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Regulation of flowering time by FVE, a retinoblastoma-associated protein.

The initiation of flowering in plants is controlled by environmental and endogenous signals. Molecular analysis of this process in Arabidopsis thaliana indicates that environmental control is exerted through the photoperiod and vernalization pathways, whereas endogenous signals regulate the autonomous and gibberellin pathways. The vernalization and autonomous pathways converge on the negative regulation of FLC, a gene encoding a MADS-box protein that inhibits flowering. We cloned FVE, a component of the autonomous pathway that encodes AtMSI4, a putative retinoblastoma-associated protein. FVE interacted with retinoblastoma protein in immunoprecipitation assays, and FLC chromatin was enriched in acetylated histones in fve mutants. We conclude that FVE participates in a protein complex repressing FLC transcription through a histone deacetylation mechanism. Our data provide genetic evidence of a new developmental function of these conserved proteins and identify a new genetic mechanism in the regulation of flowering.

Arabidopsis↗

A genetic link between cold responses and flowering time through FVE in Arabidopsis thaliana.

Cold induces expression of a number of genes that encode proteins that enhance tolerance to freezing temperatures in plants. A cis-acting element responsive to cold and drought, the C-repeat/dehydration-responsive element (C/DRE), was identified in the Arabidopsis thaliana stress-inducible genes RD29A and COR15a and found in other cold-inducible genes in various plants. C/DRE-binding factor/DRE-binding protein (CBF/DREB) is an essential component of the cold-acclimation response, but the signaling pathways and networks are mostly unknown. Here we used targeted genetic approach to isolate A. thaliana mutants with altered cold-responsive gene expression (acg) and identify ACG1 as a negative regulator of the CBF/DREB pathway. acg1 flowered late and had elevated expression of FLOWERING LOCUS C (FLC), a repressor of flowering encoding a MADS-box protein. We showed that acg1 is a null allele of the autonomous pathway gene FVE. FVE encodes a homolog of the mammalian retinoblastoma-associated protein, a component of a histone deacetylase (HDAC) complex involved in transcriptional repression. We also showed that plants sense intermittent cold stress through FVE and delay flowering with increasing expression of FLC. Dual roles of FVE in regulating the flowering time and the cold response may have an evolutionary advantage for plants by increasing their survival rates.

Arabidopsis↗

Biological effect of radiation-degraded alginate on flower plants in tissue culture.

Alginate with a weight-average molecular mass (Mw) of approx. 9.04 x 10(5) Da was irradiated at 10-200 kGy in 4% (w/v) aqueous solution. The degraded alginate product was used to study its effectiveness as a growth promoter for plants in tissue culture. Alginate irradiated at 75 kGy with an Mw of approx. 1.43 x 10(4) Da had the highest positive effect in the growth of flower plants, namely limonium, lisianthus and chrysanthemum. Treatment of plants with irradiated alginate at concentrations of 30-200 mg/l increased the shoot multiplication rate from 17.5 to 40.5% compared with control. In plantlet culture, 100 mg/l irradiated alginate supplementation enhanced shoot height (9.7-23.2%), root length (9.7-39.4%) and fresh biomass (8.1-19.4%) of chrysanthemum, lisianthus and limonium compared with that of the untreated control. The survival ratios of the transferred flower plantlets treated with irradiated alginate were almost the same as the control value under greenhouse conditions. However, better growth was attained for the treated plantlets.

Alginates↗

Composition and antimicrobial activity of the essential oil of Artemisia absinthium from Croatia and France.

The essential oils obtained by steam distillation from the aerial parts of two populations of Artemisia absinthium, from France and from Croatia, were analyzed by GC and GC-MS. The oils of A. absinthium of French origin contain (Z)-epoxyocimene and chrysanthenyl acetate as major components while the oils of Croatian A. absinthium contain mainly (Z)-epoxyocimene and beta-thujone. Analysis of oils before and after anthesis showed some quantitative differences. Analysis of separated leaves and flowering heads showed only few differences among these organs. As they contain no thujone, antimicrobial screening was performed on samples of French origin and showed that A. absinthium oil inhibited the growth of both tested yeasts (Candida albicans and Saccharomyces cerevisiae var. chevalieri).

