The effect of daylength on the transition to flowering in phytochrome-deficient, late-flowering and double mutants of Arabidopsis thaliana

The effect of daylength on flowering was investigated in the following mutants of Arabidopsis thaliana : phytochrome B deficient ( hy3=phyB ); phytochrome chromophore deficient ( hy2 ); late-flowering ( co, gi. fca and fwa ); the hy2 and hy3 , late-flowering double mutants and the hy2, hy3 , late-flowering triple mutants. The hy mutants flower with fewer rosette leaves than the Landsberg erecta wild type under both long day and short day conditions and express this effect to a different degree in all late-flowering mutant backgrounds and under both daylengths, with the exception of fca under short days. The number of cauline leaves and days to flowering is less affected by the hy genotype. The hy2, hy3 double mutants flower with even fewer rosette leaves than the hy2 and hy3 monogenic mutants, suggesting an inhibitory role for phytochrome B and other stable phytochromes on flowering. The complex interaction between phytochrome, daylength and the effect of the late-flowering genes on the various parameters that describe the transition to flowering in Arabidopsis is discussed.     

Flowering responses to light-breaks in photomorphogenic mutants of Arabidopsis thaliana, a long-day plant

Flowering response and plant form of photomorphogenic mutants (hy1, hy2, hy3, hy4 and hy5) of Arabidopsis thaliana (L.), a long-day plant, were examined in long and short days. There were only slight differences among genotypes including Landsberg wild type with respect to the flowering time under long days. The effect of 1 h light-(night)-breaks of far-red, red, blue and white light given in the middle of the dark period of plants grown under short days, was studied. Effects of far-red light applied at the end or the beginning of the main photoperiod on flowering and plant form were also examined. The light-breaks with all the above mentioned light qualities promoted floral initiation of all the genotypes including the wild type in terms of both the flowering time and the number of rosette leaves. In general, far-red light was most effective. It is possible to classify the hy-mutants into 3 groups by their responses to light-breaks under short day conditions: (a) Mutants hy2 and hy3, which have a reduced number of rosette leaves, and flower early. Red light is as effective as far-red light. The wavelength of light-breaks is relatively unimportant for flowering response. (b) Mutants hy4, hy5 and Landsberg wild type, which have a greater number of rosette leaves, and flower relatively late. The effectiveness of light-breaks is in the following order, far-red, blue, and red light, which is in reverse order to the transformation of phytochrome to the P_fr form. (c) Mutant hy1, which behaves anomalously with respect to relations between flowering time and number of rosette leaves; late flowering with reduced number of rosette leaves. Red, blue and far-red light are effective, but white light is ineffective for reducing the number of rosette leaves. When far-red light was given in the middle of the night or at the end of the main photoperiod, it markedly reduced the number of rosette leaves compared to those grown under short days for all the genotypes, while when applied at the beginning of the main photoperiod far-red light did not affect the number of rosette leaves. Different effects on the plant form dependent on the time of treatment with far-red light-breaks are also discussed.     

Pleiotropic effects of flowering time genes in the annual crucifer Arabidopsis thaliana (Brassicaceae)

Variation in flowering time of Arabidopsis thaliana was studied in an experiment with mutant lines. The pleiotropic effects of flowering time genes on morphology and reproductive yield were assessed under three levels of nutrient supply. At all nutrient levels flowering time and number of rosette leaves at flowering varied among mutant lines. The relationship between these two traits depended strongly on nutrient supply. A lower nutrient supply first led to an extension of the vegetative phase, while the mean number of leaves at flowering was hardly affected. A further reduction resulted in no further extension of the vegetative phase and, on average, plants started flowering with a lower leaf number. At low nutrients, early flowering affected the timing of production of siliques rather than the total output, whereas late flowering was favorable at high nutrients. This may explain the fact that many plant species flower at a relatively small size under poor conditions. Flowering time genes had pleiotropic effects on the leaf length, number of rosette and cauline leaves, and number of axillary flowering shoots of the main inflorescence. Silique production was positively correlated with the number of axillary shoots of the main inflorescence; the number of axillary primordia appeared to have a large impact on reproductive yield.     

