花内雄蕊分化及其适应意义

对花内雄蕊存在显著分化的现象进行了分析与归纳, 总结了花内雄蕊分化的各种主要形式及其繁殖适应意义。“花内雄蕊分化”是指花内雄蕊与雄蕊之间存在显著分化的现象, 这一概念可以把二强雄蕊、四强雄蕊和异型雄蕊等以往单独进行研究的相关性状结合起来, 并明确区分了几种新的花内雄蕊分化形式, 以期更准确全面地认识这些相关性状的适应意义与进化。该文将花内雄蕊分化区别为花丝的分化、花药的分化、雄蕊合生的分化、雄蕊运动的分化、退化雄蕊5大类。花丝的分化主要是花丝长度的分化, 如四强雄蕊、二强雄蕊和单强雄蕊; 花药分化主要指花药颜色、花药与花粉粒大小和花药开裂时间等的分化; 雄蕊合生的分化主要体现为花内部分雄蕊合生而部分雄蕊离生; 雄蕊运动的分化指的是花内雄蕊在运动时间或方式上存在差异, 造成雄蕊处于不同的成熟阶段和位于不同的空间位置; 退化雄蕊则是花内部分雄蕊失去了生产花粉的繁殖功能, 通常也发生了花药形态上的巨大改变。异型雄蕊不仅存在花丝和花药的形态分化, 还存在着明显的功能分化, 是分化程度很高的一类特殊的花内雄蕊分化形式。一些特殊的繁育系统, 如异长花柱和镜像花柱等在种内不同个体上存在着不同形式的花内雄蕊分化。花内雄蕊分化在花内造成了多个不同的花药位置, 在很大程度上影响了雌雄异位程度, 对植物自交与异交水平、花内雌雄功能干扰等有着潜在作用; 花内雄蕊分化形成的多个不同空间位置的雄蕊还增加了对多种访花者的吸引与适应潜力, 有可能影响到访花者的类型与访花行为, 得以适应多种传粉者。此外, 花内雄蕊分化可将花粉逐渐分批次分发给访花者, 提高花粉散布效率, 可看成是“花粉呈现理论”所指的花粉装配与分发机制之一。现有的实验研究发现, 花内雄蕊分化能够吸引传粉者、保护正常花药和花粉、促进花粉散发(降低花粉竞争)、实行延迟自交和降低花内雌雄功能干扰等。花内雄蕊分化还缺少系统研究, 有些雄蕊分化形式如单强雄蕊和雄蕊运动的分化还没有针对性的实验揭示其适应意义, 鸭跖草科和某些豆科植物的雄蕊三型分化等现象也缺少进化适应意义的研究。花内雄蕊分化对植物雌性和雄性适合度可能不同的影响、如何与访花者相互作用、如何与其它花部特征一起影响植物繁殖过程等, 可能是这一领域值得今后优先研究的课题。     

木棉黄花个体的适应意义

木棉(Bombax ceiba)花大、红色, 多为鸟媒传粉, 但也存在频率极低的黄花个体, 有蜜蜂访花。本文对比研究了海南岛木棉红花、黄花的花部特征及传粉过程, 以解释黄花个体的适应意义。木棉两种花色表型的花蜜组分(葡萄糖和果糖)及挥发性物质(烷烃类、酯类、酚类等)没有显著差异, 但黄花花瓣的反射波长范围均在鸟类与蜜蜂视觉范围内, 可同时吸引鸟和蜜蜂。木棉黄花的雌雄异熟程度较小(花粉活性时期与柱头可受期重叠较大), 可在传粉媒介访花次数有限的情况下同时输出或接受花粉。另一方面, 黄花有着更大程度的柱头探出式雌雄异位, 降低了较小程度雌雄异熟可能导致的自交与雌雄功能干扰。木棉存在明显的花粉限制, 且黄花的坐果率(1.08 ± 0.56)%显著低于红花(3.27 ± 0.93)%。我们推测, 黄花个体可能通过增加木棉访花者类群多样性、降低雌雄异熟程度而提高了木棉的繁殖成功率, 也通过吸引蜜蜂而降低了蜜蜂对红花鸟媒传粉的干扰。     

