新疆党参的花部综合征与次级花粉呈现

次级花粉呈现是植物提高花粉输出和接受的效率与准确性以及促进异交的一种繁殖策略.新疆党参(Codonopsis clematidea)具有典型的次级花粉呈现特性.本文采用野外观测、授粉实验与室内显微观测等方法,对该物种的花部综合征及其次级花粉呈现过程进行了详细观测,并对这些特征的适应意义进行了探讨.结果表明:新疆党参钟状下垂的蓝色花冠及黑黄色蜜腺,不仅避免了雨水冲刷雌雄蕊、稀释花蜜以及风移出花粉等不利影响,还增加了对传粉者的吸引,延长了花粉活力及柱头可授性持续时间.次级花粉呈现属于亚顶端花柱呈现者,呈现机制为沉积机制.该特性限制了单个传粉者带走的花粉量,延长了花粉呈现时间,使更多传粉者参与传粉过程,增加了雄性适合度.其雄性先熟及雌雄蕊空间位置变化,避免了雌雄功能干扰,为传粉者取食花蜜、输出花粉以及柱头接受异源花粉提供了通道和机会.其泛化传粉系统的主要传粉昆虫为林野熊蜂(Bombus silvarum)、草地熊蜂(B.paradoxus)和树长黄胡蜂(Dolichovespula sylvestris),其较长的单花花期(6 d左右),昆虫访花高峰期与花粉活力最高时期及柱头最佳授粉期相吻合,以及较大的泌蜜量、较高的花蜜糖浓度和较长的泌蜜时间等特征,可提高传粉者的访花频率,增加成功传粉的机会和传粉效率.在气候多变的天山山区,该物种特殊的花部综合征和次级花粉呈现机制对提高其传粉效率并促进异交繁殖成功具有重要意义.     

有花植物触敏柱头的闭合行为及其适应意义

有花植物触敏柱头的闭合行为一直是传粉生态学和进化生物学研究的热点问题之一。植物花的多裂片柱头在传粉者接触后迅速闭合的现象主要存在于玄参目的一些类群中。早在上个世纪初,植物学家就对花的柱头闭合现象进行过初步的研究,认为柱头闭合主要是由于花粉在柱头上萌发时吸收水分导致柱头细胞膨压降低而引起的;并对柱头闭合的适应性提出了一些假说。近年来,许多学者研究证实柱头闭合能促进花粉萌发及花粉管伸长,或减少花粉散出与柱头接受花粉间的相互干扰,提高传粉者访问过程中花粉的散播量,并且柱头的状态会直接影响传粉者的取食行为。到目前为止,对柱头闭合的机制及其避免自交的假说还存在许多争议。关于触敏柱头的闭合行为,尤其是其适应机制及其生态学意义还有待更加系统和深入的研究。本文概述了触敏柱头的研究进展,并对研究中尚存在的一些问题进行了讨论。     

贯叶连翘的开花动态与繁育系统研究

野外观察贯叶连翘的开花进程和花部形态特征,运用花粉萌发、杂交指数、花粉-胚珠比等方法测定其繁育系统。结果显示:贯叶连翘雌雄异熟,柱头先花药成熟,雌雄蕊无明显异位。单花花期4～5d。仅在开花当日有昆虫传粉,蜜蜂为主要传粉者。花粉在花药开裂1h后活力最大,萌发率达40.10%,花粉在柱头萌发3h后接近子房。根据杂交指数(OCI)推测其繁育系统属于异交,部分自交亲和,有时需要传粉者。花粉-胚珠比(P/O)则表明贯叶连翘的繁育系统为兼性异交。贯叶连翘结实率低,可能与花粉活力,花粉管的生长速度及花粉在柱头的竞争有关。     

