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。     

山莨菪(茄科)的传粉生物学

茄科的多数种类具有自交不亲和的特点, 主要通过异花传粉结实; 但是, 一些物种或者物种内的部分种群或者个体却高度自交亲合, 转变为自交的繁育系统。该科植物山莨菪(Anisodus tanguticus)主要分布在青藏高原, 开花较早, 比其他晚开花的植物种类更加缺少有效的异花传粉昆虫。我们选择了位于不同海拔高度的2个种群进行比较研究, 主要目的是检验该物种的繁育系统是否在极端环境下由于传粉者的缺乏而发生了部分改变。研究发现,山莨菪的花不完全雌性先熟, 柱头和花药间的平均距离随着花开放时间的延长而不断缩小, 但两者在多数花的单花花期结束时并没有发生接触。因此, 山莨菪花主要表现为适应异花传粉的雌雄异位特征。然而, 少数花 (4.9％)的柱头和花药发生接触, 为“自动自交”的传粉解除了空间隔离。2个种群的多数个体存在自交不亲和机制, 应具有异花传粉的繁育系统; 但是部分个体具有明显的自交亲和能力, 为自交提供了生理基础。高海拔种群的传粉昆虫主要是厕蝇(Fannia sp.), 它们在不同植株间的活动能够保证异花传粉结实; 同时该种群的部分个体存在“自动自交”。低海拔种群的主要访花昆虫是蚂蚁, 它们在花内的活动导致花粉在同一朵花内传递, 而引起“协助自交”; 而异花传粉昆虫厕蝇的访花频率则较高海拔种群低。两个种群的结实均由于异花传粉者不足而受到传粉限制。因此两种不同类型的自交机制为该早期开花植物异花访花昆虫的不足提供了一定程度上的繁殖补偿。     

短柄五加开花及传粉生物学研究

本文对短柄五加（Ｅｌｅｕｔｈｅｒｏｃｏｃｃｕｓｂｒａｃｈｙｐｕｓ）的花期物候、开花式样、访花昆虫种类、访花行为及访花频率进行了观察,对花粉活力、柱头可授性、花粉胚珠比以及传粉效果和繁育系统做了检测。初步结果表明,短柄五加以根茎进行无性繁殖,１个居群为１个无性系。虽自交亲和,但昆虫的访花活动可显著提高座果率。昆虫传粉主要在无性系内异花间进行,属于广义的自花传粉。传粉者为多种不同的昆虫,包括马蜂和胡蜂在内的蜂类、蝇类和甲虫。其花期物候、花粉活力、柱头可授期和蜜汁分泌期等特征使其能够适应环境,从而获得较高的座果率。平均座果率为６５％,饱满种子率为４５．５％。自然条件下未发现实生苗,广义的自花传粉导致的种子质量低下可能是其原因,也是限制短柄五加分布范围的重要因素。     

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”     

毛菍传粉生物学研究

为探讨毛菍(Melastoma sanguineum Sims.)传粉生物学特性,对海南霸王岭国家级自然保护区林缘、林内和林隙3个生境内毛菍的花期、花部形态、访花昆虫等进行观察,同时对其繁育系统相关指标进行了测定。结果表明：毛菍无花蜜分泌;花瓣对访花昆虫有吸引作用;访花昆虫主要有蜂类、食蚜蝇类和丽金龟类;主要传粉昆虫有木蜂、熊蜂,其中木蜂传粉活动最为活跃,为最有效传粉昆虫。毛菍具典型的异型雄蕊,长、短雄蕊在传粉过程中具有明确的功能分化,推测这是毛菍保证传粉成功的策略。毛菍为自交亲和的异交种,需要昆虫传粉;人工自交和异交均具有较高的坐果率;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;其繁育系统是兼性自交。昼夜温差较小和相对湿度较高的林内和林隙生境有利于延长群体花期;光照充足、温度相对较高的林缘生境,花朵发育较快,柱头可授性的时间稍早。花粉、传粉者和环境条件是制约毛菍野外传粉效率的主要因素。     

灌木铁线莲繁育系统特征及其花下垂现象的适应性意义

该研究对兰州市北山地区自然居群下的灌木铁线莲(Clematis fruticosa Turcz.)的繁育系统特征进行了实验分析,并通过对其花朵进行人工上举处理,研究了花朝向改变(花朵上举)后访花昆虫、花粉数量和质量变化以及结籽率和种子特征的变化,以此反证花朵下垂的适应性意义。结果表明:(1)自然状态下,灌木铁线莲的P/O值为9579.5,属于专性异花授粉;套袋实验结果显示,灌木铁线莲自交亲和,兼具异交和自交的混合交配系统,但不存在无融合生殖现象。(2)灌木铁线莲的主要传粉者为东亚无垫蜂(Amegill aparhypate)和中华蜜蜂(Apis cerana);人工上举处理改变花朝向后,两种传粉昆虫的访花频率和单花停留时间与自然条件下没有发生变化,但因雨水冲刷造成其花粉数量显著下降,且阳光直射使花粉活力显著下降,说明花朝向改变会降低其雄性适合度。(3)花朝向改变后,灌木铁线莲结籽率显著下降,种子宿存花柱长度降低,但并没有影响种子大小和千粒重,说明花朝向改变降低了雌性适合度,并影响到灌木铁线莲种子传播。研究结果证实,灌木铁线莲花期花朵下垂现象对有效传粉昆虫影响不大,但可以显著提高植物雄性适合度和雌性适合度,并作为一种有效的适应策略保障了其繁殖成功。     