Antifungal Agents↗

New triterpene esters from flowerheads of Arnica lonchophylla.

From the flowerheads of Arnica lonchophylla Greene ssp. lonchophylla Maguire (Asteraceae), a variety of mono-, di- and trihydroxytriterpenes of the oleanane, ursane, lupane and dammarane types were isolated and their structures elucidated by EI and CI mass spectrometry and extensive NMR spectroscopic analysis. Most of the compounds are esterified at position 3 with lauric, myristic, palmitic and stearic acids. Several ester derivatives of known triterpenes represent new natural products. The triterpenetriols 20(30)-taraxasten-3beta,16beta,21alpha-triol (arnitriol A) and lupane-3beta,16beta,20-triol, both isolated in the form of their C3-fatty acid ester derivatives, possess hydroxylation patterns which have not been described previously. Besides small amounts of triterpenetriol esters, the lauric, myristic, palmitic and stearic acid esters of arnidiol, faradiol, maniladiol and calenduladiol were also identified in the flowerheads of A. montana (Arnicae flos Ph. Eur.), A. chamissonis ssp. foliosa and A. angustifolia ssp. attenuata. A search for sesquiterpene lactones in the flowers of A. lonchophylla resulted in the identification of small amounts of helenalin and 11alpha,13-dihydrohelenalin.

Arnica↗

Hydroxyl radical and hypochlorous acid scavenging activity of small centaury (Centaurium erythraea) infusion. A comparative study with green tea (Camellia sinensis).

Small centaury (Centaurium erythraea Rafin.) is a herbal species with a long use in traditional medicine due to its digestive, stomachic, tonic, depurative, sedative and antipyretic properties. This species is reported to contain considerable amounts of polyphenolic compounds, namely xanthones and phenolic acids as the main constituents. Although the antiradicalar activity of some pure polyphenolic compounds is already known, it remains unclear how a complex mixture obtained from plant extracts functions against reactive oxygen species. Thus, the ability of small centaury infusion to act as a scavenger of the reactive oxygen species hydroxyl radical and hypochlorous acid was studied and compared with that of green tea (Camellia sinensis L.). Hydroxyl radical was generated in the presence of Fe3+-EDTA, ascorbate and H2O2 (Fenton system) and monitored by evaluating hydroxyl radical-induced deoxyribose degradation. The reactivity towards hypochlorous acid was determined by measuring the inhibition of hypochlorous acid-induced 5-thio-2-nitrobenzoic acid oxidation to 5,5'-dithiobis(2-nitrobenzoic acid). The obtained results demonstrate that small centaury infusion exhibits interesting antioxidant properties, expressed both by its capacity to effectively scavenge hydroxyl radical and hypochlorous acid, although with a lower activity against the second than that observed for green tea. Green tea exhibited a dual effect at the hydroxyl radical scavenging assay, stimulating deoxyribose degradation at lower dosages.

Antioxidants↗

Paternal leakage sustains the cytoplasmic polymorphism underlying gynodioecy but remains invasible by nuclear restorers.

Cytoplasmic male sterility (CMS) in plants often results in gynodioecious populations, composed of hermaphrodites and male-sterile females. All models of gynodioecy assume maternal inheritance of the cytoplasmic alleles and postulate a variety of negatively frequency-dependent mechanisms to maintain the cytoplasmic polymorphisms observed in many natural populations. However, in some plant species, mitochondria are transmitted at least occasionally by pollen, a process called paternal leakage. We show that even a small amount of paternal leakage is sufficient to sustain a permanent, stable cytoplasmic polymorphism. Because only hermaphrodites provide pollen in gynodioecious species, the effects of paternal leakage are biased and occur more often from the non-CMS male-fertile haplotype to the CMS male-sterile haplotype. We also show that a nuclear restorer disrupts the polymorphic cytoplasmic equilibrium, leading to fixation of both the CMS allele and the restorer. Although a dominant nuclear restorer fixes, it fixes much more slowly than in the standard CMS models. Although a stable cytonuclear polymorphism is possible with "matching alleles" nuclear restoration, oscillations to low frequencies present a risk of loss by drift. Paternal leakage enhances the stability of joint cytonuclear polymorphism by reducing the chance that a CMS allele is lost by drift.

