Selection on flowering time in Mediterranean high-mountain plants under global warming

Under climate warming, plants will undergo novel selective pressures to adjust reproductive timing. Adjustment between reproductive phenology and environment is expected to be higher in arctic and alpine habitats because the growing season is considerably short. As early- and late-flowering species reproduce under very different environmental conditions, selective pressures on flowering phenology and potential effects of climate change are likely to differ between them. However, there is no agreement on the magnitude of the benefits and costs of early- vs. late-flowering species under a global warming scenario. In spite of its relevance, phenotypic selection on flowering phenology has rarely been explored in alpine plants and never in Mediterranean high mountain species, where selective pressures are very different due to the summer drought imposed over the short growth season. We hypothesized that late-flowering plants in Mediterranean mountains should present stronger selective pressures towards early onset of reproduction than early-flowering species, because less water is available in the soil as growing season progresses. We performed selection analyses on flowering onset and duration in two high mountain species of contrasting phenology. Since phenotypic selection can be highly context-dependent, we studied several populations of each species for 2 years, covering their local altitudinal ranges and their different microhabitats. Surrogates of biotic selective agents, like fruitset for pollinators and flower and fruit loss for flower and seed predators, were included in the analysis. Differences between the early- and the late-flowering species were less than expected. A consistent negative correlational selection of flowering onset and duration was found affecting plant fitness, i.e., plants that bloomed earlier flowered for longer periods improving plant fitness. Nevertheless, the late-flowering species may experience higher risks under climate warming because in extremely warm and dry years the earlier season does not bring about a longer flowering duration due to summer drought.     

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.     

Studies on the Photoperiodic Responses in Short-Day Plant Helianthus tubevotus

Plants of Helianthus tuberosus, variety white tuber, were treated with various daylengths of 4, 6, 8, 10, 11, 12, 13, 14, 15, 18 h. for 25 days as soon as six leaves on a plant appeard Irradiation for 6–13 h per day induced the plants to form flower buds and flowering, daylength with 14 h or longer kept the plants in vegetative growth. The experiments showed that this variety of Helianthus tuberosus required short days for flowering and the critical daylength was about 13 h. The plants were treated with short days for different durations. At least 17 days were required, Formation of flower buds and flowering had positive correlation with the number of short days over 17 days. After short-day induction, the shorter the daylength is, the more the flower buds inverted. Long-day treatment after an appropriate period of short days wouid reduce the number of flower and induce new vegetative branches from flowering granehes.     

The gibberellic acid biosynthesis mutant ga1-3 of Arabidopsis thaliana is responsive to vernalization.

The Arabidopsis mutant ga1-3 contains a deletion in an enzyme that catalyzes an early step in the synthesis of gibberellic acid. It has been shown that ga1-3 mutant plants cannot flower under 8-h short-day (SD) conditions, even after vernalization. In this article, we present data demonstrating that the ga1-3 mutation does not block the response to vernalization in intermediate photoperiods or in long-day conditions in a late-flowering, vernalization-responsive background. Thus, GA may not have a direct role in the vernalization response in Arabidopsis, but it may be required for an alternate pathway that promotes flowering in noninductive photoperiods.     

Inhibition imposed by developing flowers on further flower-bud initiation in Chamelaucium uncinatum Schau

In Chamelaucium uncinatum, an Australian woody perennial, flower initiation ceases under continuous inductive short-day (SD) conditions after the first flowering flush. The developing flowers were found to be the prime cause of the cessation in flower initiation. Removal of flowering shoots or lowers as soon as the buds appeared resulted in continuous flower formation. Pruning the plants below the young flower buds at the same stage also caused increased flower formation at the tips of the new growth. If pruning was delayed until flower buds were approx. 3 mm in diameter, however, nor further flower initiation took place and the plants, although still under inductive conditions, shifted to vegetative growth. The inhibiting factor is translocated from one branch to another. At least a six-week rest period (a vegetative growth period under long-day conditions) is needed before the plants are able to respond to further SD stimuli.Abbreviations LD long day - SD short day - SE standard error     

