the development of fresly fruits

8/9/2019 The Development of Fresly Fruits

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Ann Rv Plant Physiol. 777

THE DEVELOPMENT

OF FSY FRUIS1

B . Coombpartmt o at hyoy Wa Actra arc ttth rty of a Omo Sot Atraa 504

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INTRODUCTON ............................ ... 08

FRUT FLESH ............................ 08

Denitins ................... 08Orgns of Frut Fesh ............... 208"Functions o Frut Fesh ..... ................ 209

DVOPMNTA PARS OF FLSY FRUTS .................. 20Methods and Interpretations ............... 20Dvlopmal Pas.............................. 22Copnents of Grwth ......... .................. ....... ...... ... ....................................... 2

C number 2Cl volum .... . . 24Cl d ......... 214

rut Ripenig ..... . . 1

COMOSIION O RI ESH .................... 2CMTON OF WTR ND SOS .............

Suppy of Slutes; Pathways .... 2

Pors o Vauos

.....

.

.

220Vauols as Lysosoms ......... ........... ..................... ..... ....... ............ 22Sugar Accumuation b Sugarcane Intedes .... 222W roth nd Wtr Accumutin: Sning 2

CONTRO MCANISMS.......................... 223The Triggerig oipening ........ 224

Etn . 22ABA 5fAA oxdon products ............. 225

CONCSIOS .. .

'rowth a ar aia a oow a a); thpho 2 horo

thy phopoi a A io·at a 24T 24 trihorophoyatai)

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208 COOMBE

NTRODUCTON

The numbers o reserh ers on bs quesons onernng he grow nddeeomen o rus re meer n omrson wh hose on oher ses o ngrowh. The ed s sowmong nd duse nd s e wonder h dug he27 yers o hs journ he subje hs been reewed ony we, he rs n 195 by P. Nsh (83) nd he seond, onernn he se ses o growhreuors, by rne n 19 (2) Physoogss re nury reun o work on sube or whh he mer s he resu o on nd ome rowh roessnd oen s ereme rbe. n he oher hnd mnuon o hresed rus omrey sme nd onsderb more reserh hs been done on oshr-es hsoo n he sube hs been reewed our mes n hs ourn one

(7, 9 1 118).A mesone n he erure on ru rowh s he wooume work on he bohemsr o rus, eded b ume nd ubshed n 197 nd 197 (3).sey reen re he wo hers on ru hysoogy wen by Bord (8)nd Nsh (8). Ths ubon remoes n omuson I mh he hd ooer he ed brody Rher w od ny emhss o seed grow bsssonnd rowh reuors nd nsed w onenre on esh deeomen n emere nd ro rus edn o some ommens bou onro mehnsms

FRU FESH

Denitons

A ru s he rodu o deermne rowh rom n nosermous oer ornoesene Broos dened rus s mured res wh or whou essorysruures nd/or seeds (12). Nsh reognzed h he bons's denon ddno wys mee he ymn's one o ru nd emhsed h he unynor ws hsoo nd he suesed, ws ssoed wh he hnes our-rng n he oues. He hereore dened rus s "he ssues whh sur heoues nd whose deeomen s deende on

he eens ourn n hese oues

(82). Ths denon hs mn dees (118). The word ru dees rom he

Ln"ruuso enjoy, rodue The ord ngsh Donr denes ru s"he edbe rod o n or ree, onssn o he seed nd s eneoeesey he er when uy nd uy I one nerres eneoe n he brodsense nd does no onne o he err he denon sus hs eew The mor use reers hereore o eshy edbe rus. h dry rus suh snus, grns, oes, nd henes, s he seed whh s he morn rodu. Themor rus o he word [re bnn omo rus, uurbs ome sonerus, mngo (3)] e eshy nd uy n emhss on he sudy o he hysoogo ru esh.

Orgn of Frut Fleh The soundn dersy o ssues whh n beome eshy hs been reerred o b oher uhors (8, 8, 8). As Fre usres, ry rs o he o

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DEVELOPMENT OF FESHY RUIS 0

PORNSDlS

Ouines of sectios of the uits o species showing the diversity o tissues whichan do ito fuit h. [Som dawigs opid om lrih (117)]

nsn s n sm sps dvpd n s Is n pss s ny dns n dvlpmnl v n ms dns n mpll n mmn s sssm w nd mny n mpnds s ldn nss ndvnss y lky pnyms sss qnly spl yndl nd n s nsns y d lys

Fs nd l ls nlss lls lnn ls s dnd s n dvsn pnsn nl ss (plp) ms y nnsmsm vy nss on mo ll ss, .pln (m) spm (nn p) nd nddm s (nn n) Tss n ls vy y l pnsn nd vs n n w nn s (nss s nly ss m sy s

'unctions of Fruit Flesh

s ( s) y ss my s d ds nd s n d my dvd nd mnd sds dssmnd 8) wn s pnn v dvn w s nd n sps y vn s sdvn nsps nd v nms n dn dsps ( ).

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0 CMBE

n ts ntrtton th rn o strkng oors (rds urs nd yows)n ut sn nd o swt nd juy sh s not sursng. h shy tssu my so

rott th sd nd nun ts nronmnt (gs, wtr, hms) but dosnot ontrbut to th nourshmnt o th grmntng mbryo s do th sd tssusFruts ormd n mortnt sour o ood or mn whn h ws hunt ndgthrr nd t s undrstnd tht rut nts wr domsttd nd modh to o gtt rogto dd ths ross.

