organic acid metabolism.pdf
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ORGANIC AC METABOLIS
By KENNETH V HMANN AD WALTER D BONNER, JRH Bo brors bre 38 sshse
IROUCTO
One o he principal dicuies in reviewig he meabolism o organicacids in plas is o deermine he boundaries o he subjec. he orgaicacids ie a a crossroads in plan meaboism. As Chibna 1) pus i, "i ishe group of subsances we reer o as organic acids ha occupy hecenra and thereore he key posiion in he carbohydrae proein and a
meabolism o plan cels addiio o uncionig in caaboic changeshe orgaic acids produced by he dar xaion of carbon dioxid apparenyac as imporan iermediaes in phoosynhesis Organic acids may be ac-cumulaed i quaniy as ed producs o meaboism; urhermore, he oxdaio of organic acids as a cose coneo wh wh Fo h eenreview aenio has bee conceraed o he organic acids hemselves andheir occurrece and r6le in caaboism and growh Ierreaionships wihhe meabism o oher compouds have bee principaly sudied ihmicroorganisms and aimal issues, hle he uncio orgaic acids in
photosyhesis is discussed i anohe review
The lieraure o 190 has been covered by o previous reviews 3)and here is aso a recen brie discussio by McKee ) The work ih oherha pan ssues has been revieed receny ad horoughy (5
. ETERMNATON AD SEPARATON
he mehods empoye or he exracion a eermiaiono various organic acids are esseiay hose discussed in previous reviews, however some vauable modicaions have bee iroduced Pucher
7 have revised and summarzed heir mehods icudng a oprocedure or he exracion ad deemaion o oal acidiy, oxalic malicand ciric acids Broyer (8 hae discussed he eecs o prereamenon he oal acidiy o barley roos and recommend he ue of reshy expressed sap or such deerminaion. hey hae used a iquid exraciomehod or the preparatio o he oranic acid racion rom such expressedsaps 9 his mehod is aso appicabe o pa isues in geneal, alhoughi suers rom a very log exracion period hr In he auhors' laboraory i has been oud ha drying causes a considerabe increase in he
keo acid racion o seedig maeri. n heir experiencehe mos reliable
mehods o preparing pla issues or anaysis are eiher: by grinding heresh, horoghly chied issue, incorporaing his maerial wih asbesos[as as proposed by Pcher or drd isue] or by lyophiiinghe issue and hen grinding i Exracion is wih a Soxhe exracor ineiher ase, and need be run only oerigh. he AOA Associaion oOcial Agricuural Chemiss mehods or he deerminao o oa
7
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76 THIMAN AND BONER
ty fut fut put v b vtt vw byB 0 )
w 2) v tt pu b up tmp w f t u tt um t mt b u uuy by umb f wk
Indvidual .Vu m mt f t tmt fm v b pp vw 7, 3 7) um &Am 8) v w tt t 24tpy y btfm t t m t t mt Pu at ), t 24py y y tt t fm t t m pt ty T mt, tut p pv t b t mt tv t m
tmt A mmt mt ut wt t tmt f um b pp b b al. 9) At mmt mt b t bt t m t t b b pp by t 20) pp byGt 2) T vpmt f t mt wu pp t b mtw
T mtmt t b by u( to and is best descrbed by Umbei ) utuy f t mt ttmt mt Pu w m b
Bu & Tu 27), w u tt t m mt w t btt w t m p f t .w mt f t tmt f u v pp 3,
4 7 28) but t m tt t u mt b byUmbt al 2) t bt r t Smy, mt t -tmt t v pp 5 29, 30)Pbby t mpt t mt p t v f t t t t wt t t ptv t t b ut m t ubum tt vv
T vu t tt, wv v ubt t mu 29 32, 33 34) A t J 33) t ubum mtu 80 p t t , but b Et 89 pt Vky bm 34) ty fu 89 p t
Tw mt f t tt tt v pp 34) Kb al 9) v pp mmt mt f fum T tmt f tt b vw by Vt a 35,3) w w tt f k t t v bf mt, mpt pvt T mt p byt
wt b 37) w tt t byt ptuy wt ptmum t but p 4 ty t by umum, pp, f, fu V & 38) v b mt f kt ut T mmt u 39) f pyuv m 40, 4) w pp t b vy b Nwmt mt pt f 7 3)t 42, 43) v 4, , 42) y t bv m
Annu.Re
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ORGANI AID ETAOLIM
tod tods a dsbd by Umbt e al 2 A d sms oav b somwa ltd s t publsd mtods a usuy usul fo ps of o vol ds ulss t uts avlabl su fo Dulu dsllo. owv mod ofsl & Caa sms poms ad soud b sp tout ompl.
s as b potd ou by may auos atual poo of ou of sp d pl ssus s of o v by ts dmtobu oly by s atual solo d dtto. ods fo qultv oto of vaous o ads a v by Ds 5. spto of vaous ads by sal ppato ad by t lassal stdsllo mod of sl wdy d sussfuy usd. A
modto as b mad lt mtod upubsd w ovs stao w doma o dao. y ts poduo obas t mtyl o tyl sts a oom tmpau us omh s emen w ydclorc cd a vd mptus
A mod fo spaao of o ds by pto omtopy as b poposd by swood 2 ad as b sussfuly usd modd fom by us & u pva omuato so e l6 v spd v boy ds by wodmso pp omopy mod s puy pplb o dov
wo. A mo l omatoap mtod w oto oft ads s suppssd y ss of fom d s b dsbd byu & Ovll T RF vlus fo o ds a v d pmt w ot s sows t py of mod. t ommuato dal w t spto d do ofo pospt sts y pap stp omtoapy wood mo lso play pms ty v bsussful sp vous o ads by ssl sm tus. dvopd s mods sould povd a vy valub toolfo mospato ad dao of oa ads.
TE ORGNIC AIDS OF LS
Succulens-T l dul ad ot of vous ssua plts pomo ow s assul mbosm d ssotd w t suult b s vd osdbl o du ls fw ys bs obsvao w s b ptdly omds a ad ot of s lvs ass du t d dss du t day Fo ltau po o 0 ee (3)] O of tppal ads of vaous suul pls w t ps s b ow
s ssul mal ad ws dpdt dtd s so dby odl ad by Pu ad P & Vy 50 5 d ts wasomd by bs & lso 2
A ss of suds o mbos of sd Bryophyllum lnulvs s md by Pu e al 52 o 55. Ts vy oou wosus fom fat tat ul las oup of pms oy lmd
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7 TIMNN AND ONNER
attto wa dotd to tmpratr otrol, ad mh of th wor waarrd ot at a tmpratr hghr tha that optmal for ad formato Byphyllum p. otal ora adty ad mal ad wr fod toamlat larly th la ad tm of B alynum drg a growthprod of 5 day; amlato of tr ad otr ad lowd o owardth d of th rod. Appartl otr ad whh prd abot 8pr t of th total orga old, amlatd a rapdly Byphylluma dd mal ad tobao ( th athor' pro td), whl trad amlatd a appromaly th am at both plat. Oalad whh a maor ad ompot of tobao, tally abt Byphyllum Th appar to b th a ral othr lt p(30)
I a tdy o th rlato bw arh ad oga ad Byhylum la, Phr al (53, 5 arrd ot prmt of two kd, owth la hartd from plat whh had b grow drg th wtr
a grho, ad th othr wth la ltrd watr th dak olght or altrat lght ad da rod h formr la wr throw at a mor or l hgh ad otat gh ad day tmpa, whh lattr wh ltrd th lght wr at 20 o 2C. ad wh thk hee erre e ve hh w v
for ha b how by Wolf (5) at adar d p Byhyllum
la kpt th dark for log prod at 20C or hghr Howr, t r hat Thoda & Rhard (5 fod hat wh Klna wa kpt thark a 25, h mal ad rmad at a hgh lvl th wor ofhr al (5) th mal ad tr ad howd th laal dral arato wth la ltrd altrat lght ad dar, bt otr adotttd a pto, t drad at ght ad rad drg thday O th othr had, Krb ggto 29) ad Bor Bor (30)fod mall ra otr ad th dar Byhyump
Phr al (53, 5 ormd ad tdd th dg that tarh
ard a mar or to tha o h orga ad, ., drg tprod o a ormato th dar tah wa d p, ad drg thprodof ad dappara h lg t, tarh rappard hthr tarh
drtly formd from orga ad (rathr tha by photoyth ,howr, far from rta Th oly dato that th mght or throbrato that la olltd th morg, . at thr pak orgaad ott ad ltrd omplt dark, howd a lo of abot ohalfo thr otal adty 8 hr, drg whh tm thr wa a mal btprood formato of tarh (6 gm pr kg frh wt) t to b -
ptd, of or, that ad wold b d p th dark at 20 to 25C, bf thr oro to tarh ral th wold b powrfl d that thrato rlatg arbohydrat o orga ad ar dd flly rrbldr atral odto f th potlatd oro of mal ad toarbohydrat th bt (3. Howr, th workr dd ot obr aormato of tarh h dark qattatly omparabl wth that obrd h lght hy tat that owg o th gat ra
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ORGANI AID METAOIM
tot onic oi un te y, potoynte cn py ony en te foton of tc Howee it ee tt te incee in wi ueto potoyntetic fotion of cboyte y be ey oet by te
o of wet to be cbe to onc ci ippenceA ee of expeient n wic teete w contoe un tek cutue of exce yyum ee peente in te n ppe oft eie 55) Hee tey cone peiou obeton [30) n tetue cte teen tt te opti tepetue fo c foton fbeow 20C. n coe to 10. Decen te cutue tepetue fo0C. to 9C eute n et ncee of c ynte n t te ete n incee in te quntty of tc conue At 20 C. , uin 2ou peio tc ece 8 we 1 . of onc c wee
foe At 9 tee ue wee tc ecee to 2 onic ci foe t 1. , 6 . tc ecee n 16 . onicci foe 66 .) Hence, tee i o ipe etion between tc nonc c ince t te ow tepetue tee iuciet tc coue to ccount fo te c foe u te uto tte "c ybe te ouce of uc of te ci in cetin ce of but it i not te onypobe ouce. Tee e ny oe inteetn fetue of t ppe butpce oe not peit tei icuion.
