Long-day induction of flowering in Lolium temulentum involves sequential increases in specific gibberellins at the shoot apex

One challenge for plant biology has been to identify floral stimuli at the shoot apex. Using sensitive and specific gas chromatography-mass spectrometry techniques, we have followed changes in gibberellins (GAs) at the shoot apex during long day (LD)-regulated induction of flowering in the grass Lolium temulentum. Two separate roles of GAs in flowering are indicated. First, within 8 h of an inductive LD, i.e. at the time of floral evocation, the GA(5) content of the shoot apex doubled to about 120 ng g(-1) dry weight. The concentration of applied GA(5) required for floral induction of excised apices (R.W. King, C. Blundell, L.T. Evans [1993] Aust J Plant Physiol 20: 337-348) was similar to that in the shoot apex. Leaf-applied [(2)H(4)] GA(5) was transported intact from the leaf to the shoot apex, flowering being proportional to the amount of GA(5) imported. Thus, GA(5) could be part of the LD stimulus for floral evocation of L. temulentum or, alternatively, its increase at the shoot apex could follow import of a primary floral stimulus. Later, during inflorescence differentiation and especially after exposure to additional LD, a second GA action was apparent. The content of GA(1) and GA(4) in the apex increased greatly, whereas GA(5) decreased by up to 75%. GA(4) applied during inflorescence differentiation strongly promoted flowering and stem elongation, whereas it was ineffective for earlier floral evocation although it caused stem growth at all times of application. Thus, we conclude that GA(1) and GA(4) are secondary, late-acting LD stimuli for inflorescence differentiation in L. temulentum.     

The Control of Apical Bud Growth and Senescence by Auxin and Gibberellin in Genetic Lines of Peas

Pea (Pisum sativum L.) lines G2 (dwarf) and NGB1769 (tall) (Sn Hr) produce flowers and fruit under long (LD) or short (SD) days, but senesce only under LD. Endogenous gibberellin (GA) levels were inversely correlated with photoperiod (over 9-18 h) and senescence: GA20 was 3-fold and GA1 was 10- to 11-fold higher in flowering SD G2 shoots, and the vegetative tissues within the SD apical bud contained 4-fold higher levels of GA20, as compared with the LD tissues. Prefloral G2 plants under both photoperiods had GA1 and GA20 levels similar to the flowering plants under LD. Levels of indole-3-acetic acid (IAA) were similar in G2 shoots in LD or SD; SD apical bud vegetative tissues had a slightly higher IAA content. Young floral buds from LD plants had twice as much IAA as under SD. In NGB1769 shoots GA1 decreased after flower initiation only under LD, which correlated with the decreased growth potential. We suggest that the higher GA1 content of G2 and NGB1769 plants under SD conditions is responsible for the extended vegetative growth and continued meristematic activity in the shoot apex. This and the increased IAA level of LD floral buds may play a role in the regulation of nutrient partitioning, since more photosynthate partitions of reproductive tissue under LD conditions, and the rate of reproductive development in LD peas is faster than under SD.     

The nature of floral signals in Arabidopsis. II. Roles for FLOWERING LOCUS T (FT) and gibberellin

Signals produced in leaves are transported to the shoot apex where they cause flowering. Protein of the gene FLOWERING LOCUS T (FT) is probably a long day (LD) signal in Arabidopsis. In the companion paper, rapid LD increases in FT expression associated with flowering driven photosynthetically in red light were documented. In a far red (FR)-rich LD, along with FT there was a potential role for gibberellin (GA). Here, with the GA biosynthesis dwarf mutant ga1-3, GA(4)-treated plants flowered after 26 d in short days (SD) but untreated plants were still vegetative after 6 months. Not only was FT expression low in SD but applied GA bypassed some of the block to flowering in ft-1. On transfer to LD, ga1-3 only flowered when treated simultaneously with GA, and FT expression increased rapidly (<19.5 h) and dramatically (15-fold). In contrast, in the wild type in LD there was little requirement for GA for FT increase and flowering so its endogenous GA content was near to saturating. Despite this permissive role for endogenous GA in Columbia, RNA interference (RNAi) silencing of the GA biosynthesis gene, GA 20-OXIDASE2, revealed an additional, direct role for GA in LD. Flowering took twice as long after silencing the LD-regulated gene, GA 20-OXIDASE2. Such independent LD input by FT and GA reflects their non-sympatric expression (FT in the leaf blade and GA 20-OXIDASE2 in the petiole). Overall, FT acts as the main LD floral signal in Columbia and GA acts on flowering both via and independently of FT.     

