铃铛子和赛莨菪（茄科）的传粉机制比较研究

通过野外交配系统试验和传粉昆虫观察,以铃铛子（Anisodus luridus）和赛莨菪（A．camiolicoides）为研究对象,探讨了山莨菪属内自交亲和系统的进化与传粉昆虫的转变。结果表明,铃铛子和赛莨菪均属于自交完全亲和的类群,但两个种的自动自交能力均不强,而且都存在传粉限制。北方黄胡蜂（Vespula rtfarufa）和石长黄胡蜂（Dolichovespula saxonica）分别是铃铛子和赛莨菪的主要传粉昆虫,自然状态下两种昆虫的传粉效率均比较高,但两种昆虫访问两种植物时传递花粉的方式不同,其中北方黄胡蜂主要促进了铃铛子的花间传粉,而石长黄胡蜂访花引起赛莨菪同一朵花内的传粉。通过与该属的另一种植物山莨菪（A．tanguticus）的传粉机制比较,发现在山莨菪属的物种分化过程中,由自交不亲和转变为自交完全亲和,传粉昆虫也发生了转变,证明了自交亲和系统存在于起源较晚的类群中。高山环境中频繁且不可预测的降雨可能降低了传粉昆虫的活动能力,进而导致铃铛子和赛莨菪均存在传粉限制。     

山莨菪(茄科)的传粉生物学

茄科的多数种类具有自交不亲和的特点, 主要通过异花传粉结实; 但是, 一些物种或者物种内的部分种群或者个体却高度自交亲合, 转变为自交的繁育系统。该科植物山莨菪(Anisodus tanguticus)主要分布在青藏高原, 开花较早, 比其他晚开花的植物种类更加缺少有效的异花传粉昆虫。我们选择了位于不同海拔高度的2个种群进行比较研究, 主要目的是检验该物种的繁育系统是否在极端环境下由于传粉者的缺乏而发生了部分改变。研究发现,山莨菪的花不完全雌性先熟, 柱头和花药间的平均距离随着花开放时间的延长而不断缩小, 但两者在多数花的单花花期结束时并没有发生接触。因此, 山莨菪花主要表现为适应异花传粉的雌雄异位特征。然而, 少数花 (4.9％)的柱头和花药发生接触, 为“自动自交”的传粉解除了空间隔离。2个种群的多数个体存在自交不亲和机制, 应具有异花传粉的繁育系统; 但是部分个体具有明显的自交亲和能力, 为自交提供了生理基础。高海拔种群的传粉昆虫主要是厕蝇(Fannia sp.), 它们在不同植株间的活动能够保证异花传粉结实; 同时该种群的部分个体存在“自动自交”。低海拔种群的主要访花昆虫是蚂蚁, 它们在花内的活动导致花粉在同一朵花内传递, 而引起“协助自交”; 而异花传粉昆虫厕蝇的访花频率则较高海拔种群低。两个种群的结实均由于异花传粉者不足而受到传粉限制。因此两种不同类型的自交机制为该早期开花植物异花访花昆虫的不足提供了一定程度上的繁殖补偿。     

人工栽培山莨菪和野生山莨菪中4种托烷类生物碱含量的比较研究

为了评估人工栽培山莨菪的药用价值,采用高效液相色谱技术对人工栽培和野生山莨菪的地上部分和根中具有生物活性的4种托烷类生物碱：樟柳碱、山莨菪碱、东莨菪碱和阿托品的含量进行了测定。结果表明无论是人工栽培还是野生植物,地上部分中4种生物碱含量均远低于根,这解释了人们为什么用山莨菪的根而不是整株人药。在栽培植物的根中,一年生山莨菪中各生物碱含量均小于二年生山莨菪,其根中4种生物碱总量与野生根相比差异不是很明显;二年生山莨菪根中,4种生物碱总量以及樟柳碱、东莨菪碱和阿托品含量均比野生的高。这说明人工栽培的山莨菪,尤其是二年生山茛菪,同野生山莨菪一样具有一定的药用价值。     