Tt te onc ci of iou ccuent pnt, t te opt o
tepetue fo ci fotion n n te k o not e in te info te oxiton of cboyte but by te ctu xtion of cboioxe, w eontte by Bonne & Bonne [30) . lo 58 n teiexpeent tee w own to e n ot ine etonp wt owcbon ioxie concentton fo 0 to 0.1 pe cent in i between c fo-ton n te cbon ioxie pt peue At ie pt peue upo 10 e ent onoe n tee wee fute ncee in o-nc c content Te jo p of te c ncee ue to cbon oxiexton w foun in c . Tey wee unbe to boi ci pouc-
tion n cbon oxiefee n t y we be tt t te tepetue ue3Q) t eiu ci fotion eeent tt foe ecty fo teoxition of cboyte o poycie. t ee inicte tt tow tepetue ci e foe piiy by cbon ioxie xtionwee t te ie tepetue te foton y be peonntyoxitie Howee, of 56) own tt wen ee e pce t tepetue in te it in concentton of cbon ioxie 50pe cent CO in te expecte ppence of onc cii ety e-ye ince te y eon in wc uccuent coony oui ten toe e f in nt tepetue it i ikey t in ntue eionic c e foe peoinny by cbon ioxe xtion
Te xtion of cbon ioxie wit te fotion of onic ci byee of ypyllum in te k w one by Tuow & Bonne 59)in cbon ioxie continin C14 ee uto owe tt of te C1402oiiny ien excie ee of aum 50 pe cent of te ctiity wecoee in te onic ci ction wic i i ce i copoe o
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0 THNN N NNE
whllf mlic ciric nd isciric cids ref. (30] rris & Sz (60 hveied similr resls wih B aum fr which he hve deiledlses. I e epermen he d h fer 1 mi. dr in
perid 0 per cen he rdicive r ws i he mlic cid frciIncidenll i is f iees h illmii did hge he i disrii grel r fer 1 min phperid he mlic sill cnsied 8per ce f he rdicive frci
Sideris a 61) ve shw dirl chnge f rgnic cid cnen inpinepple eaves s in he sef BophUm e mr pr f he chngews in mle (7 per ce) ccmpied similr smller chnge incire (37 per ce)
Hrders servins 62) n he mrphlg f BophUumin dieren
phperids re f ieres here ri he d h pls kep in lngds lse heir scclen hi he leves ecmig hin d lrger i reThe scclece cld n e resred high slii i he sil. This seemsver sggesive in ligh f he experimens descied ve:i m e hlgd cdiis cse he dispprce f rgnic cids nd h helss f scclece is resl f his chnge
cher a (2 3 63 in heir sdies f leves hve ls fllwed heehvir f rgic cids in he se isse I is remarkable that in hisisse here ws i rs he leves n mrked dirl vrii
cid ce here ws pH chge similr h i he leves. Thhere re sme scclens sch s Msmahmum h d n ehvein he sl mnner hs lred en served (30 j he srpising fch he rgic cids f he sem d fllw he sl per indicesh he lk o e organ cid melism kes plce whll wihin helef nd h here is ver lile rspr side he lef. In his cnneci
i is ineresig h in he experimens wih vris mns f nire ndmmnim nitroen (see elw) he sem isse respded el s didhe lef isse
osuus-There hve ee everl ivesigis delig wih heeect of nutrton on the oran cds o eaves. Very a (65) andcher a (63 6 hve sdied he eec n he cmpsii f ccassus pous d B aum f he frm in wic nirge is spplied Grwing whle plns r he leves nd sems n clre slinswhich ontaned the sae aont o ntroen (rngig frm 100 per cennire irge 100 per cen mmni nirge wih severl inermediesges) he shwed h n hese pls l rgnic cids decresed wihincresig mmim irgen. The drp in sme idividl cids ws mchgreer in Bophum h in cc. The sgges h he lwes mle
level fnd i he 100 per e mim nirge plns epresens hemn reqired fr espirin. The d n assus re priclrl in-eresing in h his is he rs recded invesigin f he rgnic cids frpls i his nicl grp. h he ls d leves f develped plswere lw in rgnic cidi he highes vles fnd eing respeivel per ce nd 04 per e f he l rgic slids I crs cc
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R MM 8
Bll n , w om o l of t l tmn coxlc, ctc, octc, mlc, n ccnc) t mjo c, as n v contn olc mc ctc n ccc c n n
mont tt ccont fo onl lf of t totl ognc ct. T to not tt o t pnc of octc c n t t n t poltt , n Bryophyllu, t compon m t lk componntt asus t w mk ncmnt of t t mc cfollowng clt on ntt ntrogn. ml xpmnt to to t mnton v n c ot lmov 66) n vn & k 6)on tocco; ntll t m lt w otn, nml tt lntgown on mmonm ntogn contn mc l ognc ct tn to gown on ntt ntogn. Blcmn Tmpmn 68) lo fon
tt clcm ntt nc t totl ognc c n ot g ncov. pkowt, Glt v 69) fon tt wt ot gown wtn wtot ntt ntogn totl ognc c nc n pnc ofntt t nc w ptl mlt ptl nknown c t lttng t gt fcton of t otl ognc c c om lo pnnt on t pnc of ntt, t t conl c owg v n plnt gown n t nc of ntt T wok lonvtgt t ct of oxgn tnon n fon tt t l of totlognc c w g t low tn t g oxgn lvl. T w no
ct on mlt fomton owv n t fomton of oxlc c wfon to v nvl wt t oxgn tnon T fct tt mt fomton w not pnnt on oxgn tnon wol m to n ft -tntton.
T nnc of cton pp omwt compl lmo ) ot tt n tocco n kokg t ct of potm on on tmtolm of ognc c pn n wt t plnt w gown onmmonm togn o on ntt ntogn T pnc of potm ct ognc c contnt o plnt grown on ntrt ntogn to ll wr
n to gown on mmonm ntogn, potm c n nc Tnvtgto lo fon tt t pnc of lt low t mont ofognc c wl clo t concl tt mmonm ltpl potm n mgnm lft ol nc t contnt of c tnc wtn pnt wl tt pl potm o oumclo ol nc t ognc c contnt.
