Temporal and intraclonal variation of flowering and pseudovivipary in Poa bulbosa

Background and Aims

Versatility in the reproductive development of pseudoviviparous grasses in response to growth conditions is an intriguing reproduction strategy. To better understand this strategy, this study examined variation in flowering and pseudovivipary among populations, co-occurring clones within populations, and among tillers in individual clones of Poa bulbosa, a summer-dormant geophytic grass that reproduces sexually by seed, and asexually by basal tiller bulbs and bulbils formed in proliferated panicles.

Methods

Clones were collected from 17 populations across a rainfall gradient. Patterns of reproduction were monitored for 11 years in a common garden experiment and related to interannual differences in climatic conditions. Intraclonal variation in flowering and pseudovivipary was studied in a phytotron, under daylengths marginal for flowering induction.

Key Results

Clones showed large temporal variability in their reproductive behaviour. They flowered in some years but not in others, produced normal or proliferated panicles in different years, or became dormant without flowering. Proliferating clones did not show a distinct time sequence of flowering and proliferation across years. Populations differed in incidence of flowering and proliferation. The proportion of flowering clones increased with decreasing rainfall at the site of population origin, but no consistent relationship was found between flowering and precipitation in the common garden experiment across years. In contrast, flowering decreased at higher temperatures during early growth stages after bulb sprouting. Pulses of soil fertilization greatly increased the proportion of flowering clones and panicle production. High intraclonal tiller heterogeneity was observed, as shown by the divergent developmental fates of daughter plants arising from bulbs from the same parent clone and grown under similar conditions. Panicle proliferation was enhanced by non-inductive 8 h short days, while marginally inductive 12 h days promoted normal panicles.

Conclusions

Interannual variation in flowering and proliferation in P. bulbosa clones was attributed to differences in the onset of the rainy season, resulting in different daylength and temperature conditions during the early stages of growth, during which induction of flowering and dormancy occurs.     

Photoperiodic flowering of Arabidopsis: integrating genetic and physiological approaches to characterization of the floral stimulus

In many plants the transition from vegetative growth to flowering is controlled by environmental cues. One of these cues is day length or photoperiod, which synchronizes flowering of many species with the changing seasons. Recently, advances have been made in understanding the molecular mechanisms that confer photoperiodic control of flowering and, in particular, how inductive events occurring in the leaf, where photoperiod is perceived, are linked to floral evocation that takes place at the shoot apical meristem. We discuss recent data obtained using molecular genetic approaches on the function of regulatory proteins that control flowering time in Arabidopsis thaliana. These data are compared with the results of physiological analyses of the floral transition, which were performed in a range of species and directed towards identification of the transmitted floral singals.     

Dual induction control of flowering in Leucanthemum vulgare

The perennial herb Leucanthemum vulgare (oxeye daisy) has a dual induction requirement for flowering. The primary induction is a typical low temperature vemalization response. Temperatures up to 15°C are effective, and the optimum is 6–9°C. Short days (SD) during low temperature exposure enhanced primary induction, but SD could not fully substitute for low temperature in primary induction. At optimum temperatures about 6 weeks exposure were required for 100% flowering, but the flowering response increased with increasing exposure up to 12 weeks, especially at higher temperatures. Seedling have a short juvenile phase of about 4 weeks. Populations with origin ranging from 59 to 69°N in Norway did not vary in their primary induction requirements. Long days (LD) were required for inflorescence initiation and stem elongation at 9°C. At 21 and 15°C some plants initiated and developed inflorescences in SD, but the inflorescences were sessile and their development strongly delayed. More than 16 LD cycles were required for normal stem elongation (bolting).     

