植物雄蕊合生的多样性、适应意义及分类学意义初探

定义并总结了雄蕊合生现象,对雄蕊合生的类型、适应意义、进化及分类学意义进行了初步的讨论和总结.本文定义"雄蕊合生"为,花内雄蕊与雄蕊之间部分或整体的结合生长,既包括愈合与黏合,也包括结合紧密的贴生和靠生.根据雄蕊合生部位的不同,将雄蕊合牛分为"花丝合生"、"花药合生"和"花丝花药均合生"等三大类.每一类中还存在着合生程度不同的子类型.雄蕊合生的结构还可能进一步与柱头或花柱合生在一起,形成"合蕊柱"一类的复杂结构.雄蕊合生在裸子植物较进化的科如百岁兰科和买麻藤科中就出现了,在种子植物中集中分布在近50个科内,且从系统关系来看,雄蕊合生各个类型可能发生过多次进化上的反复.在被子植物中,花丝合生类型更多地与较早进化的类群联系在一起,且多出现在离瓣花中;花药合生及花丝花药均合生则较晚进化,多与有着明显花冠筒的合瓣花伴随出现.这暗示着,雄蕊的合牛结构有可能与花部其他特征一起影响了植物的繁殖过程,具有一定的适应意义.目前,还没有实验研究针对雄蕊合生各个类型开展实验以揭示其发育机制和适应意义.理论上,花丝合乍能增强雄蕊的强度,有时候还形成了围绕子房与花柱的杯状、管状或环状结构,能对子房和花柱有着保护作用且承受传粉者在花内移动的压力:花丝合生还可将雄蕊固定在一个较为稳定的位置,使得花药接触传粉者身体的部位相对固定,减少了花粉浪费.花药合生能将花内花药都集中到同一个位置,花粉接触传粉者身体的同一部位,从而降低花粉损耗和提高异交授粉的准确性.花丝花药均合生可以同时具有以上两种合生方式的适应意义,而且还极大改变了雄蕊的结构与空间位置,改变了花药与柱头之间的空间位置(雌雄异位),对花内自交可能性和雌雄功能干扰有着潜在的影响.雄蕊合生各个类型的适应意义及其对访花者类型与行为和植物繁殖策略的影响,还需要开展实验进行针对性的研究.由于雄蕊形态与结构较为稳定,雄蕊合生的不同方式以及合生程度可以作为科与种的分类参考.     

光果甘草二体雄蕊的发育及其适应意义

甘草属(Glycyrrhiza)植物具“9 + 1”二体雄蕊, 其中9枚合生雄蕊的上部花丝分离, 分离的花丝在发育过程中存在由早期长、短交错的二组排列方式转变为后期以雄蕊管最长的1枚雄蕊为中心向两边渐次缩短的倒“V”形排列。为了解这种雄蕊发育动态、分化现象及发育成熟后的适应意义, 该文以光果甘草(G. glabra)为实验材料, 比较了雄蕊发育过程中的形态变化、成熟花粉的理化性质及在传粉中的作用。结果显示: 雄蕊发育早期长、短两组雄蕊在花药大小与形状上存在分化, 但后期伴随着花丝的快速生长与花粉的成熟、散出, 花药大小与形状趋于一致; 花粉组织化学成分及授粉成功率无差异, 但成熟花粉的数量和花粉活力存在差异; 去雄处理虽然使访花者在一天内的两个访花高峰期的访花频次降低, 但结实率高于自然对照, 说明以异交为主的花去除雄蕊后, 降低了雌、雄蕊间的功能干扰, 提高了传粉昆虫的授粉率; 发育早期长、短交错排列的二组雄蕊到成熟期时发生的倒“V”形排列的转变, 使不同数量与活性的花粉分布在花内不同空间, 最大化接触访花者, 实现了资源节约, 提高了雄蕊的雄性适合度, 即在有限的空间内用最节约的雄性资源投入、使传粉空间与传粉几率最大化的方式, 来提高雄性功能。     