DVOPMNTA PATTNS O SY UTS

Method and Interpretations

h onom mortn o ruts nd th dstntnss o thr domntrosss h d to mny growth studs. Unortunty th morty o thsstuds r dut or yss o h omonts o growth S msurmnts on r o mtd us d msurs o ontrtos o drnt omounds t nothng bout umuton r rut unss r dt on rut s(rrby rsh nd dry wghts) r rodd I ths msurs r obtnd onth sh rt d th numrs o sh s r nown thn on n ntrtth dt n trms o th rt nd durton o th umuton o both wtr ndomonnts o dry mtr r .

h usu thnqu or gthrng suh dt s to t sms o numbr o

ruts t ntrs durng growth Sm s nuns h ury o sssmnto th outon but dos not rmt ssssmnt o th dgr snhon

twn nddus n hr domt hr r thr mthods whh rduor od ths rom () to st outon whos domnt s synhronous( mthod tht rqurs n obt nondstrut msur o unomty) b tos ndtors o domt nd to y dt y rr to dd ts(orn nyss s usu) to us nondstrut mthods o msurmnto nddu ruts n stu th rgrd to smutnous nd nondstrutmsurmnts n md o s s h s oo, do, dothr rtrs Voum n b stmd rom nr dmnsons or msurddrty by ud dsmnt th us o quds suh s wtr rsnts duts

but th thnqu usng n nrt gs k hum s hs bn usd or th msur-mnt o th oum o who nms (0) wrrnts nstgton th nowdgo domnt gnd by ths tys o msurmnts th usunss o smrgs n b grty mrod

Asd rom th robms o msurmnt, th robms o ntrrtton rqurommnt Som o thm n strtd by rrn to th doub sgmod uro, sy oum gnst tm so runty shown by drus nd brrs (Fgur It hs bn ustomry to dsgnt th rst nd sond rd growth hss s

tgs I nd III nd th ntrnng sow growth hs s tg II 80 ortunty th oudrs tw ths stgs r dtrmnd rtrry th s nodton o thr rs oston on t umut ur nor on th doub-humd rt ur A mthod or dtrmng th oston o nds urs hs

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DVLOPMNT OF LSHY RUITS

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A doub sigmd grwth rve f vlume versus me expessed n cumuatve andate bases. ay methds have been used t desrbe his ye f urve: () the assameh begn n 194 by nns 20) (b) esignan f a remnary hase he dvs the lag hase n w ars ; (d) he alan f hases angt eaks an rghs n he grwh ae 7).

n dd mmm lood mod o Qnd (9), n dly v n l on .

n om wy non ym n dopd p om y n lol dl om onono n md y dnn ddonl 0 nd IIA nd IB ( F ) Blly wo onv mod vn npon pod o lon, pon o mo pd ow , podo don nd n nd T pop n dmnd om v(F ) o om lol mm nd lol mnm n dvv o modv T low ow p ( II) on nl dlon o y nd nnn on o ond yl w non mn

npon Two mod old d o qny ln: mlvv moo on old ll nvl wn o mm n ond dvv v v dn nd mn n d-mnd n Qnd mod d o ov.



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6/22

OOMBE

oter dicuty i iterpretatio is te actua reatiosip used Quite dieretiterpretatios ca emerge if data ae cacuated o dieret asesper uit dryweigt fres weigt protei voume of water per ce etcor if rates are made

reative to a previous referece eg reative growt rates 7) or te data trasformedi oter ways.

Developmental Pattes

e review y oard 8) provides a good accout of te ature of fruit growtcurves. Some of tose avig a sige sigmoid curve are appepear date pieappeaaa avocado strawerry orage tomato ad meo ose avig a douesigmoid are a stoe fruits g accurrat rasperry ueerry grape a oivee growt curve of A fruit is descried as tripe sigmoid a) ordoue sigmoid fruits iterestig data ave ee compied o te duratio ad rateof growt fo te tree stages i fruit of dieret spcies 8). Witi a species tefruit of utivars wic ripe eary ave a sort sowgrowt pase rom a ompari-so of tree grape cutivars a growt pases occupied 8 to 8 days except Stage of a earyripeig cutivar wic was oy 5 days i egt ad Stage ofa ateripeig cutivar wic occuped 5 days 79). rom comparisos of growtcurves o te erres o cv. uscat o exadra te rst growt cyce was oudto e reasoay costat ut te sowgrowt pase vaed from 8 to 48 daysdepedig o te eviromet ad weter te iorescece was primary or sec

odary te autor upuised).Some iterpretatios of growt curves ca e queried e growt data egtad diameter) for te itci are said to prtray a sige sigmoid curve (0 ut ivoumes are cacuate ad seed voume is sutracted a doue sigmoid curve isfoud e curve of te cassica sige sigmoid fruit te appe sows a distictaerratio at a eary stage accordig to oe set of measuremets ere is oreaso wy simiar discrepacies soud ot occur commoy or istace y daiymeasuremet of grape eries i situ we ave demostrated tat te seco rapidgrowt pase egis wit a sigiat tasiet oe days) sput of gowtCoome ad Bisopupuised)tus cormig a earier suggestio 0).