ctov ) pot tt t ctt contnt of vo pntw nc potm o popt, w W t ) fon popt to p t mont of totl ognc c n tocco
Iljn 3) t t lton twn ognc c n clcm cloon lv of ppl n c. Iljn 4) n Cool ) v nvtgt tlton twn nognc lt n orgnc c n gowng n mtlv Totl ognc c n ctc c w colt wt t montof pot n xpnng lv t mlc c w mo clo lt toclcm j fo xmpl n xpnng n t lv of gl ), tgt vl fo mlt w fon wn t mxmm mont of clcm
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82 HA AD BO
ws fon ref e osevon of cmn n empemn 6 oveIjn's exensve nses on speces show e o cs, mc ncc, ncese o w ge n w e vncng seson In gene e
cs e coee w e cm conen, espec fo pns gowngon ck so mn eowng cses no ppece econ ece& ppemn 6) ve me sm s on common gcspeces, fon e coeon eween vos ons n e ogncc conen e mos neesng fee of e wo, wc ncenses fo o ognc cs, oc, mc, cc, n nknown cs, wse g mon of nknown ognc cs n ese pn specesn moscses monng o pe cen o e o, o moe
n eeng expemens w occo (N. tabaeum, N ustiea, n N
glauca) Kosov & Nkoov fon epo oms ene o vemoe cc c n e po Snce gowng conons e no men-one s of nees o now ow mc ese eences wee e o vons n g, empee, n fees
ce & cke 8 se e cnges n ognc c composonn excse occo eves ce n we n n nognc n ognc ssoons Ce n we n n nognc ss e o e fomo ofa w a ospodg dcas mala, w oa ad -
mne essen consn. Ce n me ge sme e fom-
on of ce, we ce on ce, sccne o fme ogo nceses n o ce n me. Ce on e cse nncese n e nknown c fcon n ncon of e ccmon os c eves ce on mone sowe n ncese n o cswc n some cses mone o moe n co e ccone fo epke of mone one n ncese n cc c n s gee e-cese n mc c mo mes of fme n mone cse ge ncese n o cs, moe n n n of he oe ess, n ccncese we mc ecese ese expemens nce mone
s wo e epece) ocs e fomon of me en s ppenson ce wo ppe o e fome fom e sccne wc"c p Une nom conons ce s ppen fome fomme ce & ce conce om s wo ose cs wce p of e coxc c cce ene e no e meosm ofe occo ef n ence enme ssems cpe of meongese cs e pesen
Some of e ove nngs wee conme n e sme oo wnoe ve of occo 34 n, n on, eves wee ce on
soce n on cee soce og o n ncese ceconen n e sme me sm ecese n me s ssme hsoce s convee e conse of e ef o ce, og f swee nee e cse, ess sm o ose one fom ce once so ve een one, .e o ce n me so vencese Te peence conse n occo eves s n een emon
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R MM 83
trated The eave ultured aetate beae aid by the ed f theuture erid i trat t al ther rgai aid teted but all theaetae take u fo the ulture uti had diaeared fr the leaftiue There were arked hage i ttal aid ad it i reued thataetate i etey xidied a lui a idiated by Krtkv &arker exeriet (9 ith radiative aetate ee Mabolsm Theereult led Vikery Abrah (34) t lude that ulture rgai aidbrig abut a greatly ireaed reirati uh a deduti i i agreet with diret eaureet whih are diued belw I regard t theiterrelatihi betwee ariu rgai aid the exeriet hw thatdurig uture i water abt tw e alate are verted t e lef itrate whih reatihi al hld true durig uture aate urig uture itrate e t tw e f thi aid r e e f alateThu whe the yte tai re ta the uual aut itrate theral ure f reati hih vert alate t itrate are idered tbe revesed. Pyatnitskii in expeme were tba leave wereultured i lti rai aid red ad te teted r rgaiaid al luded that itrate i fred at the exee alate but thatthe reee f ageiu i i reuired fr thi t take ae Tee bervati ae urted b iilar exeriet rebikii i thatduig uig the ireaed irate tet wa rrtial t the falate additi the frati f itrate wa further tiuated by theeee f a hhate buer 8) Thi authr a hwed that girdlig the te aued a iderable ereae i thee tw aid eeiallyalate
Wd al (83, 8 85) ad Cruikhak & Wd 86) have wed thergai ai tet uda gra ikuyu gra ad at thrugh a erie ivetigat erig the etabli hee leave durig tarvatiThe behavir f the variu etabite wa eetialy the ae i theleave f all hee i air Hwever the ehavir f ali ad itri aidiered arkedy fr that deribed by Vikery al 87 88 i earlirivetigati with rhubarb ad tba the grae tarvati i itrge rdued little r ireae i the alate ad itrate tet airhwever alate auulated u t a late tage f tarvati ad the dereaed whie itrate a ireaed t a axiu whih ae ealer thathat f alat ad the dereaed Thi wa aaied by a veral nreae i the ttal aid I tba the ther had durig tarvati iair the ttal id reaied eetially the ae while aate dereaedideaby ad a rredig aut itrate wa red 8)
rhubarb there wa a dereae i ttal aidity a well a i alate ad itrateWd al feel that their exeriet hw that i grae rgai aidad artiularly alate are t derived i areiable aut r itrg-eu ur uh a aarti aid ie therwie the rteli wihaaie tarvat hud have rdued alate eve i ioge Theyre hwver that the iterveri rgai aid i thee at
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M R
tak pa va a tay ad y whh y td t tg ta va ad ad a ad, a ha ppd y vauwk th pat ad dud ath t Thy a uggt
that th tg data tw ga, ta, ad hua ay d y aug that a tay ad y pat aa ad that th d ut f vaat dvdua zyatvt
It w ad that Waug, a fa ak a 886 (8 fud a duavaat ad tt f vau uut av Th vatha d ad tdd t a vaty f av (0, , ut adf t t th ha tt a vtgat f th phHwv, th ag vd that a dd at, whhay ay t vdd y dua vaat ga ad tt,u wdy uut av ty, u & tutz (60 havaayzd ay ta, ad tat av whh hav tatd t a5 dak at pd th p f C fwg a pd fpuat Thy fud f ta, that wth ay 43 p t ad47 p t f th wa th u ad a ad fat ptvyI ta 88 p t ad tat 85 p t f th C wa fud tha ad fa u ad ha t gay udd afaay, ut a tv uvy (64) hw that t wdy dtud pat tu Hwv, a a t tut tha p t f thdy wght Dtat ad dug th dvpt f th tapat datd that v thugh th ad pt y a autt ay gadd a f th t atv tat Th wud agwth th av dg ad a wth th f Cav & ( Ff dak at
Vd ad h aat (2 t 7 hav ad a vaua tut t u kwdg a dd at th tud
aya hgh pat pay t thy hv tad adtudd a aat aya, wh atvt a aatd wth aa dhydga atv wth TP th yt appa t atayz twat dpdty th f whh hav hw t vTh at a: a th v ayat f pyuvat wth thfat f aatat, th w kw d ad ka at,ad ( th v dut f aatat t aat a at whhu a hydg dat Czy II Th zy yt appaty vy wdpad, f Vad (6 ha hw t t pt 3
dt p f hgh pat, t, tu, d, ad av Ctha& Vad (7 hav ad a zy ppaat f pay twhh fd tay ad th p f ktguta ad ada dd th at aga at a ay Ifzy II addd t h yt, th ta at pdut dudt t ad th data hw th vy wdpad dtut fdak a dd g ha hgh pat
Annu.Rev
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MM 8
FUIT
While there have bee ay vetigatio oered th the iolatioof orgai aid fro fruit (ee etio o OUENE ND IIBUTN),
detailedtudie o the bioheitry of orgai aid foratio ad etabo-li i fruit are fe Thi i regrettable i vie of the fat that ay fruitaaretly oe a very ative orgai aid etaboli I additio theaout of orgai aid reet are frequetly largeh hould ilifythe aalytial aet of uh tudie The tadard deteriatio ethod[revieed by Bolliger (10 11] are geerally adequate ithout iroodiatio for uh aterial
ihard 98 fro tudie of the buddig of variou varietie o dier-et roottok ha reorted that the roottok eert a deite iuee o
the itri aid otet of itru fruit here have bee fe tudie of the iuee of utritio o orgai aid otetlike thoe reorted above ithtobao A ireaed itri aid otet i graefruit due to ageiu orotaiu fertilizatio ha bee oberved 99). ilair aey 00have foloed the orgai aid otet of develoig Valeia orage or aeightoth eriod Ther reult idiate tat the foratio of orgaiaid i oleted early i the develoet of thi fruit ad that there ilittle hage durig aturatio I the graefruit the ai hage i dereae i free tri aid ad ireae i itrate The tudie of ilair ad
oorker o other itru fruit (1707 1775 are uarized brieyi the Tale uder CITIC ACID Peyaud (101 0 ha ade a o-iderabe tdy of orgai aid relatiohi durig rieig of grae ofhih the aor aid are tartari ali ad itri fortuately there ilittle to idiate the re of tartari aid hih he foud to rie to a ratherhgh erao a he grae re I t aear that ali aid dereaemaredly durig rieig eeially if the teerature i high (0ay (10 foud the ae thi true to a leer ett for tartari aidPeyaud' ork ha bee diued by aue (04 ho diagree th oe
of Peyaud oluio a to the origi of the three aor aid The latterauthor oider that durig the eig eriod tartari ad ali aidarie fro lati aid hile itri aid arie fro aeti aid I vie of therarity of lati aid lat thi hyothei, ad erha the vie of boththee autho ill require further eerimetal veritio
I ale averier & aqui (105 foud that ali aid hih i0 to 5 er et of the total aidfall durig rieig to about a third ofthe value i the early ue fruit The itri ad uii aid ad 0eq reetvely er liter of uie hoed o dereae at al durig th
hole eaohih ea of oure that the total aout ireaed i ro-ortio to the volume of tiue I earthe ame autor (10 foud urriigly large aout of itri aid I table variete it aear itraid aout for oly 5 er et of the total aid (the ret beig alibutoe varietie gro for reig have ore itri tha mali t oitedout that the juie of thee ould otai 8 g itr aid er literhile thela (o doubt iteded to over adulteratio allo oly 05 g
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HA AD BOE
GENERA BHEMTRY OF RAN N ANT
s as mod oduo s o mpossbl o osdoga as pa ssus ou ag o aou amos sp o spao ad mabolsm. Ma oks a ssd mpoa o oga ads og mabolsm 07) ad a bo ad ll s o s subj 4). Baus ) p
ss a dagam o aous mabol posss ssss l ooga ads o ol omao oso ad bado oamo ads bu also al poso spao dog aspoaob pospolaos ao ad lbao o abo dod adammoa ad sss o sp ogous ll sbsas. Hoaua obsaos o a su aosps g pa sss asa.