管花秦艽的传粉生态学研究——兼与同域分布近缘种的比较

在青藏高原东北部连续两年观察了晚秋开花植物管花秦艽Gentiana siphonantha的传粉生态学特征,并在此基础上进一步比较分析了与该物种同域分布且亲缘关系较近、但开花较早的麻花艽G straminea之间的传粉生态学特征.管花秦艽的花发育过程表现出雌雄异熟和雌雄异位的特点,不存在花内的自花传粉,套袋隔离的花不结实也支持这一结论;株内自交的高结实率表明该物种是自交亲和的.盛花期每植株平均有15朵开放的花,雄性和雌性阶段的花比例为1.2:1;自然条件下产生种子必须依赖传粉媒介;苏氏熊蜂是最有效的传粉昆虫,且访花过程中埘雄性和雌性阶段花不具明显的偏向性;株内连续访花的频率高达87.8%,从而导致同株异花传粉自交的广泛存在.与同域分布的麻花艽相比,管花秦艽的单花花期、雄性和雌性期持续时间缩短.但盛花期开花数量明显增加.令人感兴趣的是尽管两个近缘种的花形态特征存在显著差异,但都是由同一种熊蜂传粉.这一特点与过去认为花颜色和花管长度是物种分化过程中与不同传粉昆虫协同进化导致牛殖隔离的假说不相符合.管花秦艽单花的访花频率和同株异花连续访花的比例都明显高于麻花艽.两个物种不同花序设计导致访花昆虫行为的改变可能是造成这一差异的主要原因.两个物种具有不同的开花时间,但仍然存在一定的花期重叠,表现出不完全的传粉生殖隔离状态.     

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。     

雄全异株植物瘿椒树（省沽油科）的传粉生物学

从开花动态、传粉昆虫、花的形态结构、繁育系统、花粉活力和柱头可授性等方面研究了我国特有珍稀植物瘿椒树（Tapiscia sinensis Oliv.）的传粉生物学特性。瘿椒树是典型的雄全异株植物,两性花中含有功能性花粉,且自交亲和,但雄花花粉活力和萌发力是两性花的10倍以上。雄株和两性植株具有相同开花物候期,花期均为5月下旬至6月上旬,单花期为4-5天,雄花和两性花的5枚花药开裂的不同步性明显延长了散粉时间。两性花雌蕊先熟,柱头可授性较长。具有适应风媒和虫媒传粉的花部特征。传粉昆虫主要为蜜蜂科（Apidae）和食蚜蝇科（Syrphidae）昆虫,访花高峰期为8：30-10：30。维持瘿椒树雄全异株的可能机制是：雄株总体上增加了异交花粉的数量和质量;两性花的雄蕊为该物种提供了繁殖保障,同时为传粉者提供了报酬。     

黄河三角洲柽柳的开花特性及传粉生态学研究

野外定位观测了柽柳(TamarixchinensisLour.)的开花物侯、开花动态、访花者的种类、行为和访花频率,并对其繁育系统、花粉-胚珠比(P/O)、花粉活力进行了检测。结果表明,柽柳单朵花的花期仅1d,P/O为390.9±81.0,繁育系统属于自交亲和、需要传粉者的类型。花的结构和开花式样适合以食蚜蝇为主的多种昆虫传粉。自然条件下的结实率为95.23%。柽柳的果实成熟期较短(15d),整个居群有长达几个月的花期,开花数目众多,因此能产生大量的种子,在适宜条件下会形成大片的柽柳灌丛。     

Breeding System of Protogynous Mandragora caulescens（Solanaceae）

We studied the breeding system of Mandragora caulescens that distributed in the northwest of Yunnan province. Our results showed that this species bloomed from late May to early June. Flowers were protogynous and lasted （9.9±2.8）d. Three types of visitors were observed. The Queen of Bombus festivces was an effective pollinator, but its visiting frequency was rather low. Although flies and ants visited flowers with high frequency, flies rarely pollinated flowers and ants were totally nectar thieves. Pollination experiments suggested the species was partly self compatible, but protogyny made autonomous selfing become impossible. Fruit set and seed production with supplemental outcrossing pollen was significantly enhanced compared with natural flowers, suggesting pollen limitation in this species. Most species of Solanaceae were strictly self incompatible, but M.caulescens was partly self compatible which was similar with three studied Hyoscyameae members. Hyoscyameae and Mandragoreae formed a monophyletic group independently as a result of two independent dispersal events from the New World into Eurasia. We considered the two tribes independently lost SI systems, and this transition related to harsh environment of alpine region. It is worth conducting further study to clarify whether the evolutionary transition is common in alpine region.     