We measuremets are precise growt curves of orgas are frequety erratic adare dicut to descrie i simpe matematica forms.

Components of Growth

e iterva from atesis to ripeess of fruit varies for dieret species rom wees strawerry) to 0 wees Vaecia orage) ut i may fruits it ocupesaout 5 wees 8). Durig ts iterva fruits icrease i voume ad weigt severatousad times. e of te arger creases is sow y avocado (300 times)ad oe of te smaer is tat of dwar squas (X 40 or orit grape (X 300 (3

4 0. e argest of fruts pump ad watermeo are ow to attaivoumes i excess of 30 iters per fruit ut do ot ave a uduy arge expasiofactor i fact o greater ta most grapes X 4 ecause of ter reativey argeovares at atesis 07).



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EEE F FE F

w p dnd b l nb ll vl,nd ll dny nb nd vl ll pn nnd b nb nd vl n nd nd dn l dvn nd ll

pn

MBR n n l ll ll dvn (3 b eea-e e e ecoe a p eeve o pooyae n b eieaic ad go coece, a i e ui a e n w eve ae n iiiaed geeay acie ( 4 icvw provd d accou o e ipoa eve akig pl n wnd l deveope ldn o e e ui 6

v dvn n l ld n nl pd w wk

n (. wv, nbl pn n dvn n pp n p n (3 n n b nd n b ( n nd, dvn nn n vd l o e ui(0) Te mos omrehs ny lv l ll dvn nd llnln n w nd dvlpn b b Snn(06 07 H pd vl b pp nd wd pd w mld d ld ppkn d ln pd ll dvn pll n e oue ue n dn dceaed ei ce epaded oe

T w iio ll i e e o a appe a aei ad oy iio

a ave ( T v nb dbln d b nb oy 4. ubig ae iiay, i p ee a 0 iio ll i ev n nd 06 lln 4 day ae (4 e ue o douig ocv b nd n ( ee oud appea o eape oppouiy vn n nb b n. ndd l ( wd lwn p b v Sln d n l dwn b b n nd 40 d l dvnd ppd l dn w d nl pn dvn. Unn ll wk p nd n p d

n n l dvn nl nb vn coeaio eee ui l ue ad ce voue ipe apicoui o wn idividua a cocuded a ce ue w pn coiuig o vaiaio i ui ie. We e opao weeded o eee , e b pnn, ce nb nd l nbd vaian ui o eay oe w l pn due aiy a age nb ll w ndd a dn w due ovn n nb ll n (

n n dvn n dn p , b l dn dvn (plnl nlnl nd nd nn b

w n l pnn nn l l p l , nd ppn nd . n pnl dvn n n l l w dvlp n ndn pn d 128; lwll n nd npn nnl dvn pdn n

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8/22

COOMBE

edererry pera grape ( Dvso appears o our ay dreos raouar ue ( 0 oud e empased a e es of fru ssuesoer a e es ae er ow de mg ad dreo of dvo.

LL V We s agreed a e mer a mpora faor frdeeopme ad oe o varao fr se ere s e do a fordvdua frs of mo pee e rease e oume maes fa e greaesoruo o e oa expaso o e ru. e proporo of e oa eae e of a grape err aer aess a a e arued o e ree ompoeof grow are as foows e mer fod; e desy fod e ume0fod or more. Some of e arges es o e foud pas our e esof rpe frs e egs ewee 0 ad 7 Jm are ommo ad

ome aes exeed mm ose of e e urroudg pomegraae seed ofeexeed mm eg . Voues rage p o 08 Jm ad gures of 06 o07 m are sa e fa a e es of ee sses ae a vome of o ao1 Jm a aes ae e ge amo of expao aeed.

maure fru ere s frequey a gade rada dameer of es dereas-g from e eer owards e osde eg pp 0. e amegdn n n n e mesoarp of apro u ere was o su red e voume was ar at e tere was o grade e voe egrade wa oe of e sape ad a s wod e expeed a e opeo dvo es are approxmaey sodame ad f e a expad o a equa-

e vl wn cnn e ey woud ave o asume sapesn 6

an ann cll n nnc y e wa eaor( (paczn pn wal aal, ncln degee of seodary wavln, r (war an rnc n c pese sde ad cll, an cnran n y e exesy of es r r rrnn lar ac cnn a b nncparal b facr such s growh rgulors nd h nvonmn

la (ac oi a bn n ccr n fru es e (0

0 ad may we e cn Bral a rawn ann o rcrlan ewee nclar vl an cll vl ( r l consde rab ad reases w a Bral rae were ae w iapo a dug age es e were n aveage pod we oe weeefore rpeess e aveage wa pod p o pod was meased . ra-me of apros w e ax 5 a e ed of age eased pody dgage y 30 pere ad a pees y 90 pere; fru es ad frs wereommeuraey arger (I ese auor suggesed a e rae rease pody preeded e rae of rease e se ad a ormoe grades may evoved s oro

ru es amuaes arge amou of varous orga ompouds ad er oerao rease w me ear owever a erease oerao ao ao for e arge amou of dry maeramuaed per fru s s due more o e arge reae w or e



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DEVEOPENT OF FLESHY FRUITS

v (wt) th pvdn th t n whch sts cn cct hsthss s dscssd t

Pcnt d wht whch ps c dnst chns dn th ntn

t In th pcp p t s hh t nthss d t hh cncnttn ttt () nd pss s cs th cs hh ptpsc hpcn dcns s h ps dvp chn n t t th t stst wth n th st cc Th th vs s st d t cctn t nd tt thn cs nd ts ( ) nts dsp s td t Gd pch s cp cs thccc cvs sh wht nd d wht t phs nd wt nptpcds d tt npt n ch wth cc (17).

