新西兰岩石百合中两种共存的繁育策略：延迟自动自交和欺骗传粉吸引

延迟自动自交和欺骗传粉吸引在被子植物多数类群中有相关报道,但是至今没有发现两种繁育策略在同一物种中共存现象。本研究通过对照试验检测新西兰岩石百合雄蕊附属物和花冠闭合运动是否分别具有欺骗吸引和延迟自交功能。研究结果表明,新西兰岩石百合黄色的雄蕊附属物拟态花粉（或花药）,约93%的昆虫访花行为源自黄色花药附属物的欺骗吸引,雄蕊附属物的报酬拟态功能有效提高昆虫拜访频率并促进异交。同时,研究发现新西兰岩石百合花期结束时花瓣闭合促使雌雄隔离距离的缩小,花瓣自然闭合的花朵平均结籽数（20.62）显著高于闭合前去雄处理花朵（11.79）。我们的结果表明延迟自动自交与欺骗传粉吸引两种繁育策略在新西兰岩石百合中共存。     

新西兰岩石百合中两种共存的繁育策略：延迟自动自交和欺骗传粉吸引

延迟自动自交和欺骗传粉吸引在被子植物多数类群中有相关报道,但是至今没有发现两种繁育策略在同一物种中共存现象。本研究通过对照试验检测新西兰岩石百合雄蕊附属物和花冠闭合运动是否分别具有欺骗吸引和延迟自交功能。研究结果表明,新西兰岩石百合黄色的雄蕊附属物拟态花粉（或花药）,约93%的昆虫访花行为源自黄色花药附属物的欺骗吸引,雄蕊附属物的报酬拟态功能有效提高昆虫拜访频率并促进异交。同时,研究发现新西兰岩石百合花期结束时花瓣闭合促使雌雄隔离距离的缩小,花瓣自然闭合的花朵平均结籽数（20.62）显著高于闭合前去雄处理花朵（11.79）。我们的结果表明延迟自动自交与欺骗传粉吸引两种繁育策略在新西兰岩石百合中共存。     

野牡丹异型雄蕊的功能分化

野牡丹科植物具有形态、大小和颜色显著不同的两种异型雄蕊。关于其异型雄蕊是否具有功能分化还一直存在争论。本文以野牡丹科植物野牡丹(Melastoma malabathricum)为实验材料, 比较了两种异型雄蕊在传粉过程中的功能作用。结果表明, 两种异型雄蕊在形态、花粉量和人工控制实验条件下的结籽数, 以及主要传粉昆虫木蜂(Xylocopa sp.)访花时的行为等方面都有显著差异, 说明两种雄蕊在传粉过程中存在一定的功能分化: 外轮紫色雄蕊中的花粉为后代提供雄配子, 而内轮黄色雄蕊中的花粉则为传粉昆虫提供食物。但两种雄蕊在花粉活性、花粉组织化学成分和结实率方面差异均不显著, 表明两者在生理上并没有分化。实验结果还表明, 除花前套袋不结实外, 自交、异交和自然对照都具有较高的结实率, 说明野牡丹不存在无融合生殖和主动自交及自交不亲和现象, 为兼性异交。     

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。     

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。     

蛛网萼开花物候、花部特征及繁育系统研究

为探讨蛛网萼(Platycrater arguta)传粉生物学特性,该研究以江西上饶广丰铜钹山国家级自然保护区、江西阳际峰国家级自然保护区蛛网萼为材料,采用野外观察和人工授粉等方法,对蛛网萼开花物候、花部形态及繁育系统进行了研究。结果表明:(1)蛛网萼为伞形花序,种群花期一般在6月中下旬至8月中旬,单花花期为2~3 d,花序花期持续10 d。两个保护区由于年均温度与光照的不同,花期进程有显著不同。(2)蛛网萼可孕花与不孕花从绿色变为白色时对访花昆虫有吸引作用,不孕花能够增加昆虫的访花频率。(3)蛛网萼花粉胚珠比(P/O)为2 520~3 100,杂交指数即是3或4。(4)繁育体系属于以异交为主,部分自交亲和,需要传粉者;熊蜂(Bombus sp.)、四条蜂(Apis florea)和油茶地蜂(Andrena cammelia)是蛛网萼的有效传粉昆虫。(5)套袋和人工授粉结果显示,蛛网萼为兼性自交和异交授粉,异株异花授粉能显著提高坐果率及结籽率,这主要与生境中有效传粉昆虫种类、数量少,花粉传递效率低,种群间基因流交流降低以及柱头自花及同株异花传粉等有关,从而表明在蛛网萼现有生境中植株的早期生殖成功受到影响。蛛网萼种群分布范围扩张以有性繁殖为主,无性繁殖作为重要补充及繁殖保障。     

