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ASPECTS OF
GRU#TH CUNTROL IN KIWIFRUI'rA
NDBLACKCUHRAN'r
A
Thesis Submitted in Partial Fulfilment of the Requirements for the Degree ofDoctor of Philosophy at
Massey University
GEORGE STEPHBN LAWBS
1979
ACKNU 1'/LEDG EM.C:N'l' S
'l'h e author ackno >.Jl edge s t h e as s i s t anc e o f Pro f e s s o r J.A . V e al e and Dr . D . J. Woolley as
·supe rv i s o rs o f th i s the si s .
Thank s are al so ext end ed t o :
Mr . W . R . Boyc e , D i rec t o r , Ho rticul tural R e s e arch C entre , L evin , for making plan t mat eri al availabl e .
Pro f e s s o r W. W. Schwab e , 1iy e Col l e g e , England , for making fac i li t i e s avai l�bl e .
Me s srs . A . N . W at son and D . And erson for t e chnic al assi s t anc e .
S p e c i al thank s are ext end e d t o my wi f e Meryl , f o r h e r t o l eranc e and support , and for typing thi_s the s i s .
TABL.c; OP COHT.!!:NTS
ACKNO iVLEDG El'.liHT;::;
ABSTRACT
Part 1. INTRODUCTION
A . l SHOOT GRm'/TH
1.1 Leaf effects 1.2 Root effects
1.3 Carbohydrate reserves 1.4 Growth regulators
B.l SAP FL011 AND SAP HOR!iiONES 1.1 Sap flow
2. Sap hormones
2.1 Gibberellin-like substances 2.2 Cytokinins
C.1 BUD DORMANCY
1.1 The development of dormancy 1.2 Bud break
1.3 Growth regulators and dormancy
C. 2 APICAL DOI\liNANCE AND CORaELATIVE INHIBITION
2.1 Growth regulators
S e c t i on 1.
EXP EHHII.C:N'£AL SHOOT GHOWTH
A B
Sho o t growth of the k iwi frui t vine Expl ant sho o t growth
1
1 2 5 7 11
18 19 21 21 23
26 28 29 31
35 37
41 41 57
Page
DI;:)CUSSION 78
Sec t i on 2. KIWIFRUIT VINE SAP 95
A Sap flow 95
B Sap growth r egul ator s 106
DISCUSSION
A Sap flow 128
B Sap growth r egulat o r s 137
Sec t i on 3. DORMANCY AND BUD BURST 145
A Winter do rmancy and bud burs t 145
B Co rrel ative inhi b i t i on 180
DL:iCUSSION
A Bud d o rmancy and bud burst 188
B "Summer d ormancy" 201
Part 2. IN'rRODUCTION 207
JUVENILITY 207
EXPEHIMENTAL 215
DISCUSSION 227
FINAL DISCUSSION 232
LITERATURE CITED 244
Figure 1 2 3
4 5
TABL� OV FIGURES
Effect of dofoli�tion on sho o t growth.
Effec t of defoliation on int ernode growth .
Effect o f dec:.::.pi tat i on and defoliat i o n on internode growth after 53 days .
Growth of kiwifrui t pl2.nt internodes .
Di stribut i on of grov1t
h
vvi thi n an int ernode .6 Standard curve for lciv-: i frui t seedling's respon se to gibberellic ac id .
7
8 9 10
Effec t of growth regulu.tor sprays on plant height .
Effec t of GA on kh�ifrui t plant height . Effect of CCC on lciwifrui t plant height . Kiwifrui t res p irat ion aft e r summer sprays.
11 Treatment effect on stem elong:tti on
kiwifruit.
12 Typi cal effec t of growth re�ulut o r s on growth of kiwifruit cut t ings.
13 Relat ionship b e t ween Cl.l tting volume and sub sequent shoot growth.
14 1 5 1 6
1 7
Effect of lAA on kiwifrui t shoot growth. Effect of GA and �CIB on shoot growth.
Ext raction of soluble sugars and reserve po lysaccharides .
Sap flo� from individual sho o ts cut 0n Day 0, in eac h successive 24-hour period.
