准噶尔无叶豆的开花物候与生殖特征

 为了研究沙漠稀有植物准噶尔无叶豆(Eremosparton songoricum)的开花物候特征及其对生殖成功的影响, 2005和2006年连续两年对其自然种群的开花物候和开花过程中的花部表型变化进行了观察, 并运用相对开花振幅、开花强度和开花同步性等开花物候指数研究了开花物候特征。结果表明: 准噶尔无叶豆在5月下旬至6月中下旬开花, 其种群、个体、花序和单花的花期分别历时26~29 d、8~10 d、5~7 d和2~3 d。单花开花进程依其形态和散粉特征可分为散粉前期、散粉初期、散粉盛期和凋谢期4个时期。其个体水平的开花物候进程(开花振幅曲线)呈渐进式单峰曲线, 具有很高的开花同步指数, 表现出一种集中开花的模式。开花物候指数与座果数之间的相关分析结果表明, 始花日期与花期持续时间存在负相关关系, 而与开花数和座果数存在正相关关系; 花期持续时间与开花数和座果数存在显著正相关关系。准噶尔无叶豆个体开花物候在很大程度上是由其遗传因子决定的, 而开花物候在年度间的变异, 可能是由于荒漠气候的差异(主要是水分和温度的差异)所引起的。作为沙漠窄域分布特有种, 准噶尔无叶豆在环境和人为干扰的双重选择压力下, 为了吸引更多的传粉者访问而达到生殖成功, 形成了大量集中开花的模式。     

大花百子莲的开花物候与生殖特性

连续2年对引种植物大花百子莲在中国上海市人工种群的开花物候、结实情况进行观测记录,从不同水平分析了大花百子莲的开花物候对其生殖的影响。结果显示:（1）大花百子莲在引种地的开花时间为6月初至7月末;种群花期历时40 d左右,单株花期在14～31 d波动,单花寿命主要为2～3 d;2年的开花振幅平均在6朵·株^-1·d^-1左右,花朵开放百分率值在3.69%~4.87%左右,平均开花式样为11~17朵。（2）大花百子莲种群内2007年比2008年的各项开花物候指标均提前6～9 d;种群开花进程为渐进式的双峰曲线;种群花期重叠时间较长,2年的平均开花同步指数分别为0.663、0.695;植株个体相对开花强度主要分布频度为50%～70%。（3）开花物候指数与座果率的相关分析表明,始花时间与开花数目、座果数、花期长度呈显著负相关关系,开花数目与座果数、花期长度以及花期长度与座果数均呈显著正相关关系,同一花序内单花所发育成果实的数量与单花开放时间呈极显著负相关关系。研究表明,大花百子莲始花时间早的植株比始花时间晚的植株花期更长,开花数目多的植株比开花数目少的植株花期更长,花期长的植株比花期短的植株座果率更高;而且在同一花序上,越早开放的单花其结实的可能性越大。作为引种植物,大花百子莲开花物候年度间具有相似性,说明其在引种地可以适应环境压力,形成稳定的开花时间与开花模式;较长的花期重叠和大量的开花数目可以吸引较多的传粉者访问,为其生殖成功奠定了良好基础。     

巫山淫羊藿开花物候与生殖特征

研究巫山淫羊藿(Epimedium wushanense)3个种群的开花物候进程和生殖特征,分析了不同种群中的花部特征、开花物候指数和结实率差异,以及开花物候指数对巫山淫羊藿生殖成功的影响。结果表明:巫山淫羊藿花期为3月初到4月中下旬,种群开花一般历时22～27d;单株花期(花序)约12～17d;单花花期一般为3～4d。3个种群的个体水平的开花振幅曲线呈单峰型曲线,均具有较高的开花同步指数,表现出一种集中开花式样。相关分析表明:坐果数与开花高峰期、终花期、花期持续时间和花蕾数均呈极显著正相关,但坐果数与始花期无相关关系。开花物候变异系数分析表明:3个种群中花期持续时间、单花持续时间、终花期、开花高峰期和坐果数差异极显著;总花数、开花振幅和结实率差异显著;开花同步指数和始花期无显著差异性。3个种群中除同步性指数外,其余指数的变异系数均有差异,此外,巫山淫羊藿中结实率受到距的长短的影响。表明巫山淫羊藿的开花物候和生殖特征受到微环境的影响,同时生殖特征还受到花部特征距的影响。     

