桂林唇柱苣苔和百寿唇柱苣苔的开花动态及花粉活力和柱头可授性的比较

对桂林唇柱苣苔的两个居群(YZ居群和DB居群)和百寿唇柱苣苔的开花动态、花粉活力和柱头可授性进行了比较研究。结果表明,桂林唇柱苣苔的两个居群花期、单花持续期存在明显差异,DB居群的花期较YZ居群晚,但花期长于后者。YZ居群的花期、单花持续期与百寿唇柱苣苔的花期、单花持续期相近。三者开花过程相似。花粉活性和柱头可授性达最大的时间、持续时间在桂林唇柱苣苔的两个居群以及百寿唇柱苣苔之间具有明显差异。     

开花式样对传粉者行为及花粉散布的影响

理解植物花的特征可以从单花特征和群体特征两个层次入手。开花式样是植物的花在群体上的特征体现, 通过在开花数目、开花类型以及花的排列上的变化, 不同的开花式样对传粉者具有不同的吸引力, 影响昆虫在植株上的活动, 使花粉运动的方向发生相应变化, 从而影响着植物最终的交配结果。此外开花式样随环境改变也会发生一些变化。本文介绍了开花式样研究的进展, 对开花数目、开花类型以及花的排列等3个方面的已有研究进行了分别阐述, 并提出开花式样研究应更多地考虑影响传粉的各种因素。     

青藏高原高寒草甸植物花寿命对传粉环境的响应

花寿命是指花保持开放且具有功能的时间长度。不同物种的花寿命具有显著的差异, 但一定程度上的可塑性反映了植物对传粉环境变化的适应性。本研究以青藏高原高寒草甸不同海拔(2,900 m和3,600 m)的11种开花植物为研究对象, 通过套袋、补充授粉以及自然授粉处理分别测量了植物的潜在花寿命、最短花寿命以及实际花寿命。分析了不同海拔植物花寿命的变异, 以及对套袋处理(潜在花寿命与实际花寿命之差)和补充授粉处理(实际花寿命与最短花寿命之差)的响应及其可塑性(潜在花寿命与最短花寿命之差)的变异。结果表明: 无论是高海拔还是低海拔, 套袋显著延长了花寿命, 而补充授粉显著缩短了花寿命, 即, 潜在花寿命>实际花寿命>最短花寿命。高海拔群落中植物的实际花寿命和潜在花寿命均显著延长, 而最短花寿命在海拔间差异不显著。相对于低海拔群落, 高海拔植物的花寿命对补充授粉处理更敏感, 花寿命的可塑性更大。本研究说明传粉者稀少的高寒环境中, 植物通过提高花寿命及其可塑性来增加授粉机会, 获得更高的适合度。     

Pollen limitation and variation in floral longevity in gynodioecious Potentilla tanacetifolia

Pollination limitation is common in flowering plants and is thought to be a factor driving the evolution of floral traits.The plasticity of floral longevity to pollination may be an adaptation of plants to pollen limitation.However,this adaptation is less critical in short-lived flowers.To evaluate pollen limitation and the plasticity of floral longevity to pollination in Potentilla tanacetifolia,a gynodioecious herb with short-lived flowers,we analyzed its breeding system,tested sex-differential pollen limitation,and compared variations in floral display size in natural populations in Duolun County,Inner Mongolia,China.Hand pollination experiments and pollinator exclusion treatments revealed that P tanacetifolia is self-compatible and non-autonomously apomictic and shows sex-differential pollen limitation.The plasticity of floral longevity to pollination was observed; the floral duration of female plants was prolonged by approximately 3-4 hours with pollination exclusion treatment.Moreover,the percentage of flowers displayed on female plants during pollination exclusion treatment was significantly higher than that during natural pollination.Under natural pollination conditions,the percentage of flowers displayed on female plants was significantly higher than on hermaphrodite plants.Furthermore,approximately 50％ of the pollen grains spread out of the anthers of hermaphrodite flowers within 2 h of anthesis; the number of pollen grains adhering to the stigmas of hermaphrodite flowers was significantly higher than that adhering to female flowers when flowers shed their petals.These results indicate that variation in floral longevity may be an adaptive strategy to pollination conditions for gynodioecious P tanacetifolia.     