宁夏枸杞的花粉呈现式样及交配系统研究北大核心CSCD

被子植物的花回馈、雌雄蕊时空分离特征和花粉呈现式样等花部特征及传粉者效率会影响雄性适合度和有性繁殖过程。宁夏枸杞(Lycium barbarum)是中国宁夏、新疆、内蒙等干旱和半干旱地区分布的特有种。该研究对新疆喀什地区宁夏枸杞自然种群的花部综合征、花粉呈现式样、花回馈、传粉者行为与交配方式的关系进行观察及统计分析,以探讨其花粉逐步呈现的适应性及其在提高雌雄繁殖过程中的意义。结果表明:(1)宁夏枸杞的单花寿命为(4.07±0.15) d,而雄性持续时间(0.07±0.01 d)比雌性持续时间短(4±0.01 d);花寿命内雌蕊长度比雄蕊长,属柱头探出式异位类型;花粉呈现式样为不完全逐步呈现。(2)花寿命不同阶段花回馈间存在显著差异。(3)意大利蜜蜂(Apis mellifera)、熊蜂(Bombus sp.)、食蚜蝇(Syrphidae sp.)是宁夏枸杞在自然居群的主要访花者,其中熊蜂和食蚜蝇是主要传粉者,但传粉者效率低,属于高移出低沉积类型。(4)宁夏枸杞的花粉胚珠比(2 448.11±448.32)及授粉实验结果均表现出兼性异交特征;自发自交及人工自花授粉花的座果率及结籽率很低,属于自交不亲和类型;自然传粉花的座果率及结籽率比人工去雄异花低,存在其较高的花粉限制(40.71%)。研究发现,宁夏枸杞花部综合征表现出雌雄异位和花粉不完全逐步呈现式样,这是避免雌雄功能及雄蕊各花药间的干扰、减少花粉同步移出及保障其雄性适合度的有效途径,但自然居群的传粉者种类限制、传粉效率低以及自交不亲性是导致宁夏枸杞花粉限制及降低雌性繁殖成功的主要原因。     

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”     

黑果枸杞两种花型的花部综合征与传粉特性

黑果枸杞(Lycium ruthenicum)是中国西北地区极端环境中分布的国家二级保护植物, 该物种在新疆南部的自然种群中出现了同型花柱类型(同位花)和柱头探出式雌雄异位类型(异位花)个体, 并且遭遇沙尘暴频繁的种群中异位花个体出现频率减少。该研究对喀什市自然种群中黑果枸杞两种不同花型植株的花部综合征和传粉特性进行比较研究, 以期探讨该物种不同花型植株在南疆早春极端环境中的花部特征的可塑性及其适应性机制。结果表明: 同位花雌雄蕊高度间无显著差异, 而异位花雌蕊高度显著高于雄蕊; 同位花花冠直径、花冠筒长、胚珠数均高于异位花, 而异位花雌雄蕊空间距离、花粉数及花粉胚珠比均比同位花高。黑果枸杞同位花个体比例(68%)高于异位花个体(32%), 种群水平及个体水平同位花花期((117.00 ± 2.25) d, (101.65 ± 1.98) d)比异位花((26.00 ± 1.00) d, (18.75 ± 1.00) d)长, 而单花水平上异位花单花寿命((4.50 ± 0.14) d)比同位花((3.13 ± 0.11) d)长。两种类型花在花早期(紫色)分泌的花蜜量均高于花后期(白色)。在紫色花阶段(花开放早期), 同位花上的主要传粉者意大利蜜蜂、熊蜂和食蚜蝇的访花频率和停留时间均高于异位花; 而白色花阶段(花开放后期)意大利蜜蜂、熊蜂在异位花上的访花频率比同位花高。在不同花色阶段, 同位花柱头花粉落置数、花粉移出率、花粉传递效率均比异位花高, 并且同位花自然坐果率及结籽率均比异位花高。在新疆南部的沙尘暴极端环境下, 同位花通过较高的自交亲和性保障繁殖, 而异交为主的异位花提高了异交率。异位花与同位花在花部综合征和花报酬上的差异, 是影响其繁殖成功的主要因素。     

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.     

