Structure, function and quality development in apples

175 Pages·2010·2.11 MB·English

by  Dražeta, Lazar R

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Structfuurnec,t ainodnq ualidteyv elopmienan ptp les
A thepsriess einnpt aerdtf ulifail lomfte hnrete quirements
frot hdee groefe
DoctoorfP hilosophy
III
PlanBti ology
at MaUsnsievye rPsailrtmsyet,No onr th,
NewZ eanlda
LazarR .D raZeta
2002
11
Dedication
Thist hseiissd e diceadtt o fmamyil yi nr ceognitoifot nhs efu ferintgeh ye nudreadn d
the tlehoyvi ene vstienmd e .
111
Abstract
Relatiobnesthwiepesn ansdtf runuccttiiuonarn pe p lesa swseerisensa s e edr oife s
experismteundtoiandel ifs rfe eants peocffrt usiq tu aldietvye lopAm heonlti.sd teiilcs mo
presetnhtdaeetds crtihmbeaej sroc ontriebluetmienSngpt esca.it atle nitdsii ornet cot ed
thmee chanuinsdmesr tlhyveia nrgi pohuyss iolroeogsnpisceasl.
Thset uodfty h ree latiboentswhseieepens d e t,s hfrauapinetd frmiunietsr ahloswt ehda t
fruigrto wtwha mso dural nawda sst roinngfllyu beyn csededsie tdr ibauntsdie oend
growtThh.ip sa ttienrnfl uetnhcseep sa tdsiitarli boufmt iinoenr( ael.csga .l cwiiumt)h in
thfreu it anidls i tktheoiul msyp aocnfrt u istt orqaugael Tihtneya .t uorfte h e
developsmteinmtwuaallisu n sv estbiyig na tasepidlptc iua tiooftnhsae u xin-transport
inhibNi-1t(-oNprah,t hyl)phatchi(adNlA Pa) Nm.Pi A c a pplicraetdiuvocenesd s el
difrfeeinattiinto hnse t aelnkh,a nfrcueiadtb sciasnsldie otdnoa ,r edusceeeanddd fl esh
growtBhec.a uosfte h sep eciofiftc hiaetc yt oifoPnNA ,t hied entity of the
devpemleonsttailm iumslo usslt i kteobl eya uxin.
Thsel owainnedgv ensttuoaplp oafcg ael ciimupmob ryat p lpeiss prodbutaeoba l y
decliintn hefenu ctioonfai ltxisyt lyeT mh.id se cliinfn nucet ionwaalssi htoywt nob e
duteoa p hysdiicsarlu opftthiveoa ns cbuulnard claeussb eyfld e sehx pansTihoen .
extoefn tx ydlyeusnmfc twiaogsno vernbeytd h set rucantdcu hraer actetri sosfu et he
surrounadp ianrgt ibcuunldtalyrep F el.e esxhp anasliisononfl uetnhcteee sx tural
propertthifeeur siT tho.sef p atdiiaslt riobfiu ntctieeorllnla auriw ri thtihfrneu iatnt dh e
shaapnemd u tudails poosiftthieo ncf elelwslehsra es seusssienndog v teelc hniTqhuee s.
radpiaatlt oefirn n teelrlcuailwraa rns o utn oirfmi ndicfaaettiuntrghe aasrt le i kteol y
impaucptog na tsrn asefrw iththifrenu iatnt dh uosns torbaeghveai our.
Thmee chanuinsdteircs tganadiinniegntds h tihse pseirsmt ihetel ucidoafct oimopnl ex
interrbeleattwtiehoepennr s o ceosffrus ietqs u aldietvye lopTmheweno tro.kff resn ew
insiighntttsho oe r igoifsn osmp eh ysioldoeegfciatcnsaid ln dtiecnsae wl inferofus t ure
research.
IV
Acknowledgements
I wtiost hh ank:a lple opalnoedr ganistahhtaaitv ohene sl pmeetd oc ompltehttiehs e sis
and hcaovnet ritbotu hwteee ldb le ionfmg y f aimly.
