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InterLnoawle Orx ygeLni miotfsA pplFer uit A thesipsre sneteidnp atrailf ullfmeint oft her equemiernts fotrh ed egreoef Docotro fP hilopshoy inP laSnctin ece at MassUenyi vseiyrt NewZ ealand ChirspthoeWril almiY erasley 1996 01 11t� HtIf � ad 'J:�:., 7iHt,1)� , &ti44aad rI� t4- iii Abstract Theo ptimautmm ohseeprf ro caor pw,it hrs epetcoot xy ge,ln eisj usabto vteh e lwoeorx yglemini( tOL L)a,tw himcahix mumb enietifsnr le taitoonf riuqtu iatlayr e acheiveadn d lwbo wehifcrhiu qtu aliicst oypm roismebdy f emrenttia.oI nnc otnrast top rveiowuoskr ,L OLisn t hsitsu wdeyr ees mtaiteodnt hbeas iosfs teya-dastte e itnrenaaltm ohsepr(eOLsL 'assw) elalse xterantamlo seps(h LeOrsL)as itth ei s inentralO 2p atrilpa reses urPa)c,lo seeq lutiirobi uwmit h ctyhsteoo tlh,ta (PC)2' meditasei mpotrapnth ysoigoilpcraolsc eessT.h es tutdeys wtehedt hLeOrL 'osf 'Cxo'Osra ngPei p'pa inn'dB raebuarplnpe'ws e raefe fctbeydt emrpaetuerlveea,t ed CO2a,n pdh ysgicioalalog e. Two toyfpLe OsL wse ried eindet:ti hfaen aoebricco mepnastion (pAoCiaPnnt)d thfere mneattoint hersho(lFdT )A.C wPa sd esibceridn t eromfsp o ltosft hietn ernal CO2 )v ersuitsnre nal anedx terna)lO 2a,n d int eromfsp o ltosf (J2)J () (JJ()2 botahm easeuo rft hrese piraqtuooertny(it R iQ)aa nedt nhoa(lE tOcHon)c erntation versPuOsza nPdC ') M2athetmiacsaollu tifooenrss t imAatCiaPnn gd basoendt he RQi(aFT QR)a,n ads ttaist'iobcoatls tprraopc'ee s duutibralfeo ers ttiimanalgl antdh iebri sa-ocrre9ct5e%cd o inefdnciene trvlasa,rd ee rsciebd. ofp osltmicactefrriiuotcfb otchut liavrtse enddt oi necarsei gshltly betewe0n° a n2d8 °aCn sdah rlpya t3 2°.eL OL'rsa ngbeedt we0e.n5 kPa and 2.2 kPpa() ;lev alufeoFsrTR Qa nFdT tEOtHne detdob eh gihetrhn af oArC PE'el.va ted l (0t o8k Paat 0 °a n2d0 °dCi)nd o sti gincniatfalfyf LeOcLts a t2 0C°,b ut PC�02 inecarsienFsT RQ anFdT EOtHo cucrrfeodfr ri uattO °CA. s maldl ecreianOs2 e upkteaa nRdQ iwilas m easufroefrdr i uitn2 t o8 Pc2oa t2 0°eN.oc ontseinst chnagesi nL OL'wse roebe srvfeoder ti hecrui lvtairnr le attiopo hny siolaoggei cal (rpelcmiatcecr,ci lmiacetriocrp, o slmtiactcefrriiuatct0 ° 2o0°rC ). Inc ontrtaoAs tC p,Al C Yi necarsmeadr ekdlwyih tt emper,ra setuulretfirnogm itdse peenncdoen b otshk ipne rmeaanncrdee i srptairoant( eb ootfwh h icchhna ge ' witht iemf riuatr iens tor)a.Cgo enesqulenyut,es LofO Lsr,ta hetrhn a LOries s reocmmeenddf oorpt imiastiomnsgpe hresb oftoshre alpeadck aagnecdson torllde amtospehs etrogre,a mtion iemr siikso ff remetnaotni. iv Ackonwledgmeents Myc heifs upieorsrvP,r eosfsNoireg Bla knsh abse neb otahni snipartiaonnd freinda,n Ida mi dnebtteohd i fmo cro mpleotfti hotinehs s iIst .hn ak mcyo spuerivsoPrrseo,sf sEorrr Hoelwe tfto r ahdivsia cneDd r S iavG anefsohpr a tient guidatnhcreo suttgsaiht ipcraollbe masn idtn ruocdimneg 'toboo tsptpri'an.Igt hank PreosfsDoornC lenldwa how ilglligyna vheit sie mt oh elwpih tm atehmtaicisasule s. Myc olalgeuaetHs o trResrhceh aave beeenn cuboroaitgnahgn pdat iednutnr gim y abesncIet .h ank DWra ktCihn(rsCio srnl Ueln vieiryts,N ewY orwkh)o e nocuarged met oe norlf otrh Pihs,D a nDdr N agiLna llufo ra lhleh atsa ugmheat b out hotricuel,tth uefriirre nidpas nhsdu ppoIar mti .ne dbtteoHd o rtResearch manamgeenDtr,Js a meMsc WhaJ,o hSnha wa nIdna F egrusfoontr h iehre lapn d fnianclia assstniaceT.h anaklsst ooD rR oger Harckoelbrlo arofanoto rint he 