ripening of fruits

7/30/2019 Ripening of Fruits

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FRUIT RIPENING

C. Bdy

Plant Physioogy Unt, CSIRO Dvson of Food Research and School of BologcalScence Macquare Unversty Norh Ryde New South Wales Australa

2109

CONTENTS

1. NRODUCON......................... 155

2. RIPENING S DEVELOPMENTALLY RE GLAED..... 162 Ripening as Senescence ........ 15622 Control by Growth Regultors........................ 172. Gene Exprein in Ripening Fruit ....................... .. 160

2 Ripening Mtants ......................... 1633 MEMBRANES AND ORGANZAONAL RESSANCE........... 1644. OXDAV REAI ONS .................... 166

e e e................. 1662 The Cimteri nd Prtein Synthesis ................ 166 Other Oxidtive Retin .. .............. 169

H ONG OF FRU........................ 169 Chemis ad Enzymlgy f Softning. .................................... 162 A Rle fr Calium ........................................... 170 Sfteni a Regultr f Ripening .. ..................................... 171

6 C ONCLDNG REMARKS ................ 171

Fu pnn hs bn vwd nub of s n hs ss hphsoo of pnn was ad fl n Coob's w on fudvlopn ) nd h subc ws vwd hoouh b ch 8)n 7. Yn Hoffns 8pp 58)on h snhss and acon ofhn s pnn an h scuss sva aspcs of h phsoo an

bochs of pnn A nub of vws daln n wh pnn o wh h bochs nd/o ocu boo of pcula fus haappad dun h ls fw s 5 52 55 58 5 7 88 25)

-2/87/55$2

Annu.Rev.P

lant.Physiol.1987.38:155178.

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156 BADY

Studies of the mechanisms that regulate ripening can be traced to the 1920swhen the dominant theory emphasized organizational resisance" Thistheoy, developed in detail by Blacman Parija in 1928 (8, sggested that

the ripening events, which were perceived as largely catabolic in nature, werethe conseuence of a breadown in the resistances that et cellular comartments contained. This theor hed swa, in a largel unmodified form, for40 years or more 2) As evidence accumulated that protein and perhasRA synthesis plaed vital roles in the induction of ripening, an alteativetheo emerged: rpening was presented in ositive tems as a rocess oftissue differentiation The techniues of mode molecular biologists haveenabed this theor t be examined in a recise a, and evience for thedirect genomic control of the ripening of climacteric fruits is no accumulat

ing rapidly onetheless there is evidence that celuar compartments aremodified through ripening, and recent evidence suggests that lipid oxidationandor phase changes ithin membranes contribute to changing metabolitedistribution ithin cells as ripening proceeds Ho novel transcriptionaland/or translational events and changes in metabolite artitioning interact andcontribute interdeendently or indeendently to riening is a matter of con

jecture It may well be that both thories are more or less correct and bothmechanisms are invovd but to differing extents in diferent fruits. Fruitripening appears to be a ellregulated geneticall determined event, but

coming as it does at the end of ontogeny in a fleshy, energyrich tissue that isdestined for disersa and disosal, it oud be surrising f the mechanismsof control ere the same for all secies he revieer naturally seesgeneralities and uniformit of attern. he reader should be ed that theemphasis given to a few conclusions based on a few species that today mayseem to oint to firm guiding principles ma tomorow seem angential orpaticular as more species and more aspects are examined

P PY Ripenng as Senescence

The distinction between riening and senescence has never been finely drawnWatada et al 4) have defined ripening as changes that occur from thelatter stages of groth and development through the earl stages of senes-cence and result in characteristic aesthetic and/or food quality They definesenescence as processes that follow physiological or hoicultural maturityand lead to death of tissue These are sensible, pragmatic definitions

although it is ossible to challenge the concept that each of the pocesses thatcontribute to the senescence sndrome leads directl to tissue death. hedismantling of the chlorolast hotosstem aaratus is a rominent feature ofthe senescence of leaves and man fruits, but it is not lethal. t is perhaps

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ENNG 17

mre accurate t suest that the senescence cesses increase the rbabiit f death fr eamle b dehdratin r micrbial inasin fr there islittle eidence that senescence includes rammed death here is n

dubt that the ntusn f the rpenn syndrme nt tssue deelpmenhastens the nset f senescence and therefre the pbablty nuy anddath Ths is clearly seen in the cmarisn f th nnriein tmatmutants and nrmal tmates 27. he nnrienin mutants, which arennclimacteric eentuall senesce lse chlrlast cmnents and cel wallhemicellulses 0 and becme mre suscetible t disease The nrmalfrut under these same chanes much earler as pat f rpenn t fllwshat senescence and rienin hae cmmn features; hweer renn n-cludes pcesses that are nt part f the senescence syndrme Pmen

accumulatin and the cell wall chanes that result in the sftenn f fru arecmmn rinin chanes that are nt usually inld in the snescenceprcesses

In the clmacterc ruits ripenn s characterzed by ethylene prductnwhich is aarently autcataltic McMurchie et al intrduced thecncept f tw systems fr ethlene prductin system s the ethylene-rductin sstem cmmn t climacteric and nnclimacteric fruits anderatin in climacteric fruits until ienin cmmences. n tese maturefruits as als in sme flwers esure t ethlene induces a lare increase

in the ethylenefrmin system 8 and this is seen as the nductin fsystem 2" ethylene. lms Laties suested that the autcatalyticnductn ma be cnseuental t chanes in ranzatinal resstance"resultin in the liftin f reiusl imsed cntrls. re recent eidence8 indicates substantial increases in the actiities f the reulatr enzymes it has nt yet been shwn definitiel that the increases in actitiesresult fm te snthesis f mre enyme but cclheimide des inhibit therespnse 8 he inductin system ethyene results in massie increasesn ethlene rductin b the tissues and rienin and then senescence

fllw. Climacteric fits are nw distinuised as thse in which sstem 2ethlene perates. The cncet f tw sstems fr the cntr f ethlenesnthesis has been a useful ne and shuld be better defined. n time we willnw if searate enes cntribue t the tw systems if tw leels freulatn apply.