It h b cd p t b t c d wht wth t

shws sn sd cv vn n ts whch hv d sd vcvs (), t ths s dsptd Th cvs d wht p t p ( )nd pch ( dstnct ccc thh th dnsns th scnd cc h reter th those o the rst

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In s c dvsn nthss nd c pnsn nthss wdpp th j dnns h ncs n sh wht sh ts.ddtn t s cnttns st c dvsn t nthss nd nncs n th cncnttn sts. h ncs n d wht p t sdtnd th s cts

Fui Rpenng

pt wk n th cts wth s pn s h s ntsvdd spct phs. Ths hs n d t th nd t pv psthvst hndn t nd ds t n n ndstndn th chssnd cnt sptn whch ts n ntstn pnt sctcs th phnn th spt cctc I, , , Th wkhs cnt tndd twds th nn n ndstnd th dv-pnt phs ts.

Rpnss s th st whn th t s st tn nd pnn s th cp vnts cntn n pnss n th t sh ts pnn ncds

snn cn nd swtnn s wth n ssctd dcn n cdt ndstnnc nd n ncs n cpnds h n th vntswhch shd ncdd sch s n ncs n th t sptn nd thnpdctn ss chph nd cntnd pnsn cs Tn s p cs dncs twn spcs n th n nd tn thsvnt Thh st wd tht ths pt t dvpnt cd cssd s sscn s p t cn th s ths wd t thvpnss st (43) Th wd "ttn s t ppd t th vntscntn n th t (. p) t Th dct h s tht n cn pp

"ttn t th tt pts s wth ccs ncdn tht th tdt pcdn th pnn phs whs th wd pnn s dstnctv ts.

Rhds hs sstd tht th dnt th wd "cctc wdndt ncd th ncptn pnn chns (8 Ths s n kpn wth Kdd

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10/22

216 OOMB

ad Wess oga usage o descre crca pase e oogey o a ru. s sese e suggeso as mer Uoruaey e cocep o ocmacec as deveoped deog rus c do o dspay suc eaures as e arp

rse resprao rae. odes argues a e socaed ocmacerc rus areeamed more careuy ey may e oud o possess e caracerscs o cmacerc rus oug o dere me scaes. s posse a s opo may eog ad ece pee o use e epessos "egg o rpeg "rpegpase ad "rpeess or "rpe.

e rpeg pase o deveopme does o sar u e ru as deveopedcosderay rom e ssues prese a aess. s dcu o smuae rus odeveop rpeg paers eore a cera sae o readess; some pas o epeg process ave ee smuaed eary eg y eyee or eeo reame

u e resug produc s o paaae ). peg oud seem o e a gee-cay programmed pase o deveopme vovg a array o eyme avydere om a o e pecedg pase s rggerg s dscussed aer

e rpeg pase s sad o e dsc rus c ave a cmacerc rse resprao eg. appes aaas pears meos ad avocados ad dsc ocmacerc rus suc as cru grape sraerry ad peappe (4 eceo o rpeg e grape erry s oever que a dsc eve s caedveraso ad mars e egg o ugar accumuao oss o acd soe scoorg ad reeed ce epaso 23 ders rom e rpeg o apes orpeaces commecg earer e ru's deveopme

e mg o e pocesses eadg o rpeess e peac ave e reemedrecey ad some eresg resus ( , 5). Durg e perod e drymaer accumuao s sma eyee produco y e ru s o oug emesocarp ssue s resposve o eogeous eyee. e severa processes com-mece ogeer e mesocarp ces eg o accumuae dry maer more rpdyeyee produco creases ad e es sars o deveop yeo coor commec-g e ssue adace o e edocarp. e creased rae o aer accumuaoce epaso c s usuay regarded as cocde dry maer reasecommeces some ees eaer Gode uee pac ad s appare uderseparae coro.

oud e srucve o epoe urer e ssue orgs o pegecause s ud ad e uraveg o mecasms appears a oraso e peac e grape may rpe rom e ousde ards a accumuaoryprocesses are more ese e s a e es a e egg o rpeg23 e rpeg o daes as dcaed y sucose eves proceeds moe apdya e dsa ed o e ru 99 Hoever dspe ese dereces me ad paceo epo os ces o e es udergo e sae proesses a egaedaer e ces o coguous ssu suc as e pedc do o

CMPITI F FRUIT FLEHe dscve eaures o ru es are a epads ad ecomes ucy seead coored Waer corues y ar e greaes eg o e u o e es,



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11/22

DVOPMNT O HY FRUT

but lrge number of other compounds also ccumulte mny of which re wtersolble The list includes sugrs especially glucose fructose sucrose) cidsmlte trtrte, oxlte, citrte, succinte protons nd other ios especilly

K+, CI-, S04=, phosphte C2+, Mg2) lcohols including inositol, esters, vonoids nd glycosides including my pigmets, phenolics, mino cids, mides,proteins, vitmins, lkloids, terpees, lipids, nd my others. certi cells relrge bodies consisting of resins, tnins, or cystls 4 Crbohydrte polymers,eg strch, predomite i some fruits but re oen hydrolyzed by the time the fruitis ripe