中国特有植物血水草开花物候与生殖特性

于2008年3-5月对分布在井冈山的血水草(Eomecon chionantha Hance )5个自然种群的开花物候进行了观察,运用开花振幅、相对开花强度和开花同步性等指数研究了其开花物候特征及其对该种生殖成功的影响.结果表明:血水草开花时间为3月下旬-5月上旬,种群花期历时24 ～46 d,个体平均开花持续时间为11～21 d,单花花期一般为3～5d;井冈山血水草种群的开花物候进程呈单峰曲线模式具有一个开花高峰期,表现出一种“集中开花模式”;与大多数亚热带植物一样,血水草具有较低的相对开花强度,分布频率集中在10％ ～30％.开花物候指数与生殖间的相关分析结果表明:始花时间与花期持续时间呈显著负相关,而与开花数和坐果率呈显著正相关;花期持续时间与开花数和坐果率呈显著正相关;同步性指数与始花时间、开花数、花期持续时间呈负相关.血水草“集中开花模式”是其在长期的进化过程中适应周围气候条件及生境的一种生殖保障.     

濒危植物长柄双花木的花部综合特征与繁育系统

 通过野外观察,运用杂交指数、花粉-胚珠比、人工授粉和套袋实验及等位酶电泳等方法,对长柄双花木(Disanthus cercidifolius Maxim. var. longipes H. T. Chang)自然种群和人工种群的开花状态、繁育系统进行了研究。结果如下：该种单花花期一般6～7 d,开花过程中柱头始终高于花药的位置,花药散粉具有先后顺序,一般2花药先散粉,其余3花药滞后1～2 d。单花花期依其形态和散粉时间可分为5个时期：即散粉前期、散粉初期、散粉盛期、凋谢期。种群开花一般历时49～55 d,但不同地点、年份之间有所差异,推测开花持续时间不同主要是环境条件所致。由于其杂交指数大于4,花粉-胚珠比等于1 250,结合人工套袋和授粉实验以及等位酶电泳结果可以确定该种的繁育系统属于异交为主,部分自交亲和,传粉过程需要传粉者。花粉竞争可能是导致该种濒危的主要的生殖生物学原因。     

金城山柔毛淫羊藿的开花物候与生殖特性

采用实地观测的方法,对南充市金城山三个不同生境中柔毛淫羊藿的开花物候特征及其生殖特性进行了研究。结果表明:柔毛淫羊藿花期为3月上旬至4月上旬,其种群、个体、花序和单花的花期分别历时30~41、22~34、9~18和4~8 d。三个生境中柔毛淫羊藿种群的开花物候进程基本相同,均呈单峰集中开花式样,因而能够吸引更多的传粉者访问而达到生殖成功。开花物候指数与坐果数之间的相关分析结果表明,坐果数与始花日期存在显著的负相关关系,与花期持续时间和开花数均呈极显著的正相关关系。揭示了药用植物柔毛淫羊藿的开花物候特征与生殖特性。     

多枝柽柳春夏两季开花物候特征与生殖特性

为了研究多枝柽柳的两季开花物候特征与生殖特性,对分布于古尔班通古特沙漠南缘莫索湾荒漠区的多枝柽柳自然种群各水平的开花物候指数、开花过程中的花部特征变化和结实特性进行了观察和统计。结果表明:(1)多枝柽柳春花期为5月下旬至6月上旬,表现为"集中开花模式",夏花期为6月中旬至9月上旬,表现为"持续开花模式"。(2)多枝柽柳春、夏花期的群体、个体、花序和单花水平的花期持续时间分别为14d、10d、3d、1d和85d、77d、2d、1d。(3)花序水平上夏花期始花日期、开花持续时间、开花数和座果数两两之间均存在显著相关关系。(4)两季花期间单花水平的开花进程与形态结构没有明显差异。(5)春花期的结实率显著高于夏花期,但两季花期结籽率差异不显著。(6)春花期昆虫访花次数显著高于夏花期。(7)两季花期的花粉活力没有明显差异,春花期的柱头可授性持续时间比夏花期长。研究认为,古尔班通古特沙漠南缘莫索湾荒漠区的生境条件恶劣,多枝柽柳特殊的春、夏两季开花结实特性在一定程度上提高了其生殖成功的可能性,是对特殊生境的适应性表现。     