同域分布的紫堇属三种植物的繁育系统和传粉差异

为研究近缘物种之间繁育系统分化对传粉环境的适应性意义,本文针对湖北地区的3种紫堇属植物（紫堇Corydalis edulis Maxim.、尖距紫堇C.shearer S.Moore.和小花黄堇C.racemosa（Thunb.） Pers.）进行了传粉生态学研究,对比分析了它们在花部特征、分布模式、花期、交配系统、传粉系统等方面的差异。3种紫堇属植物常见伴生现象,花期有一定重叠;花色、距长、花蜜量等方面均有显著差异;尖距紫堇交配系统为自交不亲和、依靠传粉者异花授粉;而紫堇和小花黄堇交配系统为自交亲和,兼有自交和异交的混合交配系统。3个物种均由蜂类传粉,且花部性器官与传粉者的接触部位一致,但访花频率差异较大。在混合群落中,传粉者访花具有较高的忠实性,花部特征差异可能是传粉者选择性访花的原因。对于紫堇属3个物种,花部特征和交配系统的分化有助于其避免传粉过程的相互干扰,交配系统与传粉环境具有一定相关性,混合交配系统可能有利于提高植物对不同生境和气候的适应性。     

天然次生林三种不同生境刺五加丁香苷和总黄酮含量的研究

采用高效液相和分光光度的方法对天然次生林林内、林隙和林缘三种不同生境下刺五加(Acanthopanax senticosus)主要次生代谢产物丁香苷和总黄酮的含量差异进行了研究。结果表明:不同生境条件下,刺五加不同部位丁香苷的含量不同。生境的变化对枝中丁香苷的含量影响最大,林内生境枝中丁香苷的含量最低,林隙生境较高,林缘生境最高。而茎基部丁香苷的含量规律为林内生境最高,林隙生境次之,林缘生境最低。三种不同生境中根和根茎丁香苷含量的变化没有明显的规律性。刺五加全株均含有黄酮类物质,其中叶和花中黄酮的含量较高。其总黄酮含量受环境的影响较大,在林内生境,刺五加叶、花和果中总黄酮的含量最低,林隙生境有所提高,而林缘生境中含量最高。叶片总黄酮含量与叶面积指数呈负相关,叶面积指数越大,叶片中总黄酮的含量越低。对于刺五加的枝、茎干、根和根茎而言, 1~3年枝总黄酮在各个生境之间没有显著的差异;而茎干中总黄酮的含量表现为林内生境显著的高于林隙生境和林缘生境,后两者没有明显的差异;与茎干不同,林内生境根和根茎中总黄酮的含量最低,林隙和林缘的较高。     

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。     

宁夏枸杞传粉生态学初步研究

田间观察检测了宁夏枸杞(Lycium barbarum L.)9个品系的开花物候、开花动态、花粉活力、柱头可授性及其繁育系统,并对花粉萌发、花粉管生长以及访花者的种类和访花行为进行了初步研究.结果表明,宁夏枸杞整个居群有长达5个月的花期,单花的花期仅为3～4 d,花粉活力可保持15 d左右,柱头可授性在72 h左右,雌雄生殖单位在持续时间上有较长的相遇期;花粉数量与胚珠比(P/O)为4 000左右,杂交指数(OCI)介于3或4,繁育系统为异交型,需要传粉者;荧光显微观察表明,参试的3个品系的花粉均能在柱头上萌发,2个有自交不亲和现象,其位置发生在花柱的上半部分和子房内.访花昆虫有17种,分属双翅目、膜翅目和鳞翅目,花的结构和开花式样适合以意大利蜜蜂和食蚜蝇为主的多种昆虫传粉.     