Regulation of flowering in the long-day grass Lolium temulentum by gibberellins and the FLOWERING LOCUS T gene

Seasonal control of flowering often involves leaf sensing of daylength coupled to time measurement and generation and transport of florigenic signals to the shoot apex. We show that transmitted signals in the grass Lolium temulentum may include gibberellins (GAs) and the FLOWERING LOCUS T (FT) gene. Within 2 h of starting a florally inductive long day (LD), expression of a 20-oxidase GA biosynthetic gene increases in the leaf; its product, GA(20), then increases 5.7-fold versus short day; its substrate, GA(19), decreases equivalently; and a bioactive product, GA(5), increases 4-fold. A link between flowering, LD, GAs, and GA biosynthesis is shown in three ways: (1) applied GA(19) became florigenic on exposure to LD; (2) expression of LtGA20ox1, an important GA biosynthetic gene, increased in a florally effective LD involving incandescent lamps, but not with noninductive fluorescent lamps; and (3) paclobutrazol, an inhibitor of an early step of GA biosynthesis, blocked flowering, but only if applied before the LD. Expression studies of a 2-oxidase catabolic gene showed no changes favoring a GA increase. Thus, the early LD increase in leaf GA(5) biosynthesis, coupled with subsequent doubling in GA(5) content at the shoot apex, provides a substantial trail of evidence for GA(5) as a LD florigen. LD signaling may also involve transport of FT mRNA or protein because expression of LtFT and LtCONSTANS increased rapidly, substantially (>80-fold for FT), and independently of GA. However, because a LD from fluorescent lamps induced LtFT expression but not flowering, the nature of the light response of FT requires clarification.     

Changes in the Pattern of Cell Division in the Shoot and Root Meristems of Lolium perenne during the Transition from Vegetative to Floral Growth

For Lolium perenne cv. Cropper, a system which resulted in 100%flowering comprised 90 short days (SD) at 4 ?C (vernalization)and 30 SD at 18 ?C followed by 8 long days (LD). The mitoticindex and G1 and G2 percentages were measured in the shoot androot apices of plants following 2, 5 or 8 LD and in SD controlssampled at the beginning and end of induction. Identical measurementswere made in plants given 48 SD at 18 ?C followed by 2, 5 or8 LD; plants remained vegetative in response to this treatmentlacking vernalization. Significant increases in both mitoticindex and meristem size occurred in the shoot apex in LD followingthe vernalizing, but not the non-vernalizing, treatment. A clusterof mitoses in the apical dome of the shoot apex was unique tothe vernalized plants given 5 or 8 LD. However, an increasein root meristem size occurred regardless of vernalization,but a significant increase in the mitotic index was limitedto vernalized plants given 5 or 8 LD. Whilst the vernalization-LDtreatment resulted in an increase in the G2 percentage in theshoot apex following 2, 5 or 8 LD, no such alteration was observedin the root meristem. Thus, the changes to the cell cycle whichcorrelated with flowering were increased mitotic indices andG2 percentages in the shoot apex at each sampling time and increasedmitotic indices in the root apex following 5 and 8 LD. Key words: Cell division, flowering, Lolium perenne L.     