濒危植物长柄双花木自然种群生殖构件的时空动态

根据野外观测数据,研究了濒危植物长柄双花木(Disanthus cercidifolius var.longipes)在不同生境条件下自然种群开花物候变异以及生殖构件的时空动态.结果表 明,不同群落类型中长柄双花木开花物候存在一定差异,在纯林中个体的平均花期最长,达75 d,而个体间开花同步性最低,仅为0.717;竹林中的个体花期最短,只有47d,而同步指数最高,达0.968;生殖枝数量及花序数均随个体年龄增长而增加,在年龄为30～35年时达最高,此后又呈下降趋势;在海拔为810m左右处,个体生殖枝数及着生的花序数均达最大值,且冠层间差异明显;纯林中个体着生的生殖枝数及每生殖枝上着生的花序数均最高,分别为411.39和7.857;竹林中个体每生殖枝着生的花序数显著低于其它群落.年龄小的个体,各时期生殖构件败育率均高于年龄大的个体;另外,生殖构件的败育与海拔高度间无相关性,而与构件发育时间、分布的树冠层次以及所处的群落类型相关.影响生殖构件败育的主要因子是光照条件.     

蒙古莸的开花物候与生殖特征

通过对蒙古莸自然种群和人工种群的开花物候进程观察,统计不同种群的结实率,分析开花物候对其生殖成功的影响。结果表明:(1)蒙古莸花期在7月末至9月中旬,人工种群较自然种群的开花物候明显提前,其中始花期提早4～5d,但两个种群的花期持续天数基本一致。(2)蒙古莸个体间具有较高的开花同步性,不同种群蒙古莸植株的开花同步性指数均大于0.80,且自然种群略高;晴好天气下绝大多数花朵都集中在上午10:00以前开放,阴雨天则延迟开放或者不开放,这种"集中开花模式"增加了传粉者的访问频率,是蒙古莸提高生态适应性、保障其生殖成功的一种表现。(3)开花物候指数与结实数之间的相关分析表明,花序水平上,开花数及花期持续时间与结实数呈极显著正相关关系。     

短命植物角茴香的开花物候与生殖特征

为了研究角茴香的开花物候特征及其对生殖成功的影响,于2015年对其自然种群各水平的开花物候指数、开花过程中的花部特征变化和结实特性进行了观察和统计。结果表明：（1）角茴香种群花期为4月下旬至5月下旬,种群水平的花期持续时间为36 d。（2）单花具有独特的花瓣结构,其内层花瓣由白色的侧裂片和黄色的兜状中裂片构成,中裂片是角茴香植物花粉二次呈现的关键功能器官;开花进程依其外层花瓣开裂大小可分为4个阶段,即外层花瓣顶端未张口、外层花瓣顶端张口5~7 mm、外层花瓣顶端张口12~14 mm和外层花瓣顶端张口3~5 mm。（3）花序水平的开花振幅曲线呈渐进式单峰曲线,开花同步指数为0.748,个体水平表现出较高的相对开花强度,其分布频度的偏斜率为0.45,主要频度范围分别集中在20%~30%和50%~70%之间。（4）花序水平的始花日期与花期持续时间存在显著负相关关系,开花数与座果数存在显著正相关关系。（5）自然状态下,角茴香结实率和结籽率分别是82.7%±0.1%和93.6%±0.8%。角茴香开花强度分布频度上出现的高、低强度的分异趋势是吸引传粉者、促进结实的有效生殖对策。     

Meta-analysis of phenotypic selection on flowering phenology suggests that early flowering plants are favoured

Flowering times of plants are important life-history components and it has previously been hypothesized that flowering phenologies may be currently subject to natural selection or be selectively neutral. In this study we reviewed the evidence for phenotypic selection acting on flowering phenology using ordinary and phylogenetic meta-analysis. Phenotypic selection exists when a phenotypic trait co-varies with fitness; therefore, we looked for studies reporting an association between two components of flowering phenology (flowering time or flowering synchrony) with fitness. Data sets comprising 87 and 18 plant species were then used to assess the incidence and strength of phenotypic selection on flowering time and flowering synchrony, respectively. The influence of dependence on pollinators, the duration of the reproductive event, latitude and plant longevity as moderators of selection were also explored. Our results suggest that selection favours early flowering plants, but the strength of selection is influenced by latitude, with selection being stronger in temperate environments. However, there is no consistent pattern of selection on flowering synchrony. Our study demonstrates that phenotypic selection on flowering time is consistent and relatively strong, in contrast to previous hypotheses of selective neutrality, and has implications for the evolution of temperate floras under global climate change.     