Temporal floral sex allocation in protogynous Aquilegia yabeana contrasts with protandrous species: support for the mating environment hypothesis

We tested one of the predictions of Brunet and Charlesworth (1995) that relative floral sex allocation will vary temporally with the mating environment and that the form of dichogamy (protandry vs. protogyny) will select for the pattern of variation in male versus female resource allocation. In many hermaphroditic plant species, allocation to female function (ovule number) decreases from early to late flowers within inflorescences as a result of resource limitation or ontogenetic changes. This pattern may obscure the effects of the mating environment and dichogamy on selection for allocation patterns in protandrous species (male allocation increases regardless). By examining a protogynous species the alternative pattern of temporal variation in resource allocation is predicted, namely that allocation to male function should decrease (or female allocation increase) throughout the flowering sequence. This pattern was observed in protogynous Aquilegia yabeana (Ranunculaceae), in which ovule number per flower remained constant whereas pollen number decreased in sequentially blooming flowers. These observations support the temporal sex allocation hypothesis of Brunet and Charlesworth (1995).     

The evolution of key functional floral traits in the early divergent angiosperm family Annonaceae

Potential key functional floral traits are assessed in the species‐rich early divergent angiosperm family Annonaceae. Pollinators (generally beetles) are attracted by various cues (particularly visual, olfactory, and thermogenic), with pollinators rewarded by nectar (generally as stigmatic exudate), heat, and protection within the partially enclosed floral chamber. Petals sometimes function as pollinator brood sites, although this could be deceptive. Annonaceae species are self‐compatible, with outcrossing promoted by a combination of protogyny, herkogamy, floral synchrony, and dicliny. Pollination efficiency is enhanced by pollen aggregation, changes in anthesis duration, and pollinator trapping involving a close alignment between petal movements and the circadian rhythms of pollinators. Most Annonaceae flowers are apocarpous, with syncarpy restricted to very few lineages; fertilization is therefore optimized by intercarpellary growth of pollen tubes, either by stigmatic exudate (suprastylar extragynoecial compitum) or possibly the floral receptacle (infrastylar extragynoecial compitum). Although Annonaceae lack a distinct style, the stigmas in several lineages are elongated to form “pseudostyles” that are hypothesized to function as sites for pollen competition. Flowers can be regarded as immature fruits in which the ovules are yet to be fertilized, with floral traits that may have little selective advantage during anthesis theoretically promoting fruit and seed dispersal. The plesiomorphic apocarpous trait may have been perpetuated in Annonaceae flowers as it promotes the independent dispersal of fruit monocarps (derived from separate carpels), thereby maximizing the spatial/temporal distance between seedlings. This might compensate for the lack of genetic diversity among seeds within fruits arising from the limited diversity of pollen donors.     

Influence of number of pollinations and pollen load size on maternal fitness costs in Collinsia heterophylla: implications for existence of a sexual conflict over timing of stigma receptivity

Costs related to pollen competition have rarely been considered, but are expected in the case of sexual conflict where male and female sexual functions have opposing evolutionary interests. In Collinsia heterophylla, delayed stigma receptivity is beneficial as it enhances pollen competition. A sexual conflict over timing of stigma receptivity has been proposed in this species as early pollination, following one-time pollinations, is advantageous to pollen donors at a cost of reduced maternal seed set (measured as seed number). In this study, we explored whether the maternal cost was still present following an additional pollination. We hypothesized that the cost is caused either by harm related to early pollen presence or by factors unrelated to harm. We performed pollinations at different stages of floral development, either one or two pollinations (24-h time lag), and varied the size of the first pollen load in the latter category. Early pollination reduced seed biomass also after two-time pollinations, suggesting a persistent maternal cost of early pollen presence. Further, pollen load size modified seed production, possibly indicating that dose-dependent harm influences the maternal cost of early fertilization. Our results strongly suggest negative effects of pollen competition on maternal fitness following early pollination, which is consistent with the existence of a sexual conflict over timing of stigma receptivity. In conclusion, we propose that much could be gained if more plant studies considered the potential for fitness costs in relation to sexual conflict, particularly those investigating pollen-pistil interactions.     

