J. Fac. Agr., KyllShu Univ., 43 (:3'4),411-418 (1999)
Gibberellin Induces Shoot Growth and Flowering in Nonprecooled
Derooted Bulbs of Tulip (T-dlipa gesneriana L.)
Marian Saniewski l , Ludwika Kawa-Miszczak 1 ,
Elzbieta Wegrzynowicz-Lesiak l and Hiroshi Okubo 2
J
I{esearch Institute of Pomology and Floriculture, Pomologlczna 18, 96-100
Skiernif"~ice,
Poland
'Laboratory of Horticultural Science, Faculty of Agriculture, KyusrlU University 46--0l.
Fukuoka 812--3581, Japan
(Rece'i'lAxl October 80, 1fUJR and accepted l'lm:ember 6, 19[)8)
Effect of gibberellic acid (GA,) at a concentration of 200mgi on shoot grO\vth and
flowering in nonprecooled demoted tulip bulbs cvs. Gudostmik and Apcldoom ",fas investigated.
In control nonprecoolec1 derooted bulbs flmver bud blasting occurred and no gro\Vth of stem was
observed. GA) greatly induced shoot gnm11l and tlowering. The stimulatory effect of gibberellin
on shoot grov,rth and llm....ering in nonprecooled derooted bulbs vms much st.ronger than in ca~e
of nonprec(x)led rooted bulbs (previously published results). GAl stimulated also stem grow1h
after excision of flower bud with upper part. of leaves and scales. 1t is probable that excision of
all roots in nonprecooled bulbs decreases the level of abscisic acid and exogenous gibberellic
acid is more effective in comparison to nonprccoolcd rooted bulbs. Or it is also possible that
better penetration and uptake of gibberellin take place after removal of roots. It seems that
gibberellin can substitute for cold requirement and that interaction of exogenous gibberellin
",ith endogenous auxin controls stem growth.
j
INTRODUCTION
Tulip bulbs, with terminal buds containing a complete flower, require a period of
12-16weeks of low temperature treatment for floral stalk elongation (De Hertogh, 1974).
It is well known that during the cooling of the tulip bulbs the amount of free gibberellins
increases (Aung and De Hertogh, 1967; 1968; De Hertogh et aI., 1971; Van Brag!, 1971;
Aung and Rees, 1974; Hanks and Rees, 1980). Hanks and Rees (1980) found two peaks of
activity of gibberellin-like substances in tulips sampled at intervals from October
(planting time) until the following April (flowering time). The first one occurred in
December or in early-January samples, before the cold requirement was completed, and
the second peak occurred around the time of rapid shoot extension and flowering found
in roots, scales, leaves, stems, daughter bulbs and in flowers. Recently, however, Rebers
et al. (1995) suggested that there was no direct correlation between cold-stimulated
growth and a change in endogenous GA status in sprouts or basal plates of tulip bulbs
during cold storage.
Saniewski and De Munk (1981) and Banasik and Saniewski (1985) concluded that
elongation of all internodes in cooled tulip bulbs is promoted by auxin produced in leaves
and flower buds, and that the auxin-release or response system was stimulated by
2
Correspondence to either author. 'Tel: +48-46-33-2021, Fax: +48-46-33-3228, E-mail:
[email protected] ~Tel: +81-92-642-2828, Fax: +81-92-642-2829, E-mail:
[email protected]
411
412
At. San'iewski et af.
gibberellins. Recently it has been suggested that the elongation of all internodes in tulips
is controlled by auxin and gibberellins (Okubo and Uemoto, 1985; 1986; Okubo et ai.,
1986; Saniewski. 1989). Okubo and Uemot.o (1986) suggested that two different gibberellins are involved in controlling the elongation of the lower and upper internodes of
tulip flmver stalk. Kawa and Saniewski (1986) showed that gibberellic acid had a strong
stimulatory effect in increasing the length and fresh weight of pistil isolated from
uncooled tulip bulbs but to lesser degree in the case of cooled bulbs, cultured in \itro.
Saniewski (1989) suggested that gibberellins produced during the cooling of bulbs play an
important role in the flower bud development, mostly pistil growth, and other gibberellins
are synthesiZf~d during shoot growth and, together with auxin, control the stem
elongat.ion in t.ulips. Hebers et ai. (1995) also suggested that. in tulip GAs are involved in
the stalk elongation response.
Exogenously applied gibberellins cannot substitute for cold treatment of uncooled
tulip bulbs but do for only partially cooled bulbs and stimulate shoot growth and flowering
(Van Bragt and Zijlstra, 1971; Rudnicki et at., 1976; Van Bragt and Van Ast, 1976; Cocozza
Talia and Stellacci, 1979; Hanks, 1984; 1985; Jones and Hanks. 1984)
In the present work the stimulatory effect of gibberellin on tulip shoot grO\\'1:h and
flowHing in n()npn~c()ol~d rlp.rooted t.ulip bulbs is documented.
MATERIALS AND METHODS
Plant materials
Tulip bulbs evs. Gudoshnik and Apeldoorn, ""ith a circumference of 10~11 cm, after
lifting were stored at 18~20"C until used for experiments bet\veen November 10~22
(nonprccooled bulbs). After the removal of dry scales the following experiments were
conducted.