4 5 45
47 49 49
51
51 5 3 5 3 5 5
59 - 61
6 4
68 7 1 7 1
74
97
Figure
18
Ai r temp8rature s
in relation
tot h e first day' s sap fl ow rat e .
19 Daily sap
flow from each main s t em o f
98
d e capi t at e d kiwi fru.i
t plan t s , cut on
Day 0. 102 2021
Rat e of
sapflo�
ateach
cutt ingdat e , for thre e d
ays aft er d e c api tat i on .
So i l and a i r t emperatur e s .
22
Ex traction o f gi bberel lins and cyt okinins from sap .
23
His t ograms o f GA-like sub s t anc es in
ble eding sap , spring
1973,at suc c e s s i ve c o l l e c tion
dat es.24
Histograms o f GA-lik e sub s tanc e s in
103 103
109
113
b l e eding s ap at int erval s aft er d e c ap i t ation ,
aft e r grad i ent elu t i on from silic a gel c o1.umn .
115 25Extrac t i on o f gibberel lins and cytokinins
from sap .
26
Standard curve s for barley endo sperm
respon s e t o GA3 d e t e rmin ed r e frac t ometri c ally
117
and c olourimetri c ally .
11927
Hi s t o grams o f GA-lik e sub s t anc e s in
b l e ed ing s ap , s pring
1975 ,at suc c e s sive
dat e s from d i f ferent pl ant s .
12028
Hi st ograms o f soybean bio as s ay o f
cytokinin s from bl eeding sap s , spring
1975,from diff e r ent plant s .
29
Hi stograms o f s oybean bio assay o f
c y t okinin s from bl e eding s a p i n suc c e s s ive
122
days , Plan t
20. 125Figure
3 0 HPLC trac e o f cytokinin extrac t o f kiwifruit
saps ,Pl an t 20 , Day 4 , an d s o ybean callus bio a s s ay result f o r e ach frac tion .
3 1 Effect o f t emperature o n kiwifrui t bud
burs t .
3 2 Eff e c t o f pruning
t .. ,;atmento n bud burs t at three harv e s t d:·.
'" es.3 3
3 4 35
Effect o f e th e phon
andpruning tre atment on bud burst .
Kiwifruit r o o t growth .
Ext rac tion o f cy t okinins from woody t i s sue .
36 Soybean b i o assay o f butanol- s o lubl e cyt okinins in kiwifruit bud s during bud burst .
37 S oyb ean b i o a s s ay of bu tano l - s o lubl e
cytokinins in buds and sap during spring
1 27
150
154
156 159 1 6 2
1 66
bud burs t . 1 6 8
38 Soybean b i oa s s ay of but ano l- s o lubl e c y t okinins in kiwi f rui t s t ern s .
39 Hi s t o gram o f cytokinin ac t i vity in
kiwifruit st em s
andbud s during do rmancy . 40 C omparison o f cy t oki·:in ac t i vity in s t em
171
1 7 3
tis sue
follo�in�di f fe rent l evels o f chill.ing . 175 4 1 Ext rac t i on o f absc i sic ac i d from buds .
4 2 C o mpari s on
ofinhibi t o r ec t ivity i n bud s
177
following d i fferent l evels of wint e r c hilling . 179
4 3 Soybean bion s say o f c�·tokin in a c t i vity in kiwi frui
i.sho o
L, ctsaffec t e d by auxin
ap plication .
188
Figure
44 Purification of blackcurrant tis su e f o r gibb ere l lin d e t ermination .
4 5 Bio a s say o f 2 5g s t em tis sue ±GA a f t e r extraction and c hromat ography .
46 His t ograms o f bio a s s ay s o f GA-like subs t anc e s in adult blackcurrant s t em .
47 Histogram s o f bioas say of GA-like sub s t anc e s in adult s t ems from fruiting bushe s .
4 8 Histograms o f bio a s s ay s o f GA-like sub stanc e s in adul t s t ems .
49 His t o gram s o f GA-lik e sub s t anc e s in
218
220
2 21
2 21
2 2 2
juvenil e-lik e s t em and ro o t s Wellingt o n XXX
.2 2 4 50 His t o grams o f GA-lik e substanc e s in Baldwin
s e edling s . 2 2 6
...