Gibberellin substitution for long day secondary induction of flowering in Poa pratensis

Plants of Poa pratensis cv. Holt initiate inflorescence primordia when exposed to short days (SD) and low temperature, but require a secondary induction by at least 4 long days (LD) for further inflorescence development and stem elongation. Single or double applications of 10 µg per plant of gibberellins A₁, A₃, A₅ and 16,17‐dihydro A₅ (DHGA₅) induced inflorescence development in a high proportion of plants in SD, but only if the plants were detillered to a single stem. Exposure to 2 LD cycles did not cause heading and flowering alone but enhanced the effect of exogenous gibberellins (GAs), bringing flowering to 100%. GA₅ and DHGA₅ were less effective than GA₁ and GA₃ in SD, especially with double applications, but were more effective than GA₁ and GA₃ when given together with 2 LD. The GAs had differential effects on vegetative growth and flowering, GA₅ and DHGA₅ causing much less leaf and stem growth than the other two GAs. Marginal induction, whether by LD or GA application, resulted in a high proportion of spikelets with viviparous proliferation. Thus, whereas GAs are inhibitory to the primary induction by SD, they can replace secondary induction by LD when vegetative growth is limited.     

Summer dormancy in perennial temperate grasses

BACKGROUND AND AIMS: Dormancy has been extensively studied in plants which experience severe winter conditions but much less so in perennial herbaceous plants that must survive summer drought. This paper reviews the current knowledge on summer dormancy in both native and cultivated perennial temperate grasses originating from the Mediterranean Basin, and presents a unified terminology to describe this trait. SCOPE: Under severe drought, it is difficult to separate the responses by which plants avoid and tolerate dehydration from those associated with the expression of summer dormancy. Consequently, this type of endogenous (endo-) dormancy can be tested only in plants that are not subjected to moisture deficit. Summer dormancy can be defined by four criteria, one of which is considered optional: (1) reduction or cessation of leaf production and expansion; (2) senescence of mature foliage; (3) dehydration of surviving organs; and (4, optional) formation of resting organs. The proposed terminology recognizes two levels of summer dormancy: (a) complete dormancy, when cessation of growth is associated with full senescence of foliage and induced dehydration of leaf bases; and (b) incomplete dormancy, when leaf growth is partially inhibited and is associated with moderate levels of foliage senescence. Summer dormancy is expressed under increasing photoperiod and temperature. It is under hormonal control and usually associated with flowering and a reduction in metabolic activity in meristematic tissues. Dehydration tolerance and dormancy are independent phenomena and differ from the adaptations of resurrection plants. CONCLUSIONS: Summer dormancy has been correlated with superior survival after severe and repeated summer drought in a large range of perennial grasses. In the face of increasing aridity, this trait could be used in the development of cultivars that are able to meet agronomic and environmental goals. It is therefore important to have a better understanding of the genetic and environmental control of summer dormancy.     

光温耦合的中国温带地区旱柳花期时空格局模拟

建立基于温度和光周期驱动的旱柳花期物候模型,旨在寻找影响旱柳花期时空变化的主要气象因子,揭示调控植物开花时间的生态机制,还可为改善柳絮造成的环境污染和花粉过敏等人类健康问题提供参考信息和依据。利用中国气象局农业气象观测网提供的中国温带地区1982-2011年49个站点的旱柳开花始期、盛期和末期观测资料及平行的逐日气象数据,分别对6种模型（简单积温模型、温度三基点模型、八时段温度模型、简单积温-日长模型、温度三基点-日长模型和八时段温度-日长模型）进行了参数率定和假设检验,根据外部检验结果,从中选出针对旱柳3个花期的最优物候模型,进而利用连续地理气象数据和最优物候模型重建了1982-2011年旱柳开花始期、盛期、末期和花期长度的时空变化特征。结果表明：光温耦合的物候模型对旱柳花期的模拟效果和外推效果优于仅基于温度的模型。旱柳开花始期和盛期最优模型均为八时段温度-日长模型,末期为温度三基点-日长模型,说明光周期和温度可能是影响旱柳花期开始、繁盛和结束时间的主要气象因子。同时,优选出的物候模型能够较准确地对不同年份和不同地区的旱柳花期进行模拟及预测。重建的1982-2011年旱柳平均开花始期、盛期和末期日期分别为4月24日、4月28日和5月3日,平均花期长度为9 d,始期、盛期和末期出现日期呈现出从海拔低到高、从南向北、从西向东逐渐推迟的空间格局。1982-2011年旱柳开花始期、盛期和末期在大部分地区呈提前趋势,呈显著提前趋势的面积分别占总面积的49.78%、50.01%和53.40%,花期长度变化差异不显著。     