Transamination and amide fomation-A mpoa lk b o-ga ds ad og mabosm s o ous asamao bu d o su aos g plas as o b gal padds 40 ad s ompl Cdagolo & Caada 08) ouda as o gamaous plas m mo a a smla
as o gumous pas pomog asamaos b aspaa a pua aspaa a ogluaa a ala adoguaa Va & La ( a sga asamaosolg gluam ad aspaag Usg usd pa plas asamad pua o ala ps o glama o aspaa oud a amds subsud o la amo
ads al amou o asamao ad ou as gal dpssdMasums o amd og spl o dug pod o asam-ao dad a ps o amds asamao posoug ospodg dabol ads aua 0 a
so po sas a ma g plas ( maal usd s o mod a apab o ag o asamaos o apa bu o o
aoma amo ads u aos a sad o a a mamum 40o 0 m opmal odos bg 4C a pH . su boad & Bs ) o gluamaapaa asamao sods ao o b ad ou b a a o g plas la smoo u ad odla ssu s auos po a asamaoa p u o ssu dass asg ag a gog pa.lbaum & o 2) a ollod asamao a dlopgoa slgs. s obj gla amopoas a ss apdlom sod da oog gmao aua (13) as sgad
omao o amo ads ad amds Pism aivm ad as agadmosad los laosp b omao o s subsas ad oga ads s as spall mal ad O o ma poblms sudd b Wood a pms alad dsussd(8 84 85) as o a amds as d om poolss
a old poblm f. )] suls la s sag las a
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ORGAN AD AOI
toge tophee podued o d but vey ttle de d loto og d tee w lge e pge wh w pe-
eded by ie lte d itteMasm dvdua ad-Te ve bee oe vey teetgobevio o te bologl oxdto o oxl d T d logbee odeed bypodut o plt etbo w we oe oedw ot edly ued up ee the dg o ke a 4 tht teo Hum quum d te eve o evel hge plt ot oxlodeydogee ekble Te plt ilude oel hubb( plt kow to toe lge utte o oxl id love d pPued eye peptio wee peped d tuded Te eye o
te o de o tt ge pt t teotably d ttepetue d p eqeet but ll the pepto hve ekbletbilty towd het Te eye bited by odu tte 67 0 but ot by yde ulde o de A deet type o oxlteoxi bee eote b o iee wo owed ttupeo o oe bought bout the odto o oxl id to ydogepeoxde bo dode T eto bed b yie o by lo oxyge oweve th w o wy puied yte te iboby yde o poo tt th eye de o tt o e at to be hoped tht egg the ope o u wo wll ly te obue poio o oxli id pt tue
The utilito o ette by tobo leve bee tuded i pel e wt boxllbeled ete b otkov & ke 79) otte exeet poto o the beed bo ppeed wteoluble ovolte to the popete o whh ugget oe o tebyl d oge epeet te lbeled bo w obted epied bo dioxide Tee utho eel tt tei expeet dtete poibily o ette utlito ge plt but do ot pov t tobe ol etbolte Evdee o te ol utiltio o ette ioeoptle h bee obted the utho botoy d will be diued the eto o GOW .
Dydoxyle d whoe peee plt tue ote beedued h bee ow by u & Do to be vey pidly oxdied todketou d we iltted to leve o Tadaa 116 Tie ed out oly i wole le tue o vey ehly peped leete to w diydoxyle d bee dded t i to be oted vew o the ppet ee o the eto tht the bove wok teuto wee iolt eet the dded dhydoxylei d d ee
the dppee o th ubte whe ltted to leve igt b by eto l to tht debe by Oo 7 whh detoiid oeted to etogluti id by bo dioide xio Diy-doxylei d edly iee oyge uptke by owgll ue.
lyol id oed we leve e expoed to lght te bee obo doxde 8 Oxyge ppetly eey o t oto t
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88 HANN AN BONNE
sappas h ak wh a apy Epms o s spao ad h oao of choophy a scussd h sco
Du h pas yas a md amou of wok hasb c a ucov vc fo pacpao of a caboyc
ac cyc pa spao h s fuama sp hs cowas a by Jams a cowoks 119 10 who dmosa hapyvc acd aos fom hoss houh a phosphoyao cyc bascaysma o h acos of yas Ths was show bay oos ad avsad cf sap ach cas by us vaos amoaphhosufocacs, whch hav b show o b spcc hbos of caboyashy w ab o soa pu pyvc acd. Thouh suds o spaoad is ihibiio Bo Wdma 1 hav dcd h covsio ofhos o pyva spach avs sma sus b obad wh hAa coop 3 h psc of mac ad socc acd hyoass h spach af was mosa ; hs wo acs as w assuccc fmac ad pyuvc acds ca b usd as a spaio subsaby sav avs I ao od hbo of spao was ocom by addd pyuva aoa hbo was ovcom by h adoo succa umaa maa o soca A hs facs a ohvy soy ss papao o a caboyc ac cyc spao h cas of h coop 3 h sus w o as ca-cu. Vaous oac acs hac h oy upa comao ofh a of Commo Thma 4 b o hbo ofspao was oy paay ovco by aous oac acds ahouhh was a sh accumuao of succa u maoa hbo aa h s vdc fo a acd cyc bu s au mas o buca fuh B & Avy hav show h psc of svahydoass b o succc dhydoas hs ssu
A appoach of a shy so has b ma by as 67 I ba oos h o upak was h sam ucos o pyuva
spao was hbd by o a hs was v by pyuvaspao was aso hbd by ooaca ad u sab cooshs oo was vd by pyuva W maoa hbo h spaocou b sabshd by a vay of oac acs d sab coos succa accumuao boh bay oos a spach avs wasfou h sc of maoa, bu h amou of accumaio was dp o h ao of h pm a h cocaio o maoaWh cas m o shy casd amos of maoa succaaf hav s accumuad o som sappad s possb
ha maoa s mabo hs ssu. Th accumuao of succacoud b hac by pyuva, fmaa o boh u coos whchwoud ca h pacpao h oav fomaio o succiaSuch a dmosao of h oav fomao of succa avs aoos s a vy so am fo h vovm of a caboyc acdcyc hh pas.