澳古茨藻传粉生物学和繁育系统的初步研究

澳古茨藻（ＮａｊａｓｏｇｕｒａｅｎｓｉｓＭｉｋｉ）是典型的水下水媒传粉植物。雄花在花粉释放前２—４ｈ花梗迅速伸长,突破膜质鞘状外被,并且向外弯曲,至花粉释放盛期几乎是水平状态,有利于释放后的花粉直接进入水流以求传播。花部组成与结构极为简化,花粉中富含淀粉粒,花粉落置柱头前常萌发出长的花粉管,形成宜为柱头捕获的花粉笺,表现出对水下水媒传粉的高度适应。有性繁殖十分发达,自交、异交和混交的结实率都在８５％以上;花粉／胚珠（Ｐ／Ｏ）为２６９０±３００,指示兼性自交的繁育系统。无性繁殖较弱,仅以易长不定根的植株片段形式存在,但仍对该种的扩散具有重要意义。作者对澳古茨藻的花生物学特征、传粉机制以及繁育系统进行了探讨,对澳古茨藻表现出的许多沉水植物所特有的特征特性：花被的简化、花粉外壁的简化作了解释,讨论了繁育系统中自交与异交的关系。     

海滨锦葵花内五柱头裂片弯曲的独自调节(英文)

海滨锦葵花柱有5柱头裂片,若传粉失败,柱头裂片向下弯曲使柱头与自身花粉接触,发生延迟自交。传粉不足条件下发生的延迟自交可能提供繁殖保障。该研究定量分析了花粉搁置和授精胚珠数对花内未授粉柱头裂片运动的影响,并测定了花粉—胚珠比及柱头可授性和花粉生活力。结果表明,花内未授粉柱头裂片的弯曲不受其他柱头接受花粉量及授精胚珠数的影响,仅响应于其自身是否接受到花粉,花粉-胚珠比值显示海滨锦葵交配系统属兼性异交;未授粉柱头经弯曲与自身花粉授触时的强柱头可授粉和高花粉生活力为自交授粉的发生提供了可能。包括柱头裂片运动在内的多个花性状有机地展示了一种新花内混合交配系统,且花内柱头裂片弯曲的独自调节为从花水平验证被广泛接受的自交进化解释——繁殖保障假说提供了可能。     

动物传粉植物花粉呈现时序的进化意义

由动物传粉的植物为了使供体花粉能够被高效地传递到受体柱头上, 进化出了多种多样的花部特征, 花粉呈现时序便是其中之一。植物主要通过包装机制和分摊机制控制花粉呈现的速度以限制昆虫一次拜访转移的花粉量。花粉呈现理论(PPT)认为: 花粉逐步呈现策略倾向于出现在传粉者数量丰富但传粉效率低的植物中, 而花粉同时呈现策略则多出现在传粉者少但传粉效率高的植物中。本文对花粉呈现时序的相关研究进行了总结, 重点介绍: (1) 限制花粉转移的花部机制; (2)花粉呈现理论; (3)花粉呈现时序的适应意义。目前的研究主要集中在花粉呈现时序对传粉动物的适应性上, 但环境因子对花粉呈现时序也有一定的影响。PPT数学模型还不能完全预测特定环境中植物的花粉呈现时序。因此, 有必要选择合适的植物类群, 从花部综合特征、传粉系统、交配系统和环境等方面进行综合研究, 进一步揭示动物传粉植物花粉呈现时序的适应意义。     

三种耧斗菜属植物柱头的特征

为了深入了解植物捕获花粉的机制和生殖策略,我们对3种耧斗菜属Aquilegia L.植物华北耧斗菜A. yabeana Kitagawa、无距耧斗菜A. ecalcarata Maxim.和秦岭耧斗菜A. incurvata P. K. Hsiao的雌蕊进行了扫描电镜和荧光显微镜观察,发现其柱头发育的特殊式样。其柱头位于花柱的腹缝面,由花柱腹缝两侧细胞发育成柱头乳突,在花期中从顶端朝花柱基部渐次成熟。伴随花柱向外弯曲,成熟的柱头表面相继呈现,逐渐增加了花粉落置的空间。这种柱头发育的式样通过延长柱头的可授期,增加了植物在不可预测的传粉环境下的繁殖机会。     