Day and night temperature responses in Arabidopsis: effects on gibberellin and auxin content,cell size,morphology and flowering time

The effect of 16 different day (DT) and night (NT) temperature combinations (DT and NT 12, 17, 22 and 27 degrees C) on rosette leaf growth, flower stem elongation and flowering time in Arabidopsis thaliana Ler was investigated. Final leaf length decreased with increasing NT due to a combination of reduced elongation period and reduced elongation rate. Final stem length increased with increasing DT due to increased elongation rate, and decreased with increasing NT due to a decrease in elongation period. Under NT 27 degrees C, however, stem elongation rate increased greatly, resulting in the same final stem length as under NT 12 degrees C. The transition to flowering was accelerated by increasing NT. A linear regression analysis was performed to clarify the relationship between final leaf length, final stem length and flowering time with DIF (DT minus NT) and/or ADT (average daily temperature). For all three variables, the effect of DIF depended on ADT and vice versa. The relationship of final stem length with DIF also depended on the temperature range. Increased cell volume in flower stems developing at DT/NT 22/12 degrees C gave rise to longer and thicker stems compared with stems developing at DT/NT 12/22 degrees C. GC-MS analysis (gas chromatography-mass spectrometry) showed that the endogenous level of IAA was 56 % higher in stems grown under DT/NT 22/12 degrees C compared with DT/NT 12/22 degrees C. Of the 12 gibberellins analysed, however, only the level of non-bioactive GA29 was affected by the temperature treatment.     

拟南芥蓝光受体CRY2的功能及其介导的光形态建成

植物具备一套复杂的由两种蓝光受体和多种信号转导下游组分组成的蓝光感应系统,通过感受光照强度、光的方向和光周期,调节自身对蓝光的应答.蓝光反应的有效波长是蓝光和近紫外光(320～400nm),故蓝光受体也叫蓝光/近紫外光受体.CRY2(Cryptochromes,CRY)是一个核蛋白,在转录水平受蓝光的调节,它的作用是增加拟南芥对蓝光的敏感性.植物蓝光调节的反应主要有向光性、抑制幼茎伸长、叶绿体迁移、刺激气孔张开和调节基因表达等.对植物蓝光反应突变体分子生物学研究进展进行了综述,对蓝光受体及信号转导下游组分在植物发育中的作用及蓝光诱发植物作出反应的分子机制进行了探讨.     

Characterization of a novel developmentally retarded mutant (drm1) associated with the autonomous flowering pathway in Arabidopsis

INTRODUCTIONThe transition from vegetative growth to reproductionis one of the most important developmental events in flow-ering plants since it is related to the competence and sur-vivability of a particular species living in a particularenvironment. The…     

Conserved structure and function of the Arabidopsis flowering time gene CONSTANS in Brassica napus

The Arabidopsis thaliana CONSTANS (CO) gene which promotes flowering in long days was recently isolated by chromosome walking. The mapping of QTLs controlling flowering time in Brassica species has identified genomic regions that contain homologues of the CO gene. Four genes homologous to the Arabidopsis CO gene were isolated from a pair of homoeologous loci in each of two doubled-haploid Brassica napus lines displaying different flowering times, N-o-1 and N-o-9. The four genes, BnCOa1, BnCOa9, BnCOb1 and BnCOb9, are located on linkage groups N10 and N19, and are highly similar to each other and to the Arabidopsis CO gene. Two regions of the proteins are particularly well conserved, a N-terminal region with two putative zinc fingers and a C-terminal region which may contain a nuclear localization signal. All four genes appear to be expressed in B. napus. The BnCOa1 allele was shown to complement the co-2 mutation in Arabidopsis in a dosage-dependent manner causing earlier flowering than in wild type under both long- and short-day conditions.     

