanJohnson,CharlesWilliamMcLaughlin,
MaryannaQuonWarner,DavidLaHartWright,Jill(2010).HumanBiologyand
Health.EnglewoodCliffs:PrenticeHall.pp.108118.ISBN0134234359.
2.West,JohnB.Respiratoryphysiologytheessentials.Baltimore:Williams&
Wilkins.pp.110.ISBN0683089374.
3.West,JohnB.Ravichandran(1993)."Snorkelbreathingintheelephantexplains
theuniqueanatomyofitspleura".RespirationPhysiology.126(1):18.
doi:10.1016/S00345687(01)002031.PMID11311306.
4.West,JohnB.(2002)."Whydoesn'ttheelephanthaveapleuralspace?".News
PhysiolSci.17:4750.PMID11909991.
5.Respiratorysystem(http://www.britannica.com/EBchecked/topic/498684/reptile/38
473/Respiratorysystem).EncyclopdiaBritannica.
6.Gottlieb,GJacksonDC(1976)."Importanceofpulmonaryventilationin
respiratorycontrolinthebullfrog".AmJPhysiol.230(3):60813.PMID4976.
7.J.Halperin,M.Ansaldo,G.N.Pellerano&C.M.Luquet(2000)."Bimodal
breathingintheestuarinecrabChasmagnathusgranulatusDana1851
physiologicalandmorphologicalstudies".ComparativeBiochemistryand
PhysiologyPartA:Molecular&IntegrativePhysiology.126(3):341349.
doi:10.1016/S10956433(00)002166.PMID10964029.
8.TheEarthLifeWeb,InsectMorphologyandAnatomy(http://www.earthlife.net/in
sects/anatomy.html).Earthlife.net.Retrievedon20130421.
9.Lighton,JRB(January1996)."Discontinuousgasexchangeininsects".AnnuRev
Entomology.41:309324.doi:10.1146/annurev.en.41.010196.001521.

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10.ThecombinationofacryingbabyandafrustratedparentorcaregiverCANBE
DEADLY(http://www.usask.ca/medicine/prevent/sbs.html).Saskatchewan
PreventionInstitute.usask.ca.
11."Respiration".HarveyProject.Retrieved27July2012.
12.Tortora,GerardJ.Anagnostakos,NicholasP.(1987).Principlesofanatomyand
physiology(Fifthed.).NewYork:Harper&Row,Publishers.pp.570572.
ISBN0063507293.
13.AllYouNeedtoKnowAboutInspiratoryMusclesPartI|SwimmingScience(htt
p://www.swimmingscience.net/2011/10/allyouneedtoknowaboutinspiratory.ht
ml)
14.AllYouNeedtoKnowAboutInspiratoryMusclesPartII(http://www.swimmings
cience.net/2011/10/allyouneedtoknowaboutinspiratorymusclespartii.html)
15.Lovelock,James(1991).HealingGaia:PracticalmedicineforthePlanet.New
York:HarmonyBooks.pp.2134,7388.ISBN0517578484.
16."Onlinehighaltitudeoxygencalculator".altitude.org.Retrieved15August2007.
17.Tyson,P.D.PrestonWhite,R.A.(2013).TheweatherandclimateofSouthern
Africa.CapeTown:OxfordUniversityPress.pp.310,1416,360.
ISBN9780195718065.
18.Diem,K.Lenter,C.(1970).ScientificTables(Seventhed.).Basle,Switzerland:
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19.Koen,ChrisvanL.Koeslag,JohanH.(1995)."Onthestabilityofsubatmospheric
intrapleuralandintracranialpressures".NewsinPhysiologicalSciences.10:176
178.
20.West,J.B.(1985).Respiratoryphysiology:theessentials.Baltimore:Williams&
Wilkins.pp.2130,8484,98101.
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21.VonEuler,U.S.Liljestrand,G.(1946)."Observationsonthepulmonaryarterial
bloodpressureinthecat".ActaPhysiologicaScandinavica.12:301320.
22.Tortora,GerardJ.Anagnostakos,NicholasP.(1987).Principlesofanatomyand
physiology(Fifthed.).NewYork:Harper&Row,Publishers.pp.444445.
ISBN0063507293.
23.FisherJW,KouryS,DuceyT,MendelS(1996)."Erythropoietinproductionby
interstitialcellsofhypoxicmonkeykidneys".BritishJournalofHaematology.95
(1):2732.doi:10.1046/j.13652141.1996.d011864.x.PMID8857934.

24.Michelle,Julia(March7,2011)."HowDoBabiesBreatheintheWomb?".
25.Sullivan,LCOrgeig,S(2001)."Dexamethasoneandepinephrinestimulate
surfactantsecretionintypeIIcellsofembryonicchickens".AmericanJournalof
Physiology.Regulatory,IntegrativeandComparativePhysiology.281(3):R770
7.PMID11506991.
26.PrematureBabies,LungDevelopment&RespiratoryDistressSyndrome(https://w
eb.archive.org/web/20070604020429/http://www.pregnancyfacts.com/articles/child
birth/prematurebabies.php).Pregnancyfacts.com.

Externallinks
Ahighschoolleveldescriptionoftherespiratorysystem(http://www.emc.maricopa.edu/faculty/farabee/BIOBK/Bio
BookRespsys.html#Diseases%20of%20the%20Respiratory%20Sys)
IntroductiontoRespiratorySystem(http://www.leeds.ac.uk/chb/lectures/anatomy7.html)
Scienceaid:RespiratorySystem(http://www.scienceaid.co.uk/biology/humans/lungs.html)Asimpleguideforhigh
schoolstudents
TheRespiratorySystem(http://www.bio.umass.edu/biology/bemis/FAOV4/Ch18.doc)Universitylevel(Microsoft
Worddocument)
Lecturesinrespiratoryphysiology(http://meded.ucsd.edu/ifp/jwest/resp_phys/index.html)bynotedrespiratory
physiologistJohnB.West(alsoatYouTube(https://www.youtube.com/playlist?
list=PLE69608EC343F5691&hl=en))

TheWikibookHuman
Physiologyhasapageon
thetopicof:The
respiratorysystem
TheWikibookAnatomy
andPhysiologyofAnimals
hasapageonthetopicof:
RespiratorySystem

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