FirsItt hlaymn,yk: s upervDirAs leoxrasn,(d Senard yL)an gP,or fPauEl.Ja a meansodn
Dr hRaircVdo lfzrot hepiart igeunitd aanncsdee lflseuspspt ohrurtgo hotuhcteo urosfe
thpirseo cjIt .h ave lgoafrtio nmte hde Hmae. n cteh,eg ya ianlm lyr espanedc t
rememnbcrea.
Thceo mpleotfti howinos r wko unlodht a vbee epno sswiibhtloeut tha es sisotfan ce
DrA lsitaHiarlD lr, NiDhea llSv iDar,C hriaaC apepllinMii,c hMJaarer lvM irDs o,u g
HopcrMorftR ,a ymonBden neMtstK ,a thy MMuirsrAsany d,rL eeao nda-rJoanneds
MrsG eralWdoiondIe.na dditsiuopnes,reb r vainceden courawgeermpeer notv ibdye d
thHeo rtResleiaabrrricManhrs S tevNeonr thMovresr ,AA ninn scaonuMdgs h Sarah
Nation.
Id eepalpyp rehceilaiptn,es piraftreiinodns Ihah inapdvr ee cefriovmem dan yg ood
peopDlrZe l:ta kKoo vaCDirDc o,rD au ardteCe a rvaDlrPh aou,lA ustDirSn t,ae nf
HentMoins,Gs e ogMiinlan Mei,sH se lBerno wMnr,D usanB ackjualP,or fMicOojl caa,
MrsC atheKreianreMi rnJ su,s Wtiilns DorKn a,t Mea guiDrrMe a,r kDuesu rer,
DrP iteeWro lswinkBearlt,o lDoDrmi ecohM irDo a,v Aed amMsr,B eAnn derson,
MrM urrOalyei rvM,r I aMnc Ivfomari,l iCelmsee nHsa,y waVrudk,c eBviiuzcm,i c,
HotceFnokmoia,nn L da urenSspoenct.ih aalnar kes dtuoe MMrasr gaLraewtla enrd
Mrs SaSnidmrpasw hoon were cmoyn tifianrc sttNsZ e ewa lanandds trosnugplpyo rted
myw orking atP ohwDar.d s
FinaIlw liyst,hoa cknowtlhefiedng aen acinpadrl eo sfsisounpaplpo rrotv ibdyte hde
AgricultMuarrakle Rtaeinsnrdegc a ahn Dde velnotpT mreu(sAtG MARDWTo)r,ak n d
IncoNmeewZ eanld-a StudSeenrtv iHcoerst,R esMeaasrscUehny,i verasnNidet wy ,
ZeanldSa ocioefPt lnyat P hysiologistsm a(dtNehZ wiSosPr pPko) s tshiabtl e.
v
Tabloef c ontents
DEDICATI.O.N. .............................i.i. ...............................................................................
ABSTRACT iii
....................... ...........................................................................................
ACKNOWLEDGEME.N.T.S. .....................................i..v.. ...............................................
TABLOEF C ONTENT.S. ..........................v. ....................................................................
LISOTF F IGUR.E.S. ............................x.. .. .......................................................................
LISOTF T ABLE.S. ...........................................x..i..v.. .....................................................
CHAPTE1RG. E NERAILN TRODUCTION
1.S1t.a teomfte hnpetr oblem . .................. .... 1
...................................... ....... . ... .. . . . . .. .. .. . .....
12..St rucotfau prpefrl uei t . . . 3
............... .................................. .................................. ........
12.1.S.t rucotruirgaoilfan psp flreu it . ..... 3
.......................... ......................... ... ......
12..S2t.r ucotftu hrseek in . .. . . .. 4
........... ........................... ........ ... ...........................
12..S3t.r ucotftuhreeex tracartpiesUsaurey . 4
..................... ..................................
12..S4t.r ucotftu hrceea rpetlilsasruye .4
........................................... ....................
12..S5t.r ucotftuhrveea sctuilsasru e. . . 4
.......... ...... ................................... .............
12.6..St rucotftu hrseee ed . . . . ....6.
................ ................................. .......... ........ .... .. .
13.T.h aet triobffu rtuqeiusta lity .6
................................................................................ ...
13..S1k.ip nr operties. . .. . ..6
................ ........ .................................. . ................... ......
13.2..Fr usiitz e . . . . 6
............................ ........................................ ................... ..........