模式植物拟南芥开花时间分子调控研究进展

植物从营养生长到生殖生长的转变是开花发育的关键,在合适的时间开花对植物的生长和繁衍极为重要,植物开花时间的调控对农业生产发展意义重大。植物开花是由遗传因子和环境因子协同调节的一个复杂过程。近年来,对不同植物开花调控的研究,特别是对模式植物拟南芥（Arabidopsis thaliana（L.） Heynh.）的开花调控研究取得了显著进展,已探明开花时间分子调控的6条主要途径分别是光周期途径、春化途径、自主途径、温度途径、赤霉素途径和年龄途径。各遗传调控途径既相互独立又相互联系,构成一个复杂的开花调控网络。本文综述了模式植物拟南芥开花时间调控分子机制相关研究的最新进展,并对未来的研究进行了展望。     

Indirect effects of FRIGIDA: floral trait (co)variances are altered by seasonally variable abiotic factors associated with flowering time

Reproductive timing is a critical life‐history event that could influence the (co)variation of traits developing later in ontogeny by regulating exposure to seasonally variable factors. In a field experiment with Arabidopsis thaliana, we explore whether allelic variation at a flowering‐time gene of major effect (FRIGIDA) affects (co)variation of floral traits by regulating exposure to photoperiod, temperature, and moisture levels. We detect a positive latitudinal cline in floral organ size among plants with putatively functional FRI alleles. Statistically controlling for bolting day removes the cline, suggesting that seasonal abiotic variation affects floral morphology. Both photoperiod and precipitation at bolting correlate positively with the length of petals, stamens, and pistils. Additionally, floral (co)variances differ significantly across FRI backgrounds, such that the sign of some floral‐trait correlations reverses. Subsequent experimental manipulations of photoperiod and water availability demonstrate direct effects of these abiotic factors on floral traits. In sum, these results highlight how the timing of life‐history events can affect the expression of traits developing later in ontogeny, and provide some of the first empirical evidence for the effects of major genes on evolutionary potential.     

The effect of far-red light on the photoperiodic regulation of carbohydrate accumulation in Allium cepa L.

Carbohydrate accumulation was induced in Allium cepa L. cv. Dorata di Parma by the addition of far-red light throughout the 18 h day. Neither an 18 h-day without far-red nor the addition of far-red light to a 10 h photoperiod could induce the carbohydrate accumulation. The accumulation of reducing sugar takes place both in the leaf-blades and in the bulb; while there is accumulation of oligosaccharides only in the bulb. The results implicate phytochrome in the promotion of carbohydrate accumulation by far-red light.     

Phytochrome control of flowering is temperature sensitive and correlates with expression of the floral integrator FT

In Arabidopsis flowering is accelerated by reduced red:far-red (R:FR) ratio which signals the presence of neighbouring vegetation. Hastened flowering is one component of the shade-avoidance syndrome of responses, which alter many aspects of development in response to the threat of potential competition. Of the red/far-red-absorbing photoreceptors it is phyB that plays the most prominent role in shade-avoidance, although other related phytochromes act redundantly with phyB. It is well established that the phyB mutant has a constitutively early flowering phenotype. However, we have shown that the early flowering phenotype of phyB is temperature-dependent. We have established that this temperature-sensitive flowering response defines a pathway that appears to be independent of the autonomous-FLC pathway. Furthermore, we have demonstrated that the phytochromes control the expression of the floral promoter FT. We have also shown that other phyB-controlled responses, including petiole elongation, are not sensitive to the same temperature change. This suggests that discrete pathways control flowering and petiole elongation, components of the shade-avoidance response. This work provides an insight into the phytochrome and temperature interactions that maintain flowering control.     

VASCULAR PLANT ONE-ZINC FINGER1 and VOZ2 repress the FLOWERING LOCUS C clade members to control flowering time in Arabidopsis

Floral transition is regulated by environmental and endogenous signals. Previously, we identified VASCULAR PLANT ONE-ZINC FINGER1 (VOZ1) and VOZ2 as phytochrome B-interacting factors. VOZ1 and VOZ2 redundantly promote flowering and have pivotal roles in the downregulation of FLOWERING LOCUS C (FLC), a central repressor of flowering in Arabidopsis. Here, we showed that the late-flowering phenotypes of the voz1 voz2 mutant were suppressed by vernalization in the Columbia and FRIGIDA (FRI)-containing accessions, which indicates that the late-flowering phenotype of voz1 voz2 mutants was caused by upregulation of FLC. We also showed that the other FLC clade members, MADS AFFECTING FLOWERING (MAF) genes, were also a downstream target of VOZ1 and VOZ2 as their expression levels were also increased in the voz1 voz2 mutant. Our results suggest that the FLC clade genes integrate signals from VOZ1/VOZ2 and vernalization to regulate flowering.     