Experiment 1
All roots were excised and bulbs were kept on petri dishes 1) with distmed water or
2) with gibberellic acid (GA,) at. a concentration 200rngl-'. In a second group all roots
\vere excised and the upper part of the scales, together with flower bud and the upper
part of the leaves at the level of the flower bud, were cut and the bulbs were kept on petri
dishes 3) with distilled water or 4) with GAl at a concentration of 200mgl-:.
Experiment 2
All roots were excised and the apices of sprouted leaves were treated as follows: a)
untreated, b) lanolin on leaf apex, c) TAA 0.1% in l;:molin on lpaf apex, rl) lanolin after
removal of 2 mm leaf apex, or e) IAA 0.1 % in lanolin after removal of 2 mm leaf apex. The
treated bulbs were kept on petri dishes 1) with distilled water or 2) with GA3 at a
concent.ration of 200 mgl-'.
In both experiments 15 bulbs per treatment were used and every experiment "vas
repeated twice. Newly appeared roots were removed daily. During the experiments the
shoot growth and flowering were measured and photographed.
Oi fJlw reUi:fl I.Hi-J l owerirlfl qfnonpTecool,~ ,-u tip o'l1.lbs
~~ffc c:t
Table 1.
of GA, 200 mgl · 0 11 shoot growth ancillowcring of nonprecoolerl
dcroot ed bulbs.
T reatmcnt.
Length of s prout (fi rst Ica f) o r "'stern
L~ l tgt.h
of intcm odc (mm) on l)e{:.
22
wit h Ilower bud (nun)
Nov.
Nov.
Dec.
30
Dec.
3
Dec.
11
U
22
23.3
28. 1
34.9
51.2
46.4
71.1
R4 .7
SU.i
11M *243.4
lsT.
2nd
3rd
Tota l
41.1 1
._--
Roots removed
c\'. Gudoslutik
'water
G.'\.
-'
32.0
25 .H
14.1
17.0
:30.9
86.G
17G.a
27.2
95.2
1f,4 .4
c\'. Apcldoom
waLer
26.9
39. 1
48.3
60 .~
n.6
66.2
GA,
3 1.1
50.5
98.1 *215.7
Roots rmrwved and flower bud togethe r with 'upper part Q{
-'
[.ua-ccs and scales excised
Gudoshnik
CY .
waler
GAl
-'
7.1
8.3
10.8
17.4
15.6
2D.6
27JJ
33.7
870
36.8
:Jl.:J
H).fj
7.4
11.1
Go .:!
6. 5
9.4
10.8
18.5
l(H>
30.2
23.5
31.5
30.0
37.0
26.5
16.7
11.4
11.7
66.3
ev. Apcldoom
water
GAOl
'No stem gro\\·1.h , aw l OUWf'.I bud blasting.
'No stem growth.
RE SULTS AND DISCUSSION
Experiment 1
The shoot growth o f nonprecoole d de rooted tulip hu lhs was very small in bot h
cultivars, only sprouting of leaves was observed and t10wer bud blasting took place (Table
I , F ig. 1). Giuuerellic acid at a concentration of 200mgl ' ca used the breaking dormancy
and indu ced th e shoot growth and flowering in nonpn:~eooled deroot ed tulip bulbs. GA~
also slirnulated stem growth after excision of the flmvcr bud and the upper part of th~
leaves at the level of the flower bud. Thus, independent of the presence or abse nce of a
flo\-ver bud in non precooled deToated tulip bulbs gibberellin stimulated stem gr owth , but
was much less effective in cases where flower bud and upper part of leaves were excised.
It. spems t.hat exogenously applied gibberellin stimulates pislil growth , and intentetion of
exogeno us gibberellin wi th' endogenous a uxin is responsible for e longatjonal growth or
stem.
Experiment 2
IAA 0.1 % applied on leaf ap ex did not. affect shoot growth of nonprecooled tulip bulbs
kept. on the water or treated wit.h GA, (Table 2, Fig. 2). It was suggested previously that
the elongation of all internodes in precooled t ulip bulbs is controlled by interaction of
auxin and gibbere llin (Okubo and Ue moto, 1985; 1986; Okubo at aI. , 1986; Sanie wski,
1989). The stimulatory e ffect of gibbe rellin on shoot g rowth in nonp recooleu de rooted
414
AI. S'aniewsh et at
Fig. 1. Effect of GA: 200 mgll on shoot grmvth and flowering oj" nonprecooled deTOoted bulbs.