'BrabeumB rowniDnigosr derre's eatrhtcwa has c onudctdeudnr git hceo suero ft he PhD nbourtte protehdee rinIa. l so PtrheoasfnskAo rtrC ameraonnDd r R andy BeaudorfMy i chainS gattUen vieirytsf otrh esiugrg etsions. Generfoniuansci las pupotw rasr eecviefdr otmh Feo nudatoinf oSrci ence Resehaa rncTde hcnoglyoa,n fdr oEmN ZAN ewZ eaaln(dn Itetrinoanwahlo) awardmeetd h 1e9 93D onS inicRrle aseha rlFcelwoshiMpy.t hkansg ot o DJrosh n Feil-dDodgsGoins,eA lhalo brnM,rH amiTsohu gahn Sdh yanHao ggeotfEt N ZA NewZ eal(annItde troinnaaflo)tre h ihre lp. Id eeplayp eprcaitteh aes issantcaen fdr nidsehiopf PtlheaSn cite nDceeap rtnmte. staaftMf as seUyni evrtsyiM.y s niceet rhantkoPs r eosfsKoernM ilea nntdh oef fice satfCfo,l leenH anloPna,mH owleH,le rKaen nedanyLd o iMsa th.e Mrany colleaghuaveesa ssstiegdie,vn a divcaen edn curoagmeed, oanmbgo tahc aedmic stfaa fnfdle olwp osrtagdusa.Itw eisha cktnool wedtghehe e lopfD rB enD adz,i e SueN icholAsnonKnain, g s,l eDJyroa ntahnC utti,nJ gontahnaD ixoSnu,z ie NewmaWni,r iApm puml,gn eM ertPse,tJ eeref rf,yC oilnT odCdh,r Riawsl sionn, MiekC urriTeo,m R obteosrnC,r aMiogw atCtas,s aonA dmraranatneKd,a te Maguei.rI na ddittoit ohnoa sboev eS,o hiaClh ouyd,Dr elwyCno okJena,eY onug anAdl isdNaoibrwl ein lgllicyo nturtieabdsp anletilssI.h avel aernmtu cthh rough thwee elkyFr iuti SecnGcreuo pa,n Cdo nrtloeld TeermaptLuarGbe ro umpe etings antdh aanllkw hoc onbturtiteotd h eod siscussions. v I agmr autle Mftasos eUynv ieirtseyn geirnasen tdce hniiacnIsaP,na niteLre,o BouletrJ,o hAnl egrB,r ucCeo lsla innJda rrHeardrg ianw hoh elpede qwuiitphm ent. It hanks attfahfte Mtahses eUyn ievrysP iltaGnrto wUtnhi pta,ir ctulRaayr ly Johnsoann,Sd h naeM ax tofhF eur iCtr oUpsn iftoc ro oslta ogrfeca iileisPte.t er Cathcesir doef TrPiagcokinan ggge noeurssluyp pplliseatdfil icm D,a ivdF lemming ofC armOerclh rada lleodmw et oh arvehsitfsr i ufto trh eex eprimeanntSdust a ratn d LeselH iumhprtyou steh Ael egrnToruns Ptac khuose. Ir seersvepeci atlh antkoms y f mailwyh oh avmea dseca riicffeosmr e t hroughout. Ic anno etx rpesisnw ordtsha ntkoms yw iefK athnr,wy hosteoe lrasnputp ota rnd encoaugmreenhtas m adoeu srt aiynP amlersNtoornta hne ncrhiinge reixepnce. ThanTkism D,a ineli,s asnaAd n derwf oyro ur ahnetdlo tp h,ee xe tndefda imloyf freinds mwahdote h set aiynP amlersNtootrnhs oe jnyoable. vi Tabloef C onetnst Abstcrta. iii ........................................................................................................... Acknowlemdengtes. . ... . . . iv ................ ....................... ....................................... Talbeo fC onten.t..s.... .......................................................................v..i.... . ..... lisotfT ables . . . . . xvii ............................. ..... ......................................................... lsito fF igeusr . . .x xi ............................................................................................ lsito fS ymb oiasn dA bberviantsi o . . .xxx ............................................... .. Chpatre1 GenrealI nrtodcution 1.1B acrkogutnodp rjeoct. . 1 .................................................................................... 1.2G enezreamdlo ideforlg aesx cnhgaien a ppl.es . ....4 .. ......... ............................ . .. 13. Optimiunmt eartnmaols pahneldr oewsoe xrgy elni mits .. . 7 ............... ................ 