13..F3r.us ihta pe. . . .... .8
...... .......................... ............................... ................... ... .....
13.4..Fl esh textu.re . . . 9
.............. ................. .... ............................................. ...........
13.4.1.C.e lwla ll .. . 9
............................ .............................................. ... ......
13.4..C2e.ls lh ape. . 10
...... ..................................................... .....................
13.4..C3e.lp la cki.ng. . .1 1
... .... ........................................... .........................
13..4.T4is.s aunead i vro luomfte h fer uit. . ...12
....... ........ .................. .... ...
13..S5t.o rqaugael ity . . . .13
................... .............. .................. .......................... ........ .
13.5.1.V.a scudleavre lo-pimnednutc antdig orno wptaht tern 13
.............
VI
13..25.X.y lefmu nctiodniaurnlaialtn ysd e asocnhaaln ge.s 15
..... ..........
13..35X..y lefmu ncti-omnianleiartcaycl u mulaantdi on
storqaugael i.ty . . ... 16
.............................. ............. ......... ... .. ............
13.6.E.a tiqnuga lity. .. . ... 18
......... .............. ..................................... ................ . .. .....
13.7..Se esde t. .. . 18
............................. ... ........................................................ .........
13..17O..v ule development . . ..19
............................................ ......... ....... .. .
13.7..P2o.l ldeenv elopment . .19
............................. .................................. . .
13.7...S3 eedde velopment ... .2 1
.................................................... .. ...........
13..47.S.e eidnsfr uidte velopmen.t . 21
............. ....... ..................................
13..7S.e5e.ads ss itoefas u xpirno duc..tion .. . ..22
. .............. . ..... .......... ... ...
13.7..C6e.l lmuelcahran iosfam u xin tarnands port
developofms etnotrq augael .it.y . . 24
. .................... .................. ......
14..Co ncepmtoudaeollffr uiqtu aldietvye lopment . . 25
................................ ....... .............
15.R.e seaorbcjhe cotftihvteeh se s.is . . 27
. .................. ..... ..................................................
CHAPTE2R.S E EDS ETF,R UIT SHAANDP FER UIMTI NERALS
21.I.n troduction . . 28
................................................... ............ ............................................
2.M2a.t erainamdle st hod.s. ... . .. . .29
...... . ... . ............................................. .................... .... ..
2.12P..l amnatt erial . . . ... .. 29
............................. .................. ........ ..... .. ... .... ..............
22..S2e.e vda riabsieleaidst yym,m eatnrfdyr uaisty mmet.r.y ...29
... ... ............. . . .
2.2.M. i3nearnaall yses . ...32
.......................................................................... .... . . ...
2.R3e.s ults 32
.................................................................................................................
23..S1e.ev da riability . . .32
....................................................... ....................... ....... .
2.3S.e2ea.ds ymmetry . . . 34
....................................... ............. ........ ........................
2.3F.r3ua.is ty mmetry . ....... .. .. 35
........................................ ...... . . ... . ......... . ........ . ....
2.43.F.r umiitn er.a.ls. . ... . .. . . .. 37
........... ........ ........... .. .. ......... ...... ........... ..... ........ ....
2.D4i.s cus.sio.n . . . . . .. ..38
. ......... ..................................... ......... ....... ........... ........ ..... .. ....... ... .
24.1.S.e evda riabilit.y .. .. . ..38
................. ............... ...................... .. ...... ............... . ...
2.4S.e2ea.ds ymme.try .. ... .. . .4 0
. ........... ... ......................................... . ...... . ...... .......
2.4F.r3ua.is ty mmetr.y . . .. .. 40
......... ....... ............. ....................... ..................... . ......
24..F4r.u miitn era.ls . . . 42
... ................... .......... ......... ..............................................
2.C5o.n clusions . . . ..43
...................................... ........................ ............................... ..... ... ...
vu
CHAPTER3 .X YLEMD EVELOP MENT IN TH E STALK OFA PPLEAND THE
EFFECTOSF AUXINT RANSPORTIN H IBITION
3.1I.tn roducti.o.n . ". ................................................................4..4.. ................
.... ... ..... .......
