开花时间对巫山淫羊藿花蜜分泌和结实的影响

运用单因素方差分析研究了巫山淫羊藿(Epimedium wushanense)开花时间对花蜜分泌、有效传粉者活动及结实性的影响.结果表明:(1)开花时间对巫山淫羊藿花蜜常备量和含糖量的影响极其显著.3个种群早期花蜜常备量较高,而花蜜含糖量在晚期较高.(2)开花时间对有效传粉者行为也具显著影响.一天中,有效传粉者的访花活动主要集中在中午;在整个花期进程中,晚期有效访花频率明显大于早期.(3)开花时间对巫山淫羊藿结实率、果实种子数及果实种子败育率的影响均显著,其中早期和中期的结实率和果实种子数大于晚期;而果实种子败育率则在晚期较高.因此,人工栽培巫山淫羊藿应注意花期调控,并选择早期果实进行实验或育种,其效果更好.     

绵枣儿的开花物候与传粉特性

该文对绵枣儿的开花物候与传粉特性进行了研究,分析了绵枣儿的开花进程、开花物候指数、开花物候指数对其生殖成功的影响,重点探讨了绵枣儿对传粉昆虫的种类及其活动规律。结果表明:(1)绵枣儿始花日期为7月中下旬,单株持续时间15 d左右,种群持续时间30~35 d。呈现出"M"型开花物候曲线,其两年的开花同步指数分别为0.862、0.885,表明开花的年际间差异较小。并且分析出植株的相对开花强度分布在30%~60%之间。(2)相关分析表明:绵枣儿的开花数目与坐果数和持续时间呈现出极显著正相关(P0.01),绵枣儿的持续时间与坐果数之间呈现出显著相关(P0.05),而其始花日期与开花数目、坐果数和持续时间并没有显现出相关性,即绵枣儿植株开花的数目越多,其坐果率越高,持续时间也就越长。(3)绵枣儿的访花昆虫总共有14种,可分属为5目10科。其中传粉昆虫有5科10种。该研究结果表明各传粉昆虫在单花停留的时间和其访花频率上呈现出显著的差异,在日访花频率上也差异明显。     

滇西北马先蒿属传粉生物学的初步研究

对云南西北部的中甸和德钦高山地区４种马先蒿属（ＰｅｄｉｃｕｌａｒｉｓＬ．）植物的传粉生物学进行了观察。晚夏开花的马先蒿与传粉熊蜂（ＢｏｍｂｕｓＬａｔｒ．）的出现在物候上有一定的相关性。其中短管、无喙、无蜜腺的肉根马先蒿（Ｐ．ｒｏｙｌｅｉＭａｘｉｍ．）和茨口马先蒿（Ｐ．ｔｓｅｋｏｕｅｎｓｉｓＢｏｎａｔｉ）是由熊蜂工蜂震动花粉、背触式传粉,这与其花冠类型的较原始性有关;长管、具喙、无蜜腺的斑唇马先蒿（Ｐ．ｌｏｎｇｉｆｌｏｒａｖａｒ．ｔｕｂｉｆｏｒｍｉｓ（Ｋｌｏｔｚ．）Ｔｓｏｏｎｇ）和云南长管马先蒿（Ｐ．ｓｉｐｈｏｎａｎｔｈａＤｏｎ．ｖａｒ．ｄｅｌａｖａｙｉ（Ｆｒａｎｃｈ．）Ｔｓｏｏｎｇ）通过熊蜂工蜂震动花粉、腹触式传粉。长管类马先蒿在植株相对矮小的情况下,通过较长的花冠管具喙向外伸出以实现授粉,这可能是与其传粉熊蜂协同进化的结果。     

黄河三角洲柽柳的开花特性及传粉生态学研究

野外定位观测了柽柳(TamarixchinensisLour.)的开花物侯、开花动态、访花者的种类、行为和访花频率,并对其繁育系统、花粉-胚珠比(P/O)、花粉活力进行了检测。结果表明,柽柳单朵花的花期仅1d,P/O为390.9±81.0,繁育系统属于自交亲和、需要传粉者的类型。花的结构和开花式样适合以食蚜蝇为主的多种昆虫传粉。自然条件下的结实率为95.23%。柽柳的果实成熟期较短(15d),整个居群有长达几个月的花期,开花数目众多,因此能产生大量的种子,在适宜条件下会形成大片的柽柳灌丛。     

