Pollinator coupling can induce synchronized flowering in different plant species

Synchronous and intermittent plant reproduction has been identified widely in diverse biomes. While synchronous flowering is normally observed within the same species, different species also flower in synchrony. A well-known example of interspecific synchrony is “general flowering" in tropical rain forests of Southeast Asia. Environmental factors, such as low temperature and drought, have been considered as major trigger of general flowering. However, environmental cues are not enough to explain general flowering because some trees do not flower even when they encounter favorable environmental cues. We propose alternative explanation of general flowering; “pollinator coupling”. When species flower synchronously, the elevated pollen and nectar resource may attract increased numbers of generalist pollinators, with a concomitant enhancement of pollination success (facilitation). However, under these circumstances, plants of different species may compete with one another for limited pollinator services, resulting in declines in pollination success for individual species (competition). Here, we present a model describing resource dynamics of individual trees serviced by generalist pollinators. We analyze combinations of conditions under which plants reproduce intermittently with synchronization within species, and/or (sometimes) between different species. We show that plants synchronize flowering when the number of pollinators attracted to an area increases at an accelerating rate with increasing numbers of flowers. In this case, facilitation of flowering by different species exceeds the negative influence of interspecific plant competition. We demonstrate mathematically that co-flowering of different species occurs under a much narrower range of circumstances than intraspecific co-flowering.     

Crop photoperiodism model 2.0 for the flowering time of sorghum and rice that includes daily changes in sunrise and sunset times and temperature acclimation

Background and AimsDaylength determines flowering dates. However, questions remain regarding flowering dates in the natural environment, such as the synchronous flowering of plants sown simultaneously at highly contrasting latitudes. The daily change in sunrise and sunset times is the cue for the flowering of trees and for the synchronization of moulting in birds at the equator. Sunrise and sunset also synchronize the cell circadian clock, which is involved in the regulation of flowering. The goal of this study was to update the photoperiodism model with knowledge acquired since its conception.MethodsA large dataset was gathered, including four 2-year series of monthly sowings of 28 sorghum varieties in Mali and two 1-year series of monthly sowings of eight rice varieties in the Philippines to compare with previously published monthly sowings in Japan and Malaysia, and data from sorghum breeders in France, Nicaragua and Colombia. An additive linear model of the duration in days to panicle initiation (PI) and flowering time using daylength and daily changes in sunrise and sunset times was implemented.Key ResultsSimultaneous with the phyllochron, the duration to PI of field crops acclimated to the mean temperature at seedling emergence within the usual range of mean cropping temperatures. A unique additive linear model combining daylength and daily changes in sunrise and sunset hours was accurately fitted for any type of response in the duration to PI to the sowing date without any temperature input. Once calibrated on a complete and an incomplete monthly sowing series at two tropical latitudes, the model accurately predicted the duration to PI of the concerned varieties from the equatorial to the temperate zone.ConclusionsIncluding the daily changes in sunrise and sunset times in the updated photoperiodism model largely improved its accuracy at the latitude of each experiment. More research is needed to ascertain its multi-latitudinal accuracy, especially at latitudes close to the equator.     

Plant reproductive phenology over four years including an episode of general flowering in a lowland dipterocarp forest,Sarawak, Malaysia

The first systematic observation of a general flowering, a phenomenon unique to lowland mixed-dipterocarp forests in Southeast Asia, is presented. During general flowering, which occurs at irregular intervals of 3–10 yr, nearly all dipterocarp species together with species of other families come heavily into flower. We monitored reproductive phenology of 576 individual plants representing 305 species in 56 families in Sarawak, Malaysia. Observations continued for 53 mo from August 1992 and covered one episode of a general flowering cycle. Among 527 effective reproductive events during 43 mo, 57% were concentrated in the general flowering period (GFP) of 10 mo in 1996. We classified 257 species into flowering types based on timing and frequency of flowering. The most abundant type was “general flowering” (35%), which flowered only during GFP. The others were “supra-annual” (19%), “annual” (13%), and “sub-annual” (5%) types. General flowering type and temporal aggregation in reproductive events were commonly found among species in various categories of taxonomic groups, life forms, pollination systems, and fruit types. Possible causes for general flowering, such as promotion of pollination brought about by interspecific synchronization and paucity of climatic cues suitable for flowering trigger, are proposed, in addition to the predator satiation hypothesis of Janzen (1974) .     