achs has aso sudd h spao of bay oos coc
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TA AD BO
colc acd on the oxyen uptake of rund barley leaf centugates, whch snot en b other acds tred. At the same tme t was shown that chlorophll dsappeared at a rate whch was dependet o the cocetraton of
lcolc acd preset 57 per cent disappearance n 10M lcolc acd)Ths author dcates that n addto to the lcolc acd oxdaton sstemthere are three other sstems noe n the oxdton of ads n eavesUlrch 13, 10) has sown that as was expected n excsed barle rootsthe Q was less than one dur the formaton of oranc acds and reatertha oe dur ther dsappearance her formato ad sappearacewas reulated b the caton-anon balance of the nutent soluto e whecatos were absorbed n excess of aios oranc acds were ormed whleorac acds dsappeared when anons were absorbed n excess of caons;
hese expements produced no edence for the fomaton of orac acdsb oxidate deamato of amno acds It s worth notn that the formato of orac acds was reduced at 5 ad almost preented at 35 consder the close parallel to the smilar behaor of succulents t may besuested that carbo doxde xaton was pmar factor ad formaton(see secton on Scnts)
'
RWH
A numbe o experments ndcate a relatonshp betwee orac acd
metabolsm and rowth Whle ts exact nature as et far from clear such arelatoshp has bee demostrated at satsfactorl low phsolocal cocetratons of the acds and edently noles ther oxdate meabolsm
n a stud of the elonato o solated Aena coleoptle sectos presence of doleacetc acd ad ucrose Commoer & hman 12) foundrowth to be hbted b odoacetate Whe potassum malate succnatefumarate or pyruate was added as well the nhbton was completely peeted t was also obsered that the oranc acd salts mentoed aboedetel promoted rowth n presece of auxn and that prereatent wth
them allowed auxin to aus susqn an inas o oxiation atese oseratos hae bee a tmes cored and etended lbaum & Commoner 11) and Albaum & Echel 12) obtae smlar esults wth whole Aena seedlns to whch the solutos were appled a theroo we Boer (13) has recently cormed the promoton of rowth ofcoleoptle sectons by malate fumarate ad pyruate Curousl eouhmaloate wa foud to hae a slar eect (12 althouh hher cocetratons maloate hbts rowth wo phenomena besdes the eect ooxdato are to be dsushedthe preento or reersal o odoacetate
nhbto and the actual promoton of rowth n a more extese studThma & Boer made th dstcto clear ad showed thatmaloate does not crease the rowth of coleoptle sectos apprecablyaboe that o cotrols althouh t does preet the odoacetate nhbtohbto by arsete and phenlmecurc salt was not preeted b orac acds and t was tentatel deduced 15 that these nhbtors mayeac at the same locus as odoacetate but wth a reater at for theenzme hs locus s probabl a ulfhdl rou he eso o wheter
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ON AD MTABOM
th i the ae o a that a whih e ogai aid theeve at i aiteeg oe o te tdy
In additio to the aid etioed above, ioiti iti ad aeti weeao tdied 144 A thee ieaed he gowth b aot p to 100 peet The gowth pooto wa ot aed by ootig ga ptaeie it oed i the abee o oe O oideabe iteet i theeated dig o ege Avey that dehdogea o iti ioitioiti ai ad gtai aid a we a aae d aoite epeet i the o!optie 1, 134 athogh oe o thee how ativatioby axi piah eave ao otai oe o thee ezye 146 Thatoeoptie epiatio poeed by way the Keb ye h bee ade vepobabe by oe 3 who howed, ite aia that aoi aid ae
the aatio o ii aid (ee etio o Respato: Thi, ooe i ia aie evide o the tioig o ii dehydogeaewhh ege Avey 14 od ot detet by the Thbeg tehiqTht te aate ad iate ieae the epiatio o oeoptiewa oed eety by Ke & Ae 47
The atio o ogi aid i ae whe the eio ae t oode oeoptie whoe gowth i owig dow Ti vaiatio o eet withage a be oeated with hage i the ogai aid otet oe patia eitivity to iodoaette ieae kedy wit age o te
eoptie 14 The aate itate ad tota ogi aid deeae teadiyoe the ae age age 148 whih i itepeted a eaig that the ogi aid ae ata potetive aget agait iodoaetate t appea48 that the gowig te o Psm eedig have a otet o totaid, itate ad aate eve hghe th that o yog oeotie
Thee epeiet hve ao gve evidee o a e aetate ioeoptie gowth 14 peee o ooetate a pei ihbito aetate etboi the apid gowth whih o de oditio o goodaeatio od be eded to a ow vae oe to that od with etio
beged i the tet otio The ihbitio wa agey eved by aette ie pea te how a ii ihbitio ad oteiibitio eetthei gowth wod ao ee at eat py to ivove aetate A ayo oe o the above wok i give i 48
The gowth of othe te i ao tiated by ogani aid idebadt & ie 150 od that tie te o oe peie howedgowth pooto by ogai aid (odi at i peee o pe etoe owth o igod tie wa pooted by ii, ai aiad tatai, Pai daiy tie by ii ad owe tie by ii
ad goi aid Peiwike ad tobao howed o ihibitio oweve ie te aid wee a teted at oe athe high oetatio 0 peet o abot 003 to 004 M a oe yteti tdy eh ight eveothe eet The avoabe iee o ii ad the ihibito eet ooi aet, popio at ll xli" gtai, ad iti adwee geea oe o the aid od be btitted o oe a a abooe
it i o iteet that Pmnas to o beet o
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2 THMNN ND ONN
n me xc ctc n scc cs tn nm tssue 151 enm tssue n Q. f 04 wc Nes Het sce t gncc fmtn e te tum tssue gve 092 It s cnceve teeett te me pygng tum tssue cnsumes mte s fst s ts fme
e seengs sw eects sm t te ve tug te esuts epte e nt s cecut s t ven n ncue sme ppent cncts. t a a, wen gwt s fwe y t engtngvescensk Kgnvv (152 fun tt succnc c stmutegt t cncenttns fm 125 X 10 t 10 M Fumc n sptccs s stmute up t 5 X10 M ut t 10 M tey epesse tegwt. Oxc c ws nty n t 10 M stppe gt cm
peteyn te pevusy mentne stuy n ts Nnce 129 fun tt te
uptke f ntte y wet ts ws nte y 24D ut tt ts n-tn ws gey pevente vese y ctc e pesm tte expements n ceptes s vey suggestve tug f cuse ce engement n st uptke my e nepenent pcesses e gnc csmy eese ntns n gene tey nt necessy pmte gwt ts Ldd () u () tt umc c X s we s mnc c nte st sptn n eeng f et
ts e epenence f ts esnse n pH Luneg sces t teeectveness f ny te unsscte c ( sectn n Rpan) In metc stuy f ecse tmt ts Henesn Stue (130 swett te 4cn cs s we s pyuvte n ctte ncese e ygencnsumtn n pesence f sucse tug tey cu nt e use ssusttutes f sucse n gt
t Cattya seengs gn n Knusn's meum tne (154ecs tt tn f pyuvc mc c t 2 mg pe te =c 2 10 puce ge seengs wt ke geen c n nge ts
tn te cnts t s me cut t ntepet te esuts f Pntvc(155 wt Nusgy seengs ecuse f te pH cnges nve enptssum sts f vus gnc cs wee e t the me Knps sutn (pH 75 n wc te pnts ee gn n stee cutue te meecme kne mmnum sts n te sme cncenttn (2 1me te meum c us te nns wee tken up pefeenty t teK ut ess py tn te NH/ Succnte ws te ny nn wcpmte gwt t s ptsum n s n mmnum st Heet seems tt s t te t pevsy scusse epements t snge
cncenttn ptcuy en t s ve 10M, e pt t gve cmpexesuts sustnce wc s gy ctve 10 M my we e smewt g cncenttn s te evees ve fun n epements wtven n ntng eect t g cncenttn my nt e n cnctwt stmutng eect t cncenttn
n styng te gt f Gyue (atm agtatm) Bnne stn (156 nte tt pnts n te pepey f n epement pt
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OANC C MTABOLSM 3
rew eer an oe n e ener. Analy o an ulure owe ae roo eree a uane w ne e row o uayule ee
n. Sma nory ee wee en y waer nuon o e rooom w un an nnam a oul e olae Aon o nnam a o eeln n al aue ey make row non Aer4 ay n o oweer e non wa no loner poue nana e a wa eompoe y e mro oa (5 . o rom aounuayule plan no n uayule eeln eer, o a e ornaoeraon no eplane e non y nnam a neereleo ome nee eaue o e elaon eween ompoun an eomeral ome nnam a w a ay a an aun Aun n
enea nh oo eonaon 8] Onar or nnam awa repoe y HaaenSm & en (159 an oep mann en(60 o e nae a an aun .e n pomon row u no ow nn ee wa oun Blaoeenk olonoa (5 n- eu s nenraonaoe 5 X M ou elow leel ome row mulaon wa oee e olaon o nnam a om uayule ue ren anleae y aler (6 o nee n onneon w e aoe oeaon mo o e a wa n e orm o e parenyl (H eer u
ee wee nan a ome ou reeURREE D TRIBUT
ere ae o many mellaneou oeaon on e ourrene o oran a ha a een ou e o preen e aa n aular orm ou ope poe a ueul ne o e aee lerauehe mao o e reeene ae aonal o oe eal w n e eDaa or ue reer o r we unle oerwe ae
CTC ACPn ei e Reeee
Su e uie mall au pese
eses u spuiu peee le lippis 9 t pe e l 3
eeses wih imeOt bley whet a sees y w i (164)
ye cee t l ui
em tiZe ys sees pe e ltle nee i emitio
MGOASchizandra hnenss ui Mai a
RByphyu yi leves pe e he y w 3u e leves ese
Annu.Rev.