两性花的雄蕊运动:多样性和适应意义

雄蕊运动指雄蕊在自身能量支持下发生的主动运动,不包括雄蕊在访花者触碰下造成的被动位移。该文总结了雄蕊的应激运动、快速猛烈弹射、缓慢运动以及级联运动等4种主要类型,分析了这些运动类型的系统分布及繁殖适应意义等方面的研究进展。雄蕊的应激运动由访花者或其他外力诱发,可能起到促进散粉和实现繁殖保障的作用;雄蕊快速猛烈的弹射运动可将花粉猛然撒向空中或访花者身上,促进了花粉的风媒或虫媒散布;缓慢运动的雄蕊可能通过在不同花期改变雄蕊的空间位置和雌雄异位程度来调节繁殖策略,或主动将雄蕊花药移至特定部位(如柱头表面)实现自交;雄蕊逐一、依次发生的级联运动较为复杂,主要分布在刺莲花科、梅花草科、旱金莲科和芸香科中,目前还缺乏实验研究;但根据"花粉呈现理论"以及其他类型的雄蕊运动研究结果,雄蕊的级联运动可以将花粉分批呈现给不同的传粉者,通过不同传粉者的分别传粉来提高花粉的输出;而且可避免已散粉雄蕊对即将散粉雄蕊的干扰,同时可能也降低了雌雄功能干扰和(或)花内自交。在芸香(Ruta graveolens)中,级联运动之后的雄蕊还会在花末期再同时向花中央运动;这种多向、多次运动方式是目前发现的最复杂的雄蕊运动类型。雄蕊运动领域值得今后开展进一步实验研究的方向主要有:1)雄蕊运动尤其是级联运动对雌雄功能干扰(性别间干扰)、雄蕊与雄蕊的"性别内干扰"等植物繁殖格局的影响;2)雄蕊运动与雌雄异熟、雌雄异位等花部特征的相互作用;3)雄蕊运动复杂类型的生理与发育机制。     

雄蕊合生植物半边莲的花部综合征与繁育系统

 为了解花内雄蕊合生结构的繁殖适应意义, 初步研究了雄蕊合生植物半边莲(Lobelia chinensis)的花部综合征、传粉特性和繁育系统。半边莲花大且鲜艳, 花瓣中下部弯折并合生成背部有裂缝的不封闭的花冠筒。5雄蕊的花药紧密合生成花药筒, 花丝中上半部也合生在一起, 只有花丝基部分离插生于花冠筒上。柱头被包裹在花药筒内。半边莲单花寿命可达5 d左右。雄性先熟, 柱头在伸出花药筒之后才具活性。花的主要访问者为蚂蚁、食蚜蝇和苍蝇类等小型昆虫。半边莲单花花粉数约为(5 200±130)粒、胚珠数约为(55±6)颗, 花粉胚珠比为94.54, 应属于兼性自交繁育系统, 但异交指数和其它特征都显示其以异交为主, 部分自交亲和。套袋和人工授粉实验发现, 半边莲不存在无融合生殖与自发自交, 但自交亲和性高。雄蕊合生(尤其是花药的合生)能把花药中的花粉聚拢在一起在传粉者的一次访问中就被同时带出, 与同样具有较低花粉胚珠比的花粉聚联(Pollen aggregation)传粉过程近似。半边莲的雄蕊合生结构(花药合生成筒、花丝上部也合生)可能与一些特定的花部特征, 如花被合生成未完全封闭的筒、雌雄异熟以及低花粉胚珠比等联系在一起, 形成了适应小型传粉者的“花部综合征”     

刺五加花的形态学及雄蕊异长现象的观察

刺五加Eleuthercoccocus senticosus (Rupr．et Maxim．)Maxim．具有花丝长度不同的三种植株类型,本文报道了其花序和花的形态结构,开花式样,异长雄蕊现象及相应的花柱长度。结果表明：长花丝长3．96～4. 44mm ,短花丝长0．45～0．93mm,中花丝长2．16～3．08mm。伴随异长雄蕊特征的其它表现在于花药大小、花药形状和颜色、花粉大小、花粉数量、柱头宽度和雌雄蕊育性。其中,长花丝对应的花药饱满、白色、花粉量大、花柱较短、雌性败育;短花丝对应的花药小、黄褐色、无花粉、花柱较长、雌性可育;中花丝对应的花药中等、花粉粒较大、雄性半不育、雌性可育。据此,刺五加被认为是单全异株的,其长花丝类型是雄性、短花丝类型是雌性、中花丝类型是两性的。另外,对三种性别形态类型的关系进行了讨论,并认为将雌株作为变种“短蕊刺五加”var.brevistaminea 显然是不恰当的。     