Restricted cell/cell communication in the shoot apex ofSilene coeli-rosa during the transition to flowering is associated with a high mitotic index rather than with evocation

Summary Plants ofSilene coeli-rosa given 5 or more long days (LDs) flowered, even when the LDs were followed by 48 hours of darkness before their return to short days (SDs). The mitotic indices of shoot apices from induced plants shortly after induction were significantly higher than the indices of shoot apices from vegetative plants. Two major mitotic peaks were observed in the shoot apices of plants given 7 long days (LDs) on day 8. One coincided with that reported byFrancis andLyndon (1979).Cell to cell movement was tested in the shoot apices of vegetative and LD treated plants using probes with a molecular size of 749 daltons (fluorescein-hexaglycine) and 847 daltons (fluorescein-leucyl diglutamyl leucine). These probes showed some movement in the shoot apices of both short day (SD) and LD treated plants, but fluorescein-leucyl diglutamyl leucine was immobile in the induced apices of 7 LD plants on day 8 at time intervals which coincided with major mitotic activity in the shoot apex. Symplasmic restriction in the shoot apex was also observed in plants given 8 LDs (i.e., plants not returned to SDs on day 7).In plants that were placed in 48 hours of darkness after the 7 LD treatment or in plants given 5 LDs, there was no strong peak in the mitotic index, even though all these LD treatments resulted in 100% flowering. In such plants no symplasmic restriction was found in the shoot. Thus the symplasmic restriction on day 8 of 7 LD plants is associated with the high mitotic index, but neither of these phenomena is an essential part of the evocation process.Abbreviations F(Glu)₂ L-glutamylglutamic acid conjugated to fluorescein isothiocyanate isomer I (F-) - F(Gly)₆ F-hexaglycine - FLGGL F-leucyl-diglutamyl-leucine - F(PPG)₅ F-the pentamer (propyl-propyl glycine) - LD long day - LDs long days - SD short day - SDs short days     

Synthesis of gibberellin GA6 and its role in flowering of Lolium temulentum

The induction of flowering by one long day (LD) in the grass Lolium temulentum is most closely mimicked by application of the gibberellins (GAs) GA(5) or GA(6), both of which occur naturally. These gibberellins promote floral development but have little effect on stem elongation. Endogenous GA(5) and GA(6) contents in the shoot apex double on the day after the LD and, for GA(5) (and we presume for GA(6) as well) reach a concentration known to be inductive for the excised shoot apex in vitro. They are, therefore, strong candidates as LD floral stimuli in this grass. The synthesis of GA(6) and an examination of its florigenic properties in L. temulentum are described.     

Gibberellins and the floral transition in Sinapis alba

The putative role of gibberellins in the transition to flowering was investigated in Sinapis alba , a caulescent long-day (LD) plant. It was observed that: (1) physiological doses of exogenous gibberellins (GA₁, GA₃, GA₉) do not cause the floral shift of the meristem when applied to plants grown in short days but have some positive effect on the flowering response to a suboptimal LD; no inhibition was observed in any case; (2) GA-biosynthesis inhibitors (prohexadione-Ca and paclobutrazol) considerably inhibit stem growth but have some negative effect on flowering only when a suboptimal LD is given; and (3) the floral transition induced by one 22-h LD does not correlate with any detectable change in GA content of the apical bud, of the leaves, and of the phloem exudate reaching the apex. Taken together, these results suggest that GAs do not act as a major signal for photoperiodic flower induction in Sinapis .     

Cellular changes induced by exogenous and endogenous gibberellins in shoot tips of the long-day plant Silene armeria