几种主要莨菪烷类生物碱在中国茄科植物中的存在

本文报道了莨菪碱,东莨菪碱,山莨菪碱,樟柳碱四种药用莨菪烷类生物碱以及一种和莨菪烷类生物碱可能有生源上联系的红古豆碱在中国茄科植物中的存在情况。结果表明:中国茄科茄族(Solaneae)中的天仙子亚族(Hyoscyaminae),向阳花亚族(Mandragorinae)以及曼陀罗族(Datureae)是此类生物碱最集中的植物群,也是今后寻找此类药物新资源应重点注意的研究对象。通过化学分析,证明矮莨菪(Przewalskia shebbearei)和青海矮莨菪(P.tangutica)的根中含有较高含量的莨菪碱和山莨菪碱,是两种有价值的原料植物。通过对唐古特山莨菪(Scopolia tangutica)的种群中所含上述五种生物碱的观察,发现这些碱在不同植物个体中的存在有变异,而且莨菪碱和山莨菪碱的含量也因产地不同而存在显著的差异。本文的研究结果,不但对进一步利用天然莨菪烷类生物碱的资源具有参考意义,而且对茄科植物的化学分类和系统安排也提供了有价值的资料。     

青海省三种茄科植物种子微形态特征

运用扫描电镜对青藏高原地区3种茄科重要药用植物马尿泡(Przewalskia tangutica)、曼陀罗(Datura stramonium)和山莨菪(Anisodus tanguticus)18个自然居群的种子微形态进行了观察.种子的种皮纹饰在3个不同的物种之间具有显著的差异,可以分为4种类型.与山莨菪和马尿泡在植株、花、叶等形态差异较大不同,种子大小、颜色、形状及表面纹饰等在两个物种间非常相似,说明与曼陀罗属相比,山莨菪属和马尿泡属间可能具有较近的亲缘关系.结果说明种子微形态这一稳定的性状可以作为青藏高原茄科这3个属间的分类依据,具有重要的系统学意义,但是分类决策必须要包括形态学、细胞学、孢粉学、分子生物学等证据.     

中国特有的两种同域分布凤仙花属植物的传粉生物学研究

中国是凤仙花属物种多样性分布中心之一,存在着大量的特有和同域分布现象。为更好地理解同域分布近缘种的传粉综合征分化以及传粉昆虫资源分配问题,该文以中国特有种黄麻叶凤仙花(Impatiens corchorifolia)和金凤花(I.cyathiflora)为研究对象,通过野外调查和实验完成传粉生物学的相关研究,比较分析了这两个种的开花物候、花形态结构、传粉昆虫的种类及访花行为和人工授粉实验等方面的异同。结果表明：(1)两种凤仙花的花距不同,花距的长度与花蜜量呈正比,但与花蜜含糖量无关。花距类型决定了两种植物的传粉昆虫种类的分化。具短花距的黄麻叶凤仙花的花蜜含量为(4.57±1.43)μL,花蜜糖含量为(45.48±2.28)%,由蜜蜂和胡蜂为其传粉。拥有长花距的金凤花的花蜜含量为(10.15±3.28)μL,花蜜糖含量为(46.12±1.48)%,除蜜蜂和胡蜂传粉外,天蛾和熊蜂也为其传粉昆虫。(2)两种植物虽然共享3种传粉者,但表现为明显的错峰访花,传粉行为和花通道的差异决定了传粉者携带花粉的部位不一致。(3)繁育系统实验表明,两种植物均表现出明显的种间杂交障碍和异花授粉偏好性。因此,花...     