The evolution of floral sonication,a pollen foraging behavior used by bees (Anthophila)

Over 22,000 species of biotically pollinated flowering plants, including some major agricultural crops, depend primarily on bees capable of floral sonication for pollination services. The ability to sonicate (“buzz”) flowers is widespread in bees but not ubiquitous. Despite the prevalence of this pollinator behavior and its importance to natural and agricultural systems, the evolutionary history of floral sonication in bees has not been previously studied. Here, we reconstruct the evolutionary history of floral sonication in bees by generating a time‐calibrated phylogeny and reconstructing ancestral states for this pollen extraction behavior. We also test the hypothesis that the ability to sonicate flowers and thereby efficiently access pollen from a diverse assemblage of plant species, led to increased diversification among sonicating bee taxa. We find that floral sonication evolved on average 45 times within bees, possibly first during the Early Cretaceous (100–145 million years ago) in the common ancestor of bees. We find that sonicating lineages are significantly more species rich than nonsonicating sister lineages when comparing sister clades, but a probabilistic structured rate permutation on phylogenies approach failed to support the hypothesis that floral sonication is a key driver of bee diversification. This study provides the evolutionary framework needed to further study how floral sonication by bees may have facilitated the spread and common evolution of angiosperm species with poricidal floral morphology.     

锦带花的花粉活力、柱头可授性及传粉者的观察

在野外实地观测锦带花(Weigela florida(Bunge)A．DC．)的开花动态和访花者,在显微镜下观测柱头和花粉的形态及花粉-胚珠比,用TTC法测定锦带花花粉的活力,用联苯胺-过氧化氢法测定柱头可授性。结果表明,锦带花的花粉活力在开花时达70％以上,且能维持3d,之后明显下降。锦带花的柱头在开花3h后多数呈白色、二裂、具可授性,柱头的可授期持续2～3d。柱头与花药之间有5．3士0．9mm的空间距离,自然条件下难于自花授粉．开花当天7．00～10．00之间访花者最丰富,开花第2天访花者减少,开花第3天时访花者更少。在锦带花上记录到的访花者隶属于膜翅目、鳞翅目和双翅目,其中,熊蜂和蜜蜂是主要传粉者。     

Enantiostyly in Chamaecrista ramosa (Fabaceae‐Caesalpinioideae): floral morphology,pollen transfer dynamics and breeding system

Enantiostyly is a form of reciprocal herkogamy, in which floral morphs present reciprocal differences in the position of sexual elements, and occurs in monomorphic and dimorphic forms. This polymorphism maximises cross‐pollination and reduces self‐pollination, being very common within the subtribe Cassiinae (Fabaceae). Nevertheless, few studies have investigated the functionality of enantiostyly, particularly in this plant group. The present study aimed to investigate enantiostyly and its functionality in Chamaecrista ramosa, a monomorphic enantiostylous shrub, in an area of coastal vegetation in northeast Brazil. Pollen deposition and capture on the body of floral visitors, the relationship of these data with floral biology and breeding system, and morph ratio were evaluated. Pollen deposition and capture occurred in specific sites of the floral visitor body, showing the functionality of enantiostyly. The floral architecture, associated with the floral visitor behaviour, resulted in indirect pollen deposition on the floral visitor body. This occurred through a loop made by the pollen upon the inner petal surface, similar that generally reported for other Cassiinae. Chamaecrista ramosa is self‐compatible, although no fruit set was observed through spontaneous self‐pollination. The occurrence and number of floral morphs was similar within clumps. Enantiostyly seems to be advantageous for this species, as it results in efficient pollen capture and deposition, reduces the chances of autogamy and maximises intermorph pollen flow.     