Cell Nucleus↗

Interaction between drought and chronic high temperature during kernel filling in wheat in a controlled environment.

Wheat plants (Triticum aestivum L. 'Lyallpur'), limited to a single culm, were grown at day/night temperatures of either 18/13 degrees C (moderate temperature), or 27/22 degrees C (chronic high temperature) from the time of anthesis. Plants were either non-droughted or subjected to two post-anthesis water stresses by withholding water from plants grown in different volumes of potting mix. In selected plants the demand for assimilates by the ear was reduced by removal of all but the five central spikelets. In non-droughted plants, it was confirmed that shading following anthesis (source limitation) reduced kernel dry weight at maturity, with a compensating increase in the dry weight of the remaining kernels when the total number of kernels was reduced (small sink). Reducing kernel number did not alter the effect of high temperature following anthesis on the dry weight of the remaining kernels at maturity, but reducing the number of kernels did result in a greater dry weight of the remaining kernels of droughted plants. However, the relationship between the response to drought and kernel number was confounded by a reduction in the extent of water stress associated with kernel removal. Data on the effect of water stress on kernel dry weight at maturity of plants with either the full complement or reduced numbers of kernels, and subjected to low and high temperatures following anthesis, indicate that the effect of drought on kernel dry weight may be reduced, in both absolute and relative terms, rather than enhanced, at high temperature. It is suggested that where high temperature and drought occur concurrently after anthesis there may be a degree of drought escape associated with chronic high temperature due to the reduction in the duration of kernel filling, even though the rate of water use may be enhanced by high temperature.

Acclimatization↗

Gender variation of sequential inflorescences in a monoecious plant Sagittaria trifolia (Alismataceae).

In protogynous plants, female flowers of early blooming plants are at a reproductive disadvantage because they cannot set fruit due to the lack of available pollen. To study this phenomenon, gender expression of the monoecious herb Sagittaria trifolia was investigated over the entire flowering season in two field and two cultivated populations in Hubei and Hunan Provinces, China. In racemes of S. trifolia, flowers open sequentially from bottom to top, with female flowers opening first followed by male flowers. This creates a temporal separation of sexes in the species. Under field conditions small plants are often male, with production of both male and female flowers increasing with plant size. Femaleness increased among sequential inflorescences since female flower production increased whereas male flower production did not. Seed production was greater in large inflorescences because they contain more female flowers, and the number of ovules increased in female flowers at basal positions within the raceme. A consistent pattern of high seed set was observed in flowers from both field and cultivated populations. About 1 % of unfertilized ovules resulted from no pollination and 2 % of the seeds produced were only partly developed due to resource limitation. In the first inflorescence of the six experimental populations, 6.7-40.0 % of individuals produced only male flowers, and female flowers of 1.9-6.5 % individuals were aborted. The occurrence of male flowers in early blooming inflorescences could be an adaptive strategy to conserve resources and enhance pollination of female flowers in protogynous S. trifolia.

Analysis of Variance↗

Hypergravity stimulus enhances primary xylem development and decreases mechanical properties of secondary cell walls in inflorescence stems of Arabidopsis thaliana.

