两种姜科花柱卷曲性植物柱头的位置与其可授性的关系

花柱卷曲性是一种见于姜科山姜属(Alpinia)和砂仁属(Amomum)植物中的独特的性二态现象,具有这一性系统的植物,其居群包括上举型和下垂型两种表型,所有个体的雌雄性别功能在时间和空间上分离。本研究通过操控授粉和花粉管生长两项实验,探讨花柱卷曲性植物个体两性功能的隔离方式及其适应意义。云南草蔻(Alpinia blepharocalyx)操控授粉实验表明,自花花粉对其异交率影响不显著(P>0.05),但可能由于自花花粉沉降导致胚珠贴现进而引起近交衰退,操控条件下每果结籽数显著减少(P<0.01)。云南草蔻和红豆蔻(Alpinia galanga)花粉管生长实验都显示无论对两种表型进行怎样的授粉处理,花粉粒萌发和花粉管生长的速率仅在柱头处于可授位置时表现出最大值,与自交和异交授粉方式无关;上举型植株上午花粉囊虽未裂但其内花粉已成熟。研究结果表明即使不考虑花柱运动,山姜属植物也具有异型雌雄异熟的特性。这一结果证实了花柱卷曲运动机制是通过互补式雌雄异位和异型雌雄异熟相结合形成的花部二态性,异型雌雄异熟促进了异交,而花柱运动的功能可能在于避免雌雄干扰。     

山姜属植物花柱卷曲性传粉机制的研究

暂性雌雄异株是一种在两性花或雌雄同株植物个体上表现出的雄蕊和雌蕊成熟时间不同而形成的暂时性雄性阶段和雌性阶段不重叠的现象。这种现象减少了自交亲和的雌雄同株和两性花种类发生自交的可能性。作系统研究了花柱郑曲性传粉机制在西双版纳姜科山姜属（Alpinia Roxb.）植物中的普遍性,这一机制使得植物通过完全的暂性异株达到了异交的目的。这类植物的种群中具有两种独特的表现型,其差异央开花行为的不同：柱头下垂型个体在刚开花时其柱头向上反卷。位于已开裂的花药上方,到中午开始向下运动,下午则位于花药下方;柱头上举型个体的柱头在开花初期向花冠内卷曲,位于未开裂的花药和唇瓣之间中午开始向上运动,当柱头上举到花药上方后,花药开裂,花偻开始散发。两种个体的开花行为是同步的,通过两种柱头反向运动授粉方式（不同时间的自花和异花授粉）具有相同的结实率,但套代的花序几科不结实,两种表型在自然种群中的比例为1：1。     

青藏高原特有植物祁连獐牙菜（龙胆科）的花综合征与虫媒传粉

雌雄异熟和异型花在一些被子植物中的共同存在被认为是一种有效避免两性花植物自交并促进异交的机制。本文报道青藏高原特有植物祁连獐牙菜Swertia przewalskii Pissjauk.的花同时具有雌雄异熟和异型花这两种特征。在此基础上,我们详细观察了该植物访花昆虫的访花行为,主要探讨这种促进异花传粉的机制是否能完全避免株内异花传粉的自交。繁育系统试验证明祁连獐牙菜单花套袋不结实,说明雌雄异熟和异型花的花特征完全避免了单花的自花传粉,自然结实必须依赖昆虫传粉才能完成。然而,该植物却是高度自交亲和的,因为人     

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。     

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。     

草豆蔻传粉生物学的研究

比较研究了花柱卷曲性植物草豆蔻 Alpinia hainanensis两种表型(上举型和下垂型)的传粉生物学特性。结果表明:草豆蔻的花柱卷曲运动节律与其他已报道的山姜属Alpinia植物基本一致,而下垂型花的花柱卷曲运动明显滞后于上举型花约2 h。草豆蔻的花柱卷曲运动中存在一些不同步的现象,甚至在同一花序上的两朵花的花柱运动节律亦会表现出不一致的现象,但上举型花的花药开裂时间却严格同步,都发生在同类型个体的柱头全部位于花药上方之后进行。草豆蔻上举型花的花蜜分泌量、单花花粉量、花粉/胚珠比率(P/O)均明显比下垂型花多,而两种表型的胚珠数、花蜜糖含量、氨基酸含量无显著差异。在草豆蔻的单花期内不同时间段进行人工授粉,上举型花在其柱头位于花药下方时进行人工授粉,其结实率明显比柱头位于花药上方时人工授粉的处理高,下垂型花则没有显著差异。在自然居群中,草豆蔻的主要传粉者是无垫蜂Ameglla sp.和两种木蜂Xylocopa sp.,但存在传粉者不足而影响结实率的现象。完全套袋、去雄和去雌(去柱头)处理的均不结实,表明草豆蔻中不存在无融合生殖、主动自花授粉和滞后自交的生殖保障现象。而人工自交和异交均具有较高的结实率,表明草豆蔻为自交亲和植物。草豆蔻的繁育系统是具花柱卷曲性运动的异花授粉的交配系统。     