华北蓝盆花的开花特性及传粉生态学研究

华北蓝盆花具头状花序 ,花冠蓝紫色 ,1朵花的花期约为 7～ 8d。边缘花比中部花大 ,但二者在花药大小、花丝长度、花柱长度、柱头直径等方面无显著差异。花粉寿命约为 4 d,花柱在花开后逐渐伸长 ,柱头微二裂、干燥型。同一花序内的花朵在 2～ 3d内逐渐开放 ,晚开的花散粉时、早开的花花柱仍未伸长 ,因此 ,花药与柱头之间有空间间隔和时间间隔 ,同一花序内的各花朵间相互传粉的机会很低。按照杂交指数 ,其繁育系统属于异交、部分自交亲和、需要传粉者。依据 P/ O数据 ,其繁育系统属于兼性异交。罩网试验显示 ,华北蓝盆花没有自发的自花传粉现象 ,自然条件下也没有无融合生殖现象 ,没有访花者访问便不能座果。在华北蓝盆花花序上记录到的访花者有 10余种 ,以双翅目、鞘翅目、膜翅目昆虫为主 ,最常见的访花者是雏蜂虻和豆芫菁。开花后不同日期 ,访花者的访花频率有所不同 :开花第 2天到第 7天的花序上 ,访花者的访花频率较高 ;开花第 4天 ,花序上访花者的访花频率最高。同一天内的不同时间 ,访花者的访花频率也有差异 ,一般规律是 :11:0 0～ 15 :0 0之间 ,访花者的访花频率较高。此外 ,气温骤降、阴天及降雨能使访花者的访花频率明显降低。     

兰科雄蕊发育多样性研究进展

兰科是被子植物中多样性最丰富的家族,其雄蕊形态和功能分化在亚科间变化明显,是该物种多样性形成及适应性传粉生物学的研究重点。基于现有研究资料,该文初步归纳了兰科雄蕊发育多样性的主要研究内容及现状,为野生兰花资源的保护与利用提供科学依据。结果如下:(1)可育雄蕊数目的减少和花粉愈合程度的增加在兰科分子系统树上呈明显的平行演化趋势。(2)兰科雄蕊数目的减少和功能分化与早期花器官发生中存在大量的滞后和缺失、次生融合与分裂现象等密切相关。(3)花药开裂时的4类散粉单元的花粉超微形态特征在亚科、族、亚族、属和种间差异明显。(4)兰科花药散粉单元可以为单花粉粒,也可以通过花药发育过程中源自绒毡层的三类黏性物质而聚合成不同的散粉单元,包括花粉鞘、弹性黏素和其他黏性物质。(5)花药发育揭示了兰亚科的花粉小块结构主要有三类(红门兰型、树兰型和过渡型),树兰亚科的不同数目(2、4、8)和形态(全缘、浅裂、深裂、孔裂)的花粉团是由于花药原基分化出的不育隔膜组织的数目、朝向和位置而形成的。(6)兰科花药发育中,花药室数目、花药壁发育类型、绒毡层细胞核数目、不育隔膜组织分化、胞质分裂类型、小孢子四分体排列形式、花粉细胞核数目等在亚科和属间差异明显。然而,由于种类繁多,现有研究资料难以为理解兰科雄蕊发育提供清晰的线索,包括雄蕊的发育模式、散粉单元的形成机制、花药发育的胚胎学特征等。因此,有必要重视兰科雄蕊发育研究,包括扩大取样范围和利用多学科技术方法和修订兰科花形态常用术语等。     

大麦小穗结构与开花特性研究──雄蕊在开闭花过程中的生长动态和结构变化

开花前,雄蕊花丝细胞中的淀粉等物质水解,细胞水势下降而吸胀,花丝伸长。随着＂小花轴＂中物质的输入,细胞进一步吸水膨大,花丝迅速伸长,花丝维管束中的导管被拉断,薄壁细胞内膜系统崩解,细胞自溶,降解物质＂撤回＂,＂小花轴＂被重新分配利用。开花后,花药表面大量失水,药壁开裂传粉。不育系雄蕊花药药隔小、发育不良,绒毡层发育和行为异常,其花药通常为空药室或花粉败育。用可育系花粉对不育系小花授粉,其小花能逐渐关闭。     