Cpper: rv. C:udoslmik, lower: C\" l\pel( loofl1. Treatment was on November 22 and the
plant') were vhotographcd on December 17, either viith leaves (Left) or after removal
of all leaves (Right) before photography. From lett to riRht., cOIltrol, distilled .vater-no
grmvth of slem and Oower bud blasting can be observed, GA1-stimulator.y effect on
shoot growth and nO'wcring, control, dist.illed water-small grmvtlt of leaves and stem
after ell\. of flower bud together ""it.h leaves ami scales, GAo-stimulatoTY effect on stem
grm;vth can be observed
Gibberellin onjlon:eTinq (Jfnonprecooled tulip bulbs
415
Table 2. Effect of LA. l\. (J.1 % treaLlllf:'llt. OIl the leaf apex (Jll shoot growth and flowering of
nonprecoo\ed derooted bulbs ev. Gudosltnik kept. in wat.er and GA" 200rngl-'
Treatment
Length of internode (mIll) on Dec. 12
Lengt.h of sprout Cfin;t leaf) or
*stcm with flower bud (nun)
Dec.2
Dec.oS
2nd
Ut:!-'.IL.
---
------
;3rl1
4Lh
Tut.]l
----
Untreated
43.7
66.9
\vaLer
GA
Lafuilin
Oll
57.5
93.4
75.7
*2Wdl
19.5
16.6
20.9
75.8
134.8
64.1
*22B.1
2Ed')
22.0
26.:l
8:3.D
167.8
10'l.5
*180.9
24.t5
18.6
18.4
62.5
12'1.0
31.8
23.6
268
86.6
WH.8
;10.(J
21.6
23.8
7:3.7
14D.l
intac:t leaf ape]·
water
49.4
63.8
GA,
69.5
D8.9
L4A ()j% on intaci lea/ape.};
water
63.1
82.7
GA,;
82.6
117.6
!,((rwiin (lfter renull.'ul of 2 mm leaf apex
water
43.8
G7.D
G.lt
60.5
.'l7.n
liLl 0. 1% aJter rernauoJ qf 21ilm leaf apex
w8ter
50.3
77.9
Ti'.2
llS.2
GA.J
7S.H
*232.fl
fl9.3
*20!J.fl
---
-
-
No st.em growth, and flower bud blasting.
tulip bulbs was much stronger than in case of nonprecooled rooted bulbs (Rudnicki et at.,
1976). In our experiments we used GAl hut GA:, GA1 , GA" GAu, GA~.I and GA1.j ,,/ere
identified in hoth non-cold cooled tulip bulb sprouts (Rebers, 1992; Rebers e/ aI., 1994b;
1995). GA 1 was the major gibberellin, \vhile G1\.], GA~ and GA: 14 were present in Imver
amounts. Rebers et al. (l~J95) suggested t.hat in tulip, gibberellins arc not the
cold-induced trigger but are involved in the stalk elongat.ion response, as \'vas shown b.y
t.he inhibition of this elongation by the GA biosynthesis inhibitors, ancymidol and
padobutrazol, and the reversal ofthis effect by applied GA (Shoub and De HeItogh, 1974;
Saniewski, ID89; Rebers el 01., 1994a). Rebers el 01. (1995) showed the higher level of
GA and it.s inactivatiull prouuct GAl. oIlly in grmving Doral stalks of cooled bulbs, and the
absence of a significant. quantitative accwnulation of GAl and suggest that GA4 is probably
involved in the floral stalk elongation of tulip .
It is possible that excision of all roots in nonprecooled tu1ip bulbs decreases the level
of abseisic acid (ABA) and/or exog0Ilous gibbnellic acid is more effective in comparison
1.0 intact non precooled rooted bulbs. The presence of ahscisic acid in tulips is well
documented eAung and Dc Hertogh, 1979; Singh ef at., 1979; Syrtanova et aI., 1975; Terry
et at., 1982) and Imv temperature treatment. decreases the amOlmt of ABA (Syrt.anova et
at., 197:3; Rakhimbaycv et at., 1978; Aung and De Hertogh, 1979). Aung and De Hertogh
(1979) suggested that probable sites of absc-isic acid biosynthesis in tulip bulb are the
developing bulblets, basal plate and roots, since t.he ABA level is high in these organR and
low in the scales and shoot. The inhibitory effect of ABA on pistil and stem growth in
tulips Vi'-as also well documented (Sanjewski et al., 19HO). It is also possible to consider
that after excision of roots in nonprecooled tulip bulbs hetter penetration and uptake of
gibberellin take place and they are more effective in stimulation of shoot grO\vth in
j
·116
Af. SanJewski et al.
Fig. 2.
Mfed ot leaf apex lA..;\ 0.1 % treat.ment on the shoot grO\\'th and
flowering of nOllprecooled demoted bulbs cv. Gudoshnik kept in
\vater and CiA:< 200mgl-' Upper row; bulbs kept. in wat.er, lower
row; bulbs kept in GAJ , From left to right; untreat.ed , lanolin on
leaf apex, fAA on Jeaf apex, lanolin after removal of 2 rmn leaf
al_)l~x, fAA af!.i-Or IHTloval of 2 rllTn leaf apex. Treatments applied on
November 19, photographed on December 21.
comparison to Ilonprecooled rooted bulbs. Our results indicate that gibberellin can
substitute for the cold requirement of tulip bulbs.
ACKNOWLEDGMENTS
The stay of Marian SanieV'{ski at Kyushu University, Fukuoka, .Japan was supported by
the in\itation fellmvship program for research in Japan, Japan Society for the Promotion
of Science.
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