Overexpression of COL9, a CONSTANS-LIKE gene, delays flowering by reducing expression of CO and FT in Arabidopsis thaliana

CONSTANS (CO) is an important floral regulator in the photoperiod pathway, integrating the circadian clock and light signal into a control for flowering time. It is known that CO promotes flowering in Arabidopsis under long-day conditions. CONSTANS-LIKE 9 (COL9) is a member of the CONSTANS-LIKE gene family, encoding a nuclear protein. The expression of COL9 is regulated by the circadian clock in the photoperiod pathway and is detected in various organs. Unexpectedly, overexpression of COL9 in transgenic Arabidopsis resulted in delayed flowering, while co-suppression lines and a transferred DNA (T-DNA) knockout line showed earlier flowering under long-day conditions. Overexpression of COL9 did not enhance the late-flowering phenotype in a co mutant background. Double overexpressors produced by overexpression of CO in COL9 transgenic lines showed an early flowering phenotype similar to single CO overexpressors. The pattern of oscillation of a number of circadian-associated genes remained unchanged in the COL9 transgenic lines. Compared with wild-type plants, the abundance of CO and FLOWERING LOCUS T (FT) mRNA was reduced in the COL9 overexpression lines. Our results indicate that COL9 is involved in regulation of flowering time by repressing the expression of CO, concomitantly reducing the expression of FT and delaying floral transition.     

Both phytochrome A and phytochrome B are required for the normal expression of phototropism in Arabidopsis thaliana seedlings

The role of phytochrome A (phyA) and phytochrome B (phyB) in phototropism was investigated by using the phytochrome-deficient mutants phyA-101 , phyB-1 and a phyA/phyB double mutant. The red-light-induced enhancement of phototropism, which is normally observed in wild-type seedlings, could not be detected in the phyA/phyB mutant at fluences of red light between 0.1 and 19 000 μmol m⁻². The loss of phyB has been shown to have no apparent effect on enhancement, while the loss of phyA resulted in a loss of enhancement only in the low fluence range (Janoudi et al. 1997). The conclusions of the aforementioned study can now be modified based on the current results which indicate that phototropic enhancement in the high fluence range is mediated by either phyA or phyB, and that other phytochromes have no role in enhancement. First positive phototropism was unaffected in phyA-101 and phyB-1 However, the magnitude of first positive phototropism in the phyA/phyB mutant was significantly lower than that of the wild-type Landsberg parent. Thus, the presence of either phyA or phyB is required for normal expression of first positive phototropism. The time threshold for second positive phototropism is unaltered in the phyA-101 and phyB mutants. However, the time threshold in the phyA/phyB mutant is about 2 h, approximately six times that of the wild type. Finally, the magnitude of second positive phototropism in both phyA-101 and phyB-1 is diminished in comparison with the wild-type response. Thus, phyA and phyB, acting independently or in combination, regulate the magnitude of phototropic curvature and the time threshold for second positive phototropism. We conclude that the presence of phyA and phyB is required, but not sufficient, for the expression of normal phototropism.     

Effect of white and far-red light on betalain formation

Light-induced betalain synthesis has been studied in three species of Amaranthaceae. The results obtained suggest that the response to short-term irradiation with white light is mediated by phytochrome, while the effects of prolonged illumination are controlled by the photosynthetic system. Betalain accumulation stimulated by prolonged far-red irradiation is controlled chiefly by the photosynthetic system, the participation of phytochrome being of minor importance.     