Specialized pollination by carpenter bees in Calanthe striata (Orchidaceae), with a review of carpenter bee pollination in orchids

Calanthe striata has nectarless flowers that are self‐compatible, but pollinator dependent. Field observations showed that the flowers were pollinated exclusively by the carpenter bee Xylocopa appendiculata circumvolans, although the bees occasionally wasted pollen by delivering to the stigmatic surface pollinaria that retained their anther caps. Fruit set ratios at the population level varied spatiotemporally, but were generally low (8.3–17.3%). Calanthe striata blooms in spring when post‐overwintering carpenter bees have not yet started foraging for brood production. It can therefore exploit an abundance of opportunistic/naïve foragers. This timing may also increase the possibility of pollinator visits, because no rewarding co‐flowering plants are available in the orchid habitats. A literature review of Orchidaceae pollinated by carpenter bees revealed that at least 14 species of Orchidoideae and Epidendroideae have evolved flowers specialized for carpenter bee pollination. They typically have shallow pink/magenta flowers with a foothold for pollinators; pollinaria are attached to the head, ventral thorax or base of the middle legs of carpenter bees when they insert their heads and/or proboscises into flowers; pollination success is generally low, a probable consequence of the deceptive pollination systems. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013 , 171 , 730–743.     

Reproductive biology of two Himalayan alpine gingers (Roscoea spp., Zingiberaceae) in China: pollination syndrome and compensatory floral mechanisms

According to the concept of pollination syndromes, floral traits reflect specialisation to a particular pollinator or set of pollinators. However, the reproductive biology of endemic, and often specialised, plants may require increased attention as climate change accelerates worldwide. Species of Roscoea endemic to the Himalayan region have striking orchid-like flowers with long corolla tubes, suggesting pollination by long-tongued insects. Until now, the reproductive biology of species of Roscoea has been poorly documented. We investigated the floral biology, breeding system and pollination ecology of R. cautleoides and R. humeana, from Hengduan Mountains, a global biodiversity hotspot in southwest China. We also tested whether floral longevity increases pollination success. Pollination experiments showed that the two species were self-compatible and depended on insects for fruit production. Over several flowering seasons we did not observe any potential pollinators with long tongues that matched the corolla tube visiting flowers in centres of distribution. The principal pollinators observed were pollen-collecting generalist bees, with low visitation frequencies. In general, members of the ginger family are characterised by short-lived (usually 1 day) flowers, but flowers of R. cautleoides and R. humeana last 8 and 6 days, respectively. Removing stigmas decreased fruit set in both study populations. Our results suggest that the original pollinators may have been long-tongued insects that are now absent from the Chinese Himalayas because habitats have responded to climate change. However, long-lived and self-compatible flowers, coupled with the presence of generalist pollinators, are traits that have allowed these gingers to reproduce and continue to persist in the alpine habitats.     

Three pollination guilds and variation in floral characteristics of Bornean gingers (Zingiberaceae and Costaceae)

The pollinators of 29 ginger species representing 11 genera in relation to certain floral morphological characteristics in a mixed-dipterocarp forest in Borneo were investigated. Among the 29 species studied, eight were pollinated by spiderhunters (Nectariniidae), 11 by medium-sized Amegilla bees (Anthophoridae), and ten by small halictid bees. These pollination guilds found in gingers in Sarawak are comparable to the pollination guilds of neotropical Zingiberales, i.e., hummingbird-, and euglossine-bee-pollinated guilds. Canonical discriminant analysis revealed that there were significant correlations between floral morphology and pollination guilds and suggests the importance of plant–pollinator interactions in the evolution of floral morphology. Most species in the three guilds were separated on the plot by the first and second canonical variables. Spiderhunter-pollinated flowers had longer floral tubes, while Amegilla-pollinated flowers had wider lips than the others, which function as a platform for the pollinators. Pistils and stamens of halictid-pollinated flowers were smaller than the others. The fact that gingers with diverse morphologies in a forest with high species diversity were grouped into only three pollination guilds and that the pollinators themselves showed low species diversity suggests that many species of rare understory plants have evolved without segregating pollinators in each pollination guild.     