Effects of fire season on flowering of forbs and shrubs in longleaf pine forests

Summary Effects of variation in fire season on flowering of forbs and shrubs were studied experimentally in two longleaf pine forest habitats in northern Florida, USA. Large, replicated plots were burned at different times of the year, and flowering on each plot was measured over the twelve months following fire. While fire season had little effect on the number of species flowering during the year following fire, fires during the growing season decreased average flowering duration per species and increased synchronization of peak flowering times within species relative to fires between growing seasons. Fires during the growing season also increased the dominance of fall flowering forbs and delayed peak fall flowering. Differences in flowering resulting from variation in fire season were related to seasonal changes in the morphology of clonal forbs, especially fall-flowering composites. Community level differences in flowering phenologies indicated that timing of fire relative to environmental cues that induced flowering was important in determining flowering synchrony among species within the ground cover of longleaf pine forests. Differences in fire season produced qualitatively similar effects on flowering phenologies in both habitats, indicating plant responses to variation in the timing of fires were not habitat specific.     

Paternity analysis-based inference of pollen dispersal patterns, male fecundity variation, and influence of flowering tree density and general flowering magnitude in two dipterocarp species

Background and Aims

Knowledge of pollen dispersal patterns and variation of fecundity is essential to understanding plant evolutionary processes and to formulating strategies to conserve forest genetic resources. Nevertheless, the pollen dispersal pattern of dipterocarp, main canopy tree species in palaeo-tropical forest remains unclear, and flowering intensity variation in the field suggests heterogeneity of fecundity.

Methods

Pollen dispersal patterns and male fecundity variation of Shorea leprosula and Shorea parvifolia ssp. parvifolia on Peninsular Malaysian were investigated during two general flowering seasons (2001 and 2002), using a neighbourhood model modified by including terms accounting for variation in male fecundity among individual trees to express heterogeneity in flowering.

Key Results

The pollen dispersal patterns of the two dipterocarp species were affected by differences in conspecific tree flowering density, and reductions in conspecific tree flowering density led to an increased selfing rate. Active pollen dispersal and a larger number of effective paternal parents were observed for both species in the season of greater magnitude of general flowering (2002).

Conclusions

The magnitude of general flowering, male fecundity variation, and distance between pollen donors and mother trees should be taken into account when attempting to predict the effects of management practices on the self-fertilization and genetic structure of key tree species in tropical forest, and also the sustainability of possible management strategies, especially selective logging regimes.     

Unravelling proximate cues of mass flowering in the tropical forests of South‐East Asia from gene expression analyses

Elucidating the physiological mechanisms of the irregular yet concerted flowering rhythm of mass flowering tree species in the tropics requires long‐term monitoring of flowering phenology, exogenous and endogenous environmental factors, as well as identifying interactions and dependencies among these factors. To investigate the proximate factors for floral initiation of mast seeding trees in the tropics, we monitored the expression dynamics of two key flowering genes, meteorological conditions and endogenous resources over two flowering events of Shorea curtisii and Shorea leprosula in the Malay Peninsula. Comparisons of expression dynamics of genes studied indicated functional conservation of FLOWERING LOCUS T (FT) and LEAFY (LFY) in Shorea. The genes were highly expressed at least 1 month before anthesis for both species. A mathematical model considering the synergistic effect of cool temperature and drought on activation of the flowering gene was successful in predicting the observed gene expression patterns. Requirement of both cool temperature and drought for floral transition suggested by the model implies that flowering phenologies of these species are sensitive to climate change. Our molecular phenology approach in the tropics sheds light on the conserved role of flowering genes in plants inhabiting different climate zones and can be widely applied to dissect the flowering processes in other plant species.     

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。     

中国北方主产地苹果冷热积累变化及其对始花期的影响

开展气候变化背景下苹果冷热积累变化及其对始花期的影响研究,对指导苹果种植及生产具有重要意义。本研究选取山东福山、山西万荣、甘肃西峰和新疆阿克苏代表中国北方苹果主产地,利用1996—2018年红富士苹果的始花期观测资料和逐时气温数据,采用动态模型、生长度小时模型分别计算逐日冷积累量(CP)和热积累量(GDH),并利用偏最小二乘回归法,对逐日冷、热积累量和各地苹果始花期进行相关分析,以明确各地苹果冷、热积累起止日期和积累量,以及冷、热积累期内温度变化对始花期的影响规律。结果表明: 我国北方主产地苹果冷积累时段集中于10月1日前后至2月中下旬或3月中旬,积累量为74.1～89.3 CP;热积累时段集中于1月下旬前后至始花期,积累量为4010～5770 GDH。西峰和阿克苏冷积累期内平均气温每升高1 ℃,冷积累量将分别增加3.8和5.0 CP;各地热积累期内平均气温每升高1 ℃,热积累量将增加725～967 GDH。与冷积累期内温度变化的影响效应相比,热积累期内温度变化主控我国北方主产地苹果始花期,且气候变暖总体有利于冷积累期内平均气温较低地区的苹果开花和生产。     

Low temperature influence on flowering in Citrus. The separation of inductive and bud dormancy releasing effects

The flowering response of Owari Satsuma mandarin ( Citrus unshiu Marc) to low temperature treatments has been determined using potted trees and in vitro bud cultures. In potted trees the chilling treatments released bud dormancy and enhanced both sprouting and flowering, but these two responses could not be separated. However, bud cultures showed no dormancy, and a specific effect of low temperature on flower induction was demonstrated. Low temperature appears to have a dual effect, releasing bud dormancy and inducing flowering. Potential flower buds have a deeper dormancy than vegetative buds, and the first stages of flower initiation seem to occur before the winter rest period.     