Control by Growth Regulators

here is idisutable eidence that ethlene s nled in the inductin frinin in fuits 8 A main cncern f the cmmercial stharest

hrticutural industry is t limit the esure f harested fruit t ethylene Anasect f uit rienin that attracts mlecular blists s the rsect thatethlene induces a mature, nnrwin tssu t radly dierentiat int a




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58 BRAD

w sttto swtch from opg to pg Bcus th tssu sdy mtu d th fom d stuctu of cs dcdd, t s thought thtth moculr stps vovd wh h swtch s throw my b tvly

fw h sysm hrfor hs pp s o whc o sudy rctobtw h sglg chmcl, thyl, d h gom 58 59. compcto s tht th tssus posd to ow vs of thy

poducd by systm thyl thoughout dvopmt. Eposu to gcoctros of hyl producs, fru ssus, spos h vrsbw spcs d wh ut muriy 5 I h mor sstv spcs,fo mpl ctoup 92 o b 26, pg s mmdtly ducd,but th mo mmtu th fut th hgh th coctto of thyl thts qurd I th lss sstv spcs for xmpl tomtos o ppls

thy rtms rduc tm bfo ripg occurs 9, 9. cipg s ot mmdty ducd tomtos by thyl ttmts,systm 2 thy sythss s ot ctvd. Ethyl sythss s omlymd by h supply of h d prcuso -mocycloproplcrboxyc cd ACC Durig rpg, s sysm hyl s cvd,ACC syths ctvty d cott of ACC ris srply 32, 8. Immtu tomtos xposd to xogous ty sow o cs syths ctvty but thy do show dffrc thr rguto: ththyl-formg ym EF s ducd w ACC syts s o I

cucumbs, F s cos dug dvlopm d ripg, whl ACCsyths d systm 2 hy sytss cs wh rpg 39. Mturg pps gduy dvop th cpcty to poduc EFE spos toppd thy .

A rl for CC syhs rgulg hy syss d rpg sdmostrd by hbo of h duco of ripg wh 2-mo-lkoy-ts--butoc cd movylglyc, VG s dmstd toprcmctrc prs 0, 2 or pps 2, . Th hbto of thducto of pg by V s ovcom by subsqu posus to

thy I D'jou ps, whch ur low mprtur sorg prio oorml ripg, Bksp Rcrdso 9 foud EF crsddug cod ptrtmt. hs frut, s tomtos, systm 2 EFE dvlopsror to sysm 2 syhs. I ctrs 28, EFE ormy cssdug pg, but hs t l mtuts cpcty xcss o tht of syths I hs fut xogous thy dcss EFE cvty.

Buf Bgth 7, 34 ppld thyl o propyl to ppls sordud hypobc codtos. Futs stod hypobcly fo shot podstd th cpcty to gt systm 2 syths thos stod fo

logr pods lost ths cpcty Apptly thr mmtur tomtos ommtu or ovmtu pps bl to produc systm 2 thy o p spos to ppld thy t s kow o th tl fcos tht

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R RIG 59

reglte the epone o rt to ethylene; poblte nclde the concentrton o othe growth regultors nd the ntegrton o eler responses togrowth regultor Thu the reponse to ethylene my depend pon the et o

genes tht e trnscbed or hve been trnscbed when the expoure toethylene occrome r eg vocdos 4, do not rpen whle ttched to the tree nd

grdlly ncree ther sentvty to ethylene wth tme er hrvest. Thsh red th concept o rpenng nhbtor or jvenlty ctor deved rmthe tree In vocdos, wth mmture pple or tomtoes hgh concentrton ethyene omehow redce the retnce o the r to ethylenend redce te tme beore sytem ethylene oprte Whether one sees ths ttrton o the resstnce o the tue to ethylene or s the need or slow

ethylenepvoked developmentl progrm crrently mtter o choce.There hv een mny ttempts to dene roles or the other wellknown

plnt growt regltor n the reglton o rpenng or retnce ctorsThs sbject w thoroughly evewed by McGson et l 9, nd moerecent tude hve provded no generl clricton. bscc cd commonlyri lt n development or duing rpenng 59, nd tretment wthcc cd my dvnce rpenng 6 86, 98, , 9. Abscsc cdrequeny pomoe he enecence o geen se 55, nd bcc cdncrese n osmotclly or slt-streed sues 56. lt tretments o plnt

dvnce rpenng t let n tomtoes 9 o there premptve ceor ole o bscc cd n ome rpeng recton. However the ce hnot been btntted or nely drwn nd y et l 4, ond noconstent reltonsh between the bscsc cd content o uts nd therrpenng or enecence.

The hypothes tht endogenous xn ply roles s rpenng nhbtors4 ttrctve Evluton o the hypothess complcted becue uxnsboth ncree ethylene prodcton nd ody tse repone to ethylene58 In ple, Mosdle Knee 0 ond pek n indole-cetc cdconcentrton sytem 2 ethylene ws ntted but they wee nble toconclde tht the two were relted.

Reech n ths re h been hmpered by the dculty o equtngreponse to extelly ppled regltors wth endogeno control by thegrowth regltor, by the ntercton o the derent regultors, nd bynlytcl dclte when ttempt were mde to monitor endogenou concentrton. he ler dcltes re recedng, bt recognton o themportnce o dstrbton wthn nd between cells nd o the desrblty omontorng eceptors 42 s well s growth regltor lmt stdes n thsre

t poble tht the nluence o ethylene receptors my be montoredlbet ndrectly by the pparent blockng o thee receptors by nonc lver




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60 BRADY

48 49 lve nb e penng of bnn 9 nd omo 66 9f e nonoc concenon nd e nhbon no eveed byeelly ppled eylene. In oo fr dc lver nhb fer

penng f ppled o prly rpened e G A cke nd C. J. Bdynpbled; ndce eylene con nvolved og erpenng peod nd no lmed o n nl ggeng of penng. ghconcenon of eelly dded clc nb e ndcon of rpenng n nmber of fr 45. In vocdo nflron wh 04 M clcm pevened noml penng 44 een w ehylene ovece he clcefec. he eec of clc w o preven ye 2 ehylene ynewo modfyng e cpcy o repond o e ge concenon ofeylene h e prodced by ye 2 ehylene