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HANN AND BONN
Pnt
OAEAEpple ( aees
Pea (1 aeesDewbey (2 hybdsBlackbey (3 aees
EMOAE9 speces
UAEE
Lemon
onghy emonLemo
Maeril
esh cefesh jceff
seeds
whole fpeeled fi
Lemon, oange and peel sapgapef
ded peel
Oange jce
Oange (alena) leaes
Lmes; medca and ice deuana
apef ce
marks
a m eq pe e; fms3 o 10 pe cen of ol acd
om o 11 m. eq pe e5 pe cen of oal acdConans none
033 ( V. faba) o pe cenPhaeou mutiou) ofdy w
4.2 o 5 pe cen of ce o .5 pe cen of jce49 pe cen of jce4 o 5.0 mg pe cc of
whoe jce9 o 1 mg pe cc of sap19 o 0.51 mg pe gmVaes nesely wh am.
. pe cen of ehe solble
acdPepn as calcm cae debed
20 o 23 mg pe c oe 90pe cen as fee acd, whchbecomes paly neazeddng gowh
Reerene
(105)(10(1)(1)
(164)
(1)
(19)(10)
(11)(1)(12)
(14)
(15)
AEAape
AVAAE
ooEAEAE
Cacao
(Baden Wnes) 431 mg pe e (1)
(1)
AEAEVaccnm
OANAAE
aw es
bean
Poo (een on cean)
Poao leaes
Poa o Swedsh soed)
Poao, Swedsh
semsbesbes
bes
o . 10 pe cen
0.45 o 064 pe cen of dy w (1)oac he only ohe acd
Among he bes soces n
20 imes as concenaed asmalc
2 o 1 .20 pe cen dy w)0.15 o 0. 149 pe cen dy w0.12 o 09 pe cen y w00 o 0.55 pe cen fesh w
05 o pe cen fesh wcommeca podon descbed
(19)
(10)
(11)
(1)
(13)
Annu.Rev.Plant.Physiol.1950.1:75-108.D
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OIC CID TBOLI
Pant Maeral
tt nhtsh vsnd btrswt
mtcc
HRkhrk cr nts rn
hus rwn
rs"wst
vs stmsn s
Reak
8 t r cnt dry wt
r cnt mximumEtctn mth
cn mthDi nyss ivn
MALC ACD
RAAWht nd bu rssSur cnLoium penne
HOODAABt nd c
OYGOAABuckwht
MAOAA
vsjuic cins
s ndtis
vs nstms
Schzandra hnensi fuRUAA
Bryphyum anum vs
u aceOAA
vrts
Pr 7 vritisDwbrry
juc
yrursh juicfri
Bckb vrtis ruit
GUOAP im bn sy vs n
bn nd ffsms
UAALmnLmn rn nd
rruitO (nc)
rrut
uc srd vs
uic
8 8 m q r cnt77 r cnt by vum t r cnt tt
crss wih
D nyss ivn
Dtid nss ivn
rinci ci
but 7 r cnt th ry wtwr in m thn n m
Prsnt
35 t 7 m q r ir mrss t r cnt tt cid crss withrinn
thd ductinth rductn 8 m q r r8 t r cnt t rnic
cdsrm t r cnt
rnic cds
Di nyss ivn
t m r cc t 8 m r cc t 88 m r m8 r cnt thr sub
cs t m r cc
Refen
7
78
88
8
8
8
(165)
(50)
(166)
87887
7
8
77
7
7
Annu.Rev.Plant.Physiol.1950.1:75-108.D
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6 HANN AND BONN
n Maa
AARDAEAEhus ph u
AEAErape Calfrna ue
ar.)rape
Grape Cnrd juerape Suen Bul
garan Wne
rape HungaranWneAAEAE
Cn ra berMEERAE
Carr
OAAAEa reen Mun
an)
bama
RAEAECanalupe meln
OMOAEeue
fermenedjue
ue
aeleaeem
leaeem
leae
marks
Calum malae ryal n
eed epderm: per enf eed eg
Dereae rpenng
Dereaed n!ly durng eae
0 0 gm. per 00 We 3.66 gm. per ered 20 gm pe ler
ed and yed gen;ey ar dely
0.3 0. per en
09 per en f al ad
/20 nenan f r
Exran med3 6 m. eq. pr en3 89 m eq per en
2 26 m eq per en m eq. per en
2 09 m. eq per e
OXLC CID
BROEFura and Sphgnum leae
RAMEAESugar anelum rnn
lu gra and eaEOAEAE
SpnaSpnaSpnaSpna 3 aree)
algn glmru
jueleaf ppng
leae
leaeleaepeeeae
le plan
Calum xalae ryal preen
00000 per en by lme03 00 pe en al
dereae ageCnan nne
208 309 m eq. per en par n 33 0 m eq per en00 800 mg. per 00 gm. dry
nly abu /3 a alum al
p 20 per en f al dry n fall
frn
89)
90)
03)
9)9
93)
)
9
80
8)8686)
86)86
86)
95
62)6)
86)
86)96)869)
98)
Annu.Re
v.Plant.Physiol.1950.1:75-108.
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OANC ACD ETAOLS
a Maera
Be vugar d bt lvBea uar lv
polMAANTHACEAE
Aeahera e lvAmahu aquacaAmaanhu lv
poygoodeA maahu gageu lv
PORTUACACA
Tau pecoa lv
UCIFRA
ul poutLGE
lm n llnd oybn
Caa orabaa gradora
TACA
lvtmlvlv
Lmon ong nd dd pl
gputValencia ong lv
VTACA
pAACA
Coton
Coton
RSRMACA
Blk tl
TRCACA
CoBA
ot
AACA
vn hoot"
d ootb
w b
lvlv
bn
mntdu
Poo n oun un)
Tomo utlvt
Tobo wtOTA
tu lv
Reark
Hgh ontnt 03 p nt29 to 323 m q p nt
7 to 9 m q. p nt
p nt o dy w.033 p nt o h wt1 1 2 p nt o dy wt
p nt o dy wt
127 nt o dy wt npt o bng tn
vgbl n Clon
Dnt n oxlt
eq . o 31 m. q p nt02 p nt o dy wt.0 p nt o dy wt
00 to 0 mg p gm
399 p n o old
ot undnt d
Eltodl ld kg.
0002 to 000 p nt o dywt =030 p nt od
1 pt n 270 o dy wt1 24 p nt o dy wt
032 to 00 p nto dy wt
0 p nt o totl d
02 p nt
thn pt n 0000 to 9 m q. p nt2 q p ntmthod o tton
to 2 q p nt
7
Reerece
1991)
1)
200)199)200)
200)
200)
19)
(6)1)200)200)
7
173)
201)
202
17
19)200)
17
194
10
9)()1)14)
Annu.Rev.Plant.Physiol.1950.1:75-108.D
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MNN ND BONNER
Pn
THER
"Malaya vegetable
"Formoa vegeable
Me Rerk
Oxalae conen ofen high [cfalo 200)J
Oalae conen gven
SOCRC CD
RCEE (Caulacean malic idened a iociic acid)
Bryphyum ynum leave lea pe cen of he dyw.