High ratio of illegitimate visitation by small bees severely weakens the potential function of heteranthery

Aims In heterantherous plants, 'division of labor' among structurally different stamens, i.e. pollinating and feeding functions, has been thought to reduce the evolutionary conflict of relying on pollen both as the carrier of male gametes and as the food for pollinators. The key to successful division of labor among different sets of stamens is the size match between stamens and legitimate pollinators, which results in the precise deposition of pollen onto specific locations on pollinator's body and facilitates cross pollination. However, the potential impact of small illegitimate insects that are ubiquitous during the pollination process on the plant reproduction in heterantherous species has been largely neglected in previous studies and never been demonstrated experimentally.Methods Here, we investigated the functions of three different types of stamens in Commelina communis. The pollinator visitation, pollen removal and deposition were compared among flowers with different types of anthers emasculated at two natural populations. Moreover, the mating systems of C. communis in wild populations were estimated using microsatellite markers.Important findings Our data showed that the main floral visitors for C. communis at the two studied populations were small illegitimate bees rather than legitimate pollinators, accounting for 77.5 and 92.2% of total flower visits, respectively. Flower manipulations in C. communis demonstrated that the two types of brightly yellow stamens separately functioned as 'deceptive attraction' and 'feeding' functions. Although the brown inconspicuous stamens of C. communis with the largest amount of fertile pollen had the potential function in offering pollen for cross pollination, the high ratio of illegitimate visitation by small bees significantly affected the dispersal and deposition of pollen from the pollinating anthers, and subsequently decreased the levels of outcrossing (t m = 0.23–0.32) in wild populations. Our work further confirmed that the size match between pollinators and the floral morphology is the prerequisite to successfully fulfill the functional differentiation among different sets of stamens in heterantherous plants. Local high ratio of illegitimate visitation by size unmatched insects could significantly weaken the potential functions of heteranthery, affecting the dispersal and deposition of functional pollen in heterantherous plants and further the whole mating systems.     

Pollination biology of the sclerophyllous shrub Pultenaea villosa Willd. (Fabaceae) in southeast Queensland, Australia

The pollination biology of the common shrub Pultenaea villosa Willd. was examined in a subtropical dry sclerophyll forest in eastern Australia. We determined floral phenology and morphology, the timing of stigma receptivity and anther dehiscence, nectar availability, the plant breeding system, and flower visitors. The shrub's flowers are typical zygomorphic pea flowers with hidden floral rewards and reproductive structures. These flowers require special manipulation for insect access. A range of insects visited the flowers, although bees are predicted to be the principle pollinators based on their frequency on the flowers and their exclusive ability to operate the wing and keel petals to access the reproductive structures. Nectar and pollen are offered as rewards and were actively collected by bees. Nectar is offered to visitors in minute amounts at the base of the corolla. In Toohey Forest, P. villosa flowers in spring and is the most abundant floral resource in the understory of the forest at this time. The breeding system experiment revealed that P. villosa requires outcrossing for high levels of seed set and that the overlap of stigma receptivity and pollen dehiscence within the flower suggests the potential for self-incompatibility.     

Pollination ecology of the Cerrado species Eschweilera nana (Lecythidaceae subfam. Lecythidoideae)

Eschweilera nana is pollinated by a guild of pollinators consisting of mostly bees. Effective pollinators are large bees able to force their way into the closed androecium to access nectar. The morphology of the flowers diminishes self pollination and promotes cross-pollination. Although many pollinators make diurnal visits to the flowers, fruit set was very low in comparison with the number of flowers produced. Breeding system tests yielded only two fruits, one produced by xenogamy and another one in the control test. The results of this study are consistent with studies of other Cerrado plants pollinated by guilds of insects and support the conclusion of other pollination studies of Lecythidaceae that fruit set is low in comparison with the high numbers of flowers produced.     