青阳参花部特征及其传粉适应性

对青阳参花（Cynanchum otophyllum）部综合特征、访花昆虫种类、访花行为及传粉过程进行了研究,结果表明,青阳参花结构复杂,两个子房基部离生、花柱联合与雄蕊形成合蕊柱,柱头表面被邻近花药的侧翼紧密包围形成5个柱头腔。青阳参的花粉形成独特的花粉块,一次传粉过程可以转运大量的花粉。东方蜜蜂（Apis cerana）是青阳参的主要传粉昆虫,其传粉包括两个过程：（1）当蜜蜂的口器或足插入着粉腺的槽口后借助蜜蜂的力量将花粉块从花上拔起;（2）当蜜蜂再次访花时将携带的花粉块插入其中一个柱头腔。花粉块里面的花粉粒住柱头腔中萌发出花粉管,然后沿着花柱道向下生长最后进入子房。在整个花期仡粉保持有相对较高的生活力,而其柱头可授性则在7天后逐渐降低。     

橙花瑞香的繁殖特性研究

瑞香属植物具有重要的药用和观赏价值,在中国资源丰富,但自然条件下低坐果率限制了该属植物的进一步开发和利用。该研究以橙花瑞香为对象,通过对其有性繁殖及传粉特性的研究,探索其自然坐果率低的原因,内容包括花部特征的测量分析,MTT染色法测定花粉活性,联苯胺-过氧化氢法测定柱头可授性,扫描电镜观察柱头、花粉的形态,传粉者观察,通过花粉胚珠比(P/O)和人工授粉实验推测橙花瑞香的繁育系统类型。结果表明:橙花瑞香的花部结构特殊,管状小花,花药两轮,雌雄蕊分离。花开后的花粉具有活性,柱头具有可授性,扫描电镜下,柱头和花粉的结构没有发育异常,且柱头上有花粉落置。橙花瑞香的传粉者主要是夜间访花的蛾类,访花频率低。P/O及人工授粉实验表明橙花瑞香的繁育系统为专性异交。橙花瑞香的坐果率非常低,自然坐果率为1.4%,人工异花授粉为23.3%,低坐果率可能是受其开花量大、异花花粉限制、资源限制以及花部结构等因素的影响。     

珍稀濒危物种格木传粉方式和交配系统的初步研究

为研究格木(Erythrophleum fordii Oliv.)的传粉方式和交配系统,在鼎湖山国家级自然保护区对珍稀濒危树种格木进行野外观察,花的微器官(花粉和柱头)用扫描电镜观察,并进行控制授粉实验和遗传分析。结果表明,格木具有颜色亮丽和散发香味的总状花序,格木花粉包被厚重的粘性物质,柱头没有易于捕获花粉的结构,风媒传粉没有坐果,推断格木应该是通过虫媒传播花粉,或者主要是通过虫媒传播。遗传分析和在不同繁殖环境的两棵目标树的坐果情况表明,格木偶尔可以自交产生后代,但异交占据着绝对的优势,鼎湖山格木种群的异交率达到90.6%。格木传粉方式和交配系统的研究有利于对格木进行保护和保育。     

Evolution of pollination system and characters of stigmas in Najadales

Diverse ways of pollination, including aerial, epihydrophilous and hypohydrophilous ones occur in the Najadales. Although the knowledge of pollination biology in this order has been largely accumulated in recent years, most researchers ignored the role of stigmas in the pollination process. Based on the previous works, we supplemented observations on stigmatic characters and pollination mechanism in nine species from four families. The flowers of Aponogeton lakhonensis ( Aponogetonaceae ), Triglochin palustre ( Juncaginaceae), Potamogeton crispus, P. distinctus, P. gramineus, P. pectinatus, P. perfoliatus, P. pusillus (Potamogetonaceae), are all found to open above or on water surface. Stigmas in these species have a large surface area. Correspondingly, the pollen grains are all found to be globular. Of the eight species, except for P. pectinatus and P. pusillus, the remaining six obviously expose their stigmas out of the perianth. Another species , Najas marina (Najadaceae), has filiform stigmas and ellipsoidal pollen grains which can germinate in water before being transferred onto stigmas in natural conditions. Pollination takes place in this species strictly under water. The characters of pollen and stigmas could be divided into two types, and the pollination system is of two kinds of mechanisms of stigmas capturing pollen in the Najadales. The species with globular pollen grains have wide, large stigmas while those producing filiform or precocious pollen grains, which are likely to be captured by stigmas during the pollen dispersal by water currents, normally have filiform stigmas. It is inferred that various water stresses might have resulted in the diversification of pollinationsystem in the Najadales.     