国家二级保护植物翅果油树传粉生物学的初步研究

对翅果油树进行了传粉生物学的野外观察和实验表明：（1）翅果油树的花具有自花传粉的结构特征,同时具有蜜腺,又具备异花传粉的特征。传粉者主要是一种家养的蜜蜂。（2）翅果油树花形态结构与传粉者的形态和传粉行为非常协调和适应,昆虫的传粉部位主要是头前部和胸部。（3）蜜腺的产蜜量大,开花后1～2天蜜腺分泌量最大。对传粉昆虫竞争中具明显优势。昆虫的访花频率明显高于同花期的其他植物。（4）翅果油树没有融合生殖现象。     

马先蒿属花冠形态的多样性与传粉式样的关系

马先蒿属(Pediculais)是有花植物中花冠形态多样化最为集中的属。该属主要的传粉者是熊蜂属(Bormbus)昆虫;在北美,熊蜂和蜂鸟是马先蒿植物一些种类有效的传粉者;也发现壁蜂(Osmia)为其传粉。不同的传粉机制要求某一特定的取食式样储藏和释放花粉。本文讨论了花冠类型的进化趋势与传粉式样和花粉形态的关系。传粉者的选择压力是决定花冠多样化的重要因素之一;花冠类型与传粉者和传粉行为紧密相关。马先蒿植物和传粉者的相互依赖与其花冠类型、功能和物候互相适应,但花冠类型与花粉形态两者之间似乎没有明显的一一对应关系。通过北美、日本和喜马拉雅不同地理分布马先蒿种类的比较研究表明,具有相同花冠类型的种类有着相同的传粉方式,花冠形态与传粉式样存在紧密的协同进化关系。     

山地五月茶的蝇类传粉研究

观测了山地五月茶Antidesma montanum的开花物候、开花动态、访花者种类和访花行为,并对其繁育系统、花粉组织化学、花粉胚珠比（P／O）、花粉活力进行了检测。结果表明：山地五月茶是雌雄异株植物;其总状花序花期可长达7天;雄花／雌花花序数目为140．33±27．79／208．33±33．65（n=6）,雌雄花的颜色为很淡的黄绿色小型花;雄花单花花期为2天,花药开裂当天有活力,花粉为非淀粉型,花粉胚珠比为3333．33±607．18;单花花蜜量可达0．34±0．03μL,花蜜含糖量3．69％±0．30％。花的结构和开花式样适合蝇类传粉。芳香的气味是吸引蝇类的直接物质：花蜜是传粉者的报酬。主要的传粉昆虫为双翅目Diptera丽蝇科Calliphoridae的Chrysomya megacephala、Chrysomyasp．、寄蝇科Tachinidae的Drino sp．和蝇科Muscidae的Spilogona sp．、Mitroplatia sp．。套网不授粉的处理不结实,表明山地五月茶不存在无融合生殖,人工辅助授粉的坐果率（39．1％）略高于自然坐果率（36．7％）,二者无显著差异,表明其坐果率主要受自身资源分配限制。还讨论了蝇类传粉与雌雄异株性系统以及蝇类传粉与热带林中小型黄绿色花植物的相关性。     

珍稀濒危植物瑶山苣苔开花生物学及繁育系统研究

对仅分布于广西金秀老山自然保护区的国家Ⅰ级保护植物瑶山苣苔进行了开花生物学和繁育系统研究.结果显示:(1)瑶山苣苔花期从8月底至11月初,开花无固定时间.单花开放过程可分为萌动、露白、盛开、凋落4个阶段,且花朵具明显的增大再生长现象.(2)单花、花序、单株水平上的开花物候都表现出开花不同步性,单花花期约5~14 d,单花序花期约11~20 d,单株花期约8~20 d.花粉活力在散粉后6 d内相对较高,但花粉在野外通常于散粉3 d内被昆虫啃食完.(3)柱头在花药散粉时明显高于花药,便于接受异花花粉,柱头在散粉后第4天具可授性,可持续5~6 d.(4)瑶山苣苔杂交指数(OCI)为5,花粉-胚珠比(P/O)为379.64±145.61,繁育系统为兼性异交,自交亲和,传粉过程需要传粉媒介.研究表明,不稳定的传粉环境可能是该物种至濒的主要生殖生物学原因,自发自交"是其在开花期间对不稳定传粉环境的一种适应.     

