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。     

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。     

荒漠孑遗植物裸果木的花部特征及繁育系统

裸果木是亚洲中部荒漠区少有的第三纪孑遗物种,也是构成石质荒漠植被群落的重要建群种之一,由于人为干扰,其自然种群处于不断衰退中。本文通过野外观察和人工授粉实验等方法,对裸果木的花部特征及繁育系统进行研究,对于揭示该物种生活史特征并探讨影响其生殖成功的因素及制定相应的保护和管理策略具有重要意义。结果表明:裸果木为两性花,花小,无花瓣,雄蕊10枚,外轮5枚雄蕊败育,胚珠1枚,开花盛期有少量花蜜并散发浓烈难闻气味,花粉活力和柱头可授性之间存在46 h的重叠期,且不完全雌性先熟,柱头和花药在多数花的单花花期结束时并未接触,因此,裸果木花主要表现为适应异花传粉的雌雄异位特征;花粉/胚珠比(P/O)为1371.67±236.21,异交系数(OCI)为3;不同花粉来源(自然授粉、自花授粉、同株异花和异株异花授粉)的花粉在柱头上均能萌发,但花粉管生长速度存在差异,同株异花授粉和异株异花授粉的花粉管生长速度较快,自花授粉的花粉管生长速度最慢且部分花粉管(43%)在到达子房时停止生长,并未到达胚珠;人工套袋实验的结果表明,裸果木不存在无融合生殖,自然结种子数低,自然授粉花的种子数远远低于人工异株异花授粉花,说明存在由于异花传粉者不足造成的传粉限制。裸果木部分雌性先熟和雌雄异位是两性花为避免雌雄功能重叠,促进异交而采取的一种花部机制,其繁育系统为兼性异交类型且需要传粉者,部分自交不亲和和传粉限制是影响该物种有性生殖成功的主要因素。     

类叶升麻(毛茛科)自花传粉机制

毛茛科类叶升麻(Actaea asiatica)有2种开花方式, 一种与同属其它植物相同, 花萼片在开花早期脱落, 花药远离柱头; 而另一种较为特殊, 能使该植物进行自花授粉, 即在花开放前, 花萼从花托上脱离, 但萼片并不张开, 部分雄蕊的花丝伸长, 将花药推入萼片和柱头之间, 收缩的花萼片将开裂的花药压在宽大的柱头上, 进而完成自花授粉。套袋实验结果表明, 类叶升麻自发自交具有很高的结籽率, 不具有孤雌生殖和风媒传粉。繁育系统估测分析结果显示, 类叶升麻自交亲和,为兼性自交的繁育系统。     

山羊臭虎耳草的延迟自动自花授粉机制

通过对山羊臭虎耳草6个花形态及其发育阶段、授粉过程的连续观察和对花粉活性、柱头可授性的连续检测,报道了一种新的延迟自动自花授粉机制,即山羊臭虎耳草花丝运动引导的延迟自动自花授粉。利用电子显微镜研究了花粉囊的破裂方式和柱头结构,同时利用苏丹Ⅲ研究了花粉的组织化学性质。结果显示:(1)第四生长阶段,当花丝引导大部分花粉囊运动至柱头上方时,花粉和柱头的可授性才均达到峰值[可授性分别为(94±6)%和(96±13)%],这种运动消除了第三生长阶段花粉活性较高[(90±8)%]而柱头可授性较低[(6±17)%]的异花授粉特征,即消除了花粉与柱头的时空隔离;自动自花授粉发生于异花授粉特征消除之后,因此山羊臭虎耳草具有典型的延迟自动自交行为。(2)第四发育阶段柱头空腔的大量柱状突起和花粉鞘的油脂性特征,增加了花粉和柱头的接触表面,并且使花粉更好地互相堆积粘附在柱头上,提高授粉效率。(3)对山羊臭虎耳草的繁育系统研究表明,强制自花授粉和人工自花授粉产种量无明显差异,2007年和2008年的自交系数高达0.92和0.91,说明山羊臭虎耳草具有很好的自交能力并且不存在花粉数量限制;人工异花授粉的产种率显著高于人工自花授粉,说明自花花粉的质量显著低于异花花粉,虎耳草延迟自动自花授粉存在生殖衰退,生殖保障系数2007年为0.92,2008年为0.91,表明在极端环境下自动自花授粉为该植物提供了繁殖保障,但极端环境下,没有观察到有效的授粉昆虫。研究表明,作为高山冰缘植物的山羊臭虎耳草,保留了开花植物异花授粉的特征,在极端环境下利用延迟自动自花授粉机制完成授粉,保障生殖。     