Inflorescence commitment and subsequent development differ in their responses to temperature and photoperiod in Osteospermum jucundum

Although Osteospermum is a species which is known to require a period of chilling to induce flowering, the precise form of the relationships between temperature and photoperiod on the phases of flowering has not been quantified. This study aimed to investigate the effects of temperature and photoperiod on time to inflorescence commitment and on the rate of subsequent floral development in Osteospermum jucundum cv. Zulu. To assess how temperature and photoperiod affected the number of days needed for inflorescence commitment, plants were transferred from a range of photothermal environments to a non‐inductive environment. The effect of temperature and photoperiod on subsequent inflorescence development was examined by transferring plants with initiated inflorescences to a range of photothermal environments. Inflorescence commitment occurred first in plants grown at a low average diurnal temperature (10.6°C), but no evidence was found to suggest that photoperiod affected the duration of this phase. Once initiated, high temperatures and long days hastened inflorescence development. The rate of progress to flowering from initiation increased linearly with photoperiod and temperature (up to an optimum of 23.5°C).     

Circadian rhythms and the induction of flowering in the long-day grass Lolium temulentum L.

Plants of Lolium temulentum L. strain Ceres were grown in 8-h short day (SD) for 45 d before being exposed either to a single long day (LD) or to a single 8-h SD given during an extended dark period. For LD induction, the critical photoperiod was between 12 and 14 h, and more than 16 h were needed for a maximal flowering response. During exposure to a single 24-h LD, the translocation of the floral stimulus began between the fourteenth and the sixteenth hours after the start of the light period, and was completed by the twenty-fourth hour. Full flowering was also induced by one 8-h SD beginning 4 or 28 h after the start of a 40-h dark period, i.e. by shifting 12 h forward or beyond the usual SD. The effectiveness of a so-called ‘displaced short day’ (DSD) was not affected by light quality and light intensity. With a mixture of incandescent and fluorescent lights at a total photosynthetic photon flux density of 400 μmol m⁻² s⁻¹, a 4-h light exposure beginning 4 h after the start of a 40-h dark period was sufficient to induce 100% flowering. The flower-inducing effect of a single 8-h DSD was also assessed during a 64-h dark period. Results revealed two maxima at a 20-h interval. This fluctuation in light sensitivity suggests that a circadian rhythm is involved in the control of flowering of L. temulentum.     

A MADS domain gene involved in the transition to flowering in Arabidopsis

Flowering time in many plants is triggered by environmental factors that lead to uniform flowering in plant populations, ensuring higher reproductive success. So far, several genes have been identified that are involved in flowering time control. AGL20 (AGAMOUS LIKE 20) is a MADS domain gene from Arabidopsis that is activated in shoot apical meristems during the transition to flowering. By transposon tagging we have identified late flowering agl20 mutants, showing that AGL20 is involved in flowering time control. In previously described late flowering mutants of the long-day and constitutive pathways of floral induction the expression of AGL20 is down-regulated, demonstrating that AGL20 acts downstream to the mutated genes. Moreover, we can show that AGL20 is also regulated by the gibberellin (GA) pathway, indicating that AGL20 integrates signals of different pathways of floral induction and might be a central component for the induction of flowering. In addition, the constitutive expression of AGL20 in Arabidopsis is sufficient for photoperiod independent flowering and the over-expression of the orthologous gene from mustard, MADSA, in the classical short-day tobacco Maryland Mammoth bypasses the strict photoperiodic control of flowering.     

Gametogenesis,brooding and planulation in laboratory populations of a temperate scleractinian coral Balanophyllia elegans maintained under contrasting photoperiod regimes

Summary

Males of Balanophyllia elegans Verrill, 1864 in Monterey Bay, California, spawn in the fall. Fertilization occurs within the females and the internally brooded embryos develop into large benthic planulae that are released mainly in the late winter. Field-collected corals were held in the laboratory under contrasting photoperiod regimes (ambient or in-phase, and 6 months out-of-phase) for 6.5 years. After two years in the laboratory corals planulated at the same time, independent of photoperiod. Corals show a cyclic pattern of later winter and early spring planulation that coincides with declining sea temperatures in Monterey Bay. Gametes appeared in the first laboratory generation at IS months and brooded embryos were found in females at 31 months under both light regimes. Planulae production by laboratory-reared corals peaked at the same time as that of the parents. These findings suggest that the temporal pattern of planulation is regulated by both endogenous and exogenous components that can be modified in some way by disturbance caused to the corals when collected or by the time of collection.     