铃铛子和赛莨菪（茄科）的传粉机制比较研究

通过野外交配系统试验和传粉昆虫观察,以铃铛子（Anisodus luridus）和赛莨菪（A．camiolicoides）为研究对象,探讨了山莨菪属内自交亲和系统的进化与传粉昆虫的转变。结果表明,铃铛子和赛莨菪均属于自交完全亲和的类群,但两个种的自动自交能力均不强,而且都存在传粉限制。北方黄胡蜂（Vespula rtfarufa）和石长黄胡蜂（Dolichovespula saxonica）分别是铃铛子和赛莨菪的主要传粉昆虫,自然状态下两种昆虫的传粉效率均比较高,但两种昆虫访问两种植物时传递花粉的方式不同,其中北方黄胡蜂主要促进了铃铛子的花间传粉,而石长黄胡蜂访花引起赛莨菪同一朵花内的传粉。通过与该属的另一种植物山莨菪（A．tanguticus）的传粉机制比较,发现在山莨菪属的物种分化过程中,由自交不亲和转变为自交完全亲和,传粉昆虫也发生了转变,证明了自交亲和系统存在于起源较晚的类群中。高山环境中频繁且不可预测的降雨可能降低了传粉昆虫的活动能力,进而导致铃铛子和赛莨菪均存在传粉限制。     

铃铛子和赛莨菪( 茄科) 的传粉机制比较研

通过野外交配系统试验和传粉昆虫观察, 以铃铛子( Anisodus luridus) 和赛莨菪( A. carniolicoides)为研究对象, 探讨了山莨菪属内自交亲和系统的进化与传粉昆虫的转变。结果表明, 铃铛子和赛莨菪均属于自交完全亲和的类群, 但两个种的自动自交能力均不强, 而且都存在传粉限制。北方黄胡蜂( Vespula rufarufa) 和石长黄胡蜂( Dolichovespula saxonica) 分别是铃铛子和赛莨菪的主要传粉昆虫, 自然状态下两种昆虫的传粉效率均比较高, 但两种昆虫访问两种植物时传递花粉的方式不同, 其中北方黄胡蜂主要促进了铃铛子的花间传粉, 而石长黄胡蜂访花引起赛莨菪同一朵花内的传粉。通过与该属的另一种植物山莨菪(A. tanguticus) 的传粉机制比较, 发现在山莨菪属的物种分化过程中, 由自交不亲和转变为自交完全亲和, 传粉昆虫也发生了转变, 证明了自交亲和系统存在于起源较晚的类群中。高山环境中频繁且不可预测的降雨可能降低了传粉昆虫的活动能力, 进而导致铃铛子和赛莨菪均存在传粉限制。     

Ants in your plants: effects of nectar-thieves on pollen fertility and seed-siring capacity in the alpine wildflower, Polemonium viscosum

Flowers of the alpine skypilot, Polemonium viscosum , are attacked by nectar thieving ants of Formica neorufibarbus gelida . Ants exert selection on flower scent, size and shape in skypilots by damaging the pistils. Here, I report on the frequency and nature of contact between ants and pollen-bearing anthers and determine the consequences of such contact for pollen performance and pollen donor paternity. In laboratory trials, ants entered flowers with full intact anthers and emasculated (female) flowers equivalently. Similarly, flower visitation rates of ants foraging naturally were not affected by the frequency of male phase flowers per plant. Ants actively interacted with the pollen-bearing anthers during 21% of flower visits, on average. The rate at which such interactions occurred was predicted by the proportion of flowers in the male phase, under a random foraging model. The effect of ants on pollen fertility was tested experimentally by enclosing ants in male-phase flowers on intact inflorescences. Adjacent control flowers were left un-occupied. Pollen from flowers with a history of ant occupancy had significantly lower germination on virgin recipient stigmas than pollen from unoccupied control flowers. With hand-pollination, sufficient pollen was transferred from ant-occupied flowers to saturate seed set. However, a model based on the relationship between seed set and compatible pollen delivery by natural pollinators indicated that ant damage to pollen should reduce paternity accruing per flower visit by 20–26% on average, in nature. Results support the hypothesis that in P. viscosum , selection on floral traits by nectar-thieving ants operates through male as well as female function.     