The mgitude of the cocetrtions is illstted by the following verges ofthe most common compods in gpe juice, expessed on mol bsis: glucose0 M, fucose 0 M, potssium 00 M, trtte 00 M, mlte 002 M,

Mg2 0 M phosphte 0 M, C2, iositol, prolie, d rginie 0 M,with totl molrity of 2 There is lttle doubt tht wter d sugrs rethe domint compouds i most fruits This ct is illustrted i Figure 3 showingthe wter cotet nd sugr cocetrtion, o res weight bsis, o the mtureedble esh o dierent fruit species The digonl represents pure sugr solutionsscled on percent wter bsis The verticl distnce from this line to ny prticulr

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12/22

COOB

fri inicaes is proporion of nonsar solis. can be seen a waer an sarpreoinae excep in e socalle veeablefris an in acclaors of lipiavocao olive aci leon an sarc rian.

e ajor fris of coerce e. rape banana orane apple ave fro o0 percen sar Grapes can aain c ier concenraions. ave easrelevels of ore an percen on a fres wei basis (0% of ry wei n esall beries of cv. orin represenin a exose olariy of 6 e qesion arisesweer ese i sar concenraons are aaine by acclaion of e oleor by loss of e solven waer. e aa of Ry for aes sow a e iniialfris o inee lose waer ( evereless e fris are a e sae ie aclain scrose on a per fri basis an s bo acclaion an eyraionconrbe o e nal concenraion is icl o apporion e relaive conrib

ion ae by ese wo processs b before e nal pase of "ree rpeninconcenraions of 37 percen sar are fon eivalen o 0. srose an 04 exose eyraion probably plas a lare pa in e nal rpenin lean oconcenraions of over 60 percen (2 e i concenraions of soles in pecarob es sar 4 percen fr w an aarin plp sar percen arrae percen fr w are probably conribe o by eyraion as well as acclaion(2 11 n eneral wen fris are overrie an cells of e es are osinriiy i is coonly el a increases in concenraion are solely e oeyraion. However ave fon a e wei of sar per fri i.e. per cellconines o increase in a rape bery rn a leas e rs wee of is pase

Desie e aboveenione excepions i sees ceain a in os fris econcenraions of soles are increasin wile e es cells are expanin iseans a bo waer an soles are acclain a e sae ie b e raeof sole acclaion is reaer. e larer e vole of a fri e lare is isry wei

Acclaion of lare oleclar wei copons sc as sarc an soraeproein is no a proinen feare of e es of any fri species is conrasswi e siain in sees r an evinary viewin is prbably reecseir relaive role e see in norisin e seelin an e es in aiin see

ispersal. ro a broa opoeneic viewpoin e ierence probably reecseir relaive opoloy e see bein soon consraine by ri isses oroerwise inibie wile e fri es is oen exeal an relaively ninibie.n ose fris werein e ry aer acclae as lile osoic eec e. oilroples in olive an avocao e are es cells are sall (6, 0.

e noion a pysical resricion inences e evelopen of ese oransis in soe respecs analoos o e esis of Willias a consrains wiin esoo aex inence e row rae an fo of leaves (125

ACCUMULATON O WATER AND SOLUTES

Waer an soles acclae ainly in e vacoles of es cells. o eyacclae oeer ere is lile experienal wor on e connecion if any

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DEVELOPEN OF FLEHY FRU 29

etween the rocee of ce exnion n ote ccmtion n 1960 (21 otte tht the exnion of eicr ce of gre ring the econ growthcyce w e to n omotic ttrction of wter foowing the ccmtion of g

i their vcoe here re two ning which negte thi hyothei Firt theri incree in growth rte t the eginning of thi cyce i not mtche y imir incree in gr ccmtion (3 n thi the gre reeme the ech(17 Secony, when gre errie were tte with tic encore to reventtheir exnion, they fie to ccmte gr; on remov of the encoreoth gr concentrtion n frit vome incree (2 hi re gget therevere hyothei, tht ote ccmtion een on exnion owever it inot eenti tht the two event e cy rete they imy my occr in theetie t ot the me time

Sppy o Sos; Pahwayshe min y of the orgnic comon which re tore in the eh notey from recent hotoynthei t i oie tht in rge tree reerve cocontrite Rei ieeki (97 ggete thi w ikey with ricot eceorito, the chief crohyrte trnocte n hoem, w ony metoie withicty y rienng ricot eh, n tht the croe which ccmte in thefritro originte from reerve in the tree hi wo eem nkey to ethe oe ore when one ccte the retve qntitie ccoring to their tricot frit ccmte 1 g croe er frit ring the 2 y efore riene