3.2M.a teriaalnsdm ethods. . . .. .. ...... .... ... 45
... ... ...... ........ . ..... . ....... ..... ... . .... . ..... .. ........ ... ......
3.2P.l1a.n mta teri.al ........ . . .. 45
.. ........ .... .... .. ... ....... .......... ...... ................. ..... ..... .. ...
3.2P.2re.p aratianodn a pplicaotfai uoxni tnr anspoirnth ib(iNtPoAr) . ..... 54
. . ... .. .
3.2.V3e.ss meela sureme.nts.. . .... . . . .46
........ . .. .... .. ... . .... ... ........... ............... ..... ...... ..
3.3R.es ults 49
....... ..................................... .......... ........ ........................... ............. ...........
3.3.S1pa.t idails tribouftt ihovene ssels . . ....... . 49
.... ...... .... ..... ... ... .. .. . .... ........ .........
33..2.F requencyd:issitzrei bouftt ihovene ssels. ....... ... ..50
....... .... . .. ... ......... ........ .
3.3D.e3v.e lopmoefnt th set alk . . . . . . ... 52
............ . ..... ....... ....... ....... .... ..... ..... .... ... .....
3.3.E4f.f eocftN PAo nfr uiatb sciss..i.o.n . .. . . . 54
. .. ..... . ... ..... . .... ...... .... ......... ...... ...
3.3.E5f.fe ocftN PAo ns eeandd fruidte velopment . .... ..5 5
.... ........... . ........ . .. ...... .
3.3.E6ff.e ocftN AP onv essdeilf ferenti.ati.on ... .... 56
... ..... ..... ............ . . ...... .. .. . ....
3.D4i.s cussi.o.n. ....... . ... . .. . . . 59
. .. .... ... ... ... . ... ..... ........ ....... ... . . ..... ... .......... ... ..... ... . ........ ...... ..
3.4.V1e.ss meela suremeanntdes s timactoendd uctan.ce. ... . .59
.... ..... ... . ... .... ....... .
3.4.S2pa.t idails tribuotfit ohnve e sse.l.s.. ...... . . 59
... . . .. ...... ......... . ........ ... .... ...........
3.4.F3r.e quencyd:issitzrei boufttihovene ssels. .. . .60
....... ........ ..... . ... .... ..... ......... .
3.4D.e4v.e lopmoefnt th set alk.. ... . . .. ... ....61
.... . . ... .. ........ . .... ... ... ... .......... . ... .... . ... ...
3.4.E5f.fe ocftN AP onf ruaibts cissi.o.n. . . . ..62
.... .. ...... ..... ..... ...... ........ .... ..... .... ...
3.4.E6f.fe ocftNP A onv essdeilf ferentiat.ion ... . . .6 3
....... .. ... .......... . ... . .... ......... ....
3.4.E7f.fe ocftN PAo ns eeandd fruidte velopm.ent . . . . 64
...... .... .... . .... ........ ....... ..
3.5C.on clusions .. . .. .. . ... . ........ 66
..... .... ...... ..... . ...... .... ....... ... ... .... .. . ................ ... .... .. . .. . . ........
CHAPTER4 .C AUSESA ND EFFECTOSF CHANGESIN XYLEM
FUNCTINOALITYIN APPLEF RUIT
4.1I.tn roduction. . ... . ... . ... . . 67
......... ....... .... . ... . ............ ....... ......... ...... ..... .... . ....... ... .. .... ...... ...
4.2M.a terianadl mse thods. . . . . .. . .... ...68
.... ... .... .... .... .... ....... .... ..... .......... ... .. ... ... ..... ..... .. ... ..
4.2P.l1an.t m ateria.l .... . .. . . .. .. .... ..68
. ..... . ... .. ..... .. .... . ....... ..... .. ....... . ....... . .... .. . .... .... ..
4.2T.e2m.p oral chianxn ygleesfm u nctional.it..y . . ....68
....... .... . .. ........ ... .......... . . ..
Vlll
4.2E.x3p.o stuirmee 70
............................. ..............................................................
4..24S.p atcihaaln ignxe ysl efmu nctionality 70
....................................................
4.2A.n5a.t oomfxy y lebmr eakage 71
................................................................. ...
4.2G.r6o.wo tfth h dei ffefrreutniitts sues 72
.........................................................