Involvement of the ethylene response pathway in dormancy induction in chrysanthemum

Temperature plays a significant role in the annual cycling between growth and dormancy of the herbaceous perennial chrysanthemum (Chrysanthemum morifolium Ramat.). After exposure to high summer temperatures, cool temperature triggers dormancy. The cessation of flowering and rosette formation by the cessation of elongation are characteristic of dormant plants, and can be stimulated by exogenous ethylene. Thus, the ethylene response pathway may be involved in temperature-induced dormancy of chrysanthemum. Transgenic chrysanthemums expressing a mutated ethylene receptor gene were used to assess this involvement. The transgenic lines showed reduced ethylene sensitivity: ethylene causes leaf yellowing in wild-type chrysanthemums, but leaves remained green in the transgenic lines. Extension growth and flowering of wild-type and transgenic lines varied between temperatures: at 20 degrees C, the transgenic lines showed the same stem elongation and flowering as the wild type; at cooler temperatures, the wild type formed rosettes with an inability to flower and entered dormancy, but some transgenic lines continued to elongate and flower. This supports the involvement of the ethylene response pathway in the temperature-induced dormancy of chrysanthemum. At the highest dosage of ethephon, an ethylene-releasing agent, wild-type plants formed rosettes with an inability to flower and became dormant, but one transgenic line did not. This confirms that dormancy is induced via the ethylene response pathway.     

Plant size and season of burn affect flowering and fruiting of the grasstree Xanthorrhoea preissii

Flowering and fruiting were assessed on 14 populations of the grasstree, Xanthorrhoea preissii Endl., occurring in the Darling Range near Perth, Western Australia. Independent of site, season of burn or year of flowering, there was a strong relationship between plant height, which varied from 0.1 to over 2 m, and the incidence of postfire flowering, which varied from 1% (winter burn) to 75% (summer burn) of grasstrees present. There was no relationship between inflorescence dimensions, or flower or fruit production on a spike basis, and plant size/age (height). When standardized for height, spring‐burnt populations produced 40% as many inflorescences as autumn‐burnt populations and 20% as many as summer‐burnt populations. Inflorescences produced by spring‐burnt plants were moderately smaller than those by summer–autumn‐burnt plants. Fruit density per spike in autumn‐burnt plants was 80% of that in spring–summer‐burnt plants. The net effect was an average of 70 000 fruits produced per 100 summer‐burnt plants, 22 000 in autumn‐burnt plants, and 14 000 in spring‐burnt plants. Ecophysiological explanations of these results and their implications for population dynamics have yet to be explored.     

Flowering of Cultivated Green and SAN 9789-Treated Chenopodium rubrum Plants Exposed to White,Blue, and Red Light

Green plants and plants devoid of photosynthetic pigments were compared with regard to their ability to flower under various growth conditions. Green plants of Chenopodium rubrum L. and plants treated with norflurazon SANDOZ-9789 (SAN) were grown on sucrose-containing media with or without hormones (GA₃, BA, IAA, ABA) under short-day photoperiodic or continuous illumination with white, blue, or red light. Green and SAN-treated albino plants produced flowers only under short-day conditions. The flowering of green plants was independent of the presence of sucrose and hormones in the medium as well as of the light quality. The albino plants produced flowers under white and blue light but did not flower in red light. The addition of GA₃ or BA to the medium induced flowering of albino plants exposed to red light. The functional interaction of photoreceptors in the flowering control is discussed.     

Floral initiation in strawberry: Spectral evidence for the regulation of flowering by long-day inhibition

Summary Floral initiation in strawberry cv. Cambridge Favourite, a facultative short-day plant, was inhibited by a daylength extension with red light (R) during the second half of a 16-hour night but not during the first half, and by far-red light (FR) in the first half but not during the second. Mixed R plus FR light was inhibitory to flowering at both times. This change in sensitivity to R and FR light in the evening and morning resembles the pattern for flower induction in long-day plants but differs from the pattern for flower inhibition in several other short-day plants, examples of which are given. These experiments afford further support for the hypothesis that the control of flower initiation in strawberry depends on the production of a flower inhibitor by leaves exposed to long photoperiods.Abbreviations R red - FR far-red - SD short day - LD long day - SDP short-day plant - LDP long-day plant     