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。     

濒危植物长柄双花木自然种群结实的花粉和资源限制

长柄双花木（Disanthus cercidifolius Maxim．vsr．longipes H．T．Chang）只零星分布于湖南、江西和浙江等省的少数县,已处于濒危状态。通过对其野外自然种群试验,研究了花粉和资源的有效性对雌性生殖的影响,并进一步探讨了长柄双花木“花多果少”的生殖机制。结果表明：不同来源的花粉对长柄双花木结果率及结籽率均产生显著影响。花粉来源而不是花粉数量对长柄双花木结实存在显著影响。自然条件下,长柄双花木“花一果转化率”约为54馏,“胚珠-种子转化率”约为83．19。施肥处理有利于花芽发育,可使植株花芽败育率明显下降,开花比例升高;同时座果率和结籽率均显著提高,而果实败育率则显著降低。随剪除叶片量的增加,结果率明显降低,但果实败育率在处理间无显著差异;剪除弱小枝条及病虫枝后,个体总开花数虽有所下降,但是结果率和结籽率均显著提高。去除花后,长柄双花木平均单果重、单粒种子重均显著降低。营养水平的时空异质性使自然种群结籽率受到资源和花粉的限制;补充资源可以通过增强花对传粉者的吸引而间接影响传粉。长柄双花木自然种群的坐果率极低,而且存在严重的“大小年”结果现象,不同的因子可能对其低水平坐果率产生相互作用。选择性败育、子房供应和雄性功能假说似乎是长柄双花木“花多果少”的生殖策略的最合理解释。     

The presence of co-flowering species facilitates reproductive success of Pedicularis monbeigiana (Orobanchaceae) through variation in bumble-bee foraging behaviour

Background and Aims The presence of co-flowering species can alter pollinator foraging behaviour and, in turn, positively or negatively affect the reproductive success of the focal species. Such interactions were investigated between a focal species, Pedicularis monbeigiana, and a co-flowering species, Vicia dichroantha, which was mediated by behaviour alteration of the shared bumble-bee pollinator. Methods Floral display size and floral colour change of P. monbeigiana were compared between pure (P. monbeigiana only) and mixed (P. monbeigiana and V. dichroantha) plots in two populations. Pollinator visitation rates, interspecific floral switching and successive within-plant pollinator visits were recorded. In addition, supplemental pollination at plant level was performed, and the fruit set and seed set were analysed in pure and mixed plots with different densities of P. monbeigiana. Key Results Pollinator visitation rates were dramatically higher in mixed plots than in pure plots. The higher pollinator visitation rates were recorded in both low- and high-density plots. In particular, successive flower visits within an individual plant were significantly lower in mixed plots. Supplemental pollination significantly increased fruit set and seed set of individuals in pure plots, while it only marginally increased seed set per fruit of plants in mixed plots. Conclusions The presence of V. dichroantha can facilitate pollination and increase female reproductive success of P. monbeigiana via both quantity (mitigating pollinator limitation) and quality (reducing geitonogamy) effects. This study suggests that successive pollinator movements among flowers within a plant, as well as pollinator visitation rates and interspecific flower switching, may be important determinants of the direction and mechanisms of interaction between species.     

Extended flowering in a Mediterranean shrub: Seasonal variability in seed quality and quantity

Fluctuating conditions throughout the year and changes in floral display may promote shifts in pollinator activity and predator pressure, influencing female reproductive output, especially for extended flowering species under seasonal climates. In this regard, flowering and fruit production were tracked in 2 different years in 2 populations of Ononis tridentata in Central Spain. Total fruit production was estimated, and fruits were harvested to obtain primary fruit investment, pollination success, predation incidence, seed production, seed weight and germination rates. Ononis tridentata combined spring mass flowering with a steady long flowering period across the summer and fall. The fewer flowers that are produced in fall were successfully pollinated, and produced fruits that were subject to minimal predation pressure relative to spring fruits. Moreover, fall fruits contained a higher number of heavier seeds and showed higher germination rates than those of spring seeds. Fall reproductive output represent 10% of annual viable seeds and thus may act as an important complement to the main spring reproductive investment. Extended flowering could be interpreted as a “bet-hedging strategy” for enduring Mediterranean unpredictable and changing environmental conditions.     

Reproductive isolation and pollination success of rewarding Galearis diantha and non-rewarding Ponerorchis chusua (Orchidaceae)

Background and Aims

Increasing evidence challenges the conventional perception that orchids are the most distinct example of floral diversification due to floral or prezygotic isolation. Regarding the relationship between co-flowering plants, rewarding and non-rewarding orchids in particular, few studies have investigated whether non-rewarding plants affect the pollination success of rewarding plants. Here, floral isolation and mutual effects between the rewarding orchid Galearis diantha and the non-rewarding orchid Ponerorchis chusua were investigated.