TWO-DIMENSIONAL POLLINATION IN HYDROPHILOUS PLANTS: CONVERGENT EVOLUTION IN THE GENERA HALODULE (CYMODOCEACEAE), HALOPHILA (HYDROCHARITACEAE), RUPPIA (RUPPIACEAE), AND LEPILAENA (ZANNICHELLIACEAE)

In most plant species with abiotic pollination systems, pollen is dispersed in three dimensions. Theoretical considerations suggest, however, that there are significant advantages for two-dimensional pollination systems over three-dimensional systems, especially if pollen is dispersed in conveyances or aggregations of large diameter. We report that two-dimensional pollination systems occur in the genera Halodule, Halophila, Lepilaena, and Ruppia, where pollen grains are not transported through the water singly, but in rafts or search vehicles. These genera possess unusual pollen morphologies which facilitate assemblage of pollen grains into search vehicles. These floating search vehicles have large diameters, thus greatly increasing probability of encountering a stigma. The grains have a hydrophobic surface that appears to mediate adhesion by an external coating of proteins and carbohydrates. Similar adaptations to two-dimensional pollination are found in the target organs, the stigmas. The long filamentous stigmas of the marine genera float, as do the indusiate stigmas of the freshwater genera, creating small depressions in the water surface. Pollination occurs through the collision of a moving search vehicle with a floating stigma. The existence of similar pollen search vehicles, stigma morphologies, and flowering phenologies in these unrelated hydrophilous genera evidences convergent evolution towards efficient search strategies in surface-pollinated aquatic plants.     

Abiotic pollen and pollination: Ecological, functional, and evolutionary perspectives

The transport and capture of pollen in ~20% of all angiosperm families occurs in air and water. In other words, pollination is abiotic and occurs via the fluid media, not an animal vector. Whereas some early concepts considered abiotic pollination to be largely a stochastic phenomenon, there is sufficient evidence to indicate that wind pollination (i.e. anemophily) and water pollination (i.e. hydrophily) have deterministic features and are sophisticated fluid dynamic solutions to the problem of pollen release, dispersal, and capture.An abiotic pollination syndrome is defined in which there is spatial or temporal separation of carpellate and staminate flowers, which are drab, a reduction in perianth parts, stigmas and anthers are exposed to the fluid, and typically unclumped pollen may be produced in large amounts. Separate pollination syndromes are defined for anemophilous (i.e. wind-pollinated), ephydrophilous (i.e. surface-pollinated), and hydrophilous (i.e. submarine-pollinated) plants. Distinctions are based on habitat and physical conditions for pollination, pollen size, shape, and ultrastructure, morphology and ultrastructure of stigmas, and outcrossing rates. For example, anemophilous pollen are spherical and small, ephydrophilous pollen are spherical or reniform and large, while hydrophilous pollen are filiform (i.e. filamentous) or functionally filiform. The pollination mechanisms and mechanics associated with these syndromes reveals a strong evolutionary relationship between plant morphology and fluid dynamics.     