琼花生殖器官结构及传粉昆虫的观察

为了探讨影响琼花Viburnum macrocephalum f.keteleeri有性生殖的因素,对其生殖生物学和传粉生物学进行了研究。研究内容主要包括琼花花部特征、生殖器官解剖结构、花粉活力、花粉胚珠比(P/O)、繁育系统、传粉形式、传粉昆虫和花粉管生长路径等。结果表明:(1)琼花聚伞花序由大型不孕花和小型可孕花组成,可孕花雌雄蕊发育正常,雄蕊5枚,雌蕊1枚,干型柱头,单子房,倒生胚珠;不孕花的雌雄蕊在发育早期正常,而在花期时退化。雄蕊退化表现为雄蕊消失、花丝缩短或消失或花药大小不一;而雌蕊退化表现为雌蕊缩小或柱头开裂;有时雌雄蕊也存在瓣化现象。(2)自然条件下,单花花粉活性在散粉4–5d后明显下降,居群花粉活力在4月25日–28日开始显著下降。(3)花粉胚珠比P/O值为12800–18700,属专性异交型;繁育系统为异株异花授粉,属虫媒传粉植物。(4)可孕花大量散粉时间为9:00–16:00,昆虫访花高峰时段为11:00–15:00,访花昆虫中以蝶类和蜂类为主,蝶类访花频率最高。(5)落置柱头的花粉萌发率较高,花粉粒授粉后1h左右开始萌发,花粉管从柱头乳突细胞的间隙穿入花柱,沿花柱中央引导组织生长,18h左右进入子房,20h左右从珠孔进入胚囊。讨论了琼花的不孕花形成、花部结构适应、花型演化趋势、低结实率和花粉管生长特点。     

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。     

Intrafloral differentiation of stamens in heterantherous fowers

Flowers that have heteromorphic stamens (heterantherous flowers) have intrigued many researchers ever since the phenomenon was discovered in the 19th century. The morphological differentiation in androecia has been suggested as a reflection of "labor division" in pollination in which one type of stamens attracts pollina-tors and satisfies their demand for pollen as food and the other satisfies the plant's need for safe gamete dispersal. The extent and patterns of stamen differentiation differ notably among taxa with heterantherous flowers. Seven species with heteromorphic stamens in three genera were sampled from Leguminosae and Melastomataceae, and the morphological difference of androecia, pollen content, pollen histochemistry and viability, pollen micro-morphology, as well as the main pollinators were examined and compared. Pollen number differs significantly between stamen sets of the same flower in most species investigated, and a correlation of pollen number and anther size was substantiated. Higher pollen viabilities were found in the long (pollinating) stamens of Senna alata (L.) Roxb. and S. bicapsularis (L.) Roxb. Dimorphic pollen exine ornamentation is reported here for the first time in Fordiophytonfaberi Stapf. The height of stigma and anther tips of the long stamens in natural conditions was proved to be highly correlated, supporting the hypothesis that they contact similar areas of the pollinator's body.     

Adaptive Significance of Floral Movement

Since Darwin observed the reconfiguration of pollinia in orchards and referred to it as a function to reduce self-pollination, diverse floral movements have been investigated and various hypotheses have been proposed to explain their adaptive significance. However, adaptive significance of floral movement in some species has yet to be fully explained. Increasing evidence suggests that some floral movements, which have previously been considered as a mechanism to avoid self-pollination, may act as a mechanism to reduce intrafloral male-female interference. In this review, we first explore insect-induced floral movement—such as secondary pollen presentation—that enhances the efficiency and accuracy of pollination. Secondly, we outline the active movements of different floral structures of pistil (style), stamen (filament, anther, pollen), and corolla, such as flexistyly, pollen sliding and catapulting, and anther rotation. Thirdly, we introduce movement herkogamy, which makes herkogamy decrease or increase via the movements of floral structures, especially decreased approach herkogamy via style curvature in the Malvaceae with a monadelphous column. Fourthly, we highlight the different adaptive significances in floral movements for understanding their evolution, including reduction in intrafloral male–female interference, promotion of outcrossing and/or avoidance of self-pollination, delayed autonomous selfing, and tolerance to harsh environments. In particular, we explore the possibility of three functions of one floral movement in one species, which differs from the generally recognized and conventional notion of one floral movement in one species with only one or two functional mechanisms. Finally, we provide perspectives on the challenges and opportunities for using demographic and molecular genetic approaches to (i) study the relative importance and evolutionary mechanism of different adaptive significances in one floral movement; and (ii) simultaneously investigate the floral movement and correlative traits of broader species in related or unrelated families to test how they evolved and the evolutionary relationship between their functions.     