Gene Expesin in Ripening Fui

he concep of penng dffeenon even w nlly mled byn ppren ncree n ynhec cvy ely n e cmcerc re. Foexple n vocdo ere were rge ncreen n e ncorporon ofpecror no proen 22 nd nclec cd 23 he nbon of penor nclec cd ynhe 28 pevened nr rpenng nd h w ken evdence h novel nlonl nd/o rncponl even occrreddng he ely ge of penng. e evdence w f fo fcoy.

o ple lbel or o nrodce nhbor e ce were ed o hnecon beween e effec of lcng 32 nd oe of penng weelwy nvolved. here w no cler evdence of he ynhe of pecfcpoen h d ole n penng. I w ggeed h mlc enyme wnewly ynezed n he ely clmcec of pe 4 b e evdencefled o dngh ne yne fro ove. In bnn ee wevdene mo o he ncemen n oen yne ely n e clmcec re reled n n ncree n oer nd he replceen of preexngpece of proen 25.

More recenly evdence for redecon of poen yne dng penng emeged Incee n e cve of 4 glcnhydole cellle n vocdo 37 nd n nvee 72 nd polyglcone n ooe22 4 45 rel fro ncree n e on of enye proen eed nologclly. he conen of ee ydrolyc enzyme nceefollowng cnge n her re of yne ndo degrdon nd no el of e cvon of preco olecle ynezed drng erledevelopen her e ynhe of ee poen ncee dcllywen penng comence or ele pevoly nble rnlon podcbecome ble. Bece e enzye normlly fncon n e cell wllcellle polyglcrone nvere or n he vcole nvere bly cold concevbly be e coneqence of developng mecn fonpong or poceng he pry rnlon prodc.

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FRUIT RPENNG 11

In vocdos ristofesen et l (38) showed tht with the dvent ofripeng there ws chge in the popultion of trnsltble mesenger NAmoleces. vtro stio of poy (A +)-NA popultions showed thee

products tht wee not pprent in trnsltions of NA fom nonripeningfrt. n etenson of this wo ucer Lties (1) identfed byimmunopecpttion 53000 dlton poduct of the in vitro tnstionsyse s he recursor of cellse ipenigspecfic cDNAs were loctedin cDNA lbry constrcted from the poly (A +)NA of ripe vocdo fruitA cDNA specific fo celllse ws identified by hybrd seection i vtotnsltion nd podut immunopecpittion (37) y hybriiztion themessenger NA fo the ceulse pecurso ws shown to increse t lest50fod dug ripenng (37)

Bennett ristofersen (13) determined the sequence of ful lengthcDNA d the Nterm sequece of the mture cellse protei ndsuggested th the pimry trnstion poduct is 5000 dlton polypepde vivo remov of sig peptde dig membne pssge yields processed polypeptde of 52800 dltons; gycosyltion prodces membnessocited form of 56500 dlons tht is timmed to the mteprotein of 5420 dltons which is presumbly locted in the cell wllregio

his wor on the ipening vocdo provides direct evidence of qulittive

chge protei sythesis durig ipening A eslt of the chnge is incese durng ipening of gene pouct which pesumbly plys role inthe softening of the fruit (62). he eperiments do ot involve slicing nd thecosequentil interctions with woundin (32). he evidence estbishes chnge i the stedy stte level of messeger NA fter ripening is inititednd the cDNA probes llow n pproch to the gene(s) nd their prmoteseqeces. t is not yet estblished tht he trsciption of he celllse geeis ctivted when ripening initites bt clerly the contol of celulsesynthesis is t pre-trnsltionl level.

Similr evidence h risen from studies of the tomto frit. Within 48hours of the initition of ipeningindicted y system 2 ethylene synthesishnges in the popltio of messeger NAs were emonstrted(57 37) A few messenge NAs dcresed in reltive intensity nd si toeight incesed in etive intesit he screeig of cDNA libies pepredfrom te poly(A NA of ripening frit confirmed the pesence of numberof messenge As tht re uique to or gety ehnced in ripening frit(56 87 13) Mnsson et l (87) found tht bout % of the homologues ofhe cDNAs they prepred fom the poly(A +) NA of ipe fruit were mospominet in mtue onpeing frit bout 2% wee most prominent npeg fit whie the emide were eqly pmiet n unrpe d pefrit Piechul et (112) detected tnscrpts for rnge of chlorplstprtens n immture green tomto fruits trnscripts of only wo of these




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12 BRAD

wee peset pe ft The cofmed ele obsetos tht pe ftcontn clooplst bosoml RN (11 13) nd tt te plstds of petomto ft ppe to em cte n pote stess ()

olyglctonse otin ncss n tomtoes dng nng (2) nn soft clts t becomes mo pote the pe ft (22) The lgencement in nt synthss suggsts tt the polygctonse messengeRN m become ey pomnent dg penng nd nbe of gopse sogt to ccteze ts messege RN nd tence ts gene () Teenzyme exsts n tee molecl foms n extcts of pe tomtoes () bttee my be only oe cte polypeptde, te ote foms epesetgint gycosytion n th snc o nctiv subunt ( 101) The most pomnent ezyme, wt sngle sbnt s been ssgned

os molecl wegts 3000 ()000 (1) 000 () nd 00(101) dltos eso d co-wokes descbed the pecptto, wtoygctuons-scic nts of n n to synthsiz polypeptde of8000 dltons () hy so obsv tht cDN to penng-encedpoly RN speces hybi slct mssng RN tht coded fo polypeptde of 8000 dltons, wc ws pecptted b te sme sem(3) Te ptte polyglctose messenge RN ws of eltely lowbndnce. to t l (130 131) immunocitt, with n ntsrum topolglctose polpeptde of 000 dltons fom tsltons of te

RN of pe tomtos By thi stmt somwht ss thn 1% o thtsltble messege RN n pe tomtoes coded fo polyglctonseh oth ots o 54000 ton cuso to tomto oyglctuose (16 1 1 100) odcts n the 5000 to 5000 dlton egon epoment mong te polypeptdes snteszed n to fom te RN fompe tomto ft whete the tslto s b whet gem (13) o etclocte lste tnslton system ( 3 tnsltes of RN fom geen ft donot he pomnent podcts o this sz DPnn t (1) peped cDN lby fom the poly RN of pe tomto ft n inst thlb nto expesson ecto olyglctose cDN clones wee dented by nologcl sceenng. 2.kb messenge RN fo polyglctonse ws detfed d ws sown to code fo 000 dltonoyt Te messege RN ws not detected n mmte geen ftws ist tct s inng commnc, n gty incs in bunncs th uit colo h vnc is now substntil tht th imy tnsct o tomto oyglctuons is olyti o bout 54000 ltonsd tt t ncldes n nsll long sgl peptde Seqece stdes ecofmed te dett of cDN pobes (W. R. tt d R. See DDellen d B Bennett D. eso, pblsed nd wll dntiction o te pocessng teps between th imy tnslton ouctnd te mte enymes.