蝶形花植物猪屎豆自花传粉机制研究

猪屎豆( Crotalaria pallida)为典型的蝶形花植物,分布极广,是路边或遭破坏生境中最常见的先锋种之一,野外观察未发现其有营养生殖的现象,主要为种子繁殖。该研究通过对自然生境中猪屎豆开花物候、访花昆虫及繁育系统的研究,旨在从繁殖的角度阐述其快速扩张的能力。结果表明：尽管猪屎豆的主要访花昆虫是蜜蜂,但蜜蜂的访花频率极低[(1.73±1.30)次/花序·h-1],且在整个花期内花药和柱头均被龙骨瓣包裹,蜜蜂访花时未成功接触柱头和花粉,不能实现传粉,因此蜜蜂不是猪屎豆有效的传粉昆虫,这与假说“蝶形花普遍是对膜翅目昆虫,尤其是对蜜蜂传粉的适应”不一致。人工授粉结果显示,猪屎豆为自交亲和种,不存在无融合生殖现象,其繁殖主要通过主动自交生产种子来实现,且在自交过程中长短花药都参与主动自交。这种自交方式不同于其他蝶形花植物的主动自交仅由短花药实现。对猪屎豆而言,长短花药均参与自交能够增加柱头的授粉几率,保证其在不利的生境中成功结籽,是其成功扩张的关键因素之一。     

黄花蔺的繁育系统研究

黄花蔺Limnocharis flava(Linn．)Buch．的蝎尾状聚伞花序上通常产生两性花2∽13朵,单花花期5～13h。自然居群中观察到,花开放后有两种甲虫访花,但外围数轮无花药的退化雄蕊限制了花粉向外散布,内轮雄蕊上花药开裂后花粉直接落在自花的柱头上。花被片闭合后内轮花被片萎蔫成液体,促进了花粉在柱头上萌发。花粉与胚珠的比值为930±180(650∽1120),属于较经济的自花传粉类型。果实成熟后花序倒伏,其顶端产生的营养芽萌发形成新的营养苗。一个基株一年内可产生6∽20个营养苗。黄花蔺自花传粉结实率达80％,每果实产生种子470～640枚。黄花蔺近距离扩散以营养繁殖形成的新植株完成,长距离扩散或占领新生境以及抵抗干旱则依靠种子。人工去雄、套袋及授粉实验表明,黄花蔺的繁育系统是以自交为主的混合交配类型,并可能是由异交向自交演化的。     

枇杷少核株系花粉管生长差异研究

以‘大五星’枇杷退化种子少核株系‘川农1号’(C1)为试材,以‘大五星’枇杷自花授粉为对照,采用荧光显微技术,对枇杷少核株系自花、异花授粉后花粉管生长情况进行观察。结果表明:(1)C1株系异花授粉后花粉能在柱头上萌发,并伸入中央花柱道,在授粉后48h左右到达花柱基部。(2)C1株系自花授粉后,花粉萌发与花粉管的伸长速度相对于异花授粉滞后,自花授粉后36h大多数花粉管到达花柱的中上部并停止生长,且伴随产生花粉管顶端形态异常、荧光异常明亮等现象,最终只有极少数花粉管能到达花柱基部。研究表明,C1株系为配子体自交不亲和,C1株系的自交不亲和性是引起其少核的重要原因之一。     

新牧Ⅰ号杂花苜蓿异交机制与自花粉的生殖干扰

通过控制试验和荧光观测等手段对新牧Ⅰ号杂花苜蓿Medicago varia Martyn'Xinmu No.1'柱头可授期与最佳授粉时间、自花粉的堵塞效应、花粉萌发与花粉管生长状况等进行研究,结果显示:(1)苜蓿柱头可授期与单花寿命具高度的同步性,在单花开放的第1天授粉,传粉效率最高,结果率达65.170%±2.01%(P<0.01),为柱头的最佳授粉时期;在单花开放后第4天授粉,其结果率仍达34.25%±6.73%.(2)自花授粉比异花授粉有着较低的花粉萌发率和花粉管生长速度;清除柱头自花粉后再授异花粉,其结荚率/花序和英内种子粒数由原来的51.760%±5.37%和2.11±0.18粒,提高到72.31%±6.24%(P<0.01)和3.46±0.25粒(P<0.01),自花粉对柱头的堵塞效应非常明显.(3)在繁育系统中,苜蓿采取了雌雄异熟和自交不亲和两套机制来避免自交,但是由于其独特的花结构,即使雄蕊先熟,在保证苜蓿异交机制中也没有起到有效的作用,而其典型的自交不亲和系统,虽然保证了苜蓿较低的自交率,却无法避免自花粉的生殖干扰.     