4.R3e.s ults 74
....................................................................... ....................................... ...
4.13E..x postuirmee 74
................................................ ...........................................
4.3T.e2m.p ocrhaaln ignxe ysl efmu nctionality 75
................................................
4..33S.p atcihaaln ignxe ysl efmu nctionality 79
............................ ........................
4..34A.n atoomfxy y lebmr eakage 81
...................................... ..............................
43..G5r.o wotfth h dei ffefrrueitntit s sues 84
.........................................................
4.531..A.b soltuitsegrs ouwet (hA TG) 84
........................... .......................
4.3.R5e.l2att.ii svgrseou wet (hR TG) 85
...................................................
4..35S.p3a.ta irraaln geomfep netta anlsd e pbauln dles
int hfel e(sPhI S) 86
........................................................................
43.5.4..Th lea yooufst e pbauln drleel attoti hvleeo c(uSl-eL ) 87
.............
4.D4i.s cussion 88
...............................................................................................................
4.14E..x postuirmee 88
........................ ...................................................................
4.4T.e2m.p ocrhaaln ignxe ysl efmu nctionality 88
.............................................. ..
4..43S.pa tcihaanlg ienxsy lefmu nctionality 92
.................................................. ..
4.4An.a4t.o moyfx ylebmr eakage 93
....................................................................
4.4G.r5o.wo tfth h dei ffefruriettn its sues 94
.........................................................
4.C5o.n clusions 96
....................................................... .................................................. ....
CHAPTE5RA. IR AND CELLM APPINOGFA PPLFER UIT
5.I1n.t roduction 98
................................ .............................................................................
5.M2a.t eriamlest hanodd s 99
............................................................. ...............................
5.12P..l amnatt erial 99
...........................................................................................
52..A2i.vr o lummeea surement 100
........................ .................................... ...........
5.2T.i3m.i ngs 120
................................................. .................................................
5.42.S.p atdiiaslt riobfiu nttieornc seplalcuelsa r 130
.............................................
5.2C.e5lm.le asurements 140
.................................................................................
IX
52.5..S1t.a ining . 140
.............................. .......................... ..........................
5.2.C5e.ldl2i m.e nsions 150
........................................... ..................... .......
52..A6d.j ustmfeonarti l ro ss . 150
..................................................... .....................
5.R3e.s ults 170
................... ........................................ ................................................. ...
53.1.T.i mings . 170
.................. ............................................................................ ...
5.23.C.e ll measurements . 180
.................................... ............................................
5.3A.d3j.u stmfeonarti l ro ss . 190
................................................... .......................
5.3S patdiiaslt riobfiu nttieornc seplalcuelsa r 121
A. .......................................... ...
54..Di scussion . 131
............................ ...................................................................... ..........
54.1.T.i mings . . 131
.................................. ................... ................................ ...........
54..C2e.l l measurements . . . 141
....................................... .... ................ ...................
54..12S..t aini.n.g. ....................1. 4.1 .......................................................
'"
54..2C.e2ld.li mensio.ns .141
...... ............................................................. ..
50.42.Ce3ls.lh aapnecd e plalc king . 151
................................. ...................
5034A..d justfmoearni ltro ss 161
................... ..................................................... ...
5 Spatdiiaslt riobfiu nttieornc seplalcuelsa r 171
AA. .......................................... ...
5.C5o.n clusions . 191
........................ ................................. ................................... ..............
CHAPTER6 .G ENERADLI SCUSSAINODNC ONCLUSIONS
61.G.e nedriaslc ussion . 120
............................................ ..................... ..............................
61..S1e.ea dsa s ourocfde e velopsmteinmtualli . 102
................... ...................... ..
61..2N.a tuorfte h dee velopsmteinmtuall.u s. 132
.... ...... ......................... ...............
61.3.C.h araocffrt ueitrti sgsruoew atsah detenninant of
fruiqtu aldietvye lopment 142
......................................... .............................
61.40.Ti ssauiaern cde allli gnmeanntfrudsi ts torqaugael ity .162
....................... .
6.G2e.n ecroanlc lusions 172
........................................................... ...................................
REFERENCES. 192
....... ............................... ......................................................................