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Ripning Mutants

F NN 63

arl mi a lat ao variti ar wil rcogi amog hortcultural cp hr occuc mpl that thr a gtic compot rgulatigth iitiatio o ripig but it o o tablh a pcic gtic cotol oth ot of ripig arl ma b o apct of a plioropc ctprhap rci though growth rglator or rcptor coctrato

Sral potao mtat of tomato appar to b pcficall rpg mutat. Two of th th ripg hbtor orin mtato o chromoom 5 a th oripig ornr mutatio o cromoom 10 r rcivmtatio that rlt i oclimactic ft (14 h uit ail to prouc tm 2 thl a al to go omal ipig Wh po tohigh cocrato of thl ur pcific coitio ri fuit vlop amot amot of lcop bt o o o apprcabl (487 14 rfuit prouc om lcop rig cc a thr i a light r ithl puct. Wt alt tatmt nr ut at lat om gticbackgrou ma vlop ar-ormal lvl of caot a om polgalactoa activt ) Both ri a nr ar tv to thl i that th prc of a thl th rpiratio rat r (141 i rin platthl tratmt iuc a icra i EFE (83) bt th fll tm 2apparatu i ot uc a thr i o mav icra A tha

Th two mutat lack th capacit to iuc tm

thl thi ath alo ac th capact to rpo to th pour i t o oalripig A arch for mutat that proc tm thl bt o ot rpi rpo o a thl or rip i th prc of thl withot tm l ma b wohwhl. Th vlopmt ofmolcl pb for rpig a tm 2 thl (prmabl Atha) woul ubtatal a uch a arch. rin a nr fal to vlop thcapaci to rp a th ar vopmtal mtat. t ollow thatripig a tag o vlopmt pccall cooll b a mall g t

aitio to ri a r which oclmactrc thr r a mbr ofmtat i which all rpg pamt occur lowl a co ovr alog po Nvr ip" (Nr a omat alll o chromoom (14gr rp (r) i a pal omat alll (73 a alcobaca i arcv alll that alllic wth (4 or poito clo to (0) nr ochomoom 0. h low-ripg mutat hav a pr a tclimactric color lowl a omtm icompltl a pc mallamout of polgalacturoa which catalz a low oig o th fuit.por to cra lvl of thl o ot hat th pig pc.

Th oripig mutat althgh rcv c th rpg ofuit i htrozgou plat. Th htrogot ip mor lowl tha homal prt a polgalactroa prouctio much rc (46. th htrgot th nr g grall a a gratr ct tha o rin but




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BDY

e nfence of e mn enes s rey nfenced by e enec bcgron.

gcer e recognzed e ve of e rpenng mns s npproc o e condonng evens ow rpenn o proceed eysggesed e syness of poygconse my be e crc even ne ndcon of rpenng nd s con d cyc roe n endcon of nrcer rpenn. s seems ney n e non ofpoygcronse syness s revey e even n omo rpenng 22 8 n ere s some evence nd re be osynesze poygcronse n some crcmsnces or some sses. esowpenng ns syesze poygcronse e presence nd encness o e poygcronse genes w ndobedy be evedrecy now moecr probes or e ene re vbe reseny epysoogc n bocemc evence sggess e mons re n rnge of gees rege e epresson of e rpenng syndrome rern n genes code or cys of one or more of e recognzed rpenngevens.

Rpenng mns ve been recognzed n oer speces A sowrpenngnecrne genoype 29 fs o prodce norm eves of eyene Wensmed by dded eyene or propyene endogenos eyene syness s

ncresed n rpenng vnced27

M. E. Person s recognzed nonccec ppe spor person commncon nd seems key w e rezon of e ve of sow or nonrpenng genoypes oerrpenng mns w be escrbed

B Z

ere s now frm evdence e syness o poygconse n ooes nd cese n vocdos occrs drn e cmcerc nd con

nes n rpe fr 7 A fncon rbosom ysem s rerd nd recovery of poysomes from rpe r s conssen w s fndng 7 ere re src onc reremens for poysome sby nd fncon7 n ese reremens re ppreny me n e cyopsm of erpe frus. s mpes e psmemm n onops mnn erfUncons es s ong s e ne synessnd presmby e processngnd eporf e wydroyzng enzymes occrs. One ms cocde ere s no brekdown n permeby brrers even n e rpe fr

Sdes of e rsrcre of rpenn vocdos nd omoes 0

ve ed o me concson. Aog some cnges occrred n esrucre of moconr e enopsmc recm n e psmemmdrn rpenn, ese cnges were ssoced w meboc evens,n prcr w e epor of e w-ydroyzng enzymes Oer




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FRUT RIPENNG

rasrucural sudis o rus hav sggsed a disurac o memrauco i ssc pos-climacric ru cells u hr ae usios as

o how accuraly srcus wer maiaied i h rp issues i whch wllegadaio 15) was advacd he avocado sudy 13) oed ha aphas chag i h plasalemma ilayr may occur i ripig r u hevidec suggesed ha he hag occur a h climacrc pak ad wasasory. s woud dsguish ay such ve om he phase disuracsh occur i sesc coyledo 94 4) ad oral issus 0) ad hahave ee ascried o hlee acio. aher simila chages we descid he microvscosiy o micosomal" mmaes o sescg apples 08)