金沙江干热河谷滇榄仁开花物候与繁育系统

通过野外定点观测,并利用套袋实验、解剖实验以及花粉形态特征观察与活力检测、柱头可受性检测、花粉/胚珠比以及异交指数等,研究了金沙江干热河谷滇榄仁自然群体花部形态特征与开花进程、繁育系统特征以及传粉媒介与环境的适应性等内容。结果表明,滇榄仁始花期在每年4月上旬,群体花期持续时间为30d左右,开花同步性高,呈典型的"集中开花模式"。单花开放时,柱头先伸长和膨大,约4d后雄蕊伸出,具有雌雄异熟的特征,一定程度上避免了自交的发生。单花开放进程可划分为花蕾期、柱头伸长期、雄蕊始露期、盛开期、盛开后期、花谢期等6个时期。除了花蕾期外,整个开花期的花粉均具有活力。花粉活力与柱头可受性之间避开了最适授粉期,但也有部分重叠。滇榄仁可能同时具有风媒和虫媒传粉。繁育系统以异交为主,部分自交亲和,需要传粉者。滇榄仁花部特征、开花物候和繁育系统为其适应干热河谷恶劣气候环境提供了一定的生殖保障和进化潜力。良好的群落环境有助于促进滇榄仁异花传粉,产生更多有效的种子,从而促进林下自然更新。     

The mechanism of delayed selfing in Collinsia verna (Scrophulariaceae)

Collinsia verna, blue-eyed Mary, has floral attributes of an outcrossing species, yet most flowers readily self-pollinate under greenhouse conditions. Here we describe the mechanism of self-pollination in C. verna via changes in relative positions of the stigma and anthers and late timing of receptivity, resulting in delayed selfing. Each flower contains four anthers that dehisce sequentially over ∼1 wk. Pollen that is not collected by pollinators accumulates in the keel petal and retains high viability (>80% pollen germination) up to the time of corolla abscission. The stigmatic surface does not become receptive until after the third anther dehisces. This overlap in the sexual phases is concurrent with a change in herkogamy during floral development. In most flowers (70%), the stigma has moved to the front of the keel and is positioned near the anthers when the third anther dehisces. Under field conditions, fruiting success of plants within pollinator exclosures was ∼75% of the fruiting success in open-pollinated plants (33% fruiting success via autogamy vs. 44% fruiting success, respectively). Collinsia verna plants in pollinator exclosures exhibit variation in autogamy rates within natural populations (range 0–80%). In addition, only half of naturally pollinated, receptive flowers examined had pollen tubes growing in their styles. In contrast, shortly after corolla abscission, nearly all flowers examined (96%) had pollen tubes in their styles. Thus we find that in C. verna, autogamy occurs late in floral development, which has the potential to provide substantial reproductive assurance, and that individuals vary in their ability to set fruit through this mechanism. We suggest that delayed selfing mechanisms may be overlooked in other species and that variable pollinator availability may play a significant role in the maintenance of mixed mating in species with delayed selfing, such as C. verna.     

Does stigma curvature promote delayed selfing? An experimental investigation in Triodanis perfoliata (Campanulaceae)

• Self‐fertilisation that is delayed until after opportunities for outcrossing have ceased has been argued to provide both the reproductive assurance benefits of selfing and the genetic advantages of outcrossing. In the Campanulaceae, presentation of pollen on stylar hairs and progressive stigma curvature have been hypothesised to facilitate delayed selfing, but experimental tests are lacking. Stigma curvature is common in Campanula, a genus largely characterised by self‐incompatibility, and therefore is unlikely to have initially evolved to promote self‐fertilisation. In derived self‐compatible species, however, stigma curvature might serve the secondary function of delayed selfing.
• We investigated delayed selfing in Triodanis perfoliata, a self‐compatible relative of Campanula. Using floral manipulation experiments and pollen tube observations, we quantified the extent and timing of self‐pollination. Further, we hypothesised that, if stigma curvature provides the benefit of delayed selfing in Triodanis, selection should have favoured retention of self‐pollen through the loss of a stylar hair retraction mechanism.
• Results of a stigma removal experiment indicated that autonomous selfing produces partial seed set, but only some selfing was delayed. Pollen tube observations and a flower senescence assay also supported the finding of partial delayed selfing. Scanning electron microscopy revealed that pollen‐collecting hairs retract during anthesis, which may limit the extent of delayed selfing.
• Delayed selfing appeared to be only partially effective in T. perfoliata. The stylar hair retraction in this species would seem to contradict selection for selfing. We suggest that caution and rigour are needed in interpreting floral traits as adaptive mechanisms for delayed selfing.

