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Gibberellin and CCC Effects on Flowering and Growth in the Long-Day Plant Lemna Gibba G31 Charles F

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Gibberellin and CCC Effects on Flowering and Growth in the Long-Day Plant Lemna Gibba G31 Charles F Plant Physiol. (1969) 44, 503-507 Gibberellin and CCC Effects on Flowering and Growth in the Long-day Plant Lemna gibba G31 Charles F. Cleland2 and Winslow S. Briggs2 Department of Biological Sciences, Stanford University, Stanford, California 94305 Received October 11, 1968. Abstract. The application of gibberellic acid (GA3) to the non-rosette long-day plant Lemna gibba G3 at concentrations from 0.1 to 100 mg/l did not induce flowering on short days and inhibited flowering on long days at ooncentrations of 1 mg/l and higher. On both short and long days GA3 concentrations above 1 mg/l caused a decrease in frond size and fresh and dry weight, but an increase in the rate of frond production and thus an increase in the # VF (number of vegetative fronds). Identical results were obtained when gibberellin A7 was used instead of GA3. The addition of the plant growth retardant CCC [ (2-chloroethyl) trimethylammonium chloride] to the culture medium on long days resulted in almost complete inhibition of flowering at 10-3 M. Vegetative growth was also inhibited to some extent. With CCC at 10-3 M the simultaneous addition of GA3 resulted in partial reversal of flower inhibition with 0.3 mg/l GA3 being optimal. The inhibition of vegetative growth as measured by fresh and dry weight was also partially reversed by GA3, but the threshold concentration for reversal of flower inhibition was at least 10 times lower than that for inhibition of vegetative growth. These results are interpreted as indicating that gibberellins are important for flowering in the non-rosette long-day plant L. gibba G3, but apparently are present in non-limiting concen- trations on short days. Gibberellic acid (GA3) is known to cause stem of the plant growth retardants CCC [(2-chloroethyl) elongation and flower induction in many rosette trimethv lammonium chloride] and AMO 1618 (2-iso- long-day plants (11,12). Induction of flowering by propyl-4 -dimethylamino-5-methvlphenyl-l -piperidine- GA3 has also been reported in the non-rosette long- carboxvlate methyl chloride) which are thought to day plant Loliumn temnulentuni (6), the long-short-day act primarily bv inhibiting endogenous gibberellin plant Bryophyllum daigremitonitianunt (22), and the biosynthesis (5, 7, 10, 15, 18). The application of short-day plant Impatiens balsamnina (14). How- CCC or AMO 1618 to the long-day rosette plant, ever, in each of these last 3 cases flower induction, Sacitolts parvifloruts, the long-short-day plant, Bryo- whether by photoperiodic treatment or GA3 applica- phyllunm daigremontianum, and also the short-day tion, is accompanied by considerable stem elongation. plant, Pharbitis .il, when grown on inductive photo- In contrast, with the exception of Lolitrn and Im- periods, resulted in complete or nearly complete in- patietns, gibberellin treatment does not result in flower hibition of flowering with only a small accompanying induction in caulescent or non-rosette long-day plants inhibition, of growth (1, 23.24). In each case the and short-day plants (all of which are caulescent) simultaneous application of GA3 reversed the inhibi- (12) and may even prove inhibitory to flowering in tion of both flowering ancd growth, but a 10 to 50-fold a few cases (8, 19, 21). On the basis of such results higher concentration of GA3 was required to reverse it appears that endogenous gibberellins may be im- the inhibition of growth than was needed to reverse portant for flowering at least in those plants in which the inhibition of flowering. flower induction is accompanied by considerable stem These results provided further evidence that gib- elongation. berellins might be important for flower induction in In recent years further evidence for the involve- long-day rosette plants and in the long-short-day ment of gibberellins in the flowering process of at plant Bryophyllumn. In addition, the results with least some plants has been obtained throtigh the use Pharbitis suggested that even in plants for which GA3 treatment does not result in flower induction, gibberellins might be involved in the flowering process. In view of these results it was felt that 1 Supported by grant GB-2846 to W.R. Briggs from plant the National Science Foundation, a grant from Research similar experiments on a non-rosette long-day Corporation to W. R. Briggs, and by a National Science might provide additional in,formation on the role of Foundation Predoctoral fellowship to C. F. Cleland. gibberellins in flowering. 