A m o aayses ipds ave e rpoed ad some o hse

wer coupld wih osrvaios o h permailiy characrisics o lipsomes prpared rom he lipids or lipid suracios. osss o memralipids om omao 5 o plasid mmra lipids 53) ad o phospholipds f app a a a ppa 9 n rp appsad o u saaio i h memra lipds o ripig aaas 3 hav allee descried. sscig lea ad lowr issue chags i ludy havee racd o icrases i eural lpdsserols ree ay acds or hydoprodeuaccompaid y chages he phosphopd casss or hayl spcis 4) A icras i lipid proids durig ripig has e

osrvd i omaoes 4 ad i hydge proid i prs 30 Aalycald liposome sudis sur rom avragig s as he cosius osvra memras are mid. hey my provide videc ha he poeialoaag chags oe or more meraes u hy cao e devecau hy o llowc o h cpaciy o h ces o cop whicreasd lkiess usig aciv cair or oher eas. he chages hahav osrvd as i aaas 5 ad appls ) phasie hdyamc au o he issues ad h capaciy f chag ha ss. capaciy or chag may also ma a capaciy or repair a aspc ha has

e mphaied y omai 1Much o he videc or a disrupo o issue comparmes duigrpg vlvs he use o ssue scs 8) Mhods usg slices cessarily ivolve a ieracio we h iiial codio o h issue ad hrspose o slicig Th iracio may dped upo cell wall popies oh osmoic gradi we issue ad ahig medium 133) or o he rao he meaolic resposs o slicg Mos reserchers who have masuedupa or om i slices have oud some chag associad wihipeig alhough somemes he chages are sligh. omaes her is

vidce or a small disiuio o poassium ewe cell cmpams, f a icrase i draulic coduciviy or a dras hcapacy or aio acd upake 5) or a crase acd u a or a dras i cyoplasmic volume 6 o o h chags suggs a




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66 BAD

catastrophic eaage between compments [although if ctoasmic potassium concentration is ctuall as low as 50 (1 the eect woud be

considerabe (157] and some of the changes become signicant onl ate inpenig whih suggests an associaton with senescence rather than withening er se

he sum of evidence is that frit cels do not suffer a massve breakdown inceur comartmentation durng rienin but that there ma well be atendenc towards a greater leakines of membranes. The evidence or theater is cuuative ad boows heavi o that peraiig to petals adcotedos No sumatio of abiguous evidene no atter how cosistenti its detio a satisfactorl estalish a onlusion ad evidene fm

anases from iosoes and from tisue sices al have some ambiguit Insitu aases b M or oter mans ma eventual rove stead statechanges in distibutio; changes in uxes ma be arder to rove.

XV

Th Rsrto lmtr

Muc o te asnaton n stue of rt enng les n attemtng toexain the resirato climacteric and its relation to other riening events

here have been erods when te climacteic se was thought to be directlreated to the aaboc needs of ipening ( and oters when it wasconsidered the rme reflection of the decine in organizational resistance 86 Both peods sem to have passed but no entirel satisfacto exana

tion of the cimacterc has emerged. he subject has been reviewed recent(1 13 as were the athwas fr eectron ow 80and no extendedeatent is reuired here

he inrease i espiratio appes a to be a oseuee of the increasei edogenous ethlee b to resut in increased AP levels and erhas in

an inrease in energ hge in the cel (5 and (cto be assoiated with ainease in ftose26-bishoshate conenaton and a subsequent nreasein gcotic ux (A. B. Bennett unulshed he contrbutions o thectochrome athwa the canide-resistant alteative athwa and theextent of b-passin of site hoshoration (4 ) remain mattersfor debate Since ATP concentrations increase though the cimacteic eod,the increase in resiration rovides a sul of hemcal energ in ecessof the demands of the tissue

Th ltr Rs d Protn Synthss

Energ baane and energ use n nongowng tissuesthe socalled maintenance eneg reuirementae ofen calcuated but in tems of cel bioogae ool understood The main sinks for maintenance energ are thought to


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U RIIG 1

e the prerato o o gradet ad pote over (11 Wheoclmacterc rt, or clmacterc t that lack the matty to rpe, reepoed o ethylee he reprao ae cree We eee wh

raw, the reprato rate all here a mlar repoe o ethylee everal or e (1 learly h repratory re a repoe oethylee ad ot a demad or eergy wt whch o yheze rpepecc eyme oehele aocaed wh he creae repaohere a creae proe yhe ( 17 J Sper, rady, Lee, plhed), a lrge compoe o whch prote over hwhe ethylee tmlate reprao, oe o he wo ma o mateace eergyproe tovercreae It etrely pole that theeergy demad or the mateace o o grade alo creae th wold

occr ethylee perrae memrae co ad creae pave vty (ee Seco 3).

oe look oadly at reprato ogrowg pla te, t ot atal al o oee ha he reprao re reglae y rae pply79 h o, eve e ampy lled wth caohydrate reevePremaly, compartmeao wh cell crtcal o th reglato It polae ta he cyaderea pway operae whe eleco owexcee he accepoegae cyochrome pahway 79 a, patclar,whe e pply ecee e, may e that ethylee

teatmet creae trate pplye y a eec o glycoly, ooopla lxe, or oherwewh a rela creae reprao ratea chemcal eergy O he potaed mao k o maeaceeergy, e over may e the mot ele o ha a creae pe ye oered a a coeece o ethylee aco hee oher evece pa ha pote over relae o he reprato rate(10

A creae prote yhe early he clacec pero ( 1147 ha ee meare may t ecae he creae appear o eted to he reprato creae () altatve a oppote tepreatoare avalale he al eect o ehylee may e to creae reprato awth avalale eergy the rate o the tao reaco are orolle yeergy pply perhap va he eergy chage, a creae polyomewol elt O he ohe ha, a erly eec o ethylee eamet ocreae he pply o meeger NA avalale or ralao, he thedemad or eergy wold creae ad a re reprao rae wold ollowA choce ewee he oppote ot preetly pole We kow ether theATP yeld rom he repraory creae or the aolte rae o poeyhe volved learly he ecod aleatve demad a tgh oplg ohe eegy e poe yhe wh he eergy yeld rom reprato hert alteatve may how h too, t ot a prereqte o he moel