LI::>'l' U}' 'l'ABLt.:S
Table
l M e an
e;rowthr esponc; e
to pruningt reatment s.
2 Bffect
of pr'.Aningand
�rowth regulutort
reatme
nt on
pl�nt crowth.3
Eff ect
of growth regulatort reatm ent on explant rr1ean dry we ieh t dL::;t ri but i o n .
4 Change s in explan t carbohyd rat e l eve l s . 5 Gibberellin- l ik e activity in s ap .
6 Cyt o kinin-l i ke act i vi ty in sap .
7
Cytokinin ac t ivi ty in s ap over five d ay s aft e r d ecapi tat i on .
8 Bud burst at d i f f
erent harve st date s . 9 Bud burst f o l l owine
prQninet reatment
10 The effect o f ro o t remo val on kiwi fru i t bud burs t .
11
12
Comparison
of cyt ok inin l e ve l s in bud s during bud bur s t .
C ompari s on o f cyt o k inin l evel s in sap and bud s during bud bur s t .
1 3 Bud movement i n re spon s e t o hormone treatment in d i fferent
seasons.1 4 Effe c t o f bud
positionon cut ting on bud burs t .
1 5 Bud burst re s ponse t o varying l ev e l s o f wint e r chi lling .
6 2
6 6 7 7 119 1 2 3
124 147 1 5 3 1 57
16 5
16 9
1 8 3
1 8 6
1 9 0
LIST
OF
PLNrESPlate
1 The effec t o f gibbere l l in on sho o t growth of
anAbbo t t kiwi fruit cut ting .
2 The effect o f a pre-bud burst appl i cat i on o f gibb e rellin
andcytokinin on sho o t growth .
Page
6 5
6 5
...
ABS'l'HACT
This study emphasized the interplay of s ev e ral typ es of growth regulator in t he control of growth and d evelopment in the kivlifru i t plant. Hence different
parts of the p lant are seen to contribut e to this control , th e p l ant appearing as a fully-integrated syst em with the shoots and roots interacting with each othe r .
The gibb erellins w ere active in the promotion of st em elongat ion , and it is consid ered stem g rowth is d ep e ndent on the photosynthate sup ply , and that this is med i at ed by the growth ree;ulators. A leaf had a dominant
effect on internod e gro·nth , and therefor e shoot length , at a v e ry early stag e of its developmen t - but the effect of an individual leaf w as very loc alis ed . The plant
roots w e re not clearly shown to be ess ent i al for breaking dormancy , bud burst , or early shoot growth . How ever it was found the roots could be supplying gibb e r e llins as one of the factors reou ired to maintain normal shoot growth . Cytok inins for early bud growth could have been suppl i ed in th e bl eedine sap either from th e roots o r the stem tissu e . R es ervations ar e exp r essed about the need for h i gh spring sap flows for plant growth . H i gh sap flows may b e a r esult of a c o mbination of c i rcumstanc es at the t im e , and not a necessary pre-requis it e to growth .
SADH and mal e i c hydrazide are effective shoot growth retardants , and may f i nd co�nerc i al ac c eptanc e , exc ept their apparent effect in enhanc ing post-harvest f ruit r espi rat ion must receive further study .
Bud dormancy was great est before l e af fall , and
dormancy was broken by about 7 0 0 hours b elow7. 2°C , although warm t emp eratures were just as eff e ctive i n overcoming this dorraant condit ion . Winter dormanc y appeared to b e bas i-
c ally the sam e type of condition as correlative inh ibition , and i t is sugg est ed th8t each i s , in large part a r e sult of an inab i l ity to utilise cytokinins or oth er growth promo ter s .
It i s considered that the juvenile-like condition in blackcurrru1t i s distinct from t rue juveni lity, and the form er did not app ear to be the r e sult of prox im ity to gibb erellins from the roots . Further , it c ould no t b e shown for true juveni l ity that i t w a s lik ely to b e du e to th e production of inhibitory l ev e l s of gibber e llin by the roots.