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”     

益智传粉生物学的研究

 益智（Alpinia oxyphylla）的花期从2月底至4月下旬;单株花期（花序）约为16～26 d,通常为23～26 d;单花花期一般为1 d。正常情况下,益智和其它山姜属植物一样,具有花柱卷曲性促进异花授粉的机制,表现出两种类型：花柱上举型和花柱下垂型,其花柱卷曲运动的节律与其它已报道的山姜属植物基本一致。但观察发现,当遭遇低温天气时(日最高气温<18℃),单花花期延长为2 d,无论是上举型个体还是下垂型个体,均只表现出一种花型——下垂型：上午开花时花柱弯向上,柱头位于已开裂散发出花粉的花药上方,直到第二天上午6∶30～11∶00间,花柱才陆续地慢慢向下运动,柱头下降至与花药等高或位于花药下方。益智的主要传粉昆虫是蜜蜂（Apidae sp.）、木蜂（Xylocopa sp.）,绝大多数的访花者的访花目的是吸蜜。益智的花蜜分泌量（8.37～15.79 μl）和花蜜含糖量（30.12%～32.83%）较高,花蜜是益智作为传粉者访花的最主要的报酬。实验结果还表明,益智花部中唇瓣对昆虫访花有显著的招引作用;益智的花蜜对蜜蜂的访花频率有显著的影响,对木蜂有一定的招引作用,但并不显著。而花粉（花药）则对昆虫的访花频率影响不大。人工授粉实验结果表明益智存在自交亲和性,无论是上举型或下垂型个体自交和异交均有较高的结实率;人工自交和异交的结实率在上举型植株中存在较大的差异,而在下垂型个体中则差异不明显;去雄套袋、去柱头套袋和完全套袋不授粉等处理均不结实,表明益智不存在无融合生殖现象和自动自花授粉现象。益智的繁育系统是异花授粉交配系统。     

雌先熟植物茄参 （茄科）的繁育系统

对分布于云南西北部的茄参（Mandragora caulescens）的繁育系统进行了研究。茄参的花期为5月底到6月初,单花持续时间为（9.9±2.8）d,其性表达方式为雌性先熟。有三种昆虫拜访茄参的花,白背熊蜂蜂王是最有效的传粉昆虫,但其访花频率很低;苍蝇和蚂蚁的访花频率较高,但苍蝇传粉效率十分有限;蚂蚁是窃蜜者。授粉实验证明茄参具有部分自交亲和能力,但由于雌性先熟,不具备主动自交能力。人工添加异交花粉显著提高了结实率和结籽数,证明茄参存在花粉限制。茄科的大部分植物是严格自交不亲和的,但茄参族的茄参是自交部分亲和,这和茄科天仙子族的部分成员类似。我们认为这两个独立起源于新世界向欧亚大陆扩散事件的族,各自从自交不亲和向自交亲和转变可能与高山地区恶劣的自然条件有关,这种繁育系统的进化模式是否具有普遍性值得进一步研究。     

雌先熟植物茄参（茄科）的繁育系统

对分布于云南西北部的茄参（Mandragoraca caulescens）的繁育系统进行了研究。茄参的花期为5月底到6月初,单花持续时间为（9．9±2．8）d,其性表达方式为雌性先熟。有三种昆虫拜访茄参的花,白背熊蜂蜂王是最有效的传粉昆虫,但其访花频率很低;苍蝇和蚂蚁的访花频率较高,但苍蝇传粉效率十分有限;蚂蚁是窃蜜者。授粉实验证明茄参具有部分自交亲和能力,但由于雌性先熟,不具备主动自交能力。人工添加异交花粉显著提高了结实率和结籽数,证明茄参存在花粉限制。茄科的大部分植物是严格自交不亲和的,但茄参族的茄参是自交部分亲和,这和茄科天仙子族的部分成员类似。我们认为这两个独立起源于新世界向欧亚大陆扩散事件的族,各自从自交不亲和向自交亲和转变可能与高山地区恶劣的自然条件有关,这种繁育系统的进化模式是否具有普遍性值得进一步研究。     

Adaptive significance of flexistyly in Alpinia blepharocalyx (Zingiberaceae): a hand-pollination experiment