Methods

Flowering phenological traits were monitored by noting the opening and wilting dates of the chosen individual plants. The pollinator pool and pollinator behaviour were assessed from field observations. Key morphological traits of the flowers and pollinators were measured directly in the field. Pollinator limitation and interspecific compatibility were evaluated by hand-pollination experiments. Fruit set was surveyed in monospecific and heterospecific plots.

Key Results

The species had overlapping peak flowering periods. Pollinators of both species displayed a certain degree of constancy in visiting each species, but they also visited other flowers before landing on the focal orchids. A substantial difference in spur size between the species resulted in the deposition of pollen on different regions of the body of the shared pollinator. Hand-pollination experiments revealed that fruit set was strongly pollinator-limited in both species. No significant difference in fruit set was found between monospecific plots and heterospecific plots.

Conclusions

A combination of mechanical isolation and incomplete ethological isolation eliminates the possibility of pollen transfer between the species. These results do not support either the facilitation or competition hypothesis regarding the effect of nearby rewarding flowers on non-rewarding plants. The absence of a significant effect of non-rewarding P. chusua on rewarding G. diantha can be ascribed to low levels of overlap between the pollinator pools of two species.     

Co-flowering plants support diverse pollinator populations and facilitate pollinator visitation to sweet cherry crops

Many food crops depend on animal pollination to set fruit. In light of pollinator declines there is growing recognition of the need for agro-ecosystems that can sustain wild pollinator populations, ensuring fruit production and pollinator conservation into the future. One method of supporting resident wild pollinator populations within agricultural landscapes is to encourage and maintain floral diversity. However, pollinator visitation to crop plants can be affected either positively (facilitation) or negatively (competition) by the presence of co-flowering plants. The strength and direction of the facilitative/competitive relationship is driven by multiple factors, including floral abundance and the degree of overlap in pollinator visitation networks. We sought to determine how plant-pollinator networks, within and surrounding sweet cherry (Prunus avium) orchards, change across key time points during the cherry flowering season, in three growing regions in Australia. We found significant overlap in the suite of flower visitors, with seven taxa (including native bees, flies, hoverflies and introduced honey bees, Apis mellifera) observed visiting cherry and other co-flowering species within the orchard and/or the wider surrounding matrix. We found evidence of pollinator facilitation with significantly more total cherry flower visits with increasing percent cover of co-flowering plants within the wider landscape matrix and increased visitation to cherry by honey bees with increasing co-flowering plant richness within the orchard. During the cherry flowering period there was a significant positive relationship between pollinator richness on cherry and pollinator richness on co-flowering plants within the orchard and the area of native vegetation surrounding orchards. Outside of the crop flowering season, co-flowering plants within the orchard and wider landscape matrix supported the same pollinator taxa that were recorded visiting cherry when the crop was flowering. This shows wild plants help support the pollinators important to crop pollination, outside of the crop flowering season, highlighting the role of co-flowering plants within pollinator-dependent cropping systems.     

Highly efficient pollination by bumblebees ensures seed production in Pedicularis lachnoglossa (Orobanchaceae), an early‐flowering Himalayan plant

Abstract Pedicularis (Orobranchaceae) is a common high altitude genus of the Himalayas that may be affected by pollination limitation. Using Pedicularis lachnoglossa from Yulong (Jade Dragon) Snow Mountain in Lijiang (Yunnan Province, southwest China), we investigated the effects of high altitude habitats on the process of pollination and seed production. Floral biology, pollinator foraging behavior, breeding system, and pollination efficiency were examined using observation and exclusionary techniques. Pedicularis lachnoglossa was found to be entomophilous and exclusively pollinated by Bombus friseanus and B. yunnanicola. Our results indicated that pollination limitation in P. lachnoglossa was not significant. Under open pollination, approximately 80% of flowers were successfully pollinated and developed to fruits, and about 38% of ovules developed to mature seeds. Bumblebee pollination is highly precise and efficient in P. lachnoglossa, because its flowering phenology and floral characters enhance the foraging of bumblebees on flowers. This study supports that animal pollination plays a crucial role in the outbreeding of the early flowering Pedicularis. The evolution of floral specification in Pedicularis has the advantages of adaptation to bumblebee pollination in adverse high altitude habitats.     