Stem elongation and flowering are two processes induced by long-day (LD) treatment in Silene armeria L. Whereas photoperiodic control of stem growth is mediated by gibberellins (GAs), the flowering response cannot be obtained by GA applications. Microscopic observations on early cellular changes in the shoot meristem following LD induction or GA treatment in short days (SD) were combined with GA analyses of stem sections at various distances below the shoot apex. The earliest effects of both LD and GA induction on the subapical meristem were an increase in the number of cells per cell file and a reduction of cell length in the meristematic tissue approx. 1.0–3.0 mm below the shoot apex. Within 8 d after the beginning of LD induction or after GA application, the cells in the subapical meristem were oriented in long files. In induced tips, cellulose deposition occurred mostly in longitudinal walls, indicating that many transverse cell divisions had taken place which, in turn, increased the length of the stem. In contrast to LD induction, GA treatments did not promote the transition from the vegetative to the floral stage. Endogenous GAs were analyzed by selected ion monitoring (SIM), using labeled internal standards, in extracts from transverse sections of the tip at various distances below the apical meristem. In control plants, the levels of the six 13-hydroxy GAs studied (GA₅₃, GA₄₄, GA₁₉, GA₂₀, GA₁, and GA₈) decreased as the distance from the apical meristem increased. Except for GA₅₃, GA levels were higher in tips of LD-induced plants, particularly in the meristematic zone approx. 0.5–1.5 mm below the apical meristem. In comparison with SD, the highest increase observed was for GA₁, the content of which increased 30-fold in the zone 0.5–3.5 mm below the shoot apex. These data indicate a spatial correlation between the accumulation of GA₁ and its precursors, and the enhanced mitotic activity which occurs in the subapical meristem of elongating Silene apices.Abbreviations GA_n gibberellin A_n - LD long day(s) - SD short day(s) We thank Dr. L.N. Mander, Australian National University, Canberra, for providing [²H]- gibberellins, Dr. B.O. Phinney, University of California, Los Angeles, USA, for [¹³C]GA₈, Dr. D.A. Gage, MSU-NIH Mass Spectrometry Facility, for advice with mass spectrometry, and Mr. M. Chassagne, I.N.R.A. C.R. Bordeaux, for the photography. This work was supported, in part, by a fellowship from the Spanish Ministry of Agriculture (Instituto Nacional de Investigaciones Agrarias) to M.T., by the U.S. Department of Energy under contract DE-ACO2-76ERO-1338, and by the U.S. Department of Agriculture grant No. 88-37261-3434 to J.A.D.Z.     

Flowering of the grass Lolium perenne: effects of vernalization and long days on gibberellin biosynthesis and signaling

Almost 50 years ago, it was shown that gibberellin (GA) applications caused flowering in species normally responding to cold (vernalization) and long day (LD). The implication that GAs are involved with vernalization and LD responses is examined here with the grass Lolium perenne. This species has an obligatory requirement for exposure to both vernalization and LD for its flowering (inflorescence initiation). Specific effects of vernalization or LD on GA synthesis, content, and action have been documented using four treatment pairs: nonvernalized or vernalized plants exposed to short days (SDs) or LDs. Irrespective of vernalization status, exposure to two LDs increased expression of L. perenne GA 20-oxidase-1 (LpGA20ox1), a critical GA biosynthetic gene, with endogenous GAs increasing by up to 5-fold in leaf and shoot. In parallel, LD led to degradation of a DELLA protein, SLENDER (within 48 h of LD or within 2 h of GA application). There was no effect on GA catabolism or abscisic acid content. Loss of SLENDER, which is a repressor of GA signaling, confirms the physiological relevance of increased GA content in LD. For flowering, applied GA replaced the need for LD but not that for vernalization. Thus, GAs may be an LD, leaf-sourced hormonal signal for flowering of L. perenne. By contrast, vernalization had little impact on GA or SLENDER levels or on SLENDER degradation following GA application. Thus, although vernalization and GA are both required for flowering of L. perenne, GA signaling is independent of vernalization that apparently impacts on unrelated processes.     

GAMYB-like Genes, Flowering, and Gibberellin Signaling in Arabidopsis

We have identified three Arabidopsis genes with GAMYB-like activity, AtMYB33, AtMYB65, and AtMYB101, which can substitute for barley (Hordeum vulgare) GAMYB in transactivating the barley alpha-amylase promoter. We have investigated the relationships between gibberellins (GAs), these GAMYB-like genes, and petiole elongation and flowering of Arabidopsis. Within 1 to 2 d of transferring plants from short- to long-day photoperiods, growth rate and erectness of petioles increased, and there were morphological changes at the shoot apex associated with the transition to flowering. These responses were accompanied by accumulation of GAs in the petioles (GA(1) by 11-fold and GA(4) by 3-fold), and an increase in expression of AtMYB33 at the shoot apex. Inhibition of GA biosynthesis using paclobutrazol blocked the petiole elongation induced by long days. Causality was suggested by the finding that, with GA treatment, plants flowered in short days, AtMYB33 expression increased at the shoot apex, and the petioles elongated and grew erect. That AtMYB33 may mediate a GA signaling role in flowering was supported by its ability to bind to a specific 8-bp sequence in the promoter of the floral meristem-identity gene, LEAFY, this same sequence being important in the GA response of the LEAFY promoter. One or more of these AtMYB genes may also play a role in the root tip during germination and, later, in stem tissue. These findings extend our earlier studies of GA signaling in the Gramineae to include a dicot species, Arabidopsis, and indicate that GAMYB-like genes may mediate GA signaling in growth and flowering responses.     