小花型菊花新品种的选育

利用南京野生的菊花脑(2n=2x=18)和南京野菊(2n=4x=36)与六倍体栽培菊杂交,或利用中国秋菊与从国外引进的夏、秋两季开花的菊花杂交,育成一组耐夏季高温高湿,花朵繁多,花色艳丽,具有香味的小花型品种。菊花脑和南京野菊在小花型菊花育种中是具有很大潜力的亲本。     

Mutation in TERMINAL FLOWER1 reverses the photoperiodic requirement for flowering in the wild strawberry Fragaria vesca

Photoperiodic flowering has been extensively studied in the annual short-day and long-day plants rice (Oryza sativa) and Arabidopsis (Arabidopsis thaliana), whereas less is known about the control of flowering in perennials. In the perennial wild strawberry, Fragaria vesca (Rosaceae), short-day and perpetual flowering long-day accessions occur. Genetic analyses showed that differences in their flowering responses are caused by a single gene, SEASONAL FLOWERING LOCUS, which may encode the F. vesca homolog of TERMINAL FLOWER1 (FvTFL1). We show through high-resolution mapping and transgenic approaches that FvTFL1 is the basis of this change in flowering behavior and demonstrate that FvTFL1 acts as a photoperiodically regulated repressor. In short-day F. vesca, long photoperiods activate FvTFL1 mRNA expression and short days suppress it, promoting flower induction. These seasonal cycles in FvTFL1 mRNA level confer seasonal cycling of vegetative and reproductive development. Mutations in FvTFL1 prevent long-day suppression of flowering, and the early flowering that then occurs under long days is dependent on the F. vesca homolog of FLOWERING LOCUS T. This photoperiodic response mechanism differs from those described in model annual plants. We suggest that this mechanism controls flowering within the perennial growth cycle in F. vesca and demonstrate that a change in a single gene reverses the photoperiodic requirements for flowering.     

Flowering Phenology and Gender Variation in Pennisetum typhoides

Because of the modular structure of pearl millet (an annual grass crop, Poaceae), different tillers of a plant share the same genotype but are subjected to different environmental conditions during their maturation. This allows investigation of the effects of tiller flowering phenology on allocation to resource-producing photosynthetic biomass, sexual functions, and thus tiller gender. All tillers of plants of two families collected from individual maternal plants (represented by 33 and 31 plants each) were analyzed. In both families, allocation to aboveground vegetative biomass decreased as flowering was delayed. On average, late-flowering tillers were 65% smaller than the first ones to flower. The proportion of biomass allocated to reproduction significantly increased with the flowering rank of the tillers, suggesting that translocations of assimilates occurred between early- and late-flowering tillers. In both families, late-flowering tillers produced significantly fewer pollen grains per stamen than early-flowering ones, and female reproductive allocation (expressed as seed mass per tiller) was also affected by flowering phenology. Tillers became increasingly female as flowering phenology progressed. This gender variation is possibly adaptive because pollination efficiency is maximized by plant height. Natural selection may favor a shift toward femaleness to maximize reproductive fitness in small, late-developing tillers.     

The gentian orthologs of the FT/TFL1 gene family control floral initiation in Gentiana