BACKGROUND AND AIMS: The xylem plays an important role in strengthening plant bodies. Past studies on xylem formation in tension woods in poplar and also in clinorotated Prunus tree stems lead to the suggestion that changes in the gravitational conditions affect morphology and mechanical properties of xylem vessels. The aim of this study was to examine effects of hypergravity stimulus on morphology and development of primary xylem vessels and on mechanical properties of isolated secondary wall preparations in inflorescence stems of arabidopsis. METHODS: Morphology of primary xylem was examined under a light microscope on cross-sections of inflorescence stems of arabidopsis plants, which had been grown for 3-5 d after exposure to hypergravity at 300 g for 24 h. Extensibility of secondary cell wall preparation, isolated from inflorescence stems by enzyme digestion of primary cell wall components (mainly composed of metaxylem elements), was examined. Plants were treated with gadolinium chloride, a blocker of mechanoreceptors, to test the involvement of mechanoreceptors in the responses to hypergravity. KEY RESULTS: Number of metaxylem elements per xylem, apparent thickness of the secondary thickenings, and cross-section area of metaxylem elements in inflorescence stems increased in response to hypergravity. Gadolinium chloride suppressed the effect of hypergravity on the increase both in the thickness of secondary thickenings and in the cross-section area of metaxylem elements, while it did not suppress the effect of hypergravity on the increase in the number of metaxylem elements. Extensibility of secondary cell wall preparation decreased in response to hypergravity. Gadolinium chloride suppressed the effect of hypergravity on cell wall extensibility. CONCLUSIONS: Hypergravity stimulus promotes metaxylem development and decreases extensibility of secondary cell walls, and mechanoreceptors were suggested to be involved in these processes.

Arabidopsis↗

Genetic analysis of early flowering mutants in Arabidopsis defines a class of pleiotropic developmental regulator required for expression of the flowering-time switch flowering locus C.

The Arabidopsis flowering-repressor gene FLOWERING LOCUS C (FLC) is a developmental switch used to trigger floral induction after extended growth in the cold, a process termed vernalization. In vernalized plants, FLC becomes transcriptionally silenced through a process that involves an epigenetic mechanism. We identified recessive mutations designated vernalization independence (vip) that confer cold-independent flowering and suppression of FLC. These mutations also lead to developmental pleiotropy, including specific defects in floral morphology, indicating that the associated genes also have functions unrelated to flowering time. We identified the VIP3 gene by positional cloning and found that it encodes a protein consisting almost exclusively of repeated Trp-Asp (WD) motifs, suggesting that VIP3 could act as a platform to assemble a protein complex. Constitutive transgenic expression of VIP3 in vernalized plants is insufficient to activate FLC, and thus VIP3 probably participates in the regulation of FLC as one component of a more extensive mechanism. Consistent with this, genetic analyses revealed that the VIP loci define a functional gene class including at least six additional members. We suggest that VIP3 and other members of this gene class could represent a previously unrecognized flowering mechanism.

Arabidopsis↗

Genotype-environment interactions at quantitative trait loci affecting inflorescence development in Arabidopsis thaliana.

Phenotypic plasticity and genotype-environment interactions (GEI) play a prominent role in plant morphological diversity and in the potential functional capacities of plant life-history traits. The genetic basis of plasticity and GEI, however, is poorly understood in most organisms. In this report, inflorescence development patterns in Arabidopsis thaliana were examined under different, ecologically relevant photoperiod environments for two recombinant inbred mapping populations (Ler x Col and Cvi x Ler) using a combination of quantitative genetics and quantitative trait locus (QTL) mapping. Plasticity and GEI were regularly observed for the majority of 13 inflorescence traits. These observations can be attributable (at least partly) to variable effects of specific QTL. Pooled across traits, 12/44 (27.3%) and 32/62 (51.6%) of QTL exhibited significant QTL x environment interactions in the Ler x Col and Cvi x Ler lines, respectively. These interactions were attributable to changes in magnitude of effect of QTL more often than to changes in rank order (sign) of effect. Multiple QTL x environment interactions (in Cvi x Ler) clustered in two genomic regions on chromosomes 1 and 5, indicating a disproportionate contribution of these regions to the phenotypic patterns observed. High-resolution mapping will be necessary to distinguish between the alternative explanations of pleiotropy and tight linkage among multiple genes.

Arabidopsis↗

Are there associations between grain-filling rate and photosynthesis in the flag leaves of field-grown rice?