The response of short day plantChenopodium rubrum L. to abscisic acid and gibberellic acid treatment applied at two levels of photoperiodic induction

Abscisic acid (ABA) (5 x 10^-4M and 5 x 10^-5M) and gibberellic acid (1 x 10^-4M) was applied to the plumula ofChenopodium plants with partly (one dark period) or completely (three dark periods) fulfilled photoperiodic requirements for flowering. Morphological and cytoogical criteria were used to investigate the time-course of the differentiation of the treated shoot apices. Both substances were ineffective in increasing the mitotic activity of the shoot apex at the suboptimal level of induction. The degree of branching was temporarily stimulated by ABA and GA treatment under these conditions. Moreover, GA caused the elongation of the shoot apex. With the completely induced plants ABA hastened flowering and the rise in branching was observed in all the treatment 48 h following the application of growth substances.     

Cell Cycle Responses to Red or Far-red Light, or Darkness, in the Shoot Apex of Silene coeli-rosa L. During Floral Induction

Twenty-eight-day-old plants of Silene coeli-rosa L. were maintainedin short days (SD) for 9 d (0–8) or exposed to 7 longdays (LD), or 7 SD with a 5 min exposure at 1700 h of each dayto far-red (FR), red (R) or 5 min FR/5 min R, or 7 dark-interrupted(di = 1700–1720 h) LD. Treatments were followed by twofurther SD. The mitotic index and G1 and G2 proportions weremeasured in the shoot apices of plants sampled at 2000 h ofeach day of each replicated treatment. Exposure to 7 LD (= 100per cent flowering) resulted in significant increases, relativeto the SD controls, in both the G2 proportion and the mitoticindex on d 0 to 3, 7 and 8. Five minute FR (= 0 per cent flowering)resulted in cell cycle responses similar to those in LD onlyfrom d 0 to 2. R and FR/R (both = 0 per cent flowering) didnot result in any increases in the G2 proportion in the apexapart from d 3 of FR/R. However 5 min FR/5 min R, and to a lesserextent 5 min R, did result in significant increases in the mitoticindex on d 0, 1, 7, and 8. diLD (= 8–10 per cent flowering)also prevented any significant increases in the G2 proportionon d 0 to 3, and 5 to 8 but the mitotic index was again higheron these days compared with control data. Thus the transitionto floral growth for 90 per cent of the plants is associatedwith changes in the cell cycle in the shoot apex measured asincreases in the G2 proportion at 2000 h of LD 0 to 3 and 7to 8. Silene coeli-rosa L., cell cycle, flowering, phytochrome, shoot apex     

The relationship between fruit growth and apical senescence in the G2 line of peas