Photoperiod and gibberellic acid – control of the cell cycle in the meristem of Silene armeria and its effects on flowering

Plants of Silene armeria L., strain S_2.1, a quantitative long-day (LD) species which is known to react to GA₃ by flowering after attaining, the'intermediate stage', were induced by two LD or by two GA₃ applications. Changes in the mitotic index and DNA content (microdensitometric estimation) of cells in the axial zone, lateral zone and rib meristem of the shoot apex were observed during the first 48 h of each treatment. Similar mitotic activation occurred in response to LD or GA₃ after a 6-8 h lag period. This was preceded by a decrease in the proportion of nuclei with a 2C DNA content, indicating that in this species the control point for the shortening of the cell cycle was essentially in G₁. A second mitotic peak was observed 16 h later in photoinduced meristems, resulting in more pronounced cellular synchronization. These further events were not seen in GA₃-treated plants where only the meristematic activity was slightly, but reproducibly higher than in the control. Thus, two successive synchronizations of cell division are a typical feature of LD induction. The data are discussed with regard to the competence of shoot apical cells to be reactivated. The essential changes for the transition to flowering depend on these differential patterns of cell reactivation.     

Effects of temperature and photoperiod on flowering in Hyptis brevipes

Few tropical species have been tested for their flowering response under controlled conditions. Hyptis brevipes Poit, is an annual herb, commonly found in wet margins of streams and ponds, being considered a weed for some perennial plantations in Brazil. Under experimental glasshouse conditions, this species proved to be an obligate short-day plant. Flowering was delayed when photoperiods longer than 8 h were given, the critical photoperiod being between 12 and 13 h. When both temperature and photoperiod were controlled, at 20°C a longer protoperiod (by almost 1 h) is still inductive compared to 25 and 30°C. The number of short-day cycles required for full induction is relatively high and dependent upon temperature; at 20°C or above, 10 cycles are adequate, but at 15°C, more short-day cycles are needed. The number of inflorescences formed as well as the floral index vary according to daylength × temperature × inductive cycle number, allowing flowering to be assessed quantitatively. Long days are inhibitory to flowering, either suppressing it completely (when symmetrically intercalated among 24 inductive cycles) or preventing the floral index from increasing.     

光周期途径核心因子CO的开花调控机制

CONSTANS（CO）基因是生物钟和开花时间基因之间监测日照长度的重要元件,在光周期途径中发挥核心功能。CO可以整合光信号和生物钟信号,诱导开花途径整合子FLOWERINGLOCUST（F即和SUPPRESSOROF OVEREXPRESSION OF CONSTANS 1（SOC1）的表达,进而促进植株开花。本文综述CO基因的开花调控机制,并结合CO基因的研究现状展望了其未来的研究方向。     

Monoterpene content in Origanum syriacum as affected by environmental conditions and flowering

A high content of essential oil in the foliage of two chemotypes of field-grown Origanum syriacum L. was found between August and October. The content of phenolic monoterpenes in the essential oil was low from November to January, while the content of p -cymene was high. The effect of environmental conditions and flowering on the two chemotypes was similar. Controlled experiments revealed that under long photoperiods the content of essential oils in the foliage and of phenolic monoterpenes in the oil increased, while that of p -cymene decreased. A similar effect was observed when an 8 h natural light day was extended by 8 h low-intensity incandescent light. When the 16 h day was composed of 12 h natural light, the effect on oil content and composition was intensified. Under short-day photoperiods, increasing temperature or light intensity enhanced the relative content of p -cymene, while that of phenolic monoterpenes and the γ-terpinene generally decreased. Flowering decreased the essential oil content in the leaves and the relative content of the phenolic monoterpenes in the essential oil. Since environmental conditions influence flowering, they had both direct and indirect effects on the essential oil content and composition.     