赤霉素对山莨菪与马尿泡种子发芽的影响

山莨菪[Anisodus tanguticus(Maxim)pascher)与马尿泡[Przewalskia tangutica Maxim)均属于茄科植物,是分布在青藏高原的特有种,因植物体含有多种生物硷,近几年已作为医药工业提取阿托品,东莨菪硷、山莨菪硷(新药“654”)等的原料而被利用。但是,由于这两种植物只零星地分布在海拔3000公尺以上的牧区,采集和运输都有困难,作为大规模生产的原料远远不能满足需要。因此,在青藏高原进行栽培以建立稳定的原料基地,实属必要。     

青藏高原东部橐吾属三种植物的繁育系统

研究青藏高原东缘分布的箭叶橐吾、黄帚橐吾和掌叶橐吾的繁育系统,箭叶橐吾和黄帚橐吾各自居群中有性生殖和克隆繁殖的权衡关系,分析三种植物的的克隆繁殖与生态适应,探讨橐吾属植物对高寒环境胁迫条件下的适应与进化。结果显示三种橐吾的花果期在6月中旬到10月初,花序为头状花序组成的总状排列。三种橐吾的杂交指数(OCI)均为4,繁育系统以远交为主、部分自交亲和、虫媒、需要传粉者;影响有性生殖成功的因素除昆虫侵食以外,黄帚橐吾和箭叶橐吾低的结实率主要原因是传粉媒介限制。限制结实的主要因素是高寒多风的环境限制了昆虫的活动,访花昆虫数量减少,从而使花粉转移效率降低,导致了低的结实率。植物采取的适应对策:大量开花,延长雌雄配子体的功能期和传粉媒介的泛化等,可见三种橐吾有性生殖的潜力依然很大。箭叶橐吾和黄帚橐吾在自然条件下主要进行有性生殖,在保护草场时,应针对其有性生殖器官采取措施。     

广西特有属——异裂菊属开花生物学及繁育系统特征

以异裂菊属5个种的实生苗为试验材料,通过田间观察,采用TTC法、联苯胺-过氧化氢法、杂交指数、花粉-胚珠比等方法,对异裂菊属的开花物候、花部特征、繁育系统及访花昆虫行为进行了研究。结果表明:(1)绢叶异裂菊的花期最早,为6~7月份;小花异裂菊和柳州异裂菊花期为6~8月份;凹脉异裂菊和异裂菊的花期为8~12月份,整个花期持续约5个多月,单花花期约为3d,单个花序花期约为5d。(2)异裂菊属的花粉在开花后1~18h具有活力,柱头的可授性持续约48h;杂交指数OCI为4,花粉/胚珠比(P/O)为1 450~2 250,繁育系统属于异交、部分自交亲和,需要传粉者。(3)传粉昆虫主要是食蚜蝇科灰带管食蚜蝇和胡蜂科黑尾胡蜂,昆虫访花高峰期在上午10:00~12:00,每次可连续访问多朵花,每朵花访问时间约为2~10s。     

三江源地区传粉昆虫的活动规律研究

随着各种环境压力的日益增大,传粉昆虫多样性及其保护研究逐渐得到重视。为了调查三江源地区传粉昆虫的基本概况,于2021年7-9月通过随机样点法和定点观测法对14类(4目12科)传粉昆虫类群的传粉活动进行了观测。共得到传粉活动记录7 247条,观察到受访植物37种(13科31属)。分析结果表明,三江源地区传粉昆虫类群的传粉活力由高到低依次是熊蜂(访花频次占总频次的70.13%)、其他双翅目昆虫(11.19%)、灰蝶(10.49%)和蛱蝶(3.08%),而其他的传粉昆虫类群的访花频次占比不足1%,传粉作用较弱;其中,熊蜂在不同植物上的访花活动会呈现不同的高峰期：如在甘肃马先蒿Pedicularis kansuensis上呈现两个访花高峰期(11∶00-12∶00和18∶00-19∶00),而在假水生龙胆Gentiana pseudoaquatica上只有一个访花高峰期(11∶00-12∶00)等。研究显示,熊蜂是三江源地区传粉活力最高的传粉昆虫类群,且其访花的日活动规律因自身特性、受访植物种类和外界环境变化呈现不同的访花高峰期。三江源地区的生态平衡离不开传粉昆虫的作用,加强对传粉昆虫的保护有...     