Phenotypic selection on flowering phenology and size in two dioecious plant species with different pollen vectors

Dioecious plants may be pollinated biotically by animals or abiotically via wind or water currents. It has been hypothesized that these two types of pollen vectors might impose contrasting selective pressures on plant flowering phenology. In the present study we describe the flowering phenology of two sympatric dioecious species with contrasting pollination modes: Mercurialis perennis (wind‐pollinated) and Tamus communis (insect‐pollinated). We estimated selection differentials and gradients for flowering time and flowering synchrony. As flowering time might depend on the accumulation of enough internal resources, we also estimated direct and indirect selection on plant size. Both species have male‐biased sexual ratios, and males are bigger and produce larger flower displays than females, but only in T. communis do males bloom earlier and for longer than females. Selection gradients suggest that selection tends to favor early‐flowering females of T. communis. There is no evidence of direct current selection on the flowering phenology of M. perennis. Intersexual differences in phenology fit with sex allocation and sexual selection theories. As we hypothesized, phenology of the animal‐pollinated species is under stronger selection than that of the wind‐pollinated species and we discuss the potential role of pollen vectors in shaping the flowering phenologies of the study species.     

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.     

青藏高原东缘高寒草甸植物群落的开花物候

利用2008年整个生长季的地面花期物候观测数据,初步研究了青藏高原东缘高寒草甸植物群落开花物候的动态、格局、参数间的联系及其与物种共存的关系.结果表明:植物群落的花期从5月初开始,8月上旬进入开花数目的高峰期,到10月初结束,遍及整个生长季节;不同物种进入花期的时间、峰值期及持续时间不同,各物种花期物候生态位相分离,但群落中大部分物种开花峰值期主要集中在7-8月;不同物种的开花峰值期时间与花期持续时间有负相关关系:开花越早的植物其花期持续时间越长;物种间的开花物候差异主要是由植物本身的特性和环境条件共同决定的,而群落水平上的开花物候格局主要受非生物因素的控制.

Abstract：

Based on the observation on the flowering phenology in the whole growth season of 2008, the dynamics, patterns, and relationships among various parameters of flowering phenology of alpine meadow plant community in eastern Qinghai-Tibetan Plateau were studied, and the potential effects of above-mentioned factors on the species coexistence of the plant community were approached. The flowering period of the plant community started from early May, came into peak period in the first ten days of August, and ended in early October, extending all over whole growth period. Different species had their different first flowering date, peak flowering period, and flowering duration, and their flowering phenologieal niches differentiated obviously. Most species had their peak flowering period centralized in July-August. There was a negative rela-tionship between peak flowering period and flowering duration: the earlier the flowering, the lon-ger the flowering duration. The differences in flowering phonology among the species were mainly determined by species per se characteristics and environmental conditions, but the flowering phe-nological pattern on community level was primarily controlled by abiotic factors.     

SELECTION ON VARIANCE IN FLOWERING TIME WITHIN AND AMONG INDIVIDUALS

We model the impact of pollinator visitation rate and behavior on the short‐term evolution of population flowering phenologies determined by the distributions of flowering times within and among individual plants. Evolution of population flowering phenologies depends on the phenotypic variances and heritabilities of the within‐individual mean and variance of flowering time. In the ecological scenarios we investigate selection does not produce a correlation of the mean and variance of individual flowering time. Self‐incompatibility causes weak stabilizing selection on flowering time that acts to reduce the within‐individual variance in flowering time. Disruptive selection due to pollinator limitation acts mostly to increase the among‐individual variance in flowering time. Stabilizing selection due to pollinator attraction, or short reproductive season, acts mostly to decrease the within‐individual variance in flowering time. Temporal autocorrelation of environmental stochasticity in pollinator visitation rate strongly selects to increase the within‐individual variance in flowering time. These predictions can be tested by measuring the causal factors described above, partitioning the variance in population phenology within and among individuals, and estimating the inheritance of, and selection on, within‐individual mean and variance of flowering time.     