Integrated metabolite and gene expression profiling revealing phytochrome A regulation of polyamine biosynthesis of Arabidopsis thaliana

In this study, metabolite profiling was demonstrated as a usefultool to plot a specific metabolic pathway, which is regulatedby phytochrome A (phyA). Etiolated Arabidopsis wild-type (WT)and phyA mutant seedlings were irradiated with either far-redlight (FR) or white light (W). Primary metabolites of the irradiatedseedlings were profiled by gas chromatography time-of-flightmass spectrometry (GC/TOF-MS) to obtain new insights on phyA-regulatedmetabolic pathways. Comparison of metabolite profiles in phyAand WT seedlings grown under FR revealed a number of metabolitesthat contribute to the differences between phyA and the WT.Several metabolites, including some amino acids, organic acids,and major sugars, as well as putrescine, were found in smalleramounts in WT compared with the content in phyA seedlings grownunder FR. There were also significant differences between metaboliteprofiles of WT and phyA seedlings during de-etiolation underW. The polyamine biosynthetic pathway was investigated further,because putrescine, one of the polyamines existing in a widevariety of living organisms, was found to be present in loweramounts in WT than in phyA under both light conditions. Theexpression levels of polyamine biosynthesis-related genes wereinvestigated by quantitative real-time RT-PCR. The gene expressionprofiles revealed that the arginine decarboxylase 2 (ADC2) genewas transcribed less in the WT than in phyA seedlings underboth light conditions. This finding suggests that ADC2 is negativelyregulated by phyA during photomorphogenesis. In addition, S-adenosylmethioninedecarboxylase 2 and 4 (SAMDC2 and SAMDC4) were found to be regulatedby phyA but in a different manner from the regulation of ADC2. Key words: Arabidopsis thaliana, gene expression profiling, metabolite profiling, phytochrome A, polyamine biosynthesis Received 19 October 2007; Revised 17 January 2008 Accepted 18 January 2008     

Variation at two flowering time genes within and among populations of Arabidopsis thaliana: comparison with markers and traits

Flowering Locus C (FLC) and Frigida are two interacting genes controlling flowering time variation in Arabidopsis thaliana. Variation at these genes was surveyed in 12 A. thaliana populations sampled in France. These populations were also screened for variation at molecular markers [12 microsatellites and 19 cleaved amplified polymorphic sequence (CAPS) markers] and at seven quantitative traits measured with and without vernalization. Seven populations were highly polymorphic at markers (H(S) = 0.57 at microsatellites, 0.24 at CAPS) and showed heritable variation for bolting time and some other traits. Five populations were genetically fixed or nearly fixed. Q(ST) for bolting time without vernalization was significantly higher than F(ST), suggesting local divergent selection. One of the two haplotype groups at FLC (FLC(A)) was very predominant (frequency of 99%). The first exon of Frigida showed elevated nonsynonymous variation, and nine loss-of-function mutations were found throughout the gene. The association between loss-of-function and earlier bolting was confirmed. Overall, 18 Frigida haplotypes were detected. The pattern of variation at Frigida was largely similar to that found at markers and traits, with the same populations being fixed or highly diverse. Metapopulation dynamics is thus probably the main factor shaping genetic variation in A. thaliana. However, F(ST) for functional (FRI) vs. nonfunctional (FRI(Delta)) haplotypes was significantly higher than F(ST) at markers. This suggested that loss-of-function at Frigida is under local selection for flowering time.     

Effects of red, far-red and blue light in maintaining growth in latitudinal populations of Norway spruce (Picea abies)

Seedlings of trees with a free growth pattern cease growth when night-lengths become shorter than a critical value, and this critical night-length (CNL) decreases with increasing latitude of origin. In northern populations, the light quality also appears to play an important role and a clinal variation in requirement for far-red (FR) light has been documented. In this study we dissected the light quality requirements for maintaining growth in different latitudinal populations of Norway spruce (Picea abies (L.) H. Karst.) using light emitting diodes for red (R), FR and blue (B) light, as 12 h day extension to provide 24 h photoperiod. At equal spectral photon flux, FR light was more effective than R light in maintaining growth, and the requirement of both R and FR increased with northern latitude of origin. One-to-one mixtures of R and FR light were more effective in maintaining growth than either FR or R light alone, indicating a possible interaction between R and FR light maintaining growth. Using the blue light as day extension could not prevent growth cessation in any of the populations, but delayed the bud set slightly in all populations. Our results suggest that phytochrome(s) are the primary photoreceptors in high irradiance responses maintaining growth in Norway spruce seedlings.