Roderick Whitfield King

Gibberellin structure and florigenic activity in Lolium temulentum, a long day plant.

Structural requirements for florigenic activity among gibberellins (GAs) and GA derivatives, including several new ones, applied once to leaves of Lolium temulentum, were examined. The compounds were applied to plants kept either in non-inductive short days (SD) or exposed to one inductive long day (LD). Inflorescence initiation and stem-elongation responses were assessed three weeks later. Among the GAs used, the range in effective dose for inflorescence initiation was more than 1000-fold, but substantially less for stem elongation. Some GAs promoted both stem elongation and inflorescence initiation, some promoted one without the other, and some affected neither. The structural features enhancing florigenic activity were often different from those enhancing stem elongation. Except in the case of 2,2-dimethyl GA4, a double bond in the A ring at either C-1,2 or C-2,3 was essential for high florigenic activity, though not for stem elongation. A free carboxy group was needed for both. Inflorescence initiation in Lolium was enhanced by hydroxylation at C-12, −13 and −15, whereas hydroxylation at C-3 reduced the effect on inflorescence initiation but increased that on stem elongation. A 12β-hydroxyl was more effective than the α epimer for inflorescence initiation whereas the reverse was true for stem elongation. Although such differential effectiveness of GAs for inflorescence initiation and for stem elongation could reflect differences in uptake, transport or metabolism, we suggest that it is indicative of specific structural requirements for inflorescence initiation.

Floral determination and in-vitro floral differentiation in isolated shoot apices of Lolium temulentum L.

Shoot apices of Lolium temulentum excised after exposure of the plants to one long day (LD) undergo floral development in vitro, whereas those excised from plants in short days (SD) remain vegetative. Floral differentiation in vitro is reasonably normal and responds quantitatively to the preceding LD induction when three conditions are met. First, excision must not precede the arrival at the apex of the LD stimulus from the leaves. This appears to begin about 22 h after the start of the LD and is completed after a further 14 h, by which time all isolated apices have become capable of initiating inflorescence differentiation, i.e. florally determined. Second, for apices excised on the day after the LD (Day II) or early on Day III, the presence of gibberellic acid in the medium is required for floral differentiation to occur in most explants. By contrast, neither kinetin (N6-furfurylaminopurine) nor indole-3-acetic acid is required or beneficial, while abscisic acid in the medium is inhibitory to both survival and floral differentiation in excised apices. The third requirement is for an adequate supply of sugar, particularly after floral differentiation begins. Sucrose is taken up rapidly to reach high levels in the excised apices, but high sugar concentration in the medium, either alone or with gibberellic acid, does not suffice for floral differentiation to begin, and there is an absolute requirement for receipt of the LD stimulus.

The relative significance for stem elongation and flowering in Lolium temulentum of 3β-hydroxylation of gibberellins

In previous experiments with many gibberellins (GAs) and GA derivatives applied to Lolium temulentum L., quite different structural requirements were evident for stem elongation on the one hand and for the promotion of flowering on the other. Whereas hydroxylation at carbons 12, 13 and 15 enhanced flowering relative to stem growth, the reverse was the case at carbon 3 (L.T. Evans et al. 1990, Planta 182, 97–106). The significance of hydroxylation at carbon 3 is examined in this paper. The application of inhibitors of 3β-hydroxylation, including C/D-ring-rearranged GAs, reduced stem growth but, in the case of the two acylcyclohexanediones, increased the flowering response when applied on the inductive long day. Later applications of the acylcyclohexanediones, made after floral initiation had occurred, were inhibitory to flowering, suggesting that subsequent inflorescence development requires 3β-hydroxylated GAs. Applications of the 3α-hydroxy epimers of GA1, GA3 and GA4 gave slightly less promotion of flowering in comparison with the 3β-hydroxy GAs, but far less promotion of stem elongation, except in the case of 3-epi-GA4, which was comparable to GA4. The 3α-hydroxy epimer of 2,2-dimethyl GA4 gave less promotion of flowering than its 3β-hydroxy epimer but almost no promotion of stem elongation. The 3α-hydroxy epimers of GA3 and 2,2-dimethyl GA4 did not act as competitive inhibitors of the stem elongation elicited by GA3 and 2,2-dimethyl GA4, respectively. These results extend the differences in GA structure which favour flowering as opposed to stem elongation, and indicate that 3-hydroxylation and its epimeric configuration are of much greater importance to stem elongation than to flower initiation in Lolium.

The differential effects of C-16,17-dihydro gibberellins and related compounds on stem elongation and flowering in Lolium temulentum

Plants of Lolium temulentum L. cv. Ceres grown under short days (SDs) can be induced to initiate inflorescences either by exposure to one long day (LD) or by single applications of some gibberellins (GAs), which also enhance the flowering response to one LD. Single doses of up to 25 μg per plant of C-16, 17-dihydro-GA5 were about as effective as GA5 for promoting flowering after one LD but inhibited stem elongation by up to 40% over three weeks. The promotion of flowering but not the inhibition of elongation by 16, 17-dihydro-GA5 was reduced in SDs or in LDs low in far-red (FR) radiation. With shoot apices cultured in vitro, 16, 17-dihydro-GA5 was more florigenic than GA3 for apices excised after one LD of 14 h or more, but less florigenic for apices excised from plants in shorter days. 16, 17-Dihydro-GA5 was ineffective compared with GA1, GA3 and GA5 for α-amylase production by half-seeds of Lolium, a response concordant with its effect on stem elongation. As with GA5, 16, 17-dihydro derivatives of GA1, GA3, GA20 and several other GAs were more effective for flowering and less effective for stem elongation than the GAs from which they were derived. Hydroxylation at C-17 and/or C-16 generally reduced the effectiveness of 16, 17-dihydro-GA5 for flowering. These results extend the known features of GA structure which favour flowering relative to stem elongation in L. temulentum. Moreover, C-16, 17-dihydro-GA5 mimics, in its daylength- and wavelength-dependence and lack of stem elongation, characteristics of the LD stimulus in L. temulentum.

Shifts in the semidian rhythm of flowering do determine night-break timing and critical night length in Pharbitis nil

Abstract.There is a semidian (≈12 h) rhythm in the flowering response of the short-day plant Pharbitis nil Choisy following 90 min exposure to either far-red light/darkness or a temperature drop (27 °C to 12 °C) given at various times in constant conditions before an inductive dark period. This semidian rhythmic response to the temperature-drop pretreatments in the light is also evident through the inductive dark period without change of phase. Furthermore, those pretreatments which increase flowering also advance the time of maximum sensitivity to red light (R) interruptions of the dark period by up to 1.5 h and shorten the critical night length. Conversely, pretreatments which reduce flowering delay the time of maximum R inhibition by up to 1.5 h and increase the critical night length by the same amount. However the phase of a circadian rhythm of flowering response had no effect on either the time of maximum R inhibition or the critical night length. Thus, the semidian rhythm determines both the time of maximum R inhibition and the critical night length in Pharbitis.