     

Stigmatic fluid aids self-pollination in Roscoea debilis (Zingiberaceae): a new delayed selfing mechanism

Background and Aims

Delayed selfing is the predominant mode of autonomous self-pollination in flowering plants. However, few delayed selfing mechanisms have been documented. This research aims to explore a new delayed selfing mechanism induced by stigmatic fluid in Roscoea debilis, a small perennial ginger.

Methods

Floral biology and flower visitors were surveyed. The capacity of autonomous selfing was evaluated by pollinator exclusion. The timing of autonomous selfing was estimated by emasculation at different flowering stages. The number of seeds produced from insect-pollination was assessed by emasculation and exposure to pollinators in the natural population. The breeding system was also tested by pollination manipulations.

Key Results

Autonomous self-pollination occurred after flowers wilted. The stigmatic fluid formed a globule on the stigma on the third day of flowering. The enlarged globule seeped into the nearby pollen grains on the fourth flowering day, thus inducing pollen germination. Pollen tubes then elongated and penetrated the stigma. Hand-selfed flowers produced as many seeds as hand-crossed flowers. There was no significant difference in seed production between pollinator-excluded flowers and hand-selfed flowers. When emasculated flowers were exposed to pollinators, they produced significantly fewer seeds than intact flowers. Visits by effective pollinators were rare.

Conclusions

This study describes a new form of delayed autonomous self-pollination. As the predominant mechanism of sexual reproduction in R. debilis, delayed self-pollination ensures reproduction when pollinators are scarce.     

Comparative analysis of late floral development and mating-system evolution in tribe Collinsieae (Scrophulariaceae s.l.)

Species of Collinsia and Tonella, the two sister genera of self-compatible annuals that constitute tribe Collinsieae, show extensive variation in floral size and morphology and in patterns of stamen and style elongation during the life of the flower (anthesis). We used a nuclear ribosomal ITS phylogeny, independent contrasts, and phylogenetically corrected path analysis to explore the patterns of covariance of the developmental and morphological traits potentially influencing mating system. Large-flowered taxa maintain herkogamy (spatial separation of anthers and stigmas) early in anthesis by differential elongation of staminal filaments, which positions each of the four anthers at the tip of the "keel" upon dehiscence. Small-flowered taxa do not show this pattern of filament elongation. The styles of large-flowered taxa elongate late in the 2-5 d of anthesis, resulting in late anther-stigma contact and delayed self-pollination. Anther-stigma contact and self-pollination occur early in anthesis in small-flowered species/populations. Thus, we found complex covariation of morphological and developmental traits that can be interpreted as the result of multitrait adaptation for early selfing and high levels of autogamy, delayed selfing and higher levels of outcrossing, or intermediate levels of outcrossing. Continuous variation in these traits suggests the operation of continuous variation in selective optima or the combined effects of divergent selection and phylogenetic inertia.     

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.     

Delayed autonomous self-pollination in the colonizer Crotalaria micans (Fabaceae: Papilionoideae): structural and functional aspects

We investigated the reproductive biology of Crotalaria micans, a colonizing species that occurs in disturbed sites. The flowers have two whorls of stamens with differential growth and dimorphic anthers. The inner anthers serve two functions: 1) to cooperate with the stylar brush in the pollen presentation, and 2) to push the remaining pollen within the keel onto the stigma at the end of the receptivity period. Flowers pass through three phases: a) male phase, when outer anthers release their pollen, but stigmas are not receptive; b) female phase, when stigmas are receptive, but still separated from flowers' own pollen, and c) an autogamous phase, when the round anthers grow towards the stigma, leading to delayed autonomous self-pollination. Because C. micans is completely self-compatible, delayed self-pollination and geitonogamy result in approximately 76% of seeds being self-fertilized.     