BACKGROUND AND AIMS: Flexistyly is a sexual dimorphism where there are two morphs that differ in the temporal expression of sexual function and also involve reciprocal movement of the stigmatic surface through a vertical axis during the flowering period. The adaptive significance of flexistyly has been interpreted as a floral mechanism for outcrossing, but it may also function to reduce sexual interference in which styles and stigmas impede the pollen export. Here these two explanations of flexistyly were tested in Alpinia blepharocalyx through a hand-pollination experiment. METHODS: Hand-pollinations were performed in two temporal morphs and consisted of two sequential pollination treatments, namely self-pollination in the morning and inter-morph pollination in the afternoon (treatment 1) or conversely inter-morph pollination in the morning and self-pollination in the afternoon (treatment 2), and two simultaneous self- and inter-morph cross-pollination treatments either in the morning (treatment 3) or in the afternoon (treatment 4). Seed paternity was then determined to assess relative success of self- versus cross-pollen using allozyme markers. KEY RESULTS: In the sequential pollination treatments, whether the stigmas of recipients are receptive in the morning is crucial to the success of the pollen deposited. When the cataflexistylous (protandrous) morph served as pollen recipient, early-arriving pollen in the morning can sire only a very small proportion (<15%) of seeds because the stigmas were then unreceptive. However, when the anaflexistylous (protogynous) morph served as pollen recipient, early pollen did gain a large competitive advantage over the late pollen, particularly when cross-pollen arrived first. Simultaneous self- and inter-morph cross-pollination indicated that outcross-pollen is more competitive than self-pollen on receptive stigmas. CONCLUSIONS: Differential maturing of male and female organs in Alpinia blepharocalyx is sufficient for selfing avoidance, obviating the need for style movements. Instead, the upward style curvature of the cataflexistylous morph in the morning and the anaflexistylous morph in the afternoon most likely represents a means of reducing interference with pollen export.     

Mating system and stigmatic behaviour during flowering of Alpinia kwangsiensis (Zingiberaceae)

 Little is known about the breeding systems of tropical perennial ginger species. In this paper, we provide information about the breeding system of Alpinia kwangsiensis in Yunnan, Southwest China, specifically self-compatibility, mechanisms promoting outcrossing, and the visitation characteristics of pollinators. Populations of A. kwangsiensis have two specific phenotypes that differ in flowering behaviour: 1) “cataflexistyle” individuals in which the stigma is held erect above the dehiscent anther when anthesis begins in the morning and becomes decurved under the anther at noon and 2) “anaflexistyle” individuals in which the receptive stigma is decurved under the indehiscent anther first and moves into a reflexed superior position above the anther as it begins to shed pollen at mid-day. The stigmatic movements in the two floral phenotypes, which occur in a ratio of 1:1 in natural populations, are synchronous and correlate with the foraging behaviour of floral visitors; pollination is effected only between floral forms. Field experiments indicate that A. kwangsiensis is self-compatible and dependent upon insects for fertilization. This newly reported floral mechanism, which we have named “flexistyly,” adds to the repertoire of devices that have evolved in flowering plants to insure outcrossing. Received November 29, 2001 Accepted January 8, 2002     

The concrete evidence of flexistyly in Plagiostachys: pollination biology of a wild ginger on Hainan Island,China

Flexistyly in Plagiostachys was first reported by Takano et al., while they provided no detailed study on pollination biology and breeding system. In this study, we tested this suspicion in one species of Plagiostachys (Plagiostachys austrosinensis). Field observations suggested that flexistyly was present in this species, and stigmatic behavior was similar to that reported for Alpinia and Amomum species. Two phenotypes (anaflexistylous and cataflexistylous) occurred in a ratio of 1:1 in natural populations. Anthesis began around 1530–1600 h and lasted for about 24 h. Pollen viability and stigma receptivity remained high throughout the flowering process. Mean nectar volume (4.15–11.30 μL) and mean sugar concentration (>32%) also remained at a high level during the flowering process. No fruit set occurred in unpollinated bagged plants. Two pollinators (Bombus pyrosoma and Vespidae spp.) and one pollen robber (Mutillidae spp.) were found as flower visitors. Fruit set following self‐pollination and cross‐pollination did not differ significantly in the cataflexistylous morph. Partial self‐incompatibility was apparent in the anaflexistylous morph. These results provide the concrete evidence of flexistyly in Plagiostachys and a more thorough understanding of its evolutionary origin in gingers.     