31种木本植物开花物候与系统发育的关系

植物物候通常被认为是由环境因素,如降水、温度和日照长度所决定,然而环境因素是否是物候唯一的决定因素仍然存在很大争议。谱系结构表征了植物在进化上的顺序,该发育时序是否对物候产生影响,当前仍然未知。在调查2016年春季新疆乌鲁木齐市最常见的31种木本植物的初始开花时间、败花时间和开花持续时间的基础上,通过分析植物开花物候的分布特征、开花物候在乔灌木间的差别、以及植物谱系距离与开花物候距离间的关系,试图揭示植物的开花物候和物种谱系(进化)顺序间的关系。结果表明:(1)新疆乌鲁木齐市31种木本植物的初始开花时间为4月18日±9d、败花时间为5月5日±12d、开花持续时间为(16±8)d;(2)乔木的初始开花时间和败花时间的标准差分别均低于灌木,乔木开花物候相对灌木更稳定;(3)乔木的初始开花和败花时间均显著早于灌木(P0.05),但开花持续时间在两者间未有显著性差异(P0.05);(3)31种木本植物间的初始开花时间距离、败花时间距离和开花持续时间距离均与物种谱系距离存在显著线性回归关系(P0.05)。综上可知:乔灌木在垂直空间上的分化使得木本植物的开花物候在植物生活型间存在不同。对植物的开花物候,除已被证明的降水、温度和日照长度等环境因素的影响外,物种进化顺序也可能造成了它在植物种间、时间和空间上的变异。     

The strength of assortative mating for flowering date and its basis in individual variation in flowering schedule

Although it has been widely asserted that plants mate assortatively by flowering time, there is virtually no published information on the strength or causes of phenological assortment in natural populations. When strong, assortative mating can accelerate the evolution of plant reproductive phenology through its inflationary effect on genetic variance. We estimated potential assortative mating for flowering date in 31 old‐field species in Ontario, Canada. For each species, we constructed a matrix of pairwise mating probabilities from the individual flowering schedules, that is the number of flower deployed on successive dates. The matrix was used to estimate the phenotypic correlation between mates, ρ, for flowering date. We also developed a measure of flowering synchrony within species, S, based upon the eigenstructure of the mating matrix. The mean correlation between pollen recipients and potential donors for flowering date was  = 0.31 (range: 0.05–0.63). A strong potential for assortative mating was found among species with high variance in flowering date, flowering schedules of short duration and skew towards early flower deployment. Flowering synchrony, S, was negatively correlated with potential assortment (r = ?0.49), but we go on to show that although low synchrony is a necessary condition for phenological assortative mating, it may not be sufficient to induce assortment for a given phenological trait. The potential correlation between mates showed no seasonal trend; thus, as climate change imposes selection on phenology through longer growing seasons, spring‐flowering species are no more likely to experience an accelerated evolutionary response than summer species.     

武汉大学樱花花期长度特征及预报方法

武汉大学樱花是武汉的一张"城市名片",开展樱花花期长度的预报工作,可为旅游部门管理工作和游客安排出行时间提供合理的参考。根据1979—2018年40年武汉大学樱园日本樱花树始花期和落花期的观测资料及同期气象资料的研究分析表明:(1)樱花的始花期和落花期在20世纪80—90年代期间有明显的提前;从20世纪90年代末开始至今,始花期与落花期变化趋势不明显,但变率较大,与全球气候变化停滞期相吻合;40年间花期长度变率很大,整体无明显的增多或减少的趋势。(2)平均始花期为3月14—15日,落花期为3月31日—4月1日,平均花期长度为18d。(3)花期长度与当年始花期日序数、开花期间平均气温、开花期间最高平均气温、最低平均气温和温度日较差平均值均呈负相关,与开花期间总降水量呈正相关。与开花期内平均极大风速值、平均降水量和日照时数等无明显相关性。(4)用1979—2015年共37年资料建立了樱花花期长度的单因子拟合、多因子回归及主成分分析模型,用2016—2018年3年资料进行检验,对武汉大学樱花花期长度进行了预报,取得了较好的试验效果。其中主成分回归模型、降水单因子拟合模型和多因子樱花花期长度回归模型预报效果最好,平均绝对误差在1.5d左右,后期将会把预报模型运用到实际的樱花花期预报工作中。     