Rate of grain filling in terms of dry mass accumulated per panicle per day was measured in field-grown rice in the dry season in the Philippines and compared to rates of light-saturated photosynthesis per unit leaf area (P(max)) measured at 350 micro l l(-1) CO(2) for 21 d after flowering. Five new plant type (tropical japonica) varieties (NPT) and one indica variety (IR72) were used and these gave some variation in rates and patterns of grain filling. A rapid grain-filling phase (RGFP) occurred approximately 10 d after flowering in most varieties. There was no consistent relationship in any variety between the rate of grain-filling and P(max) and chlorophyll content, both of which remained mostly unchanged throughout grain filling. Significant declines in the amount of total leaf protein and ribulose bisphosphate carboxylase-oxygenase (Rubisco) occurred, but these did not occur at the same time as the RGFP in all varieties. A decrease in the ratio of chlorophyll a/b preceded these changes and a transient rise in chlorophyll content was also observed in four varieties at this time. There was no significant change in leaf non-structural carbohydrate content during or following the RGFP. It is concluded that the decline in Rubisco and protein content in NPT was not reflected in photosynthetic activity. Hence in these field experiments Rubisco accumulated to a level in excess of photosynthetic requirements, serving as a store of nitrogen for grain filling.

Carbon Dioxide↗

Characterization and genetic analysis of a lettuce (Lactuca sativa L.) mutant, weary, that exhibits reduced gravitropic response in hypocotyls and inflorescence stems.

A lettuce (Lactuca sativa L.) mutant that exhibits a procumbent growth habit was identified and characterized. In two wild type (WT) genetic backgrounds, segregation patterns revealed that the mutant phenotype was controlled by a recessive allele at a single locus, which was designated weary. Hypocotyls and inflorescence stems of plants homozygous for the weary allele exhibited reduced gravitropic responses compared with WT plants, but roots exhibited normal gravitropism. Microscopic analysis revealed differences in the radial distribution of amyloplasts in hypocotyl and inflorescence stem cells of weary and WT plants. Amyloplasts occurred in a single layer of endodermal cells in WT hypocotyls and inflorescence stems. By contrast, amyloplasts were observed in several layers of cortical cells in weary hypocotyls, and weary inflorescence stem cells lacked amyloplasts entirely. These results are consistent with the proposed role of sedimenting amyloplasts in shoot gravitropism of higher plants. The phenotype associated with the weary mutant is similar to that described for the Arabidopsis mutant sgr1/scr, which is defective in radial patterning and gravitropism.

Flowering Tops↗

Sorbitol uptake is regulated by glucose through the hexokinase pathway in vegetative peach-tree buds.

In peach trees (Prunus persica L. Batsch cv. Redhaven), sorbitol is a primary photosynthetic product and may play an important role in the budbreak process. Surprisingly, before budbreak (from January to early March), the concentration of sorbitol in the xylem sap decreases, while that of hexoses (glucose and fructose) increases. The aim of this work was to study the control of sorbitol uptake into vegetative buds by hexoses. Sorbitol uptake was selectively inhibited by hexoses at low and physiological concentrations and this effect was both reversible and concentration-dependent. In addition, the active uptake of sorbitol significantly declined in the plasma membrane vesicles-enriched fraction purified from glucose-treated vegetative buds, suggesting that the inhibitory action of glucose was at the membrane level. Finally, among several glucose analogues tested, only hexokinase substrates (2-deoxyglucose and mannose) were able to mimic the glucose effect, which was completely blocked by the hexokinase inhibitor mannoheptulose. These results represent the first steps towards a better understanding of polyol transport control in plants.

Biological Transport↗

Post-anthesis development of inferior and superior spikelets in rice in relation to abscisic acid and ethylene.