Gentians are herbaceous perennials blooming in summer through autumn. Although they are popular ornamental flowers in Japan, the regulation of their timing of flowering has not been studied. We identified and characterized gentian orthologs of the Arabidopsis FT/TFL1 gene family to elucidate the mechanisms of flowering initiation. We isolated three gentian orthologs of FT and TFL1, denoted GtFT1, GtFT2 and GtTFL1. Since up-regulation of GtFT1 and GtFT2 as well as down-regulation of GtTFL1 promoted floral initiation in gentian plantlets, these genes affected floral initiation in a similar way to Arabidopsis FT and TFL1. The expression levels of GtFT1 and GtFT2 in leaves of late-flowering gentian increased prior to floral initiation, whereas GtTFL1 was highly expressed in shoot apical meristem at the vegetative stage and decreased drastically just before flowering initiation. Comparison of gene expression patterns showed that GtFT1 expression increased earlier in early-flowering than in late-flowering gentian, whereas the timing of the increase in GtFT2 expression was similar in early- and late-flowering plants. The GtTFL1 expression in early-flowering gentian was extremely low throughout the vegetative and reproductive stages. These results indicated that either the up-regulation of GtFT1 or the down-regulation of GtTFL1 may determine flowering time. Furthermore, we found that early-flowering but not late-flowering gentians have a 320 bp insertion in the promoter region of GtTFL1. Thus, the negligible expression of GtTFL1 in early-flowering lines may be due to this insertion, resulting in a shortened vegetative stage.     

Seasonal change in a pollinator community and the maintenance of style length variation in Mertensia fusiformis (Boraginaceae)

Background and Aims

In sub-alpine habitats, patchiness in snowpack produces marked, small-scale variation in flowering phenology. Plants in early- and late-melting patches are therefore likely to experience very different conditions during their flowering periods. Mertensia fusiformis is an early-flowering perennial that varies conspicuously in style length within and among populations. The hypothesis that style length represents an adaptation to local flowering time was tested. Specifically, it was hypothesized that lower air temperatures and higher frost risk would favour short-styled plants (with stigmas more shielded by corollas) in early-flowering patches, but that the pollen-collecting behaviour of flower visitors in late-flowering patches would favour long-styled plants.

Methods

Floral morphology was measured, temperatures were monitored and pollinators were observed in several matched pairs of early and late populations. To evaluate effects of cold temperatures on plants of different style lengths, experimental pollinations were conducted during mornings (warm) and evenings (cool), and on flowers that either had or had not experienced a prior frost. The effectiveness of different pollinators was quantified as seed set following single visits to plants with relatively short or long styles.

Key Results

Late-flowering populations experienced warmer temperatures than early-flowering populations and a different suite of pollinators. Nectar-foraging bumble-bee queens and male solitary bees predominated in early populations, whereas pollen-collecting female solitary bees were more numerous in later sites. Pollinators differed significantly in their abilities to transfer pollen to stigmas at different heights, in accordance with our prediction. However, temperature and frost sensitivity did not differ between long- and short-styled plants. Although plants in late-flowering patches tended to have longer styles than those in early patches, this difference was not consistent.

Conclusions

Seasonal change in pollinator-mediated selection on style length may help maintain variation in this trait in M. fusiformis, but adaptation to local flowering time is not apparent. The prevalence of short styles in these populations requires further explanation.     

The early-flowering mutant efs is involved in the autonomous promotion pathway of Arabidopsis thaliana.

The transition to flowering is a crucial moment in a plant's life cycle of which the mechanism has only been partly revealed. In a screen for early flowering, after mutagenesis of the late-flowering fwa mutant of Arabidopsis thaliana, the early flowering in short days (efs) mutant was identified. Under long-day light conditions, the recessive monogenic efs mutant flowers at the same time as wild type but, under short-day conditions, the mutant flowers much earlier. In addition to its early-flowering phenotype, efs has several pleiotropic effects such as a reduction in plant size, fertility and apical dominance. Double mutant analysis with several late-flowering mutants from the autonomous promotion (fca and fve) and the photoperiod promotion (co, fwa and gi) pathways of flowering showed that efs reduces the flowering time of all these mutants. However, efs is completely epistatic to fca and fve but additive to co, fwa and gi, indicating that EFS is an inhibitor of flowering specifically involved in the autonomous promotion pathway. A vernalisation treatment does not further reduce the flowering time of the efs mutant, suggesting that vernalisation promotes flowering through EFS. By comparing the length of the juvenile and adult phases of vegetative growth for wild-type, efs and the double mutant plants, it is apparent that efs mainly reduces the length of the adult phase.     