Interspecific variation in pollen–ovule ratio is negatively correlated with pollen transfer efficiency in a natural community

The pollination efficiency hypothesis has long been proposed as an explanation for interspecific variation in pollen–ovule (P:O) ratios. However, no empirical study on P:O ratios has directly and quantitatively measured pollen transfer efficiency (PE). Here, we use a PE index, defined as the proportion of pollen grains removed from anthers that are subsequently deposited on conspecific stigmas, as a direct and quantitative measure of PE. We investigated P:O ratios, pollen removal and pollen deposition in 26 plant species in an alpine meadow, over three consecutive years. Our community survey showed that nearly 5% of removed pollen was successfully deposited on conspecific stigmas. The PE index ranged from 0.01% up to 78.56% among species, and correlated negatively with the P:O ratio across years. This correlation was not changed by controlling for phylogenetic relationships among species, suggesting that the interspecific variation in P:O ratios can be attributed to the probability of pollen grains reaching a stigma. The results indicate that the pollination efficiency hypothesis can help to explain interspecific variation in P:O ratios.     

The control of incompatibility in distylous Pulmonaria affinis Jordan (Boraginaceae)

In P. affinis, pin pollen is shorter on average than thrum pollen. Pins have more coronal hooks on stigmatic papillae than thrums, but gaps between papillae are relatively smaller for thrums. Pin stigmas receive more pollen than thrum stigmas. Thrum stigmas receive more (dissortive) pin pollen, but pin stigmas are assortively pollinated. Pollen only germinates when trapped below papilla coronas. On thrum stigmas, most trapped pollen is pin. Pollen germination is better on thrum stigmas than pin stigmas, and thrum stigmas show a close relationship between numbers of legitimate pollen grains, numbers of germinating grains, and numbers of pollen tubes in the style. There is no inhibition of illegitimate pollen germination. Illegitimate pollen tubes are inhibited in the style. Incompatibility operates by a combination of dissortive pollination, dissortive pollen trapping, and stylar pollen tube inhibition. All heteromorphic features differing between pins and thrums are implicated in the inhibition of within-morph fertilization in thrums.     

Surface and submarine pollination in the seagrass Zoster a marina L

COX, P. A., LAUSHMAN, R. H. & RUCKLESHAUS, M. H., 1992. Surface and submarine pollination in the seagrass Zostera marina L. Hydrophilous plants can be divided into three ecological categories depending upon whether their pollen is transported above, on, or under the water surface. A mixed mode of submarine and surface hydrophilous pollination occurs in the seagrass Zostera marina L. In the surface mode of pollination, pollen rafts or 'search vehicles' which superficially resemble snowflakes, form at low tide and are transported on the surface of the sea by winds and water currents. Some of the search vehicles collide with the floating female stigmas, effecting pollination. In the submarine mode of pollination, small pollen masses resembling whisk brooms travel beneath the surface of the water. Although we failed to observe a submarine pollination event, SEM analysis of stigmas from subtidal populations confirms that submarine pollination does occur in Z. marina . However, observations of stigmas positioned at and below the surface of the water show surface pollination to be highly efficient. Electrophoretic evaluation of both subtidal and intertidal populations indicates significant genetic variation between populations. Given the high flux rates of surface-borne pollen and pollen viability in excess of 5 h, it is likely that surface-borne pollen is a major source of gene flow in Zostera marina populations.     

蓖齿眼子菜繁殖多样性研究

 繁殖是影响水生植物生存竞争、分布以及群落结构动态变化的关键因素。通过野外观察和室内试验,较详细地介绍了蓖齿眼子菜(Potamogeton pectinatus)的繁殖策略。研究结果表明:蓖齿眼子菜有性繁殖中具有水表和水下气泡两种传粉方式;种子发生量较大,但种子发芽率低,不到6%,种子繁殖对种群恢复贡献不大,其主要作用在于远程传播和保持持久的种子库。蓖齿眼子菜具有广泛而高效的无性繁殖系统,可以通过断株、根状茎、地上块茎、地下块茎、地上茎节以及叶腋基部进行繁殖,其中最重要的是地下块茎和根状茎,次年种群恢复主要来自于地下块茎和根状茎。此外,对采用蓖齿眼子菜进行沉水植被恢复与重建提出了建议。     