高山植物扁蕾的延迟自交机制

扁蕾(Gentianopsis barbata)具有鲜艳的花和显著的腺体,并且花开放的前5 d柱头和花药始终处于不同的位置(雌雄异位),这些花综合征表明该植物应为异花传粉。为检验这一假设,我们对青藏高原植物扁蕾的海北站种群进行了3年的传粉生物学研究实验。与花综合征所表明的繁育系统相反,两年的野外观察发现昆虫的访花频率十分低,不去雄并隔离昆虫处理也能产生大量种子,说明这一种群的繁殖主要是依赖于自花传粉。尽管利用种子结实评价的柱头可授性从花开放4 d后开始下降,但随着花的发育进程,雄蕊的伸长能使得花药与柱头完全接触。实验也证明,柱头可授性和花粉活力都超过5 d,说明花药和柱头的接触能够发生自花授粉。扁蕾的这种自花传粉机制应属于典型的延迟自交类型。自花授粉发生在单花花期快要结束前,自交之前仍然保持异交传粉机制,这种延迟自交避免了自交与异交竞争造成的花粉或者种子折损,并为扁蕾在青藏高原极端环境下由于访花昆虫缺乏造成的异交失败提供了繁殖保障。     

雌雄异位和花部行为适应意义的研究进展

雌雄异位和花部行为影响植物的传粉和交配格局。二者的适应意义一直是植物生殖生态学和进化生物学研究的热点之一。该文概述了近年来雌雄异位和花部行为适应意义的研究进展,包括对雌雄异位适应意义的最早期、最传统的认识——避免自交和当前的主流观点——避免雌雄干扰;花内性别干扰的形式及其例证;花部行为在避免雌雄功能干扰、避免自交、实现延迟自交、应对不良环境等方面的适应意义。讨论了使雌雄异位程度减小的锦葵科一些植物中的单体雄蕊柱头运动可能涉及到的适应意义。针对目前在雌雄异位和花部行为适应意义研究上存在的不足,结合国际上研究二者适应意义的发展趋势正在从前期以描述性的研究为主转变到应用现代分子生物学手段(尤其是分子标记,如SSR、SNP、AFLP等)和传统方法相结合,以及随着田间实验和控制条件能力的发展正由经典的野外调查、观察为主过渡到包括花操纵在内的人工试验和野外试验相结合,提出了今后研究中应注意的问题。有必要借用多学科的手段,以可作为研究雌雄异位和花部行为的有着研究基础的植物为材料,设计和操纵不同的对比试验,系统而深入地探讨雌雄异位和花部行为的适应意义。     

早春短命植物黑鳞顶冰花的花粉呈现时序及其适应性

花粉的时序呈现是指植物花药按一定次序释放花粉的现象, 被认为是对传粉者访问频率的一种适应。在传粉者充足的环境中, 植物通过限制花粉1次被移出的数量, 使花粉供体能作为多个父本, 从而提高雄性适合度。该文从开花习性、花部特征、传粉者及繁育系统等方面对早春短命植物黑鳞顶冰花(Gagea nigra)的花粉呈现时序及其适应性进行研究, 结果表明, 黑鳞顶冰花单花期约5-7天; 白天开放, 晚上闭合, 花药次序开裂, 呈拉链式散粉, 散粉期4-6天。黑鳞顶冰花以异交为主, 部分自交亲和。蝇类和食蚜蝇为主要传粉者, 访花频率为(0.141±0.078) flower∙h^-1。在雄蕊的时序散粉过程中, 雌蕊持续生长, 经历了从低于雄蕊到等高、再到高于雄蕊阶段。在等高阶段, 单花早晚的开闭, 使得雌雄蕊紧靠在一起, 促成了自动自花授粉。在传粉者缺乏的环境中, 黑鳞顶冰花的花粉时序呈现延长了散粉期, 在等待传粉者和分摊风险方面具重要作用。这种花粉渐次呈现的策略, 在新疆的早春开花植物中可能广泛存在。     