Effect of season of burn on shoot recovery and post‐fire flowering performance in the resprouter Stirlingia latifolia R. Br. (Proteaceae)

Abstract Stirlingia latifolia R. Br. is a proteaceous undershrub that is widespread in open woodlands and heathlands of the south‐west botanical province of Western Australia where it is subject to frequent fires, both natural and the result of fuel‐reduction burns. Shoots are completely destroyed by fire but regenerate rapidly by resprouting numerous new shoots from the root crown. Flowering is strongly triggered by fire with only sparse flowering in fire‐free periods. A study was undertaken to compare regrowth and flowering in populations of S. latifolia burnt in summer or autumn with populations burnt in spring as well as in unburnt population. Post‐fire flowering was recorded in 92% of plants burnt in summer/autumn compared with 73% of plants burnt in spring and less than 3% in populations that had not been burnt for more than 2 years. Plants burnt in summer/autumn resprouted an average of 8.5 shoots from their root crown, of which 93% developed an inflorescence. In contrast, spring‐burnt plants averaged only 5.9 shoots per plant with only 64% of these bearing an inflorescence. Ability to produce flowers was found to be related to plant age, with young individuals producing fewer or no inflorescences following spring burns in comparison with the more prolific flowering of similarly aged individuals following summer/autumn burns. Summer/autumn‐burnt plants also produced significantly longer inflorescence‐bearing shoots bearing a greater numbers of flower heads than those burnt in spring. Possible explanations for these results are given in the present report.     

The floral transition is not the developmental switch that confers competence for the Arabidopsis age-related resistance response to Pseudomonas syringae pv. tomato

Age-related resistance (ARR) is a plant defense response characterized by enhanced resistance to certain pathogens in mature plants relative to young plants. In Arabidopsis thaliana the transition to flowering is associated with ARR competence, suggesting that this developmental event is the switch that initiates ARR competence in mature plants (Rusterucci et al. in Physiol Mol Plant Pathol 66:222–231, 2005). The association of ARR and the floral transition was examined using flowering-time mutants and photoperiod-induced flowering to separate flowering from other developmental events that occur as plants age. Under short-day conditions, late-flowering plant lines ld-1 (luminidependens-1), soc1-2 (suppressor of overexpression of co 1-2), and FRI ⁺ (FRIGIDA) displayed ARR before the transition to flowering occurred. Early-flowering svp-31, svp-32 (short vegetative phase), and Ws-2 were ARR-defective, whereas early-flowering tfl1-14 (terminal flower 1-14) displayed ARR at the same time as Col-0. While svp-31, svp-32 and Ws-2 produced few rosette leaves, tfl1-14 produced a rosette leaf number similar to Col-0, suggesting that the development of a minimum number of rosette leaves is necessary to initiate ARR competence under short-day conditions. Photoperiod-induced transient expression of FT (FLOWERING LOCUS T) caused precocious flowering in short-day-grown Col-0 but this was not associated with ARR competence. Under long-day conditions co-9 (constans-9) mutants did not flower but displayed an ARR response at the same time as Col-0. This study suggests that SVP is required for the ARR response and that the floral transition is not the developmental event that regulates ARR competence.     

Factors Affecting the Flowering of Andropogon gayanus Kunth. Responses to Photoperiod, Temperature and Growth Regulators

Andropogon gayanus is a short-day plant with a critical daylengthfor flowering between 12 and 14 h. Flowering is more intenseas the daylength is shortened from 12 h to 8 h, initiation isalso accelerated by increasing durations of short-day treatment.Plants flower more readily with increasing age. The maximumflowering response can be induced by short-day exposure of onlyone expanding leaf. A fixed number of short days distributedamong different groups of leaves on the plant gave less floweringthan the same number given to only one group. Approximately25°C was optimal for flowering. Root removal treatmentsin short day reduced both growth and flowering responses. Application of the growth regulators indol 3yl-acetic acid,abscisic acid, gibberellic acid or dimethylaminosuccinamic acid(B9) was inhibitory to flowering in all cases but gibberellicacid and B9 applied simultaneously had no inhibitory effect.Growth hormone treatments failed to stimulate flower developmentin long day. These results are discussed in relation to the internal mechanismscontrolling growth and flowering.