Evolution of floral scent in relation to self-incompatibility and capacity for autonomous self-pollination in the perennial herb Arabis alpina

Background and AimsThe transition from outcrossing to selfing is a frequent evolutionary shift in flowering plants and is predicted to result in reduced allocation to pollinator attraction if plants can self-pollinate autonomously. The evolution of selfing is associated with reduced visual floral signalling in many systems, but effects on floral scent have received less attention. We compared multiple populations of the arctic–alpine herb Arabis alpina (Brassicaceae), and asked whether the transition from self-incompatibility to self-compatibility has been associated with reduced visual and chemical floral signalling. We further examined whether floral signalling differ between self-compatible populations with low and high capacity for autonomous self-pollination, as would be expected if benefits of signalling decrease with reduced dependence on pollinators for pollen transfer.MethodsIn a common garden we documented flower size and floral scent emission rate and composition in eight self-compatible and nine self-incompatible A. alpina populations. These included self-compatible Scandinavian populations with high capacity for autonomous self-pollination, self-compatible populations with low capacity for autonomous self-pollination from France and Spain, and self-incompatible populations from Italy and Greece.Key ResultsThe self-compatible populations produced smaller and less scented flowers than the self-incompatible populations. However, flower size and scent emission rate did not differ between self-compatible populations with high and low capacity for autonomous self-pollination. Floral scent composition differed between self-compatible and self-incompatible populations, but also varied substantially among populations within the two categories.ConclusionsOur study demonstrates extensive variation in floral scent among populations of a geographically widespread species. Contrary to expectation, floral signalling did not differ between self-compatible populations with high and low capacity for autonomous self-pollination, indicating that dependence on pollinator attraction can only partly explain variation in floral signalling. Additional variation may reflect adaptation to other aspects of local environments, genetic drift, or a combination of these processes.     

木槿与野西瓜苗花特征和繁育系统的比较研究

为比较分析柱头裂片运动的野西瓜苗和不运动的木槿在花特征和繁育系统上的差异,对其花性状、花粉/胚珠比和不同授粉处理的座果率进行了测定。结果表明:(1)木槿与野西瓜苗在花冠大小、花瓣长及底宽、花萼长与宽、雄蕊长、最上轮雄蕊高度、雄蕊柱长及其底径、花柱长度和雌雄异位间均存在十分显著的差异(P<0.01),但二者的最下轮雄蕊高度间无显著性差异(P>0.05)。(2)木槿的花粉/胚珠比为874.90±20.79,繁育系统属兼性异交类型;野西瓜苗的花粉/胚珠比为24.72±0.68,繁育系统属专性自交类型。(3)木槿自然套袋的座果率为0%,人工自交为2.04%,人工杂交为35.85%;野西瓜苗自动自交和人工自交的座果率均为100%,人工杂交为95.30%。木槿与野西瓜苗在花性状、花粉/胚珠比及花部行为等方面形成了与其繁育系统(兼性异交vs.专性自交)相适应的花特征。     

THE EVOLUTION OF INSECT-MEDIATED SELF-POLLINATION FROM AN OUTCROSSING SYSTEM IN CALYPTRIDIUM (PORTULACACEAE)

Calyptridium monospermum (Portulacaceae) comprises two biological races visited by different groups of insects. One race, pollinated by a bumblebee, Bombus vosnesenskii (Apidae), is outcrossing, while the other race is pollinated casually by a variety of other insects whose visits result in insect-mediated self-pollination and geitonogamy within the inflorescence. Insect-mediated selfing is also the mode of pollination in the closely related species, C. umbellatum. While all plants of both species are self-compatible, insect visitation is necessary for good seed set. Selfing populations of C. monospermum exhibit many floral adaptations for insect-mediated self-pollination which parallel those of the selfing C. umbellatum, while retaining vegetative characteristics typical of C. monospermum. These floral traits involve flower color and odor, style and stamen orientation, size of stigmatic surfaces, number of pollen grains produced, density of open flowers on inflorescences, and presence or absence of protogyny. Artifical pollination experiments show that these floral adaptations are effective in increasing the probability of insect-mediated self-pollination. Insect-mediated selfing in C. umbellatum and in some populations of C. monospermum is encouraged and probably made necessary by the inconstant pollinating behavior of the insects which visit these plants, as shown by the large percentage of non-Calyptridium pollen recovered from the bodies of insects captured while visiting these plants.