Photoperiodic control of flowering in Dactylis glomerata, a true short-long-day plant

Flowering requirements of three Scandinavian cultivars of Dactylis glomerata L. have been studied in controlled environments. At temperatures ranging from 9 to 21°C optimal flowering required 10 weeks of exposure to short days (SD) followed by exposure to long days (LD). Only a few plants flowered in continuous LD and no primary induction took place in any daylength at 24 or 27°C. However, at a temperature of 3°C primary induction occurred also in 24 h LD, but more than 20 weeks of treatment were required for 100% flowering. The critical photoperiod for secondary induction was about 12–13 h, depending on the latitude of origin of the cultivar. A critical number of 12 to 16 LD cycles was required for 100% flowering, although some plants flowered after only 4 LD. A high proportion of viviparous proliferation resulted from marginal LD induction. Initiation of floral primordia did not take place in SD but required a transition from SD to LD. These results demonstrate that D. glomerata is a true short-long-day plant.     

Characterization of flowering time diversity in Miscanthus species

Miscanthus sinensis, M. sacchariflorus and their hybrids have been identified as leading candidates for the provision of bioenergy production across several continents. Flowering time is an important trait affecting biomass yield as well as certain quality attributes, such as moisture content at harvest. The aim of this study was to ascertain the level of diversity available to breeders and potential for hybridisation of different accessions in a large collection of Miscanthus. We also sought to determine trends in flowering time within and between species with respect to environment and origin of collection data (where known), whether flowering order was maintained across years, and the extent of uniformity of flowering in different genotypes. Flowering time was observed weekly in 244 genotypes of two Miscanthus species (M. sinensis, M. sacchariflorus) and inter‐specific hybrids including M. x giganteus over 3 years and using 4 clonal replicates of each genotype on a trial planted near Aberystwyth (Wales, UK). Differences in flowering time across the entire collection ranged from 160 to 334 days (June to November) and photoperiods between 7.8 and 16.6 h, with associated accumulated temperatures of 161 to 865^oCd. More than two thirds of the collection flowered by the end of each growing season. M. sinensis individuals were the earliest genotypes to flower and showed the greatest diversity with respect to the onset of flowering. Flowering times in genotypes of known origin in Asia could be partially explained by growing season rain fall, degree days and mean temperature. Uniform flowering was identified in some genotypes. This will be important for the development of genetically diverse seed‐based crops. Rank order of flowering was shown to be consistent across Western Europe, thereby justifying single site trials as the basis of germplasm characterisation for wider geographical deployment.     

温度和光周期对斑翅食蚧蚜小蜂发育与繁殖的影响

斑翅食蚧蚜小蜂是橡副珠蜡蚧的重要寄生性天敌之一。研究了温度和光周期对斑翅食蚧蚜小蜂发育、寄生及繁殖等方面的生态学特性。研究表明:温度和光周期对斑翅食蚧蚜小蜂种群增长影响明显。在32℃下不能完成世代发育,在温度为18-27℃范围内随温度升高发育加快,18℃发育历期最长(54.0d),27℃最短(22.8d);世代发育起点温度和有效积温分别为12.76℃和307.62日度。高温和低温均不利于斑翅食蚧蚜小蜂寄生,21℃时寄生率最高为(36.0%),30℃时寄生率最低(4.5%)。结合发育历期、体长、产卵、抱卵、寄生率等参数,该蜂发育繁殖的适宜温度为21-27℃度范围内,30℃以上不利于小蜂的发育及存活。斑翅食蚧蚜小蜂对长光的刺激比较敏感,随光照时间增长,发育加快、产卵量明显增加,长日照条件(LD16∶8h)的发育历期最短(24.8d),短日照条件(LD10∶14h)的最长(27.8d),产卵量在长日照条件(LD16∶8h)时最多(119.6粒),短日照条件(LD10∶14h)时最少(86.2粒)。结合发育、产卵、寄生等参数,长日照条件(LD14∶10-LD16∶8h)有利于种群增长。