14. Reseaorjbcceht iavnesdst u rctuorfte h teh esis .. .10 ............................................ 1. 5R eefrences . .. . . . . .I 3 ............................................................................................. Chpatre 2 LtieruarteR eviwe 21. Itnroduct.i.on. . . . 14 ..... .... .................................................................................... 2.2R esptiorrmaye tabo.l.ism . . . . . 15 .. ........................................................................ 2..21I nrtoudctni o . . . . 15 ....................... ................. ............................................ 2..22A eriocrb epsirati.on . 17 ....... .................................................................... 2.21. 2G.lyc.o.l.y..s...i.s. ........ . . 17 . ..... ... . ... .. .. . ......... .. ......................... 2.2.2T.r2i caarcbiodxa yncltdyihe ccell eect trraonsnsy psotretm . 20 2.2.2C.y3a nider-er-sepS.i.irsa.tt.a.i.no.t.n. ... 23 .. ...... ...... ... ........ ....... 2.2.2R.e4s piqruaott(oiRryeaQ n)ntt dhe eff eocrfte sd Ou2c ed anedl evCOa2. t.e..d.. ....... ..............2.3 .. . .. ... . ............ . .... ...... . . . 22..2.R5e spirraaatntetdi he oeefn cf otrfse dOu2ca endd elevCOa2o tnae edr roeb"ipc.i .r..a..t..i.o.n .2 5 . ... . .. .. .. .... ......... vii Modeo fa ctioofrn e duceadn de levateodn 2.2.2.6 O2 CO2 aerobriecs piration .. 26 .............................................................. Effecotfr educeadn de levateodn g lycolaynsdi s 2.2.2.7 O2 CO2 TCAc ycle . 28 ............................................................................. 2.32 .Anaerorbeiscp iration . . . 31 .......................................................................... Functoifao nna erobic raensdbp iiorcahteimoinc al 2.2.3.1 pathway.s . 32 ...................................................................... ........ Regulaotfai noane robic r.espiration 34 2.2.3.2 .................................... Responosfce rso ptosm etaboolfia tnease robic 2.2.3.3 respira.t..i...on. ... .. .. 36 .. . ... .................. ...................................... . 2.2.R4e psiartnir oatoefcs or pasn dc hnagewsit hp hyisogliocaagle 38 ............. 23. Aerocb-ianraoebriecsr paitotrrnyasi tni o . . .. ..14 ..................................................... 2.3.I1n tdruociton. .... 41 ....................................................................................... 2.3.E2x itnicotpno ionfat n aerroebsipc iratio.n.. . .41 ......................................... 23..3A naerocboimecpn satpiooinn t .42 ............................................................. 2.43 .Effeocftt e mpaeteruo rnL OLs . .44 ............................................................ 2.3.E5f feocfet el vated onL OLs .. .45 CO2 ......................................................... 23..6E ffeocfpt h syiogliocaagloe nL OLs ... .64 ................................................. 24. Effeocfdt esc reasaendde e lvated one thlnyeeb isoynetshis O2 C02 anda ctino 47 .................................................................................................... ....... 2.5G ase xcnhgaeo fb uylp klnato rgans . .94 ................................................................ 2.15 .Intdruocnt io ... .. . ........ .. . .49 .......................................... .. . . . ........................... 2.5.L2a wosfg adsi ffusi..on .. . ... ... 50 ........................................ .......................... 2.35 .Thes kiansa b arrtioge ards i ffuasnidpo atnh,w afyosgr a s excahnge .. ... . ... 53 ....................................................................................... Thes trucotftu hrseek i.n .. 54 2.5.3.1 ...................................................... Permeaonftc hese k itnog ases 55 2.5.3.2 ............................................... Routfeosgr a se xchatnhgreo utghshek in 56 2.5.3.3 ............................. 2.45 .Gasd ifsfiuoinnf riuftl esh . . 58 .................................................................... 2.6R eferen.c.es. . .. .. .. . . 61 ......................... .................................................................... viii Chatper GeneraMla teirlasa ndM ethods 3 31. Inrtoudcotni 77 ...................................................................................................... 3.2F rusiutp palneydt hyplerertnaeet mse nt . . .. .7.7 ..................... .... ... .......................... 3 Furistu pp.l.y . . . ... . . . 77 . ........................................... ................. .................... 3 Ethyelp eertneramtent. ... .. . ..... 78 ... .... ............................ ........ ... .................. 3.3C onotllretde mperfaurtisutotrar eg e ... .. ... 78 ............. ........ ........ ............................. 3..31T emperema otnutrionrgi. . .. ... .79 ................ ............................................. ... 3.3.R2ea ltvieh uimdyim tontiori.ng. .. ... . . . .80 .. ........ ... . .... .. ........ .... ..................... 3.4G asm ixgi,cn otnorldla etmoesrpsehy staenmtd r meeantts .. . 82 ........................... 3 Gasn ilixn.g. ...... .. ......... . . ... 82 .. ..... ........... .. ............ ... ...... ................... .. ....... 3 Conotlrdla etmoeserps hsytem .. ...... . . . 84 ......... . ............ .... .......................... 3..43C onotlerlda tmosept ehraetrnmtes. ... ... . . 84 .... . . ........ ..... ............................. 35. Measruemeonfit ne trnaatlm osepsh. .e.r.. . .. ....... . 85 .... .. . ......... ..... ........ .. ................ 3.15 .Surfacchema bre.s .. .. ..... . .85 .......... ..................... ... ........ . ............................ Gast ightonfse usrsf ace ..................................8... 6.. 3.5.1.1 Equiliborfsa utrifocanhc aem bertshi etnt eorn al 3.5.1.2 atmpohseorefa plpe.s . . . .. 87 . ............................... ........ .............. Result.s. . ......... . . 88 35..1.3 ........ . ................. ...... . ........ ............. ....... ........ 3 Dircetsm apilnfgor mt ehc orcea vi.t..y.. .. .... . 99 ... . . . ........... .. ........... .......... 37. Mesauredv raibalse .. ....... . .. . . ....99 ..... .......... . .... . . . ................................. ...... ..... 37..1 Gasa nalysis. . .. .. ..... . . ... 100 .............. ............ ... ..... ................. ...... .... ........... . 3..72C ailrbaitosntn adrads ... . . . . . . 101 ............ . ................... .................................. CO2a nd standa.rds. .. . . . 101 37.2..1 . .............. ... ............................... Ethvl(eC2nH4e)a n de tha(nCe2() H standa.r.d..s ..120 3.7.2.2 .. ......... . . Ethanaocle,t aldcethhvavdclee ,t aantdee t hylene 37..2.3 analv.s..is.. ........ . . ...120 . ....... . ................ ............................. ........ 