Pollen packaging and dispensing: adaption of patterns of anther dehiscence and flowering traits to pollination in three Epimedium species

Pollen presentation theory (PPT) predicts that plant species typically pollinated by frequent and wasteful pollinators ought to be much more parsimonious and only gradually release pollen compared to plant species pollinated by infrequent pollinators that are efficient at delivering the pollen they remove. To test PPT, we compare the pollen presentation schedules and pollination systems in three related Epimedium species, having different pollinators. Results showed that differences in anther dehiscence and flowering traits resulted in different pollen packaging schedules. For E. sutchuenense and E. franchetii, a special ‘roll‐up’ movement of the anther wall during anther dehiscence increased pollen removal compared to the dehiscence pattern in E. mikinorii, which lacked the ‘roll‐up’ movement. Investigations revealed that honeybees had a higher pollen removal rate and lower stigmatic pollen load compared to bumblebees. In accordance with PPT, E. sutchuenense presents pollen sequentially and slowly for the frequent and wasteful honeybees. In comparison to E. sutchuenense, E. franchetii had a faster presentation rate and was adapted to the efficient and infrequent bumblebees. However, E. mikinorii was pollinated by both bumblebees and honeybees at high frequency and had the fastest pollen presentation. This pattern could reduce pollen wastage by honeybees and might be an adaptation to its short flower longevity (less than 1 day), to increase the chances of pollen deposition on stigmas. The study indicates that pollen presentation schedules can be a consequence of interactions among anther dehiscence, flowering traits and pollination environments for a given species.     

Effects of floral sexual investment and dichogamy on floral longevity

Aims Floral longevity, the duration that a flower remains open and functional, varies greatly among species. Variation in floral longevity has been considered to be optimal strategy for resource allocation under different ecological conditions, mainly determined by the rates of pollination and cost of flower maintenance. However, it is unclear whether an intrinsic factor, floral sexual investment, constrains evolution of floral longevity. The theoretical model also predicts that dichogamy favors long-lived flowers, but empirical studies to test this prediction remain unexplored.Methods To examine the effect of floral sexual investment on floral longevity, we measured flower size together with pollen and ovule production in 37 sympatric flowering plants in a natural community. The duration of the female and male phase in 21 protandrous species and floral longevity of the other 16 adichogamous species were documented in the field.Important findings Floral longevity varied from 1 day to 15 days, while pollen number per flower varied from 643 to 710880 and ovule number per flower from 1 to 426 in the 37 species. Flower size was correlated with pollen production as well as ovule production. Floral longevity was positively related to pollen production but not to ovule production. Consistent with the prediction that dichogamy favors long-lived flowers, we found the floral longevity of protandrous species was significantly longer than that of adichogamous species. In the protandrous species, pollen production per flower was observed to be positively related to male duration, while ovule production was not related to female duration. Our analyses of variation in floral longevity and sexual investment among different species suggest that the floral sexual investment could be an intrinsic factor contributing to the selected floral longevity, particularly the male phase, and that high pollen production could potentially increase pollen removal, i.e. male productive success.     