The purpose of this study was to test the hypothesis that the interaction between abscisic acid (ABA) and ethylene may be involved in mediating the post-anthesis development of spikelets in rice (Oryza sativa L.). Two rice genotypes were field-grown, and the changes of ABA, ethylene, and 1-aminocylopropane-1-carboxylic acid (ACC) levels in spikelets during grain filling and their relationships with endosperm-division and grain-filling rates were investigated. The results showed that earlier-flowering superior spikelets exerted dominance over later-flowering inferior spikelets in endosperm cell-division and grain-filling rates. The two genotypes behaved the same. Later-flowering spikelets had higher levels of ethylene and ACC than earlier-flowering spikelets. The ethylene evolution rate was significantly and negatively correlated with the cell division and grain filling rates. By contrast to ethylene, later-flowering spikelets contained a lower ABA content/concentration and showed a low content ratio of ABA to ACC than earlier-flowering ones. The cell-division and grain-filling rates were significantly and positively correlated with both ABA contents and the ratio of ABA to ACC. Application of cobalt ion (inhibitor of ethylene synthesis) or ABA at an early grain-filling stage significantly increased endosperm cell division rate and cell number, grain-filling rate, and grain weight of inferior spikelets. Application of ethephon (an ethylene-releasing agent) or fluridone (an inhibitor of carotenoid synthesis) had the opposite effect. The results suggest that antagonistic interactions between ABA and ethylene mediate endosperm cell-division and grain-filling in rice. A higher ratio of ABA to ethylene in rice spikelets is required to maintain a faster grain-filling rate.

Abscisic Acid↗

Adaptive evolution in the photosensory domain of phytochrome A in early angiosperms.

Flowering plant diversity now far exceeds the combined diversity of all other plant groups. Recently identified extant remnants of the earliest-diverging lines suggest that the first angiosperms may have lived in shady, disturbed, and moist understory habitats, and that the aquatic habit also arose early. This would have required the capacity to begin life in dimly lit environments. If so, evolution in light-sensing mechanisms may have been crucial to their success. The photoreceptor phytochrome A is unique among angiosperm phytochromes in its capacity to serve a transient role under conditions where an extremely high sensitivity is required. We present evidence of altered functional constraints between phytochrome A (PHYA) and its paralog, PHYC. Tests for selection suggest that an elevation in nonsynonymous rates resulted from an episode of selection along the branch leading to all angiosperm PHYA sequences. Most nucleotide sites (95%) are selectively constrained, and the ratio of nonsynonymous to synonymous substitutions on branches within the PHYA clade does not differ from the ratio on the branches in the PHYC clade. Thus, positive selection at a handful of sites, rather than relaxation of selective constraints, apparently has played a major role in the evolution of the photosensory domain of phytochrome A. The episode of selection occurred very early in the history of flowering plants, suggesting that innovation in phyA may have given the first angiosperms some adaptive advantage.

Evolution, Molecular↗

Hd3a, a rice ortholog of the Arabidopsis FT gene, promotes transition to flowering downstream of Hd1 under short-day conditions.

Heading date 3a (Hd3a) has been detected as a heading-date-related quantitative trait locus in a cross between rice cultivars Nipponbare and Kasalath. A previous study revealed that the Kasalath allele of Hd3a promotes heading under short-day (SD) conditions. High-resolution linkage mapping located the Hd3a locus in a approximately 20-kb genomic region. In this region, we found a candidate gene that shows high similarity to the FLOWERING LOCUS T (FT) gene, which promotes flowering in Arabidopsis: Introduction of the gene caused an early-heading phenotype in rice. The transcript levels of Hd3a were increased under SD conditions. The rice Heading date 1 (Hd1) gene, a homolog of CONSTANS (CO), has been shown to promote heading under SD conditions. By expression analysis, we showed that the amount of Hd3a mRNA is up-regulated by Hd1 under SD conditions, suggesting that Hd3a promotes heading under the control of Hd1. These results indicate that Hd3a encodes a protein closely related to Arabidopsis FT and that the function and regulatory relationship with Hd1 and CO, respectively, of Hd3a and FT are conserved between rice (an SD plant) and Arabidopsis (a long-day plant).

Amino Acid Sequence↗