37..3C onevrsoifou nn iftoasrm uontosfg aisn s amlpes . 150 ............................ Calculoafct ioonnc entroa/t,idoinsss oiinv ed 37..3.2 02 equiliwbirtiphua mr tpiraels soufrO e2s 160 ........................... Convermtoilfnerg a ocnttsio c ocnenatitrons . 10 7 37..2.3 ............ ......... Ratoefst ransoffpe hrv sig%icgaals .es. . . 107 3.7.4 ....... .................. Ix Raotrfee spiration . 3.2 .4 .1 ........ ................................................... 107 Ethyprloednuerc attieo .n . 3.7 4.. 2 ................................................1 10 3..75E stimatfuirinstgk ipnem reantcoge a ses ..........................................l ll Permeteaotn hcae ne .. . 1 1 1 3.7.5.1 .. . .................................................. PermetCaoOn 2ac neOd2 . . 121 37.5 .. 2 ..... ..... .................................... 3.8M easuorfoe tsh feurria tt tribu.tes 131 ........ ........................................................ 3..8I Frutiits fsiunreem ss 131 ......................................................................... 3..82 Frutiotts aoll usbloliecd osn t.e nt . 141 ..................................................... 3.8.S3t aricnhd. e.x. ... . . . . .. .. . 141 .... ..... ... .. ................. .... ........... . ....................... 38..4F rubiatcg krnodus kicno lro u . .141 ........................................................ .. 3.8.F5r usiutar cfaer e.a. ..... .. . .... 151 ... .. .... .. .. ..... .. ....... ....... .............................. 38..6C ortitciaslps oureo asniddte yn si.ty. . 151 ..... .......................................... 3.9D isciuosns . 161 ........... ....................................................................................... 31.0R eferceens. ..... .. .... . . 122 .... .... .. .......... ...... .. ............ ............................................. Chpatre 4 DeteirmntaioonfL oweOrx ygeLni imtfso r ApplFeru it 4.1 Abtsract 152 ...................................................................................................... 4.2I tnrotdiuocn . . .. 162 .................... ......... ................................................................ 4.3T hoeretbiackcaglrn od.u.. ..................... ....................................................... 128 4.4M ateriamles tahnod.d s.. ..... .. ... . . 123 .... ... . . ...... .... ....... .......................... .............. 4.14 .Frusiutp p.l.y. . ..... .. .. .. .123 . ........ ........... . ................ ........ ................ .. ....... 4.4.C2A t reatm.e.nt.s .. . 123 ... ................ . ........................................................ 4.4.G3a mse asuremenatnla yasnid.s . ...... 133 ........ .... ... ..... ............................. 4.4.R4e spirateitohny lpaernnodced t uiroant. e..s . 143 ... ................................. 4.45. Skinp ermnecae . .. . 143 ............................................................................ 4.4.T6i sspuoeri otasyn cdo rttiicsadsleu inets .y. . . . 153 . .. ......................... ....... 4.4.E7x piemreanldt esiagnnad n asliys .. 163 ................. .................................. 4.5R esul.ts. . .. ... .. ... 173 . ....... ......... ..... . ........ . ......................................................... 4..51P o�a saf nuctioofp�n a npd� .. ...................................................... 173 2 2 2
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