Selecting models to predict the timing of flowering of temperate trees: implications for tree phenology modelling

Classical budburst models (Spring Warming, Sequential, Parallel and Alternating) are unable to fully predict external data, partly because of the methods of optimization used to adjust them. The purpose of this study was to examine different assumptions of budburst models and select those which are best supported by the data, defining new models able to predict external data. Eight models, each differing in one assumption, were fitted and tested using external data. The dataset used to test the models was deduced from aeropalynological data at two stations in France. The results show that some of the models proposed are able to accurately predict external dates of flowering of most of the studied species. The assumptions of those models have been individually tested and shown to improve the models accuracy. Robust estimates of the best predictor models of 12 tree species are presented. The analysis of hypothetical provenance transfer of two species, Buxus sempervirens and Platanus acerifolia, between the two study sites, shows that P. acerifolia estimates are similar in both environments whereas B. sempervirens estimates are variable. This result, which agrees with the genetic characteristics of both species, shows that local adaptation of phenology can also be studied through modelling approaches.     

Contrasting flowering phenology and species richness in abiotically and biotically pollinated angiosperms

Abstract Biotic pollination is thought to correlate with increased interspecific competition for pollination among plants and a higher speciation rate. In this study we compared patterns of flowering phenology and species richness between abiotically (wind) and biotically pollinated plants, using phylogenetically independent contrasts. We compiled phenological data from eight local seasonal floras, in which we found geographically overlapping sister clades. Of 65 documented origins of wind pollination, we were able to use up to 17 independent contrasts. In contrast to previous studies we found no difference in global species richness between wind- and biotically pollinated sister clades. Regarding phenology, we found wider phenological spread in biotically pollinated clades, earlier flowering onset in wind-pollinated trees, but no difference in duration of flowering between pollination modes. These results corroborate previous views that niche space is more constrained for wind-pollinated species, and that niche partitioning is less important between wind-pollinated plants compared to plants pollinated by animals.     

影响柑桔树开花结果性能的生态生理因素

本文报道1988年和1989年春夏之际异常和正常的桔园小气候环境对柑桔树开花结果物候期、抽梢生长、幼果发育、生理落果、当年果实产量和品质等性能的影响。结果表明,柑桔树的座果率受许多生态生理因素制约,但第一次生理落果主要与开花期桔园的小气候环境有关,第二次生理落果则与树体营养生理代谢密切相关。据此提出了预测预报柑桔树第二次生理落果的指标及其防御措施。     

Phosphorylation-mediated signalling in flowering: prospects and retrospects of phosphoproteomics in crops

Protein phosphorylation is a major post-translational modification, regulating protein function, stability, and subcellular localization. To date, annotated phosphorylation data are available mainly for model organisms and humans, despite the economic importance of crop species and their large kinomes. Our understanding of the phospho-regulation of flowering in relation to the biology and interaction between the pollen and pistil is still significantly lagging, limiting our knowledge on kinase signalling and its potential applications to crop production. To address this gap, we bring together relevant literature that were previously disconnected to present an overview of the roles of phosphoproteomic signalling pathways in modulating molecular and cellular regulation within specific tissues at different morphological stages of flowering. This review is intended to stimulate research, with the potential to increase crop productivity by providing a platform for novel molecular tools.     

Asynchronous flowering and within-plant flowering diversity in wheat and the implications for crop resilience to heat

Background and Aims

Self-pollination dominates in wheat, with a small level of out-crossing due to flowering asynchrony and male sterility. However, the timing and synchrony of male and female flowering in wheat is a crucial determinant of seed set and may be an important factor affecting gene flow and resilience to climate change. Here, a methodology is presented for assessing the timing and synchrony of flowering in wheat, Triticum aestivum.

Methods

From the onset of flowering until the end of anthesis, the anther and stigma activity of each floret was assessed on the first five developing ears in potted plants grown under ambient conditions and originating from ‘Paragon’ or ‘Spark-Rialto’ backgrounds. At harvest maturity, seed presence, size and weight was recorded for each floret scored.

Key Results and Conclusions

The synchrony between pollen dehiscence and stigma collapse within a flower was dependent on its relative position in a spike and within a floret. Determined on the basis of synchrony within each flower, the level of pollination by pollen originating from other flowers reached approx. 30 % and did not change throughout the duration of flowering. A modelling exercise parameterized by flowering observations indicated that the temporal and spatial variability of anther activity within and between spikes may influence the relative resilience of wheat to sudden, extreme climatic events which has direct relevance to predicted future climate scenarios in the UK.