Up and down: stamen movements in Ruta graveolens (Rutaceae) enhance both outcrossing and delayed selfing

Background and Aims

Stamen movements directly determine pollen fates and mating patterns by altering positions of female and male organs. However, the implications of such movements in terms of pollination are not well understood. Recently, complex patterns of stamen movements have been identified in Loasaceae, Parnassiaceae, Rutaceae and Tropaeolaceae. In this study the stamen movements in Ruta graveolens (Rutaceae) and their impact on pollination are determined.

Methods

Pollination effects of stamen movements were studied in Ruta graveolens, in which one-by-one uplifting and falling back is followed by simultaneous movement of all stamens in some flowers. Using 30 flowers, one stamen was manipulated either to be immobilized or to be allowed to move freely towards the centre of the flower but be prevented from falling back. Pollen loads on stigmas and ovule fertilization in flowers with or without simultaneous stamen movement were determined.

Results

Pollen removal decreased dramatically (P < 0·001) when the stamen was stopped from uplifting because its anther was seldom contacted by pollinators. When a stamen stayed at the flower''s centre, pollen removal of the next freely moved anther decreased significantly (P < 0·005) because of fewer touches by pollinators and quick leaving of pollinators that were discouraged by the empty anther. Simultaneous stamen movement occurred only in flowers with low pollen load on the stigma and the remaining pollen in anthers dropped onto stigma surfaces after stamens moved to the flower''s centre.

Conclusions

In R. graveolens pollen removal is promoted through one-by-one movement of the stamen, which presents pollen in doses to pollinators by successive uplifting of the stamen and avoids interference of two consecutively dehisced anthers by falling back of the former stamen before the next one moves into the flower''s centre. Simultaneous stamen movement at the end of anthesis probably reflects an adaptation for late-acting self-pollination.     

MORPHOGENESIS OF THE STAMENLESS-2 MUTANT IN TOMATO. I. COMPARATIVE DESCRIPTION OF THE FLOWERS AND ONTOGENY OF STAMENS IN THE NORMAL AND MUTANT PLANTS

The single gene recessive mutant stamenless-2 (sl₂/sl₂) differs phenotypically from the normal (+/+) only in the stamen structure. Stamens of the mutant plants were laterally free, twisted, shorter, paler in color, possessed abnormal pollen, and bore naked external ovules (E.O.) on the adaxial surface near the junction of anther and filament. Mutant plants grown in the field during summer produced flowers in which a number of carpel-like organs (‘carpelloid stamens‘) with few or no E.O. replaced the stamens. On the other hand, plants grown in the greenhouse during winter possessed flowers with greater number of yellow and pubescent stamens and many E.O. Study of stamen ontogeny revealed that at initiation (up to 100 μ in length) stamen primordia of normal and mutant plants resembled each other. Thereafter the development of stamens in the two genotypes could be distinguished.     

Comparative structure and pollen production of the stamens and pollinator-deceptive staminodes of Commelina coelestis and C. dianthifolia (Commelinaceae)

BACKGROUND AND AIMS: Flowers of Commelina coelestis and C. dianthifolia provide pollen alone as a floral reward, and rely on visual cues to attract pollinators. Three stamen types, all producing pollen, occur in each of these species: two cryptically coloured lateral stamens, a single cryptically coloured central stamen and three bright yellow staminodes that sharply contrast with the blue to purple corolla. The objective was to compare the stamen structure and pollen characteristics of each of the three stamen types, and to test the hypothesis that the staminodes are poor contributors of viable pollen for the siring of seed. The pollination roles of the three stamen types and the breeding systems of both species were also explored. Methods: Light, fluorescence and scanning electron microscopy were utilized to examine stamen morphology and pollen structure and viability. Controlled hand pollinations were used to explore the breeding system of each species. Filament and style lengths were measured to investigate herkogamy and autogamy. KEY RESULTS: Pollen from all stamen morphs is viable, but staminode pollen has significantly lower viability. Pollen polymorphism exists both (a) between the lateral and central stamens and the staminodes, and (b) within each anther. Lateral and central stamens have thicker endothecia with a greater number of secondary cell wall thickenings than the staminodes. CONCLUSIONS: Both species are entomophilous and facultatively autogamous. Lateral stamen pollen is important for cross-pollination, central stamen pollen is utilized by both species as a pollinator reward and for delayed autogamy in C. dianthifolia, and the staminodes mimic, by means of both colour and epidermal features, large amounts of pollen to attract insects to the flowers. Pollen from all three anther morphs is capable of siring seed, although staminode pollen is inferior. The thin staminode endothecium with fewer secondary thickenings retards staminode dehiscence.     