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8 BRD

In avocados Tucer & Laes 4 delneated wo regons of theclmactec perod tha are assocated wh changes n pren snthess. Thental pt of h ise-in hs as vokd by extal ethylnwas

assocad wh a hreefold ncras n olsomal A and po(A+) RArovees r un fresh weght and presumabl a hrfold ncres n proenuover. The second phase was assocaed wh a dcln n polsomerecover and wh he penngspecfc enze cellulase represenng annceasng propoton of the pren snhesed Subsequen sudes haeesablshed tha he snhe of cellulae s regulad a a tep precedngtanslaon Secon 2.3), bu he mechansms leadng o he nceases ntranslaton of he genec or "hoeeepng essengrs hav no bensough n rpe avocado fu, he enngspecfc messenge As accoun

for onl a sall pt of he oal about 4% for celulae) and ths eesalso to be the case n omaoes (55

Evdence fo an ncease and then a declne n poen snthes also coefom esmas of he ncoporaton of pecursors no pron n ssue slces5, 25 22 28) and from emate of h recover of polsomes fo ntactfus 43, ). Th evdence wh slcs s ve conssn n several specs,u here ae pblems of ntepretaon assocaed wh changes n endogenous dluton of eogenous pecursors, wh amno acd recclng, and whnteacon wth the esponse o slng Measung polsome content 34) avods hese comlcaons bu s subjc o he assumpon that recoveres re consant as he chems of h fus changes durng rpenng ToalNA analses o suppor th polsom stmas ndd n tomatoes,hee s an ncease n the proporton of rbosomes rcovrd as polsomswhen the clmacterc commnces 3), and an aparent dclne n rbosomerecover lat n penng The resuls sugges an ncrease n he rate of protensnthess durn h clmaterc rs followed b a dclne lar n rpenng;agan NA analss e reurd o consoldae he concluson

lage amount of crcumsanal evdnce lns he ehlnepovoed rsn espraton to an ncrease n pron uover Such a stuaon fts thedecrpon of hc sruggle for homeosass ofred b oman 25), but swell o ememe ha othe deenaton sgnals for eample phtocromeacvaon 2), provo a larg genrc ffec on proen snhs as wll asa specfc effect nvolvng some ranscrps Changes n cell stucue andfuncon ma b more easl accommodad f the genral ra of rplamnof macrmoecules s ncreased The wo phases detected n avocados 4)ma occur commonl, but the do no mpl an absolue seggaon n tme

of he generc and specfc responses n tomatoes thee are earl and lateepondng chnges n mssenger As 55), and h el hanges ma bewthn the frs "generc ncrease n proten snhess.




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4 Other Oxiative Reactions

FRT RPG 169

Eeio of pohe fe by oe i ow oceo o oxygencnot be eiy epine by eect o either the cyneetve or

rei reptoy phwy 135) foc tteton o othr otveevet with ower it for oe he nhbon ofpeng b ux(46, 10) n he peroive esuton o inoe-et 78 hvebeen pose s mpon eguos o peng bu unmbguous evece fvor of hi thei h bee ifct to obi Aempt hve bee me oree peoecyze recio o ethyee yhe 47 50 n hihcocetrtion of free ric cvener hve bee how o ihibit ehyeeprocio fri ie (2) howeve whehe o not ee cs hve irec roe in ehyee bioyhe remi obf 18) Eviece hsupeoxe cs ae nvove n ecos whose poucs peub membe fcton n owe h been metione prevoy (Secton 3; 42 90n my b vt o f ynpvoke epiroy cimcteric i the for te h obvo pe the cmcercf ehm 81) h propo h cyoknn which ry nisenesene agens unon s ee svenges, mpyng h freeric py criic roe n ic senescene Choops efferetitio choophy esco e p of he peing snome

my f Mtie 8 h ete th peoie or ipoyeeae voved n chloophyll cbolsm

THE SOFTENNG OF FRUS

he soeing pocess is ieg p o he pei of mo fris Ith immene commerci imporc be he pohrvet fe of he fi o age exen me by icre soness whch bi wth it ncree in phyc me r hni n incree eesusepb. o he physoogs n bohems he soe recons e

iific bece hey vove ce w che h hve o pre efseescece n h ppe o occu ony i he i pehps biozone. t o ccent tht ce w hyroyzi ezyme hve fetrepomety recet ie of the moecar biooy of ipei

1 Chemst and Enzmology of Softennghe chemic che vove n oftei the ezyme cotbt othe che hve bee eviewe rece 10 70 wi be briefy meioe hee nesi o he che h ocu r ripe

me bece nowee of he trcre of he w i mre fi ofthe ezyme ht mify he w vey me Hppy both ececere bein ree an more ee te erte

Annu.Rev.Plant.Physiol.1987.38:155178.


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16/24

70

Soening is omaly accompanied by an increase in the concentration ofsoluble pectic poysacchade 0, 0 The incease in soble onic acidresidues is oen corelated wth an incease in the polyuonde hydoyzng

enzymes esecaly endopolygaactronase [poy ,-D-gaacturonideglycanohydoase, EC3 2 1.15] In tomatoes thee s evdence o a decrease n te average size of the souble polyuonide chans late in pning(69; although hemiceluoses ae aso noaly degaded dung penngdegadation f the polyonides is nt dependent on changes n hemcellosecomponents (69 n tmates t thee s a easnable ceaton betweene amount of polygaactuonase, the ate o sofening, and pectn degadaton4 alhough nfomation he pectins emains spase hee is asoacorelation in avocados , , peas 2, and tomatoes 0 between eappearace of poygalaconase ctvity in penng fit and a loss ofectron density in he midde lamela region of cel was In a number ofothe t tee is a coeaton n me between the appeaance endopolygaacuonase actvty and the ntaton o sotenng , , , , 2.An ncease n pectn subty s nt adeqate evdence o endopoygaactuonase actvty, as ths may occ by the action of other enzymes o by achge in orgaic acids other cheatng agents o pH in the t ce was.n apples (0 and sawbeies , pectn solbty inceases bt endopoly

gaacuonase activity is not deteced but see 82 appesellulase actvity is amst awas fond n t, and the evel of actvtyoen nceases geaydng penng 0, 7. A pecse ole or celulase,ts cnbuton to stening, o even its subsate in the wals have yet to bedfned A numbe of other gycsidases occ in was and some lkexylaase in papaya was 0, ncease dng ipenng, whch uggests, butdoes not establish that they pa a oe in soening In tomatoes, thenorpening mants and 0 0 have been useful n dsngshingchanges n te was that ae asscated speccay wth penng from tose

hat occu n the absence o penng and may be cnsdeed a pa o thesenescence yndome

2 A e fo aiu

igh concenatios f cacm known to nhibit and smetes preventut penng and t ead to e t . Whle ccm ndbtedly hassuble effects on cel membanes n membane poteins 07, 8 and as asecond messenge , the hgh cncenatns cacu equed to deaypening o inhbit enescence sggest a goss efect pehaps in the wal