Relative impact of mate versus pollinator availability on pollen limitation and outcrossing rates in a mass‐flowering species

Plant mating systems are driven by several pre‐pollination factors, including pollinator availability, mate availability and reproductive traits. We investigated the relative contributions of these factors to pollination and to realized outcrossing rates in the patchily distributed mass‐flowering shrub Rhododendron ferrugineum. We jointly monitored pollen limitation (comparing seed set from intact and pollen‐supplemented flowers), reproductive traits (herkogamy, flower size and autofertility) and mating patterns (progeny array analysis) in 28 natural patches varying in the level of pollinator availability (flower visitation rates) and of mate availability (patch floral display estimated as the total number of inflorescences per patch). Our results showed that patch floral display was the strongest determinant of pollination and of the realized outcrossing rates in this mass‐flowering species. We found an increase in pollen limitation and in outcrossing rates with increasing patch floral display. Reproductive traits were not significantly related to patch floral display, while autofertility was negatively correlated to outcrossing rates. These findings suggest that mate limitation, arising from high flower visitation rates in small plant patches, resulted in low pollen limitation and high selfing rates, while pollinator limitation, arising from low flower visitation rates in large plant patches, resulted in higher pollen limitation and outcrossing rates. Pollinator‐mediated selfing and geitonogamy likely alleviates pollen limitation in the case of reduced mate availability, while reduced pollinator availability (intraspecific competition for pollinator services) may result in the maintenance of high outcrossing rates despite reduced seed production.     

Pollen limitation,species’ floral traits and pollinator visitation: different relationships in contrasting communities

Failures in the process of pollen transfer among conspecific plants can severely impact female reproductive success. Thus, pollen limitation can cause selection on plant mating systems and floral traits. The relationships between pollen limitation and floral traits might be partly mediated by the quantity and identity of pollinator visits. However, very little is known about the relationship between pollinator visits and pollen limitation. We examined the relationships between pollen limitation and floral traits at the community level to connect them to community ecology processes. We used 48 plant species from two contrasting communities: one species‐rich lowland community and one species‐poor alpine community. In addition, we calculated visitation rates and ecological pollination generalization for 38 of the species to examine the relationship between pollinator visitation and pollen limitation at the community level. We found low overall levels of pollen limitation that did not differ significantly between the alpine and the lowland community. In both communities, species with evolutionary specialized flowers were more pollen limited than species with unspecialized flowers. Species’ visitation rates and selfing capability were negatively related to pollen limitation in the alpine community, where pollinators are scarcer. However, flower size/number, ecological generalization of plants and flowering onset had greater effects on pollen limitation levels at the lowland community, indicating that the identity of the visitors and plant‐plant competitive interactions are more decisive for plant reproduction in this species‐rich community. There, pollen limitation increased with flower size and flowering onset, and decreased with ecological generalization, but only in species with evolutionary specialized flowers. Our study suggests that selection on plant mating system and floral traits may be idiosyncratic to each particular community and highlights the benefits of conducting community‐level studies for a better understanding of the processes underlying evolutionary responses to pollen limitation.     

The reproductive assurance benefit of selfing: importance of flower size and population size