Seu and Peen in place of caula-
Ehever en an malicCEEDebey 2 hybd) 5 pe cent of oal oganc
acidBlackbey 3 vaieie) Coniues 65 o 85 pe cen
of oal oganc acidNCEE
oo (Green Moun juice Small amoun peenain)
UCCIIC CID
RMNEEuga cane juice
RUCEE
u re leaveCEE
pple juiceVTCEE
rape ouher Bulgarian ne)
rae Hunarian
ine)
MEERE
Caro
THER
Calabah cuae
femeneduice
ode
Small amoun preen
peen (by olaon)
5 pe cen of otl acid
Whie: 0524 gm per lieRed 01 m per lierMehod and yield given ;
conen omewha highehan in ne of oer counie
72 r cen of ol acid
preen (by ola io)
RRC CD
NCEEShzn ne
LMINE
Tmnu n
frui
frui
rincal acid
10 pe cen of dr , a freeacid plu pe cen a potium acd al
Reerene
203)
199)
50, 566)
166)
6)
6)
10)
162)
166)
105)
192)
19)
19)
20)
65)
205)
Annu.Rev
.Plant.Physiol.1950.1:75-108.Downloadedfromwww.annualre
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by
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ORGANI CID METABOISM
Pn
VACAGape Califona) (75
a.)Gape
Mateal
Grape jue
Gape Conod ueGape uhen Bul
gaan wnes)Gape (10 Hungaan
wnes)
Remark
Deas wth penng but
less so than mliceeases dung hot weate
ut less so than malBeomes esteed as ue
nena ted 0.26 gm pe Whte: 204 m pe lteRe 230 gm pe teContent soewhat lowe than
n wnes of othe oun
PYRUVI CI
IICEE bz
Onon
My aua an d y g
gm pe
CLOROGN AONOLEE
Sweet otatoSONEE7 spees
OMPOITEspees m 9 genea
eaves and
the parts
a 2 gm o 5 b
Pent n al bon spees dd msopa; snget tand paenta nne n emys ypay n epdems
Pesent n a, especaly
epdem and e adnng ea-ven
ACC AC
ROACEAAppe (6 aees es uie
VACAGape Southen Bl
gaan wnesGape ungaan
wnes)UBFA
Caot fementedue
o 8 m. eq p ie opses aout pe ent oftotal ad
hte 0920 g pe ieRed 26 gm. p lteMethod and eds gen
7 pe ent o tot ad
Refeen
9
0
(26
9(92)
9
(207)
20
(29)2)
2
)
92
9
94)
Annu.Rev.
Plant.Physiol.1950.1:75-108.D
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00 HIMNN ND ONNER
CIC CID
Plant Matea Reark
AEAE
Gap (So gaa )
MEAE
Sga caAEAE
Gp (Hgaa)
MEAEAE
app
AMEAESga ca
AAAEAE
W: 0432 gm p Rd: 076 gm p
NNIC CID
jc Sma amo p
Mo ad yd g
O IDS
jc
apoadc
Saccac acd 03 p cby g of oa acdp, dmd by ao o aco
coc 005 p c y o;gycoc ma amo
aacoc 07 o 1 33 gmp 100 gm od
a (P ae- bakfola)
c acd CH oad 7 gm p kg
EMAE
a
d co
o pa Oaoacc 30 o 40 mg pkg a oo dca ak
o pa Oaoacc 100 mg p k at oo
Reerenc
(192)
(162)
(193)
()
(162)
(212)
(23)
(3)
(35)
AEAAEAE
Dhaetalum cymoum("gba)
Mooooacc acd fod (214 2 1)o b oc pcp gy
MAEGay
E AMLECaaba ca
oo obb
a
od
coc acd ao oa d
acamcay cama ca 20
p c of acoob
Camc p pay pay a pay
oocac ad macoccd a d
(16)(161)
(161)
(204)
Annu.R
ev.Plant.Physiol.1950.1:75-108
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ORGANC ACD MEABOM
TOTAL ACIDS
P
AMNEAE
Suar caneLu peee
WheatBe rs (a sp
OLONAEAEBuckheat
ENOAEAEBeet
Spnac
ANOAEAEchzandra hinensis
AULAEAE
Me
juceleaf clippngs
leavesleaves
leavesstems
leaves
petolleavestioles
Bypyu yu eaves
OAEE
Apple
Pear
EMNOA
uce
juice
ima bean, pea, alalfa leavesand soybean eenouse
ima bean, pea, alalfa stemsand soybean eenhouse
AEAEemon uceLemon oange an peel
gapeuLemons tngernes
oanges limes andmeos
Orange Valena eaves
Graperut
VTAEGae 7 Calonia
varietes
ek
Incease when canes sprout00 m eq pe 00 gm y tn ecease t age
22 to 33 m eq. pe cent0 to m. eq . per cet
34 to 360 m eq. per cent206 m eq. per cent
406 to 42 m eq. per cent
202 m. eq pe ent6 to 30 m eq pe cent226 to 23 m eq pr cent
e cent as many r malc and tri
e secton on EEMNAON AND EAAON
See uner andAL D
See uner C aA D
6 to 26 m eq per 00 gmdry wt.
to 29 m eq pe 00 gmry wt. usually less tan nleaves
456 to 0 mg pe cLess than in pulp
orrelated wt vamn ontent
Al soluble n water exceptoxal
Fee ac aUs an pH rses sfruit enages
Change ih ripeng
0
efeee
(62(63
(6(6
(6(6
(186)
(186)(186)
(186) .
6
50
05
06
(6
6
(0(
26
3
(
90
Annu.Rev.
Plant.Physiol.1950.1:75-108.D
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02 IA AD BR
Pt Matrml Reaks Refee
G Hg Mo y g 93
MLVACEECoo 0 o 09 p STERCUICEE
Co A od o ox r
SONCEoo 0 o 70 m p 6
m 0 o 3 m p 6CRBICE
Cop mo 6 o 99 m p 6
m 9 o 30 m p 6OPOIE
97 o m p 6
ITRTR ITD
HIBN, A ., Potein Meoi i the Pt, Y N H 939
EE- nn 6 59-94 9. ICKRY, N CER G W A. Rev Bioche. 9 9 90
McEE H N hytoogt 48 No -3 99 WOOD G hyio. Reu. 96 966 RNEIN, , Adce i Poe hty - 97 CHER G W. WKEN A J N ICKERY, H , Id Eg. Che. Aa
d, 4-6 9 ROYER, C C, L N E F G t Phyo 69-9 99 C T , OYER T t Phyo, 17 9630 9
0 ONR, H M., J. Aoc O Ag Cheit, 0- 9 ONER H M J Aoc Oc Ag heit, 0 66- 9 HROO F Bioche J 6 96
. N, N, Ucheye Zpki Mkov Godt. Pegog t, 390 Kh ee. h. 4 9 ZHE D M Viodeie i Vogduo SSS.R. No 9-0 - 90
hi. Refet. Zhr, 7 9 RNN G J Aoc O A hit 24 39 96 EN, E A. ch. . 28 73 967 R, C G t Phyio 3 9 SOINEN H N RIO, , A. h 9 07-9 979 REB, H A S D , N N , Biohe. J. 0- 900 EER, H. Mitt ebiete Le Hyg. 9 303 9 GO, H Joe uiico 2, No 633 9 O, C N HRRR Mitt eite Lee. Hyg, 3-6 963 HER G W J. Bio h, 333 9 RINON . N SCC C J Bio he, 3 93
Annu.Rev
.Plant.Physiol.1950.1:75-108.Downloadedfromwww.annualre
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ORGANIC ACD METABOISM 03
25. EMAN D, ADY H A. AND JONN J Ind En. hm, Anal 16 556 (9
26. ME W W R H AND SAE F, Manometrc Technqeand Rated Method for th Std o e Metabom, 22 pp urgessPblihn Comany, nneao, 99
2 EC F AND us R Bohm et Boph , -82 (928. CE G W AN CEY H , Ind En hem, nal Ed, 25
(929 KE H A. AD EON Bochem I 3338 (93 ; 38 263
(930. ONNE W AND ONNE Am 1. Boan, 3 (983 COA S 1 Bo hem, 23 (9532 MRTS AN EONAD H Z Phol hem, 208-2 (93
33 JONON W A. Bochem I 0653 9393 CEY H AN ARM M D 1. Bo m, 180 35 (9935 ANEN A MO A. A ANSNDMAN Somen Kemteht [
4 6 (936. ANE IMO DMN D XNNE J k em
-90 933. K, H A. Bochem I 303 (9238. ILLNO F AND ANEA G Bo. o tal bo per, 2- (939. Lu, G D. Bohem. I 295 (9390. EDN E AN O H Bo hm, 133 5859 (90
. ELA S AND NL N, 1 Bo hm 138 3 (92. HLL . AN RMEY . 1. Ao r het 52530 (953 AE S A SMMEON W H 1 Bo hem, 138 535-5 (9 CLL, P., A CARN, E , Bohm 1 -30 (925. EN G. Bl. trav oc. pharm Bodeax, 5-62 (966 ENON A A AAM A CLV, GOOAE T C, HA A AND
SEA W, 1 Am hm So (n pss) . W H, A EE T traln 1. Sc Reearh [AJ1 98
(988 HAE C S AD EWOOD F A Natre n pess
9 NORDL A. Medd nork armac Selk 298 (9250 CE G W 1 Bo. hem 145 5-23 (925 . CE G W AN CEY H 1. Bo hem., 145 52532 (9252 CE G W EAENWO C S GNE W D AND CEY H
lant ho., 9 (953 CE G W AENWO C S, GNE W D, AN CE H
Pant Phol, 360-6 (95. CE G W AENWO C S GNE W D, AN CEY H
ant ho, 93 (955 CE G W AENWO S GNE W D, ND CE H
ant Pho., 3 23-32 (9856 O J Pnta, 15 52-6 (9325. ODAY D A CA n Boan, 8 89203 (9
58. ONNE W D J Stde on the oranc acd o pnt, 03 p (Doctoa e- Cal n of enoloy aadena Ca, 96
59 OW AN ONNE , Ah Bohem, 509 (98
Annu.Rev.