2 Present address: Biological Laboratories, Harvard The non-rosette long-day plant used in this study University, Cambridge, Massachusetts 02138. was Lemina gibba G3 which previously has been 503 504 PLANT PHYSIOLOGY shown to exlhibit a qualitative long-day flowering of magnitude. It has been assumed that autoclaving response with a critical daylength of about 10 hr has a similar effect in the present study. However, (3). The present paper reports the effect of GA3 the concentrations of GA3 listed are those that were and the plant growth retardant CCC on flowering actually incorporated in the medium and do not and growth in this particular non-rosette long-day represent estimates of the effective concentrations. plant. It has also been reported that CCC may be some- what unstable to high temperature treatment (2). Consequently, each of the experiments to be pre- Materials and Methods sented has been repeated with the GA3 and CCC added to previously autoclaved medium by sterile The aquatic flowering plant Lemna gibba L., filtration. In each case there was complete qualita- strain G3 was grown aseptically in 125 ml Erlenmeyer tive agreement with the results obtained when GA3 flasks with 50 ml of E medium. The E medium is and CCC were autoclaved with the medium. There- based oni the MA medium of Hillman (9) and differs fore, it is clear that the results to be presented are only in the addition o,f 30 ,uMNM ethylenediamine.etra- due to the GA3 and CCC in the medium and not to acetic acid (EDTA). All experiments lasted 11 days any breakdown products that might possibly have and were carried out in model MB-54 growth cham- resulted from the autoclaving. bers '(Percival Refrigeration and Manufacturing Company) at a temperature of 28±3- 1° and a light Results intensity of 600 to 700 ft-c at plant level. The light source consisted of 4 cool-white VHO fluorescent The application of GA3 to L. gibba G3 when lights (Sylvania F48T12-CW-VHO) supplemented growing on short days did not result in flower in- with four 25 watt incandescent bulbs. Short-day duction at any of the concentration,s that were tried conditions consisted of 9 hr light followed by 15 hr (Fig. 1). At higher concentrations GA3 inhibited of darkness (9L:15D), while lon,g-day conditions growth somewhat as indicated by a slight drop in the consisted of continuous light. A more detailed de- fresh and dry weights with a minimum at 10 mg/l scription of the medium, culture conditions and gen- GA3. Furthermore, at GA3 concentrations of 10 eral experimental procedures used in this study has mg/l and higher the fronds were noticeably smaller been given earlier (3). and less gibbous than in the short-day control. How- The total frond numnber of a culture was deter- ever, at these higher GA3 concentrations, the rate of mined by counting all fronds, no matter how small, frond production was increased, and there was an whichl visibly projected beyond the margin of their increase in the # VF. mother frond. For the evaluation of flowering the flowering percent (FL %) was determined by divid- 100 ing the numiiber of fronds with flowers or flower SD TREATMENT primordia by the total number of fro,nds examined 80 100 o and multiplying this value by 100. Flowering was 0 0 also evaluated by determining the total number of z VF 80 m vegetative fronds (# VF) in a culture. Providing LLJ 60 ~~~~~~~~l'& G) the growth rate for different treatments is approxi- 0 ) 0.04. mately the same, there is always an inverse relation- n 40 0.4 60 mx ship between the FL % and # VF values with the z (;~~~~~~~~~~~)0 LL- z change in the # VF often being more dramatic 20 0.2 _---FW8., 0.02 40 0 than the corresponding change in the FL %. CCC was dissolved in the medium and used at 0 - t - 0 20 M. 0l concentrationls from 10- to 10-3 It was donated 0 0.11 iO 100 by the Agricultural Division of the American Cvan- (SDC) amid Company, Princeton, New Jersey. GA3 (K GA CONCENTRATION, mg/l salt, 75 %, Nutritional Biochemical Company) was FIG. 1. Influence of GA3 on flowering and growth dissolved in the medium and used at concentrations on short days. from 0.001 to 100 mg/l (corresponds to actual molarity of 2.17 X 10-9 to 2.17 X 10-4 M). When The application of GA3 to plants growing on CCC or GA3 were used they were present in the long days had an inhibitory effect on flowering at medium for the full duration of the experiment. concentrations of 1 mg/l and above (Fig. 2). At It is well known that gibberellins are somewhat these higher concentrations there was also a definite unstable to heat (20). Hillman (8) investigated the inhibition of growth with a substantial decrease in effect of autoclaving for 10 min at 15 psi on the GA3 fresh weight resulting from the fronds being rather activity for the growth of dwarf pea seedlings. His small and only slightly gibbous. The dry weight, results indicated that autoclaving the GA3 with the however, showed only a slight decrease at higher medium resulted in a loss in activitv of about 1 order GA3 concentrations.
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