When one consides the potenta ctivity o the polygalactonae and pectinetyl estease in ft tisses, the attack on wal poyonide appeas to bevery limted and i tomatoes there s evidence that cacum limits wal



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FRT RPG 7

ydoy ( Clam tat aum ould or o rdutd n rpnng frut av n ad, ut t lam ar ad onnadquat tnqu and tr a nd o valuat alm dru-ton twn t vauol and t wall y ung ound aton and anayalmtod

53 Softning a a Rgulator of Rining

ltoug t unlky tat all yroly a an ntatng ol n trpnng of lmatr frt (4 gnotyp tat otn rapdy gnrayrpn rapdly and o frut appar to n rapdly T ntratonptwn wa and ntralular vnt av n ltt tudd Rpng otn n a a r of ndpndnt nt, prap oordnatd y ym 2

tyn ut otrw opratng n parall ordng o t vw, tpron of ym tyln rult, among otr tng, n nduonof t ynt of wallydrolyng nym, w n tu au t wal oon t n and po rom ytoplam o allydrolyngnzym ar t full tnt of t ntraton twn yoplam and walTr , ovr, t polty tat t ytoplam nund y a angn t mtry and t pyal tat t wal all ydrly appa todra t alty of t all to nd a rang of pron (67 o ononqun of onng may t la, for nraon t pa-

malma, o prtn rvouy old on t wa. ond polty l n t rla of arydrat olgomr fom t wall S ogomrmay funton a ltor 27 and ay tmulat tyln poduto adot tr pon t pol tat ty av an ntnfyng rl on attr ta o rpnn? nvaon y mrooranm on advan oratn rpnng tyln produton undoutdly ontru to t pro- ut an t of t prdut or onqun of wall rakdown avnot n ludd Cang n all propt ll modfy war rlaon nfrut tu and t w ontrut to out rdtruto tw u

ompartmnt

6. CNCLUDNG MK

Trougout mot tud of t pyology of frt rpnng rarr avn ampd and frutratd y t lk o organ Blk a mdt applalty of many of mtod ta lw dd tploraton of ntdary mtaolm Tr , of our, om u nknowng wt t potntal of a ll w nym wat pona

avtut n a ulky, nngrowng tu tund don to mannanonly" raton nym and t potntal not anr noug Tltatu rammd w po of ang n t amount o t or a




18/24

172 BRADY

nym a mauy a nnmn chand bu alway h wasquvoaton as to t mann o t an. Sustat onntatons andnzym nvonmnt w alway unnwn. T mau u a t bul

as ovom y usn tssu sls ut t s nonsns to tapolatmasumnts om a apdly pan sl to a ulky oan n "mantnanonly a.

Rcn ya ha n m lfn f h fan md by tssuulk. Indd t at tat t ulky tssus a nally at omonous nll typ may pov an advanta n t us f hycal analycal mhdsu as NM and X-ay pos It s antpatd tat dtald and ousdanalyss wll yld fm nomaton on aspts su as vauola ytoplasmand spa pH and on onntatons and ans tat ou ov tm n

maj mab Smlaly cafully ald lym analy wll yldnfman n n syns n h nac ulky u an t w uo mmunoloal and nulotd pos wll sow ans n t syntssand loaton o patula maomoluls nt v on pnn nts ss wll av "susts o "s onsstnt to "mpls andmo postv statmnts tan s psntly bl.

Th bh an un and dffcul m w f nn.On bcau w and n t d o apd poss and t s usul toontmplat dtons dult aus poss s now so apd tat sums

wll b an by knwld a h w ua n Th m na wn t num o ns o avoado llulas and tomato polyalatuonas ll nown n t pomot squns o pnnlatdns wll vald and wn t onsquns o avn mo o w n nn f h n can b ad. Th m may b na wnt m"sm " and ym " tylnmakaly usul tms o adad and a halfcan b lacd by a dcn n mlcla m fat unls t onpt. o an an tat pnn ntaand dvlopmntally ulatd Soon w may kno a lttl o at tos

wods mply

tt t

Ali, Z M . , Brady, C. J. 1 982. Prfiation and chaatesation o the poygatrnaes of tomato frts. Aust Plan Physio. :778

. Apeba, A Wang, S Brgoon, A. Bake ., Lieeman, M 8 1 Ihion the onveion ACC tethyene y stct anogs inbos

of eecto tnsfe, uncoupes o oxidive phosphoylton n ee raicvengers Pan Physo. 677479

3. A S Mzh Y. 983. Sess i-

ded ipenng of the nonrpening tomato, Lyopron untum mtannor Physo. Pa

4 Awad, M, Young, 7.Posthaest varaton n ease, poygaatronase and petin methy esteasein avoad (Pra amana Mi. vFuere) fuits in elaio to resrton

nd ehyene dution. Pan Physo6430&-8

5. Bke E Anderson D, Hsck W. 8. Poein syness in




19/24

mao frui percap issue drng ripening. Chracersics of amno ac in-orporaion Plant Phyiol 120:1679

6 n R A . -amenain f amino acis in tomafrt pcar tiue. See Ref 52 , pp.3 1-2

7 Bge 1984. ngs in snstv-ity fo ehyene ing strage f append banana fuis uner hypobrc con-ions. Sc Hrtic 24 1 5 164

8 Brber R. hompson J E. 19Ntl lipis giify nsarated acylans in senscing memanes J. Exp.Bot. 4266

9 Barley I M 194 Lpd mtabosm

n pening apps. hytochemt85791 0 Baley . M. Knee M 1982 The

hemstry of textural changes n ftung sorage Food Chm 945

Bathgae, B., Pn, M., esonD Gooengh, P W 195 Pladanges drng he conveson o chlolt to chromoplt n rpenng tomoes. anta 1619-20