(97, n for wecroe tree thi wo rereent n int into frit of morethn one kg croe er y he reerve of n ricot tree re not ecrie, ty nogy with the ech tree (16 they re ch to e nikey to meet thi ortof emn for more thn 2 or y o exten thi nogy, the rte of hotoynthei in ech eve incree to high eve t the time when ccmtion in thefrit i gretet (15 hi i not to y tht reerve cnnot n o not contritegrevine wich were efoite efore owering cceee in mtring mcro (126

ing n horty er nthei the ovry h oor iity to ttrct ntrient

tht i, it i wek ink(4

Bece of thi the y of ntrient to the ovryt the time of initi etting er to een on the over tt of ntrient inthe nt hi henomenon i itrte y the etting of gre (22 2 n ofrthenocrc e (4). nce the frit h commence growth it eveo niity to ttrct ntrient, n ierent imittion to ccmtion then oerte(2 45 he moiing iity of hormone yntheie y the ee n frt heen referre to (27 85

hoem i the mot ikey trnort th to the frit n ce w the th toeh vcoe he oiiity of rge cncentrtion grient of gr h eenregre roem for which mechnim mt ccont t recent work

ggeting tht gr my occr in the ce w otion of torge tie (40 496 co men tht gr concentrtion ong the th from eve to fritvcoe how mer cttion thn w toght t h o een ggete tht

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0 COOM

ug mov fom h choop o h phom in h f vin vi c w(7), hough h ympic ou h om uppo (14).

Concpu ony wo bi nd b picud in h phwy of ug fom

v o fu h h h phom in h v (o oh ouc nd hcond h mmbn uoundng h vcuo of h h c. Thu nomphwy migh b phooynhi in choop, diuion of ug o h phomin h f civ oding ino h phom movmn ong phom down concnion gdin o vcu bund in h h, diuion hough ou pcof h c, nd ny, civ "oding ino h c vcuo.

nn' nyi of h npo ino nd ccumuion of cbohyd bywh ndopm ( pomi o povid ufu mod fo h imi pocn fui c bing in mind h ccumuion no ino ch gn u

imp ug in h ouion wihin xpnding vcuo. h wo oh nof ch which imping upon h pobm of ou ccumuion by fui hvcuo: udi of h popi of vcuo nd onop nd h yoomhypoh, nd mchnm of ug ccumuon n ugcn nnod pnchym.

Properties 0/ Vacuoles

Rfnc o vcuo i conpicuouy cing in h iu on h ucu ofpn c On cn book by mnion hm nd hwod onop ndvcuo no id in h indx (46) Th of i i dpob bu

undndb in viw of hi k of inn ucu nd h dicuy ofpimnng wih hm. uch of ou undnding of hi dynmic popih in fom udi of ion u, pciy in oo c.

I i d h vcuo povid og pc fo ou Fuh, bcu ofh pnc of ou nd w nd h pmbiiy popi of h pmmm ndo onop, vcuo h ouc of pu cing gin hc w ding o ugdy nd u ngh (114) Th vcuo connincud w, woub ion nd mocu, nd vou inoub incuion.Th juic pd fom iu wh hghy vcuod c domind by vcuo ouon.

Th omoic pu of vcuo p vi fom 35 o 150 b nd in hvcuo of ip fui h ng bwn 0 nd 4 b (114) Th pH i wy hn 7 nd in fui h i i ofn hn 4 bcu of h cidic compoundccumud (114). Th onop i uni mmbn (71) nd i pmbiiypopi di fom ho of h pmmm (

Th incuion in fui h vcuo of mny din yp boh cynnd mophou. hough h cy of pcic compound, .g. ccium ond poium bi hy pp n numou fom. Th mophou odi of gggd o poymd compound uch nnin nd n (114 Thc conning h bodi on cd hough h h. I h bndmond h in pimmon h h nnin c nc no ony in iz h fui ipn bu o in numb 55) Pumby h c conining vcuo

Annu.R

ev.Plant.Physiol.1976.27:2072

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DEVELOPMENT OF LESHY RUITS 1

ui hve highe etti the ptiu mpu t eighing e t ggetg n uneven ititn mpn etween vue

T penmen hw ey in tvee etin gpe k (4); t etintge evepmet thee i mi ppee iig m ieee iegee pigmenttn (tyniin) etween e

Vcule Lme

Tee i g eiene tt ye (ie enen gnee ntiningyyti enye) n pnt e ( 1) tey n n e 0Mte ppe tt ve e type pnt yme hng ighyeepe inten eetin pty neqene tey epn vty neme the tte mi t tnit 0 Met e een evie

epting vue m yet e n m pt hghe pnt e.g zet tip g pei titie eve ye ve een ite witthem Neeiy tee et e e all vaul bun et pp has coe from te ding of id he p emveety m te ge e Nitea e (. Mtie pee t ue the wyme whe eeing t ihemi unti vue the mphgi tte (1) He ugget tht yme my evep m gh enpi etm it gee wit te ime. eie igintem g ie my e eete it thee gee thu ing uthe ezyme n utte. Te evepent te ve/ye ee teee

t y te tnmet enzye ut ne inten eetn (ineti!) In pti Mte h empze the iity e t eueteytpi tei y eitn n pgyti Tee pee pie eei ieving n nting tne te penmen we et-e m et te )

Seneene tgt t nve pgytti tity wt te e ytpening n nt tug the iity t ynteze ptein i intne 1) e peing e ege eneene ee e), utt t t gne tt te ge n p inpt te nt e ve phgytti pe. Meni invving e n enye n n tetnpt eem e ey. Tee i yte eviene intt e (9111) n ti eene eimente tn (88 tt i yla ae te with tpt I the wk with eimete ti enzyme weethught t ite with the ie ue the tpt Thee i evene tt yet ve whih e knw t umute gnine t ighett hve pe giie tpt ytem ppetie the ytemh it prnt eegy iin i nei n peiity e eeetemne ( emy it pete t the tnpt.