The evolution of floral biology in basal angiosperms

In basal angiosperms (including ANITA grade, magnoliids, Choranthaceae, Ceratophyllaceae) almost all bisexual flowers are dichogamous (with male and female functions more or less separated in time), and nearly 100 per cent of those are protogynous (with female function before male function). Movements of floral parts and differential early abscission of stamens in the male phase are variously associated with protogyny. Evolution of synchronous dichogamy based on the day/night rhythm and anthesis lasting 2 days is common. In a few clades in Magnoliales and Laurales heterodichogamy has also evolved. Beetles, flies and thrips are the major pollinators, with various degrees of specialization up to large beetles and special flies in some large-flowered Nymphaeaceae, Magnoliaceae, Annonaceae and Aristolochiaceae. Unusual structural specializations are involved in floral biological adaptations (calyptras, inner staminodes, synandria and food bodies, and secretory structures on tepals, stamens and staminodes). Numerous specializations that are common in monocots and eudicots are absent in basal angiosperms. Several families are poorly known in their floral biology.     

芍药花粉活力和柱头可授性的研究

用TTC法测定了芍药花粉的活力和寿命,用联苯胺—过氧化氢法测定了其柱头的可授期。结果表明,栽培芍药单瓣花植株的花粉活力和重瓣花植株的花粉活力在开花初期(1～2d)无明显差别,但重瓣花植株的花粉活力比单瓣花植株的花粉活力下降快;通常情况下,花粉寿命约为7d。栽培品种的柱头有个体差异,单瓣花植株有的开花后2h即可分泌粘液,而重瓣花植株的柱头在开花后,有的一直不分泌粘液,有的在开花后两天才分泌粘液;单瓣花植株的柱头可授性比重瓣花植株的强,一般分别在开花后6d和3d之内具可授性。移栽的野生芍药植株的花粉活力比栽培品种下降的慢,其寿命也较长,一般为15d;柱头的可授性和粘液分泌比栽培植株强。     

Does the relative importance of resource competition and architectural effect in floral variation vary with stages of floral ontogeny?

Variation in floral allocation within inflorescences has been attributed to resource competition and/or architectural effect.The two hypotheses were extensively studied and both hypotheses were partly ...     

七叶树花粉活力和柱头可授性变化的研究

采用花粉离体萌发法研究不同蔗糖浓度、硼酸浓度和不同温度对七叶树花粉萌发的影响及花粉活力测定,用联苯胺-过氧化氢法测定柱头可授性.结果表明:七叶树花粉萌发的最佳培养基是12％蔗糖+30 mg/L硼酸,花粉萌发的最适温度为25℃.雄花在开花当天花粉活力最高达75.69％,并在开花当天的上午10:00时,花粉活力最强,10:00～16:00花粉活力保持较高活力,是最佳授粉时段.两性花的柱头可授期持续时间较长,为8～9 d,开花3d达到最强,开花1～4 d柱头可授性保持较高水平,为授粉的最佳时间段.因此,从七叶树的花部特征、花粉活力与柱头可授性及花粉萌发的条件看,在长期的自然选择下七叶树在花部结构和开花生理上都是相配合的,以保障生殖成功.     

The adaptedness of the floral phenotype in a relict endemic, hawkmoth-pollinated violet. 1. Reproductive correlates of floral variation

This paper examines the relationship between quantitative variation in floral morphology (sizes of petals, spur and peduncle) and maternal reproductive success (seed production) in Viola cazorlensic (Violaceae), a narrowly endemic violet of south-eastern Spain pollinated by day-flying hawkmoths (Sphingidae). This plant is characterized by broad intraspecific variation in size and proportions of floral parts. Floral morphology does not influence significantly the probability of fruit set. Among flowers setting fruit, spur length and size of petals have no significant effect on seed production, but capsules from long- and short-peduncted flowers contain significantly more seeds than capsules from flowers with intermediate peduncles. Individual plants differ significantly in average floral characteristics. Plants with comparatively long and short peduncles tend to produce more seeds than those with intermediate ones, even after accounting statistically for individual differences in flower production. These findings are interpreted as evidence of disruptive selection on peduncle length during the study season. Floral variability in this species may be explained by the combined action of disruptive selection on peduncle length (the character most variable among individuals) and little, if any, stabilizing selection on spur length and size of petals