Diplostigmaty in plants: a novel mechanism that provides reproductive assurance

Differentiation of female sexual organs in flowering plants is rare and contrasts with the wide range of male reproductive strategies. An unusual example involves diplostigmaty, the possession of spatially and temporally distinct stigmas in Sebaea (Gentianaceae). Here, the single pistil within a flower has an apical stigma, as occurs in most flowering plants, but also a secondary stigma that occurs midway down the style, which is physically discrete and receptive several days after the apical stigma. We examined the function of diplostigmaty in Sebaea aurea, an insect-pollinated species of the Western Cape of South Africa. Floral manipulations and measurements of fertility and mating patterns provided evidence that basal stigmas function to enable autonomous delayed self-pollination, without limiting opportunities for outcrossing and thus avoiding the costs of seed discounting. We suggest that delayed selfing serves as a mechanism of reproductive assurance in populations with low plant density. The possession of dimorphic stigma function provides a novel example of a flexible mixed-mating strategy in plants that is responsive to changing demographic conditions.     

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。     

Transgenic GFP as a molecular marker for approaches to quantify pollination mechanism and gene flow in Arabidopsis thaliana

Arabidopsis thaliana is commonly regarded as a self-pollinated plant. We observed that the stigma in each flower of A. thaliana cannot be pollinated by its own pollen in the early phases of the flowering process, when the anthers had dehisced but the filaments were still too short for the pollen to be deposited on the stigma. In the later stages, after elongation of the filaments, self-pollination can occur. After artificial pollination of the flower of a wild plant with GFP transgenic pollen grains in earlier stages of flowering, GFP expressed within epidermal cells was detected in some of the offspring (26.1-57.1 %). Wind-mediated pollen dispersal was poor but is likely to exist in natural habitats, while insects were observed visiting flowers of A. thaliana in natural and experimental populations. We constructed an experimental population consisting of 28 GFP transgenic plants and 240 wild plants and examined gene flow in the population. The result was that the distance of gene flow was limited to 0.5 m. 22 offspring with expressed GFP were found in 28,299 filial individuals examined, which suggested a relatively low outcrossing rate (0.74%). We conclude that outcrossing in populations of A. thaliana is mainly due to insect pollination. The data on gene flow could be useful to assess the ecological hazards of experimental transgene combinations.     

濒危植物凹叶厚朴的花部综合特征和繁育系统

为了解凹叶厚朴(Magnolia officinalis subsp.biloba)濒危的生殖生物学原因,通过野外观测,运用杂交指数、花粉-胚珠比、花粉活力和柱头可授性观测、人工授粉和套袋实验等方法,对凹叶厚朴种群的开花生物学、繁育系统进行了研究。结果表明:凹叶厚朴单花花期一般为4～5d,单花花期根据形态和散粉时间可分为4个时期:蕾期、初开期、盛开期和凋零期;种群于4—5月开花,一般开花历时30d左右,但不同地点、年份间有所差异;种群无开花高峰,单株平均开花数量为171.4～188.8朵,平均开花振幅为5～6朵·株-1·d-1;凹叶厚朴杂交指数为4,花粉胚珠比为5728.86,结合人工套袋和授粉实验结果可确定,繁育系统为异交,传粉过程需要传粉者;花粉活力较高(94%),且花粉活力可维持至散粉后的第6天。散粉后,柱头可授性显著下降,花粉高活力期与柱头可授期有5～6h的重叠;自然状况下结果率和单果出种百分率低,异株异花授粉可明显提高结果率和单果出种百分率,二者相对自然状况分别提高14.25%和43.66%。表明自然状态下传粉效率低、柱头最佳可授期短、同株自花授粉败育严重是其濒危的主要生殖生物学原因。     

珍稀濒危植物细果秤锤树开花生物学特性和繁育系统

为了解珍稀濒危植物细果秤锤树（Sinojackia microcarpa）开花特征和有性繁殖,对其花部形态特征、开花动态、花粉胚珠比（P/O）、柱头可授性、花粉活力、套袋试验、访花昆虫及访花频率进行观测。结果表明：（1）细果秤锤树的杂交指数（OCI）为4,单花期5~7 d,种群花期可持续20 d左右。花粉与胚珠比为4 093.21±498.56。开花后第3天的花粉活力最高（76.21%）,而开花后第7天时花粉活力较低（18.37%）。细果秤锤树柱头最适可授期在开花后第2天。（2）套袋试验表明,细果秤锤树存在部分自交不亲和性,同时不存在无融合生殖,传粉昆虫是其完成生殖过程所必需的,且异株授粉能够提高其坐果率和结籽率。细果秤锤树的访花昆虫有3目5科7种,主要访花昆虫有黄胸木蜂（Xylocopa appendiculata）、熊蜂（Bombus sp.）、胡蜂（Vespa sp.）、中华蜜蜂（Apis cerana）、黑带食蚜蝇（Episyrphus balteatus）、其中熊蜂平均访花频率最高,达（8.67±0.21） 次·h^-1。对该物种开花生物学特征与繁育系统进行深入研究,有利于进一步探究其濒危机制,为后续珍稀濒危植物的研究提供理论依据和参考。