It emin t e een hw eevnt the egin tui ve e t te

e e in te e ipening t Te nept e t nptie wt te e w e een pt w t epin g tin yge ntene e ee net etn) It i pie tht the vitin

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22 COOM

btwn cll in thr vacolar comoition (mntion at th n of th rviosn) ld xlain y irnc in thir nym comlmnt

vagari in th vomnt of ach larg cntral vacol 69 Sur Aumutin by Surne Inteodes

h psn, locaton, an vaation in nym activiti n ndn maolic caailiti a ti though thy o not rov that articlar athayorat o ar a rining rit ar concn v n mny ndns inicat larg chang in nzym actvity aociat with th vnt rining Ad smmy nmn aot 4 cla o nym in vloing rit han vn y Dlly (32 T slnss s nmn s lmd b a lac oknolg o th intraclllar location o nm an b th man othr ncrtain

ti aociat with th nymology of frit ti (5 An imortant aition tothi work i th crition of cro hohat ynthta an cro ynthtain th gra brr along with othr nym invov in th mtabolim o cabohy-rat (50) h nym ar imilar to tho which occr in garcan whnintrno cll ar accmlating ar 4 hi lattr ti lack man of throblm aocat wt rt an cnabl og ha bn ma innraning th mhan f sugar toae (2, 41). The infomon wnscls atntion sn it n sv s sl mdl ti on th accmlationo gar b frit h.

Whil th garcan intrno anchyma cll ar longating th accmlatingmchanim aar to b: cro in th cll all sln i hyroly by a slld nvs ls nd s vs plsmlmm y sp nrgpld mnsms in th ml mpmn (yplsm) th ss ahohorlat an cro hohat i ynthiz th cro my o sshohat i tranrr acro th tonolat by an nrgcol mchanim. hican roc againt a concntration grant. An invrta in th vacol convrtcro to glco an rcto (4 Whn tnion groth to an th inrnoms i a larg incra n th ontation of vacolar cro. haccmlating mchanim now i rom th rcing in that th wall invrta

a bon nm an thr i no invra activity in th vacol At all tagthr i a ntral invta in th tolam hich inv r cro ocrng

n aition to th val of thi work a a mol thr ar two othr ning ointrt n i that whil intrno cll ar longating an accmlating garirct corrlation it btwn oth rat an total aci invrta activit thi i intrrt in ms comonnt a th cll wall nvs hichnls th vll o ntr o ss into th cll an ( vlinvrta c s n mn an rirt cro ro toragto trctral matrial. h con ning which ma b rlvant to th rining

o frit i that xogno ABA gratly incra inva ynthi nhibitorsds ssd s smly s pd sm s formation of invrta mRNA 3 imilarly a rlationhi ha bn fon btninvrta activit an growth o xci tomato root an AA act both (1.



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DEVEOPMEN O LESH RS 223

Wa Gwt and Wat Aumuatin: Snin

Tr ar w pnmna in r dvlpmn w xprin dpnd a lain par n war amlain an n bavir ll wall ri grw (llxpaio ad ui oi (domabiiy imi o oig i laioo ow va onidably bw pi om do no o i ar am (avado gown ommially A nmb o j bo or dring al ipi v, by wi im majoy o xpaio a b aomplid (banaa, o ri rap bry on a biin o ondgow yl may w bo ip m aily mam o diama omabiliy o ingl bi i i w av o a oig pd rnwd wlln in brri by a avra 7 ay (mb an Bip,npb.

ollowg ao a ui o ll xnion a i ug p wall nin abv a rain rial val prmabily ll mmbanaa mi a ri ux o wa, ad iniiaio and mainnan mag an baig o bod i wal, i . a biomially mdia, vio-lai w maral in wall (3, , 6 r lar axin i rid.Dng a vlmn ll xnio wld m o b oolldby a u piu bom oud duig ipig w uill xpa a on o o wiou xpaio I la a r mb a ag in wall mabiliy id wio iml o g pi a b d o o omao (10). A aoiaio a b ugd

bwn i ad lla aiviy i pa 52) ad lyoi aiviy iomao (122,

o olud i io, i ao o p a r impoanribr ripnnl amlan xpanin, an ninga a wn mng an nrl. nrl manim ma b inrla b v o no o b ily oiid

CONTRO MECHASMS

Fui ad d av b a appyigou o omo bauir nain ar nrally niabl rar an in vaiv i anxrain an piain pblm ar ln. r i g ppr r ypi a iniial grw ri i pnn rmn prd by vlpn 27 8 86 r ri i wi ar n pnn naiad grw (.g. panarpi Waingn navl an pinappl banana, rn grap am yp o omo ar oug o b invovdin prvin abiin and prmin ri rw b ar prap yni by ri i il (67, 82 85 xin, gibbrlin, ykinin, abin,an yln av b o b pr on in lag amn ad o a qn pa r a pi ( l rm xiin pa an ramnwi xn rw ban w a omp ar mim yrii pomoig gow a omim a bi o a o T al