高山植物全缘叶绿绒蒿在不同海拔地区的传粉生态学研究北大核心CSCD

为了探明高山植物全缘叶绿绒蒿(Meconopsis integrifolia)的繁育系统特点和其对高山气候环境的繁殖适应特征,我们沿海拔梯度选择了5个样地(样地1(4 452 m)、样地2(4 215 m)、样地3(4 081 m)、样地4(3 841 m)、样地5(3 681 m))对其传粉生态学进行了连续2年的观察试验。结果发现,样地1、2的全缘叶绿绒蒿的单花寿命显著长于样地3、4和5。花开放早期柱头高于花药,之后花药不断伸长,并在开放中后期与柱头接触,说明全缘叶绿绒蒿具有不完全雌雄异位的花部特征。自然状态下,柱头可授能力持续期约8天(雌蕊先熟),但花药于开花第5天才散粉,花粉寿命约2天,说明全缘叶绿绒蒿为雌雄异熟,但存在一定的重叠期。人工授粉试验表明,全缘叶绿绒蒿自交部分亲和,且不存在无融合生殖现象。各样地中自然对照的结实率显著低于人工异交处理的结实率,说明自然状态下全缘叶绿绒蒿存在一定程度的传粉限制。传粉昆虫观察发现,样地1和2的传粉昆虫主要是蝇类,样地3、4和5的传粉昆虫主要是蝇类和蓟马(Thripidae spp.),蝇类在不同植株间的活动能够保证异花传粉结实,同时,蝇类和蓟马在花内的活动会引起"协助自交"。全缘叶绿绒蒿约65%的观察个体存在"自动自交"。蝇类在各样地的访花频率存在显著差异,样地1访花频率最低,样地2访花频率最高。各样地的结实由于异花传粉者的不足而受到传粉限制。两种不同类型的自交机制恰恰为该植物异花传粉者不足提供了一定程度的繁殖补偿。全缘叶绿绒蒿不分泌花蜜,当环境温度降低时,采取为昆虫提供保温庇护场所的方式来吸引传粉者。     

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.     

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.     

In the right place at the right time: Parnassia resolves the herkogamy dilemma by accurate repositioning of stamens and stigmas

Background and Aims

Spatial (herkogamy) and temporal (dichogamy) separation of pollen presentation and stigma receptivity have been interpreted as reducing interference between male and female functions in hermaphroditic flowers. However, spatial separation leads to a potential conflict: reduced pollination accuracy, where pollen may be placed in a location on the pollinator different from the point of stigma contact.

Methods

To understand better how herkogamous flowers resolve this conflict, a study was made of a subalpine herb, Parnassia epunctulata, the nectariferous flowers of which exhibit sequential anther dehiscence (staggered pollen presentation) and stamen movements; usually one newly dehisced anther is positioned each day over the central gynoecium, while the older stamens bend away from the central position.

Key Results

The open flowers were visited by a variety of pollinators, most of which were flies. Seed set was pollinator-dependent (bagged flowers set almost no seeds) and pollen-limited (manual pollination increased seed set over open pollination). Analyses of adaptive accuracy showed that coordinated stamen movements and style elongation (movement herkogamy) dramatically increased pollination accuracy. Specifically, dehiscing anthers and receptive stigmas were positioned accurately in the vertical and horizontal planes in relation to the opposite sexual structure and pollinator position. By contrast, the spatial correspondence between anthers and stigma was dramatically lower before the anthers dehisced and after stamens bent outwards, as well as before and after the period of stigmatic receptivity.