Pollination Rates and Pollen Tube Growth in a Vulnerable Plant,Liriodendron chinense (Hemsl.) Sarg. (Magnoliaceae)

The limited seed production of insect-depended plant, Liriodendron chinense was once considered to be pollen-limited and insufficient cross pollination. In this study, we counted pollen grains deposited on stigmas in three populations in Guizhou, Hunan and Zhejiang provinces of China respectively. Over 61.9% stigmas were pollinated. From 1994 to 1996, the mean number of pollen grains on each stigma ranged from 4.4 to 42.6, much more than ovules(2) in each pistil. Based on observations of three years, both the pollination rate and pollen quantity on stigmas significantly affected seed set. When flowers opened without stamens dehiscencing at the early stage of anthesis, stigmas received considerable quantity of pollen grains. Pollen grains from different sources were able to germinate and pollen tube growth rates were not greatly variable. It is very likely that pollen grains arrived first would fertilize eggs. Since only several pollen tubes went through the stylar canal, the potential pollen competition may exist. In this case, there would be strong selection on floral syndrome which benefit early insect visits. Pollen grains from the early visits would have a greater chance to fertilize ovules than those from later visits, which implies that cross pollination is the predominant breeding system of this plant. The conclusion was also confirmed by following four artificial experiments. Three treatments, including flowers bagged, netted or with the perianth removed, all reduced seed set clearly, but flowers with the stamens removed (emasculation)had a higher seed production by open pollination. As the rates of deposited stigmas in three populations were 6～8 times more than full seed set, we considerthat lower seed production in this plant may not mainly be due to pollen limitation.     

The plot thickens: does low density affect visitation and reproductive success in a perennial herb, and are these effects altered in the presence of a co-flowering species?

Plants may experience reduced reproductive success at low densities, due to lower numbers of pollinator visits or reduced visit quality. Co-occurring plant species that share pollinators have the potential to facilitate pollination by either increasing numbers of pollinator visits or increasing the quality of visits, but also have the potential to reduce plant reproductive success through competition for pollination. I used a field experiment with a common distylous perennial (Piriqueta caroliniana) in the presence and absence of a co-flowering species (Coreopsis leavenworthii) in plots with one of four different distances between conspecific plants. I found strong negative effects of increasing interplant distance (related to conspecific density) on several components of P. caroliniana reproductive success: pollinator visits to plants per plot visit, visits received by individual plants, conspecific pollen grains on stigmas, outcross pollen grains on stigmas, and probability of fruit production. Although P. caroliniana and C. leavenworthii share pollinators, the co-flowering species did not affect visitation, pollen receipt or reproductive effort in P. caroliniana. Pollinators moved very infrequently between species in this experiment, so floral constancy might explain the lack of effect of the co-flowering species on P. caroliniana reproductive success at low densities. In co-occurring self-incompatible plants with floral rewards, reproductive success at low density may depend more on conspecific densities than on the presence of other species.     

INCOMPATIBILITY AND THE POLLEN ECONOMY OF JEPSONIA PARRYI

Jepsonia parryi (Saxifragaceae) has heterostylous flowers and is strongly self-incompatible. Pin flowers have long styles, large stigmas, short stamens, and numerous, small pollen grains with finely sculptured walls. Thrum flowers have short styles, small stigmas, long stamens, and fewer, larger pollen grains with coarsely sculptured walls. Pin plants and thrum plants occur in a 1:1 ratio in field populations. Although the insect pollinators of J. parryi transfer ample compatible pollen to pin and thrum stigmas to account for full seed production, much of the pollen deposited on stigmas is incompatible. Analysis of the pollen deposits on stigmas collected from field populations indicates that compatible “legitimate” pollination of pin and thrum flowers is essentially random and is not obviously aided by floral dimorphism. It is suggested that although heterostyly had a positive adaptive value in the past evolutionary history of Jepsonia it is no longer adaptive under the present pollination regime, although it is maintained because of its strong genetic fixity.