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224 OOMB

pue f y he e d e f hme (27 . A pu- e h ee mde f e ewe GA d ABA he f whf he pe ey (2

e h ueed h h eeed y he y f hme uee mveme f ue he fu (27. Th uefu ep h e f u uded hh ukey u wehve pue f he mehm f f eh f hme Thee pew e deveped hee eue hey e ump u eue hee emy exee evew ve (26 27 6. Hweve me mme emde he f fu pe epey ep

The Trggerng f Renng

ke he ey wh f he fu he eve ved pe ppey d deped he peee f he ed A h ee d efe pe fwy e e mmum mu f p devepme d vve he he ype f ezyme ve he f h ype f heud e e f f pe fm vy he ee eve h f pefmed ezyme mpy hed pemey f meme.[She h efeed he evdee f he e (.] h ud e mdedy phyhme. Thee ue whh hve ee mde f he h f he ep f pe w e meed eyee ABA d AAxd pdu

HY Ehyee h ee w e vved fu pe (6 3d h ee eded y my he fu pe hme (6 74 92. Th mye ue f me fu u uepe ee f fu. he peey f e h u vey w pdu f ehyee du heep f pe ue e mued pdue e mu fehyee y my dee eme u hee e ewee heeeve d he m f pe (24 Du h w ee ed u he pmye f ehee he e f m fu whh h hhe ee ee yp exmpe f he vveme f ehyee. pyee ke ehyee

mued he epy me m fu u ehyee pdu dd ee u pe mmeed (; my he ee f edue m-phe peue he me f pe f m fu wee eed hemu f ehyee pee ( 2. h ppe eed h ehyee pdu e p f pe u de e h eeueed h he peee f weue ue vved ehyeeyhe eued he pep f m f pe u (04 huhm eu ed wh e f vd e epeed em f eeduppy f ye (

ee wk Gde Quee peh uefu p he py f

epe f he ep f e he e hd d f he umuf dy me he he he e ehyee e h ee fud de wh he e we pu eeed 20 dy ee d ueedh ep he h ehyee hud e uh f he e f he



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EVEOPMEN O EH R 5

epanin in vlume (7, 5. The eec ehephn applied a deren ime nhe prmin ripening peache ( l l are n a varance wih hi generalpicure. Ehyene reamen ha al been und mulae dry maer npu andi inca n h vul cu canain wr (SO, S l

Thi hrmne ha ben prped a a pible rigger penng n meui (4 44 66 76 77) Th vidnc i l in h ripning gape i allw ABA cncnrain in bry h dclind adil a h aid gwhcycl prgreed reaching a lw value abu day ber ripning prcebcam vidn. lvl hn bgan incra and hi inca cninud unilripening ven were well eablihed wherar i dclind Tramn whichelad haened ripening had parallel e n hi incra in ABA (4 Th

amun increae wa grae (n a reh weigh bai in he kin While iwa increaing here wa a cle crrelain beween he cncenrain ABA andhee here bing l ml h p ml ABA (mb npblidTremen o gaps aene rpenng when applied aer endgenuABA had reached lw level wherea earler reamen wa neecve (.

i prped ha he preliminary declin ABA a lw cncenrain i annegral par he cnrl yem and ha l perm a change n meablm. Such a cncep ha uppr in r inance he uggeed rle r ABA incnrlling drmancy bud and ed ( imilarly in aurng cn emby grmiain and an aciaed incae in proa aciviy wre rrlad

wih a lwrd cncnrain ABA, achivd by wahing pimn n hinhibiin pra aciviy by ABA and i reimulain by acinmycin D led h cncluin ha mR r pra wa bing prducd bu ha ABA wainhibiing i ranlain (54

I OX ID TION PRODUCT h dlay in h ipning ma caud byrage a lw air preur ha ben und b due he lw preur ygnand n h lw cncenrain ehlne ( iilarly h dlay in hripening avcad lice by wahing wa abued a rericed upply ygen( l1. rnkl und ha he rpenng hle green pear wa haned by rea

men wih IAA idain prduc r by an amphre ygen. Thugh hereamen mulaed ehylene prducin h cncluded ha h prmin ripenng culd n be aribued he prducin hylen e ugge in acha h AA dan prduc mulae ripenng and ehylene ynhe (5 Thcnvern ne cmpund AA rm an nhibr a prmr rui rpenngby a ingl piive acin uch a idain i an araciv mechanim and hehyphei derve urhr udy. The dual eec AA dependng n he cn-cnain applid will nd b an in accun (11

CONCLUSIONS

The develpen ehy rui i een rm an evluinary pin viw a heacquiiin characeriic which arac bird and her animal hu aiding eeddiperal.

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226 COOMBE

h aaiv i may oigia om aiay ay a o h ioovidd h i o coai om iib ay h ica i o ivovd

h i bom hy by a igad ad had dvom ivovig cdiviio ad c aio, accmaio ad cooig, ad oig Digdvom h i a ag ad bvy a

Amaio o wa i a adju o acmaio o dy ma.wo o a oidd o b o dama imoa: a e wa

extenion nd wter nd ote inx into the voe he roertie o theooa ad h ggig o h oc a o aica i.

OE AE PROO h odi o h CNR aioa Cooim o 28, r 195) id a Re gaio d a aao d

i w oay ow o m wh aig hi viw. hi bition rovide n imornt ddition to mode itertre on othrvet hyioogy nd inde ome mention o deveoment hyioogy o rit t doe nohowv a h agm dvod i h viw.

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22/22

2 COOME

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the development of fresly fruits

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