In the G2 line of peas (Pisum sativum L.), senescence of the shoot apex (which precedes leaf senescence) only occurs in long days (LD) though flowering is independent of photoperiod. It has been suggested that the photoperiodic control of senescence in G2 is mediated through different rates of seed growth. In LD seed growth is more rapid than in short days (SD) and this places a greater nutrient drain on the plant. In addition, more flowers develop into fruits in LD than in SD: 32% of flower buds abort in SD while almost none abort in LD. Senescence is associated with early seed growth and does not occur in deflowered or deseeded plants. Seed development is completed in 30d in LD while it takes 40d in SD, though the seed weights are similar. The maximum rate of fresh-weight gain of all the growing seeds of eight fruits on a plant in SD (1,440 mg/d) does not reach the maximum rate of weight gain of a similar fruit complement in LD (1,720 mg/d). The appearance of senescence symptoms in the shoot apices of LD-grown G2 plants occurs, however, prior to the time of the greatest rate of seed-weight gain. In LD, four fruits with a combined maximum growth rate of 1,250 mg/d are sufficient to cause the appearance of senescence symptoms. This is a lower combined seed growth rate than in SD where senescence does not occur. The seeds in up to 12 fruits can be growing at any time in SD with a combined maximum seed-growth rate (1,660 mg/d), only slightly less than the maximum in LD, with no sign of senescence. It is concluded that the different rates of seed growth occasioned by different photoperiods bear no relation to senescence. However, photoperiod does alter the spatial relationship of the shoot apex and the filling fruits. In LD apical growth becomes slower as fruiting proceeds so that the distance between the filling fruits and the apex is decreased to only two nodes while in SD, because of the delayed fruit development compared to LD, the spatial separation between the fruits and the shoot apex is nine nodes. Even if the growth rate of the plant had remained constant in LD it is calculated that an equivalent fruit complement would still be located three nodes further from the apex in SD than in LD. This increased spatial separation of fruits and apex in SD compared to LD probably alters the source/sink distribution of photosynthate and leaf derived hormones so that larger amounts are available to the apex in SD than LD. Also any senescence factor exported from fruits is less likely to reach the apex in SD. In continuously deflorated plants of G2 the two uppermost expanded stipules enclose the apex in SD while in LD they open out. The effect is reversible. Thus photoperiod probably affects the apex and its growth, directly, i.e. independent of fruit development, and this is accentuated by the differing spatial relationships of the apex and fruits resulting from different fruit growth rates under the different photoperiodic conditions.Abbreviations LD long day(s) - SD short day(s)     

Uncoupling light quality from light irradiance effects in Helianthus annuus shoots: putative roles for plant hormones in leaf and internode growth

An attempt has been made to uncouple the effects of the two primary components of shade light, a reduced red to far-red (R/FR) ratio and low photosynthetically active radiation (PAR), on the elongation of the youngest internode of sunflower (Helianthus annuus) seedlings. Maximal internode growth (length and biomass) was induced by a shade light having a reduced R/FR ratio (0.85) under the low PAR of 157 micromol m(-2) s(-1). Reducing the R/FR ratio under normal PAR (421 micromol m(-2) s(-1)) gave similar growth trends, albeit with a reduced magnitude of the response. Leaf area growth showed a rather different pattern, with maximal growth occurring at the higher (normal) PAR of 421 micromol m(-2) s(-1)), but with variable effects being seen with changes in light quality. Reducing the R/FR ratio (by enrichment with FR) gave significant increases in gibberellin A(1) (GA(1)) and indole-3-acetic acid (IAA) contents in both internodes and leaves. By contrast, a lower PAR irradiance had no significant effect on GA(1) and IAA levels in internodes or leaves, but did increase the levels of other GAs, including two precursors of GA(1). Interestingly, both leaf and internode hormone content (GAs, IAA) are positively and significantly correlated with growth of the internode, as are leaf levels of abscisic acid (ABA). However, changes in these three hormones bear little relationship to leaf growth. By implication, then, the leaf may be the major source of GAs and IAA, at least, for the rapidly elongating internode. Several other hormones were also assessed in leaves for plants grown under varying R/FR ratios and PARs. Leaf ethylene production was not influenced by changes in R/FR ratio, but was significantly reduced under the normal (higher) PAR, the irradiance treatment which increased leaf growth. Levels of the growth-active free base and riboside cytokinins were significantly increased in leaves under a reduced R/FR ratio, but only at the higher (normal) PAR irradiance; other light quality treatments evoked no significant changes. Taken in toto, these results indicate that both components of shade light can influence the levels of a wide range of endogenous hormones in internodes and leaves while evoking increased internode elongation and biomass accumulation. However, it is light quality changes (FR enrichment) which are most closely tied to increased hormone content, and especially with increased GA and IAA levels. Finally, the increases seen in internode and leaf GA content with a reduced R/FR ratio are consistent with FR enrichment inducing an overall increase in sunflower seedling GA biosynthesis.     