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0 THIMANN AND BONNER
BRR, H ND T, (son communcton6 DR, C P, ON, H Y, ND CU H H Q Pnt Phyol. 3869
(948
6 RDR, R ND BOD 0 Pnt 469504 (943 ; f RDR, R, NDN, J, Nch Akd W. ottngen, No , -4 (943 PCHR, G W NOH, C. S, GNTR, W D ND VKRY, H B,
Pnt Phyol 05-7 (9474 PCHR, G W, ND VCKRY, H B, Pnt Phyol. 7-83 (9465 VKRY, H B, PHR, G W WKN, A ], ND NORTH, ,
Conntt Ag xpt St 44 (9406 VLDRO A V, So Sc., 576 (94567 N, H J ND WK E Sol Sc Soc Am Poc., 35 (94768 BLCKN, G E ND LN W G Ann otny, 4, 53387 (940
69 PKOTZ, P, GLRT, S G ND SH W, Sol Sc, 58 95-303(94470 VDRO, A V, Sol Sc. 377-85 (945 KUCV P A hm eet Zh 5455 (940; dy Belok
Sel'Skohoz nt -0 (9397 W E D ND PTR, A H K Atln J xptl o Med S
34 (94073 LN, V S, Stbe Akd W Wen Mth.ntw e. bt 875
(94774. LN, V S, lo 599 (944
75 OOL, B , Plnt Phyol, 43-4 (9487 PR, E C ND pLN, C 0 Pnt Phyol, 8 438 (94377 OTO, D NK, G, . ndw Vecht Bg., 93-9 (9478 PHR, G W., ND VCKRY, H B J o. Chem 78 557-75 (9499 KROTKO G ND BRKR, H Am J Botny, 35 5 (94880 PYTNTK, M P Compt end d US.S, 55-58 (9408 PYTNTK, M P, Compt nd d c USS, 59 (9408 GNKI S 0 l Botn No. 4, dy on Int d Sc USS
Sr IV, 07-35 (94083 W . G, Atln J. xpt. Bo. Med Sc., 33 94
84 WOOD, RCKHNK, D H ND CHL, H tln J xptlBo Med Sc., 3753 (943
85. WOOD, J G, RR, F V, ND PDLO, C Atln J. ptl o. MedSc 37-43 (944
86 RCKHNK, D H ND W J G tln J tl Bo Med S, 43-4 (945
87 VCKRY, H B, PCHR G W., WKN, A ], ND NORTH, C Connectct Ag. Expt. St. Bl, 44 (939
88. VKRY, H B PCHR, G W W A J, ND NORTH, C. S.,Connectt Ag xpt St . 399 (937
8 WRR, 0 Unte ot nt ngn, 53-50 (880 SCHRTZ P, Pln 8- (339 LRK, H nt hyol 534 (939 VNNLND, B, ND LHR, R Z ch ochem 9-3 (9493. GOLL, M C. ND VNNLND, B J Bo. Chem., 33-34 (94794. SK . F. J Bo Chm. 178 354 (94
Annu.Rev.
Plant.Physiol.1950.1:75-108.D
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ORGAI AI EAOIS 0
95. VENNEAND, . GLLU, ., AND SECK, J. F J o hm 201314(1949)
96. VENNELAND ., J o hem 5919 (199)
9 EITlAML, . , AND VENNELAND ., J o hm. 1333 (194998. ICARD, A V. Trp Ar eon) 9 354-61 (1940)99 STE, W, ro Forda ae Hr o 60h Ann Meen 55-62 (194)
100 SINCLAR W , AND AMEY, , oan Ga 148 (194)101 EYNAUD, E e 9 1780, 2712 (1946)102. ENAUD, E. Inds ar e amen. 6788 (197)103. FLNZY, Ann aron 6064 (1948)104. AUE, ., Jorne noe 313 1948)105. AERNER, . AND ACQUIN, P, omp. rend, 133-4 (1947)106 AERNE, AND ACQUN , omp rend 1393-94 (1948)
10 STEAR, F STUT, P AND RETN n hyso 409-(1940
108 EDANL, AND ARANDANE, G o so. o. sper 828(1940)
9. VANN A AN LAN Z 21-1 (1941)1 10. AUANEN, J o he. 68788 (1946)1 1 1 ENARD AN UR, R J o h 01-8 (194)112 U G AND EN, o hem 9 19-26 (1943)113 UANEN, N Aa hm. and 127-39 (1948)1 14. FRANKE, W, SCUMANN, F., AND ANEREE, , Z. hso h. 2-42
(19431 15 LM, AND EER, ro. Konnk Nedernd Akad. Weenshp
9 10961100 (1946)1 16 Z AN MAN, N G. Dokady Akad ak ..R 62 255-58
(1948
117 CA, S., J o. hm 9 23, 25 (195)118 ENN, A AND ALIN, f xp. on ' p: en 0
476 (1948) ; 9 14 (1949)
119 AME, G AND AME, W , ew hyoos 9 266-0 (190)10 AME W O AME G. AND UNTIN," . ohem 5889
(191)121 XMACR, F. AND UD, . ohm. Z 1-6 (1936)122. NNER . AN WILDMAN, S G, Ah ohm 9751 8 (1946)123 NNER, Arh ohm 31 126 (198)124 MMNER, , AND IMANN V J Gen hso 2996 (1941)125. ERER, J, AND ERY, G S., R, Am J. oan 1118 (1944)126 ATE, G G Arh ohem. 2 28499 (1949)127 AE, G G Arh ohe 815 (1949)128. ACLI, . , Am J oan 183-92 (1944)129 ANE, n 9 174-76 (1949)
130 ENDEN AN AUER, Am f oany 52835 (1944)131 ENNELAR, AN EN, ew hyoos 6492 (1943)132. URNER . S ., AND ANLY, . ew hoos 19-1 (1949)133. ADARAN, AND VENKATEAN, R ro Idn A ad . [
34559 (1941)134. ERER, , AN ER G S., R J oan 299 (1943)
Annu.Rev.Plant.Physiol.1950.1:75-108.D
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0 HMANN AN BONNE
3 , N B AND SN GUTA Ann Bioem Ep M In 694)
36. URNR, J AND ANY, V Naur, 969 94)
3 UNDH H., Arki. Boan [3 3 944)38 OLNIKO A, Doky Aka. Nauk S.S.S.R 33 948)39 LRCH, A Am J Boany, 63 94)40 RICH A, Am J Boany 9 0 94)4 LAUM, G., AND OMMONR, B Bio Bu. 343 94)4 LAUM H. G, AND ICHL B Am J Boany 30 8 943)43 BONNR, J, Am J Boany 333 949)44 HIMANN, V AD BONNR D JR, Am J Boany 35 8 948)4 HIMANN V, AND BONNR D JR Am. J Boany 36 4 949)46 DN S G AND BONNR J Ar Biom 4 383 94)
4 LY, S AND RY G JR Am J Boany 46 949)48 HIMANN, V Bio Bu 9 96306 949)49 BONNR D, JR, AND HANN, V, Am J Bony, 37 66 90)0. ILDRANDT, A , AND IKR, A. , Am J Boany, 3 48 949) IH A . AND IRT, A Bioem 4 943) BLAOHHNKII A V AD OONIOA A Y omp. rn. aa
URS.S, 4 4403 94)3. UNDRH H., K Lanbruk-ko An 339 949)154 O S Bu 11 14 194 ONTOCH, V E omp rn. aa UR.SS, 36064 944)
6 BONNR J AND GATON A W Boan az 898 944) BONNR, J Boan Ga., 343 946)8 HIMANN, K V, i T ormos 436 incu, G, ND HIMNN, V,
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Cem brc 5880 944
Annu.Rev
.Plant.Physiol.1950.1:75-108.Downloadedfromwww.annualre
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UniversidadeFederaldeVicosaon03/25/14.Forpersonaluseonly.