Stamen dimorphism in Rhododendron ferrugineum (Ericaceae): development and function

The function of stamen dimorphism in the breeding system of the alpine shrub Rhododendron ferrugineum was studied in two populations in the French Alps. This species has pentameric flowers with two whorls of stamens: an inner whorl of five long stamens and an outer whorl of short stamens. We studied the development of stamens from buds to mature flowers (measurement of the filament, anther, and style lengths at five successive phenological stages) and compared the size and position of reproductive organs at maturity in control and partially emasculated flowers (removal of long-level stamens) to determine whether the presence of long-level stamens constitutes a constraint for the development of the short-level ones. Stamen dimorphism can be observed early in stamen development, from the bud stage of the year prior to flowering. At this early stage, meiosis had already occurred. Emasculation of the long-level stamens induced the short-level ones to grow longer than in normal conditions. We also performed seven pollination treatments on ten randomly chosen individuals in each population, and the number of seeds following each treatment was recorded. Results from these treatments showed that R. ferrugineum produced spontaneous selfed seeds in the absence of pollinators. However, no seed was produced when short-level stamens were emasculated and pollinators excluded, suggesting that long-level stamens are not responsible for selfing in the absence of pollinators and that reproductive assurance is promoted by short-level stamens.     

MORPHOLOGICAL STUDIES OF THE NYMPHAEACEAE. XI. THE FLORAL BIOLOGY OF NELUMBO PENTAPETALA

The floral biology of Nelumbo pentapetala (Walter) Fernald, the American lotus, native to Texas, was investigated. Anthesis occurs over three consecutive days with flowers opening each morning and closing around noon. First-day flowers are protogynous with the perianth parts partially expanded so that pollen-covered insects which are attracted by floral color and the intense “fruity” odor (diffused with the aid of increased floral temperature) are directed on to the flattened receptacle (= carpellary receptacle) from which the receptive stigmas protrude, thus accomplishing pollination. During the second morning anther dehiscence begins and insects which visit and forage within the flower become covered with pollen and typically crawl over the still receptive stigmas achieving “facilitated” self-pollination (indirect autogamy). By mid-morning of the second day the stigmas dry and become non-receptive to pollen. During the third day of anthesis perianth and staminal parts quickly abscise and over the period of a few weeks the receptacle and enclosed fruits mature. In most populations studied, Hymenoptera (e.g., Lusioglossum spp., and Apis mellifera) were the most abundant and effective pollinators. In some populations, however, Coleoptera (e.g., Chauliognathus) were also numerous and effective pollinators. It is suggested that the overall floral structure (e.g., large numbers of stamens, masses of pollen, staminal appendages) are adaptations which facilitate the pollination of Nelumbo by beetles.     

Is There ‘Anther-Anther Interference’ within a Flower? Evidences from One-by-One Stamen Movement in an Insect-Pollinated Plant

The selective pressure imposed by maximizing male fitness (pollen dispersal) in shaping floral structures is increasingly recognized and emphasized in current plant sciences. To maximize male fitness, many flowers bear a group of stamens with temporally separated anther dehiscence that prolongs presentation of pollen grains. Such an advantage, however, may come with a cost resulting from interference of pollen removal by the dehisced anthers. This interference between dehisced and dehiscing anthers has received little attention and few experimental tests to date. Here, using one-by-one stamen movement in the generalist-pollinated Parnassia palustris, we test this hypothesis by manipulation experiments in two years. Under natural conditions, the five fertile stamens in P. palustris flowers elongate their filaments individually, and anthers dehisce successively one-by-one. More importantly, the anther-dehisced stamen bends out of the floral center by filament deflexion before the next stamen''s anther dehiscence. Experimental manipulations show that flowers with dehisced anther remaining at the floral center experience shorter (1/3–1/2 less) visit durations by pollen-collecting insects (mainly hoverflies and wasps) because these ‘hungry’ insects are discouraged by the scant and non-fresh pollen in the dehisced anther. Furthermore, the dehisced anther blocks the dehiscing anther''s access to floral visitors, resulting in a nearly one third decrease in their contact frequency. As a result, pollen removal of the dehiscing anther decreases dramatically. These results provide the first direct experimental evidence that anther-anther interference is possible in a flower, and that the selection to reduce such interferences can be a strong force in floral evolution. We also propose that some other floral traits, usually interpreted as pollen dispensing mechanisms, may function, at least partially, as mechanisms to promote pollen dispersal by reducing interferences between dehisced and dehiscing anthers.