Response to Gibberellin Structural Variants Shows that Ability to Inhibit Flowering Correlates with Effectiveness for Promoting Stem Elongation of Some Plant Species

The flowering response of three plant species Fuchsia hybrida, Pharbitis nil, and Spathiphyllum "Petite" has been examined after treatment with synthetic and natural gibberellins (GAs) including GA1, GA3, GA5, 2,2-dimethyl GA4, various of their 3a-hydroxyl epimers, the methyl ester of GA3, a 15b-hydroxyl GA3, and some 16,17-dihydro derivatives. Of necessity, application techniques differed between species and dose responses cannot be compared. However, comparisons across species were possible on the basis of their differential responses to the various GAs. Flowering was inhibited and, in an inverse way, stem elongation promoted when GA3 or GA5 was applied to Fuchsia or, at high doses, to Pharbitis. The increased stem growth was apparently responsible for inhibition of flowering because epimerization of the hydroxyl at C-3 of GA3 or modification of GA5 to 16,17-dihydro GA5 created compounds that were growth inactive and that no longer inhibited flowering. However, the response of Pharbitis was more complex because both these GAs and their derivatives promoted flowering at doses subthreshold for growth. For Spathiphyllum, only promotion of flowering was evident with any GA, and the structural variants were slightly active or inactive. Thus, there are at least two antagonistic actions of GA on flowering and for Pharbitis either inhibition or promotion can be shown. The possibility is discussed that the inhibitory action of GA on flowering involves diversion of assimilate away from the shoot apex and into the elongating stem.     

The nature of floral signals in Arabidopsis. I. Photosynthesis and a far-red photoresponse independently regulate flowering by increasing expression of FLOWERING LOCUS T (FT)

Arabidopsis flowers in long day (LD) in response to signals transported from the photoinduced leaf to the shoot apex. These LD signals may include protein of the gene FLOWERING LOCUS T (FT) while in short day (SD) with its slower flowering, signalling may involve sucrose and gibberellin. Here, it is shown that after 5 weeks growth in SD, a single LD up-regulated leaf blade expression of FT and CONSTANS (CO) within 4-8 h, and flowers were visible within 2-3 weeks. Plants kept in SDs were still vegetative 7 weeks later. This LD response was blocked in ft-1 and a co mutant. Exposure to different LD light intensities and spectral qualities showed that two LD photoresponses are important for up-regulation of FT and for flowering. Phytochrome is effective at a low intensity from far-red (FR)-rich incandescent lamps. Independently, photosynthesis is active in an LD at a high intensity from red (R)-rich fluorescent lamps. The photosynthetic role of a single high light LD is demonstrated here by the blocking of the flowering and FT increase on removal of atmospheric CO(2) or by decreasing the LD light intensity by 10-fold. These conditions also reduced leaf blade sucrose content and photosynthetic gene expression. An SD light integral matching that in a single LD was not effective for flowering, although there was reasonable FT-independent flowering after 12 SD at high light. While a single photosynthetic LD strongly amplified FT expression, the ability to respond to the LD required an additional but unidentified photoresponse. The implications of these findings for studies with mutants and for flowering in natural conditions are discussed.     

玉米赤霉烯酮在冬小麦短日春化中的作用

应用酶联免疫技术,研究了冬小麦品种燕大１８１７ 在不同光周期条件下,体内内源玉米赤霉烯酮（ＺＥＮ） 和ＡＢＡ 含量的变化。结果表明,冬小麦品种燕大１８１７ 具有短日春化性,苗期经４ 周ＳＤ 处理后转到ＬＤ 下能正常抽穗。ＳＤ 诱导春化的幼苗茎尖和叶片中ＺＥＮ 含量高于未经诱导的ＬＤ 幼苗,而在两种光周期条件下生长的幼苗中ＡＢＡ 含量变化趋势并无明显差异。马拉硫磷（ ＭＡＬ） 抑制ＳＤ 幼苗体内ＺＥＮ 含量高峰的出现,也部分抑制抽穗;外源ＺＥＮ 可部分抵消ＭＡＬ对植株的影响,降低茎尖内源ＡＢＡ 含量,并有促进小麦拔节的作用。说明在冬小麦短日春化过程中,ＺＥＮ 具有促进作用,而ＡＢＡ 没有直接影响。