VARIATION IN INDIVIDUAL FLOWERING TIME AND REPRODUCTIVE SUCCESS OF AGALINIS STRICTIFOLIA (SCROPHULARIACEAE)

Field studies on two populations of Agalinis strictifolia were conducted over a 3-year period to investigate the relationship between flowering time of individuals and plant size, flowering duration, flower and fruit production, fruit predation, and growth rate. Seasonal patterns of pollinator visitation were compared with those of individual flowering time, flower density, percent fruit production, and mean seeds/fruit. In general, early and middle flowering individuals (as determined by either first flowering date or peak flowering) were larger, flowered longer, and produced more flowers and fruits than late flowering individuals. Early and middle flowering individuals (based on first flowering date) also grew faster than late flowering individuals. Although early and middle flowering individuals produced more fruits, fruit predators did not damage a disproportionate number of fruits compared to late flowering individuals. Patterns of bee visitation showed no association with seasonal patterns of flower density, percent fruit production, mean seed/fruit, or individual flowering time. In populations of A. strictifolia, it would seem that biotic or environmental determinants of growth rate (hence size and reproductive success) may be more important in generating variation in individual flowering time than patterns of pollinator visitation or fruit predation.     

Flower herbivory and seed predation in Silene vulgaris (Caryophyllaceae): effects of pollination and phenology

Pollination and seed predation were studied in Silene vulgaris populations during two seasons, one with much lower pollinator abundance than the other. Among the pollinators, noctuid moths of the genus Hadena also acted as seed predators. Nectar-foraging female moths oviposited in flowers, and their larvae consumed flowers and seed capsules.
Despite a lower percentage of pollinated flowers in the year of low pollinator abundance, similar numbers of flowers set fruit in both years, because fewer flower buds and flowers were eaten by Hadena larvae during the year of low pollinator visitation. The number of seed capsules preyed upon was also lower in the year with low pollinator abundance, resulting in a higher seed set. The positive correlation between the percentage of pollinated flowers and the percentage of seed capsules destroyed was also observed when comparing flowers opening in different parts of the season.
Early flowering plant individuals had the same pollination success but suffered higher seed predation than late flowering ones. Selection for maximized pollination success through synchronous flowering, is probably the main reason for the compressed flowering period in 5. vulgaris , but the high level of predation early in the season may further increase the reproductive success of synchronous flowering individuals.     

Effects of flowering plant density on pollinator visitation, pollen receipt, and seed production in Delphinium barbeyi (Ranunculaceae)

Variation in flowering plant density can have conflicting effects on pollination and seed production. Dense flower patches may attract more pollinators, but flowers in those patches may also compete for pollinator visits and abiotic resources. We examined how natural and experimental conspecific flowering plant density affected pollen receipt and seed production in a protandrous, bumble bee-pollinated wildflower, Delphinium barbeyi (Ranunculaceae). We also compared floral sex ratios, pollinator visitation rates, and pollen limitation of seed set from early to late in the season to determine whether these factors mirrored seasonal changes in pollen receipt and seed production. Pollen receipt increased with natural flowering plant density, while seed production increased across lower densities and decreased across higher flower densities. Experimental manipulation of flowering plant density did not affect pollinator visitation rate, pollen receipt, or seed production. Although pollinator visitation rate increased 10-fold from early to late in the season, pollen receipt and seed set decreased over the season. Seed set was never pollen-limited. Thus, despite widespread effects of flowering plant density on plant reproduction in other species, the effects of conspecific flowering plant density on D. barbeyi pollination and seed production are minor.     

Temporal variations in the linkages between plants and flower visitors and the pollination success of Primula modesta along the snowmelt gradient

Although pollination networks between plants and flower visitors are diverse and flexible, seed production of many plant species is restricted by pollen limitation. Obligate outcrossers often suffer from low pollinator activity or severe interspecific competition for pollinator acquisition among co-flowering species. This study focused on seasonal changes in plant–flower visitor linkages in an alpine ecosystem and examined whether and how this seasonality affected the seed-set of Primula modesta, a self-incompatible distylous herb having long-tubed flowers. First, we recorded the linkages between plants and flower visitors along the snowmelt gradient. Then, pollination experiment was conducted to estimate the degree of pollen limitation over the course of flowering season of P. modesta. Flower visitors were classified by their tongue length based on the morphological matching with P. modesta flowers. As the season progressed, plant–visitor linkages became more diverse and generalized, and the visitation frequency to P. modesta flowers increased. In the later part of the season, however, the seed set of P. modesta was significantly reduced due to severe pollen limitation, presumably because of increased competition for long-tongued pollinators among co-flowering species. The present study revealed that pollinator availability for specialist species may be restricted even when plant–visitor linkages are diverse and generalized as a whole. In the case of P. modesta, morphological matching and competition for pollinators might be the main factors explaining this discrepancy.