Relationships between flowering phenology,plant size and reproductive success in shape Lotus corniculatus (Fabaceae)

Over three years the flowering phenology of individuals of Lotus corniculatus has been studied in relation to fruit set and seed predation to determine the relationships between four components of flowering time, plant size and reproductive success. Timings of first and peak flowering, and duration and synchrony of flowering differed between individuals in the same years. Between years, timing of first flowering was highly correlated for the same individuals, and was closely correlated with plant size and duration of flowering–larger plants flowered earlier and for a longer period. Peak flowering and synchrony were not correlated between-years for individuals.Fruit production and seed predation were correlated with some of the components of flowering phenology in some years, but not in others. The inconstancy of these relationships suggests that directional or stabilising selection is not acting consistently on the aspects of reproductive success studied in this work. The inconstancy of selection may result in the rather asynchronous flowering phenologies of individuals of L. corniculatus observed.We emphasize the importance of studying different components of flowering phenology in relation to individual plant size over several seasons. This work has shown that plant size not only has a direct effect on individual plant fecundity but also can influence flowering time and hence indirectly affect reproductive output.     

Inter and intraspecific variation on reproductive phenology of the Brazilian Atlantic forest Rubiaceae: ecology and phylogenetic constraints

The reproductive phenology of seven species of Rubiaceae from the Brazilian Atlantic rain forest was compared to evaluate the occurrence of phylogenetic constraints on flowering and fruiting phenologies. Since phenological patterns can be affected by phylogenetic constraints, we expected that reproductive phenology would be similar among plants within a family or genus, occurring during the same time (or season) of the year. Observations on flowering and fruiting phenology were carried out monthly, from December 1996 to January 1998, at Núcleo Picinguaba, Parque Estadual da Serra do Mar, Ubatuba, S?o Paulo State, Brazil. Nine phenological variables were calculated to characterize, quantify and compare the reproductive phenology of the Rubiaceae species. The flowering patterns were different among the seven species studied, and the Kruskal-Wallis test indicated significant differences in flowering duration first flowering, peak flowering and flowering synchrony. The peaks and patterns of fruiting intensity were different among the Rubiaceae species studied and they differed significantly from conspecifics in the phenological variables fruiting duration, fruiting peak date, and fruiting synchrony (Kruskal-Wallis test). Therefore, we found no evidence supporting the phylogenetic hypotheses, and climate does not seem to constrain flowering and fruiting patterns of the Rubiaceae species in the understory of the Atlantic forest.     

北京地区木本春季物候特征与生物学特性的关系

植物物候是植物生活史中的重要性状,也是指示气候与自然环境变化的重要指标,现已成为全球变化领域的研究热点之一。传统物候研究多假设物候由气候因素决定,如气温、降水、光照等,并主要从植物物候的年际变化角度探讨了气候因素对物候特征的影响。然而,不同物种的物候存在较大差异表明植物物候还与自身生物学特性（如系统发育和功能性状）有关,但植物生物学特性如何影响植物物候仍缺乏深入研究。基于北京地区44种木本植物1965-2018年的展叶始期和开花始期观测资料,以展叶始期和开花始期的3类物候特征（平均物候期、物候对温度的响应敏感度和物候期的积温需求）为例,探究植物物候特征与系统发育和功能性状的关系。首先,利用系统发育信号Blomberg’s K和进化模型检验植物物候特征是否具有系统发育保守性,并通过系统发育信号表征曲线直观表达植物物候特征的进化模式;之后,利用广义估计方程分析植物生活型、传粉型与物候特征的关系,以揭示不同植物的资源利用方式及生存策略的差异。研究发现：（1）除展叶始期的温度敏感度外,其余物候特征的进化均受随机遗传漂变和自然选择力的共同作用,可推断物候特征具有系统发育保守性,即亲缘关系越近的物种物候特征越相似。（2）开花始期的系统发育信号强度比展叶始期更大,表明繁殖物候的系统发育可能比生长物候更保守。（3）植物展叶始期及其积温需求与生活型密切相关。灌木比乔木的展叶时间早、积温需求少。植物开花始期与传粉型相关,风媒植物开花显著早于虫媒植物。研究成果有助于深入理解物候变化的生物学机制,对于丰富物候学的理论研究有重要意义,同时对植物保护也具有重要的指导价值。     

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.