Mechanisms that limit pollinator range in Ericaceae

Studies were conducted in 2001-2003 at Valdai National Park (Novgorod region) and at the Zvenigorod biological station of Moscow State University. The morphology of flowers, flowering dynamics and composition of insect visiting flowers of Ericaceae species: Andromeda polifolia, Chamaedaphne calyculata, Ledum palustre, Oxycoccus palustris, Vaccinium myrtillus, V. uliginosum, and V. vitis-idaea L. were studied. Some species of insects visiting flowers were excluded from the list of pollinators on the basis of observation on their behavior. L. palustre was visited mainly by flies where as other investigated species were visited mainly by bumblebees. In some cases bumblebees were the only visitors of the investigated plants. Mechanisms that protect flowers from flies and short-tongued solitary bees visits and ensure a best pollination by bumblebees are various among different species of Ericaceae. Efficiency of nectary protection also differs among different plant species and is defined by particularities of their habitats and flowering phenology. As far as all species of this family during the flowering are dominants in typical habitats, a competition for the pollination with species of other families in most cases is megligible. Flowering periods of V. vitis-idaea and V. myrtillus in forest ecosystems overlapped weakly. Moreover, V. myrtillus is pollinated mainly by bumblebee queens where as pollinators of V. vitis-idaea are bumblebee workers, solitary bees and horse flies. The other investigated plant species inhabit only oligotrophic peat bogs. Thery are pollinated by bumblebees but periods of flowering are not overlapped and consequently follow one after another. L. palustre and V. uliginosum flower simultaneosly but they are pollinated by different pollinators.     

木蜂的访花行为和传粉作用

木蜂作为蜜蜂科的重要类群,是热带和亚热带地区显花植物常见的传粉者.木蜂具有访花的季节时间长、访花植物种类多、能耐高温或低光照、能嗡声传粉等特点.这些访花特点是木蜂作为热带植物、尤其是温室作物、夜间开花植物和部分茄属植物的重要传粉者.近年来国外已经证明木蜂能有效为蓝莓、西番莲、荷包豆、温室西红柿和温室甜瓜授粉,但我国有关木蜂为多数农作物传粉的重要性和有效性的证据较少.本文综述了近年来木蜂的访花行为和传粉作用的研究进展,为进一步保护、管理和利用木蜂资源提供理论依据.     

Breeding system of Anemone shikokiana and the influential factors of fruit-setting ratio in different habitats

Anemone shikokiana (Makino) Makino is a rare perennial herb distributed only in China and Japan in restricted patches of two different habitats: the shrubs of mountaintop and the conifer and broadleaf mixed forest. Our data showed that the fruit-setting ratio is much lower in the latter than in the former. Given that the reproductive strategy is very crucial for fruit-setting, to reveal the main reasons that influence the fruit-setting ratio, breeding system must be known. Therefore, firstly, P/O and OCI were analyzed and results showed that breeding system of A. shikokiana was self-compatible and mainly outcrossed. Secondly, controlled pollination tests were carried out showing that neither apomixes nor wind pollination occurred. Further results on the type of the breeding system confirmed that A. shikokiana was self-compatible and mainly outcrossed by insects. Lastly, diameters of flowers and frequency of pollinators in two habitats were compared. Results showed that flowers were bigger and frequency of pollinators were higher in shrubs of mountaintop than those in conifer and broadleaf mixed forest, while numbers of pollens and ovaries had little difference. Thus, the overall analysis of the results indicates that the paucity of insect pollinators is the most important reason for the fruit-setting ratio of A. shikokiana lower in the conifer and broadleaf mixed forest than in the shrubs of mountaintop; however, such an implication needs to be further verified with additional work.