Autonomous selfing can provide reproductive assurance (RA) for flowering plants that are unattractive to pollinators or in environments that are pollen limited. Pollen limitation may result from the breakdown of once-continuous habitat into smaller, more isolated patches (habitat fragmentation) if fragmentation negatively impacts pollinator populations. Here we quantify the levels of pollen limitation and RA among large and small populations of Collinsia parviflora, a wildflower with inter-population variation in flower size. We found that none of the populations were pollen limited, as pollen-supplemented and intact flowers did not differ in seed production. There was a significant effect of flower size on RA; intact flowers (can self) produced significantly more seeds than emasculated flowers (require pollen delivery) in small-flowered plants but not large-flowered plants. Population size nested within flower size did not significantly affect RA, but there was a large difference between our two replicate populations for large-flowered, small populations and small-flowered, large populations that appears related to a more variable pollination environment under these conditions. In fact, levels of RA were strongly negatively correlated with rates of pollinator visitation, whereby infrequent visitation by pollinators yielded high levels of RA via autonomous selfing, but there was no benefit of autonomous selfing when visitation rates were high. These results suggest that autonomous selfing may be adaptive in fragmented habitats or other ecological circumstances that affect pollinator visitation rates.     

Reproductive assurance varies with flower size in Collinsia parviflora (Scrophulariaceae)

A central question in plant evolutionary ecology is how mixed mating systems are maintained in the face of selection against self-pollination. Recently, attention has focused on the potential reproductive assurance (RA) benefit of selfing: the ability to produce seeds via autonomous selfing when the potential for outcrossing is reduced or absent. To date, there is little experimental support for this benefit under natural pollination conditions. In addition, the RA hypothesis has not been tested experimentally in a species displaying morphological variation for traits expected to influence the mating system, such as flower size, which affects both attractiveness to pollinators and ability to self autonomously. Here, we document significant among-population variation in flower size in Collinsia parviflora and show that pollinators preferred large flowers over small flowers in experimental arrays. The pollinator community varied among three study sites, and two small-flowered populations had lower pollinator visitation rates than one large-flowered population. We compared seed production between intact flowers (can self) and experimentally emasculated flowers (require a pollinator) on large- and small-flowered plants. As predicted by the RA hypothesis, small-flowered plants show a greater RA benefit of selfing than large-flowered plants; emasculated, small flowers produced very few seeds, relative to intact, small flowers or either emasculated or intact, large flowers. We also show that the RA benefit is pollination-context dependent, differing between small- and large-flowered test sites, likely due to a combination of pollinator discrimination against small flowers and differences between test sites in the pollinator community. This paper is the first experimental evidence showing a trait-dependent RA benefit of selfing under natural pollination conditions.     

克隆生长对被子植物传粉过程的影响

克隆植物与其传粉者的相互作用是植物繁殖生态学的重要研究领域之一。植物克隆生长与有性繁殖通常相伴进行, 往往产生较大的花展示与复杂的克隆空间结构, 通过传粉过程对有性繁殖过程产生影响, 共同决定植物的适合度。本文回顾了克隆生长对被子植物传粉过程影响的国内外研究进展, 从植物克隆大小、花资源空间配置、克隆构型与种群遗传结构四个方面讨论了克隆生长对传粉过程的影响及其生态学与进化生物学意义。早期研究预期, 随着克隆增大, 同株异花授粉水平增加, 因而通过增大自交率或花粉阻塞效应降低植物的适合度。但是, 后来的一些模拟与野外实验研究发现, 传粉者在同一克隆内访问的花数量并不会随克隆增大而一直增加, 访花行为也主要发生在分株内; 而且分子标记的自交率组分分析也表明自交主要发生在分株内。另一方面, 人工模型模拟以及传粉者访问行为研究表明, 当花朵数量相同时, 与所有花集中生长在同一分株上相比, 将花朵分散在多个分株上的克隆生长方式不会增加, 反而降低了同株异花授粉的发生水平。如果花序内花雌雄同熟, 花朵同时提供与接收花粉, 克隆生长会使植物接收到更高比例的异交花粉, 在提高后代质量的同时不增加同株异花授粉概率。这是从传粉生物学角度对植物克隆生长习性进化的一个全新的解释。今后, 克隆植物传粉生物学研究需要针对传粉者与克隆生长之间的相互作用建立理论模型, 探究克隆大小、克隆构型、花资源空间配置模式对传粉者访问频率和行为、花粉散布、交配格局的影响。同时, 需要在自然种群中, 尤其是克隆与非克隆的近缘类群、同一物种克隆与非克隆种群开展比较研究, 利用更高效的分子标记来研究克隆生长的生态与进化意义。     