Conclusions

It is shown for the first time that a combination of movement herkogamy and dichogamy can maintain high pollination accuracy in flowers with generalized pollination. Staggered pollen and stigma presentation with spatial correspondence can both reduce sexual interference and improve pollination accuracy.     

Potential Autonomous Selfing in Gesneria citrina (Gesneriaceae), a Specialized Hummingbird Pollinated Species with Variable Expression of Herkogamy

Species with mixed mating systems often demonstrate vadable expression of breeding system characteristics and thus represent the opportunity to understand the factors and mechanisms that promote both outcrossed and selfed seed production. Here, we investigate variation In levels of herkogamy (variation in stigma-anther separation distance) In a Puerto Rican population of hummingbird pollinated Gesneria citrina Urban. There is significant variation in herkogamy levels among individuals of this species and stigma-anther separation is negatively associated with the ability to set fruits and seeds in the absence of pollinators. The variation in levels of herkogamy may represent a mechanism to ensure the production of some self-fertilized progeny in the absence of hummingbird pollinators. We also describe a novel breeding system in G. citrina, where stamens elongate over time to reach stigma height, but stamen elongation is accelerated by pollination. These results suggest that once the flowers are pollinated, stamen elongation may favor increased pollen removal and siring success, while the reduction in stigma-anther distance no longer imposes the risk of interference between male and female functions. We discuss our findings of breeding system variation in the context of pollination system evolution in an island setting (Antillean islands).     

Potential Autonomous Selfing in Gesneria citrina (Gesneriaceae), a Specialized Hummingbird Pollinated Species with Variable Expression of Herkogamy

Species with mixed mating systems often demonstrate variable expression of breeding system characteristics and thus represent the opportunity to understand the factors and mechanisms that promote both outcrossed and selfed seed production. Here, we investigate variation in levels of herkogamy (variation in stigma-anther separation distance) in a Puerto Rican population of hummingbird pollinated Gesneria citrina Urban. There is significant variation in herkogamy levels among individuals of this species and stigma-anther separation is negatively associated with the ability to set fruits and seeds in the absence of pollinators. The variation in levels of herkogamy may represent a mechanism to ensure the production of some self-fertilized progeny in the absence of hummingbird pollinators. We also describe a novel breeding system in G. citrina , where stamens elongate over time to reach stigma height, but stamen elongation is accelerated by pollination. These results suggest that once the flowers are pollinated, stamen elongation may favor increased pollen removal and siring success, while the reduction in stigma-anther distance no longer imposes the risk of interference between male and female functions. We discuss our findings of breeding system variation in the context of pollination system evolution in an island setting (Antillean islands).     

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.     

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.     

独叶草的不育雄蕊、花被片和可育雄蕊对传粉昆虫和传粉的影响

为确定独叶草（Kingdonia uniflora）不育雄蕊、花被片和可育雄蕊对传粉昆虫和传粉的影响,分别进行去除不育雄蕊（A组）,去除花被片（B组）,去除可育雄蕊（C组）和对照组（D组）处理。结果表明：昆虫访花频率分别为A组0．4次／h、B组0次／h、C组0．9次／h、D组2．2次／h;被授粉花朵百分比分别为A组29．09％、B组40．38％、C组70.91％、D组91．67％;被授粉心皮百分比分别为A组17．77％、B组20．94％、C组为40．58％、D组为75．27％。与对照组相比,处理后被授粉花朵百分比、被授粉心皮百分比下降幅度均为A组〉B组〉C组。结合昆虫访花频率与不育雄蕊蜜汁的分泌量呈正比,可确定不育雄蕊产生的蜜汁是吸引昆虫访花的最主要因素。花被片为昆虫取食提供停歇的平台,可育雄蕊与花被片产生的色差及可育雄蕊提供的花粉,是吸引昆虫的辅助因素。     

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。