1 2 BenAie R Ksle, N Fenkel C99 Uascual changes n the ceals o pening apple and pea fuit

Pl l 221 Benne, A B Chroffersen, R 96 Synthei a d roceing of ce-ae fom pning aad fri. Planthyol 1:80

14 Biae J. B Yong R. E. 1 9 1 Tevcao par n The ochemt ofFrut and Their Producs, ed. A Clme, 216 Lonon: Academ788 p.

1 iae, J B , Yng, R 98 Rspton n penng n fuisretrospectn prospect. See Ref. 5 1 , pp 1-9

16Biggs, M S., Tieman, D M ., andaA K 1985 Rpening assciae genepesion n omato fit pecap HortScence 205 (Abst.)

ggs M. Haman, R W, HanaA K. 1986 Canges in genc expressonng ma u penng Plant hysol 8 1 :9540

1 8 Blacma . aja P 1928 An-ytca studies n pant respiraton Ih e repiraon of a popton of eneent rpening apple. Prc Soc Lonon Ser 104125

nsip . M , csn D G98 Develpment ethyne isynhe n ehyenenuc rpenng nlt 'anou (Pyru cm-un) rng the col eqiement foening J. A. So. Ho. S. 1 1 020-2

FRUIT RIPENING 3

20 Boochv, A alevy A. ., Shtzk, M. 1982 Senecence and he fu-y f ose peta membranes relatonshpo phospholip meabolism lant hy

l 99992 1 Brady, J 19 Changes accmpnying goth and enecence and ffec ofpygca tress. n Chem anohet f Hebage, W.Buler R W Baiey 2315. ndn Academc 455 pp

22 Brady, C J. , MacApne G McGlson W B., Ueda Y 982 Poygac-nas in tmat fus and the nc-in pening Aut. J. ant yol 911-

2 Brady, C J . , McGaon W. B. Pear

sn J. A. Medm S. K. Kpe-vich E. 985 neacins betwenhe amount and molecuar frms f py-galactuonae, calcum an frmnes intomato frt m Sc. Hr Sc1 1025458

24 Brady, Medm, S. K, McGlas-son W. B. A Z M. 19 Diffeen-tl accumuton of the ocuar foof plygalactuonase in tomato muants. o ohm. 77-14

25 Brady Oonne, P. B H 976On the sgnificance of ncreased pten

ss in s Plant Phyl 01-1026 By C. OCnn, P. B

Smydu, J . , Wade N L 970 Per-meabliy sugar accmuaton andrespatin ate n ipenng banana ts.A . c. 2 1 1450

Brecht J K Kader A A 194Ethylene pduction by t smeslpening nctn runu peragenypes m. Soc. Hort Sc10966

2 Brecht, J. K Kader A. A. 194 Regu

laon of ehylee podcon by ipningnecarine fui Prunu pera as nenced by ehylene and ow empera-ue m oc Hoti Si 969-7

29 Brcht J. K Kaer A A Ramming W 194 escpton an posha-ve physoogy of some sowrpenngnectaine genoypes J Am. oc HortcSc 109596600

0 Brennan, T Frenel C. 9 Involemen of hydogen peoide n the reguation of senescence in pears. lant hy-0 941 1-1

1 Besche R W, Hbsn, E 9Role of cacum n cheatng nregulang the egton of tomtofrt tissue by poygalactonase Joo ohe. 61460

2 Ber 194 Ehyleneenhanced 1

Annu.Rev.Plant.Physiol.1987.38:155178.D

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5. Bug S P Bg E. A. 967. Moeua equiemens or the ioogcal actvy o tyle. ln Phyl 5

36 Cld R. D . Wllams A A. HoadG V Banes C R. 198. The eecso amnoethoyvnygycne on mar poss hg Cox'sOang Pppi Appls J A 3577381

37. hisoersen R E Tucke M. L . ,Lais G. G. 198 Cllas g pssio ipeg avoado, ere Hss ; heaccumuaon o messenger NA anpoi as demonsaed y complemeny NA hybrzato mmnodio ln Ml B 338592

8 hrsoersen R E W E LaiesG G 1982 G expession durng rpng n avocao. lan 555257

39 Coome B G 1976 Th dvopmto shy Ann Rv ln hl27:20728

0 roo, P R. Grrso D 98. Ulasc o omao it png dhe ole o polygalacuonase soezymes n wal graao lnhl 72:0889

Delaenn D . Alexnde D. G , Ben A. B 1986 Mour og omao u polygaacuronase: analysso poygacuonase mRNA evels d-g nng. . Nl. A. .UA 83604

4 Dhidsa R J hdsa P P ho A. 1 9 8 1 ssn: orlawih inceased eves o memranepeealiy and lpd peoidationnd dasd evels o spro smtas an caalas J p B 393101

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5 Feguson I. B 98. Calcm n plntsenescence and pening. ln llEnvrn 77789

6 Frk C Dyk, R 3 Ai nhion o png in Barlt ln h/ 5 1 :69

7 Frenke C Esn M 977 Ehylenevoto as rat to hngs i hyopos n rpnng tomato itsrn 255253

8. renke . Gaison S. A. 976 niaion of yopn ynhei th tomaomnt rn as nnd by oygn nd

hyl naos en 1 09. reke . Klen, Diley D. R

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50. enkel Mua M 98. Possi ol o t l all oidaive acvy n hyne voion S R 52

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2 hs Y Chautz E eds. 98.hyl hl Phyllnd Apple Ape The Hague Matiu NjhDr. W u 8 pp .

5. alrd T. 96 8. Asecs o lipidmtaosm in hghr pans Thdentcation and uantiave anaysso lipds om he pup o p d postmateic aes. hhems 71915

5 Goodogh P W Posse M Yong K 19 85 NADP-lk maienzyme nd mat mabolsm in age

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