Change in floral orientation in Anisodus luridus (Solanaceae) protects pollen grains and facilitates development of fertilized ovules

? Premise of the study: Some floral traits could be selected by pollinators and nonpollinator agents, and studying the floral traits shaped by physical agents could reveal adaptive mechanisms to the environment. We explored the adaptive significance of the change in floral orientation, from pendulous flowers to erect fruits that have a persistent calyx, in Anisodus luridus, a perennial native to the Qinghai-Tibet Plateau (QTP). ? Methods: We examined the effect of flower orientation on pollinator visitation rates, pollen deposition efficiency, pollen quantity, and pollen viability to estimate whether pendulous flowers have improved male fitness. We then measured seed production and seed germination rate to assess whether erect fruits have enhanced female fitness. ? Key results: Pendulous flowers did not have any preferred pollinators or increased pollen deposition. In artificially erected flowers, the number of pollen grains greatly decreased after rainwash. Pollen germination experiments indicated that pollen damage by water and exposure to solar radiation is serious. In the persistent calyx that holds water within it, the temperature inside the calyx changed slower than in the calyx that had the water removed. After supplemental pollination, the seed number, seed set, and seed mass of fruits that had water removed from the calyx were reduced greatly in both years. ? Conclusions: We conclude that the change in floral orientation could enhance male and female fitness of A. luridus and is effectively adaptive to the alpine environments, indicating a strong selection by the combined pressure from various abiotic nonpollinator agents in shaping the floral traits of this alpine plant.     

The role of a mixed mating system in the reproduction of a Mediterranean subshrub (Fumana hispidula, Cistaceae)

Knowledge about mixed mating systems can improve our understanding of the evolutionary dynamics of reproductive systems. Here we report a study of the pollination system (floral and reproductive biology, pollen limitation and stigmatic pollen load, floral visitors and inbreeding depression) of Fumana hispidula, a Mediterranean subshrub belonging to a species group with a strong selfing tendency. Autonomous self-pollination, hand self- and hand cross-pollination, open pollination and controls yielded fruits (0.28, 0.65, 0.68, 0.38, and 0.67, respectively); however, autonomous self-pollination resulted in the lowest fruit set. All individuals were fully self-compatible but we found great inter-individual variation in autonomous self-pollination ability, which was probably related to a variable expression of herkogamy degree. Inbreeding depression was low (0.040 for fruit set, 0.015 for seed set and ?0.026 for seed mass). The pollen supplementation experiment did not reveal pollen limitation and pollinators were seen visiting the flowers during the observation periods. These results support the idea that F. hispidula has a mixed mating strategy, which represents a successful reproduction mode in their patchy habitats.     

Pollination Biology of Distylous Rubiaceae in the Atlantic Rain Forest, SE Brazil

Abstract: Data on pollination biology constitute important clues for the comprehension of pollen flow and genetic differentiation in plant populations. Pollinator type, availability and behaviour may modify morphological and mating patterns in populations of typically distylous species. This study investigates the pollination biology of four distylous species of Rubiaceae in the Atlantic rain forest, SE Brazil. Data on flowering phenology, floral lifespan, stigmatic receptivity, pollen availability, nectar volume and concentration, and pollinator activity were collected. The species studied flower sequentially throughout the wet season, and produce terminal inflorescences which bear small, tubular, diurnal, nectariferous flowers. Despite these similarities, some of the species studied are pollinated by different groups of pollinators, probably due to their distribution, availability of flowers and corolla length. On the other hand, pollinator specificity does not seem to be so important for distylous species. Long mouthparts, like those of most of the recorded pollinators, may reach lower sexual organs and, together with the self- and intramorph-incompatibilities observed, be sufficient to perform legitimate pollination and maintain levels of intermorph mating.