Pollen limitation,species’ floral traits and pollinator visitation: different relationships in contrasting communities

Failures in the process of pollen transfer among conspecific plants can severely impact female reproductive success. Thus, pollen limitation can cause selection on plant mating systems and floral traits. The relationships between pollen limitation and floral traits might be partly mediated by the quantity and identity of pollinator visits. However, very little is known about the relationship between pollinator visits and pollen limitation. We examined the relationships between pollen limitation and floral traits at the community level to connect them to community ecology processes. We used 48 plant species from two contrasting communities: one species‐rich lowland community and one species‐poor alpine community. In addition, we calculated visitation rates and ecological pollination generalization for 38 of the species to examine the relationship between pollinator visitation and pollen limitation at the community level. We found low overall levels of pollen limitation that did not differ significantly between the alpine and the lowland community. In both communities, species with evolutionary specialized flowers were more pollen limited than species with unspecialized flowers. Species’ visitation rates and selfing capability were negatively related to pollen limitation in the alpine community, where pollinators are scarcer. However, flower size/number, ecological generalization of plants and flowering onset had greater effects on pollen limitation levels at the lowland community, indicating that the identity of the visitors and plant‐plant competitive interactions are more decisive for plant reproduction in this species‐rich community. There, pollen limitation increased with flower size and flowering onset, and decreased with ecological generalization, but only in species with evolutionary specialized flowers. Our study suggests that selection on plant mating system and floral traits may be idiosyncratic to each particular community and highlights the benefits of conducting community‐level studies for a better understanding of the processes underlying evolutionary responses to pollen limitation.     

Dioecy inHonkenya peploides var.major (Caryophyllaceae)

Sexual difference and pollination system inHonkenya peploides var.major were investigated. The present analyses indicate that sex expression of this plant is functionally dioecious, rather than androdioecious, as described previously. Its flowers are largely pollinated by small bees, hover-flies, flies, and ants. Female flowers were morphologically characterized by conspicuously longer styles and smaller petals, and had no pollen grains, while male flowers had shorter styles, larger petals, and longer filaments, and produced viable pollen grains. Although the male flowers possess many ovules in an ovary, they set fruits at a very low frequency in natural conditions and did not produce fruits by pollination experiments. We suggest that the dioecy of this species has been derived from hermaphrodite through gynodioecy, and not directly from hermaphrodite.     

Contribution of small insects to pollination of common buckwheat, a distylous crop

Crop pollination by animals is an essential ecosystem service. Among animal-pollinated crops, distylous plants strongly depend on animal pollination. In distylous pollination systems, pollinator species are usually limited, although flowers of some distylous plants are visited by diverse animals. We studied the pollination biology of common buckwheat ( Fagopyrum esculentum ), a distylous crop mainly pollinated by honeybees and visited by many insect species, to evaluate the effects of non-honeybee species on pollination services. We focused on insects smaller than honeybees to determine their contribution to pollination. We applied pollination treatments with bags of coarse mesh to exclude flower visits by honeybees and larger insects and compared the seed set of bagged plants with that of untreated plants for pin and thrum flower morphs. We found a great reduction of seed set only in bagged pin flowers. We also confirmed that small insects, including ants, bees, wasps and flies, carried pin-morph pollen. These small insects transfer pollen from the short anthers of pin flowers to the short styles of thrum flowers, leading to sufficient seed set in thrum flowers. Consequently, small, non-honeybee insects have the potential to maintain at least half of the yield of this honeybee-dependent distylous crop.     

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。     

Change of floral orientation affects pollinator diversity and their relative importance in an alpine plant with generalized pollination system,Geranium refractum (Geraniaceae)

Floral orientation may affect pollinator attraction and pollination effectiveness, and its influences may differ among pollinator species. We, therefore, hypothesized that, for plant species with a generalized pollination system, changes in floral orientation would affect the composition of pollinators and their relative contribution to pollination. Geranium refractum, an alpine plant with downward floral orientation was used in this study. We created upward-facing flowers by altering the flower angle. We compared the pollinator diversity, pollination effectiveness, and pollinator importance, as well as female reproductive success between flowers with downward- and upward-facing orientation. Results indicated that the upward-facing flowers were visited by a wider spectrum of pollinators (classified into functional groups), with higher pollinator diversity than natural flowers. Moreover, due to influences on visitation number and pollen removal, the pollinator importance exhibited by the main pollinator groups differed between flower types. Compared with natural flowers, the pollination contribution of principal pollinators (i.e., bumblebees) decreased in upward-facing flowers and other infrequent pollinators, such as solitary bees and muscoid flies, removed more pollen. Consequently, stigmatic pollen loads were lower in upward- than in downward-facing flowers. These findings reveal that floral orientation may affect the level of generalization of a pollination system and the relative importance of diverse pollinators. In this species, the natural downward-facing floral orientation may increase pollen transfer by effective pollinators and reduce interference by inferior pollinators.