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。     

海南凤仙花不同海拔种群的传粉生物学

凤仙花属(Impatiens)植物具有极为广泛的多样性和类型各异的特化传粉者, 被誉为“双子叶的兰花”, 受到众多传粉生物学家的关注。本文以海南特有种海南凤仙花(Impatiens hainanensis)为研究对象, 对3个不同海拔梯度的种群进行了开花生物学特性和开花物候、花器官结构、花粉活力和柱头活性、传粉者种类和访花行为及繁育系统的比较研究。结果表明: 单花花期4.10 ± 0.46 d, 雄性期和雌性期分别约为3.15 ± 0.24 d和0.95 ± 0.36 d; 种群开花峰期在8月初, 高海拔种群的花期高峰相对滞后。低、中海拔种群花粉活力呈现先升高后下降的趋势, 以开花第2 d花粉活力最高; 高海拔种群花粉活力随开花时间推移逐渐下降; 柱头活性随开花时间的推移而增强, 高海拔种群开花各天次均较低、中海拔种群低。主要传粉昆虫为黄黑无垫蜂(Amegilla leptocoma)和绿条无垫蜂(A. zonata), 低、中海拔种群以黄黑无垫蜂为主, 高海拔种群以绿条无垫蜂为主。未观察到自动自花授粉和无融合生殖现象, 人工授粉能明显增加坐果率(75-90%), 自然坐果率在高海拔种群相对较低(40-60%), 说明存在较强的传粉者限制。海南凤仙花的保护需要同时关注其有效传粉者的保护, 促进有效传粉昆虫在不同海拔种群之间的往来, 保证种群间的花粉流与种子流, 维持海南凤仙花的种群遗传多样性与有效种群大小。     

荒漠短命植物异喙菊的小花异形性及繁殖特性

异喙菊(Heteracia szovitsii)是分布于新疆北部荒漠、具短寿命花和多态果实的菊科一年生短命植物。本文对该物种的开花式样和花部特征以及繁育系统与昆虫传粉特性进行了研究,以期明确其头状花序中小花的形态差异及其与多态果实间的关系、揭示其在短的花寿命内的繁殖特性。结果表明:该物种头状花序中发育成多态果实的3类小花在数量、冠毛的有无以及舌片长/宽、子房宽、花柱裂片长和子房喙长之间均存在不同程度的差异,且3类小花冠毛的有无以及子房宽和喙长与多态果实的形态一致,说明多态果实的数量与形态在小花发育过程中就已完成分化。花序中的小花每天集中在上午开放,使花序在功能上犹如一朵花,可快速增大花展示和对传粉者的吸引,以使异交在开花后短时间内完成;3类小花的自花花粉均可在花柱裂片上萌发并长出花粉管,说明该物种自交亲和。3类小花不去雄套袋后均能结实,且自然授粉后的结实率显著高于不去雄套袋的,说明该物种能自主自花授粉,且异花授粉促进了结实。3类小花通过雄性先熟以及次级花粉呈现中的泵/刷机制,将花粉呈现在花柱上部和未展开的柱头裂片外侧刷状毛上以及裂片顶端,延长了花粉呈现(雄性阶段)持续的时间,减少了花内雌雄功能间的干扰,有利于传粉者异花传粉,提高了雄/雌性适合度;同时,柱头裂片展开时可将自花花粉落置到裂片上,以完成自主自花授粉。主要传粉者六显甜花蜂(Halictus sexnotatulus)的访花时间与花序开放时间、花粉活力最高期和柱头最佳授粉期高度吻合,有利于雄性阶段花粉的输出以及雌性阶段柱头接受花粉,保障授粉在开花后短时间内快速有效地完成以及异交成功。在新疆北部荒漠的春季环境中,具短寿命花的异喙菊既可在低温多风导致传粉者不足或活动受限时,通过快速自主自花授粉及自交亲和机制来提供繁殖保障,也可在传粉者活动正常时,通过集中开花、雄性先熟及次级花粉呈现等机制来快速完成异交,提高雌/雄性及后代的适合度。     

苹果园周围生境对凹唇壁蜂种群的维持作用

【目的】凹唇壁蜂Osmia excavata Alfken是我国重要授粉昆虫之一,在北方苹果树授粉有极其重要的作用。如何通过管理措施,维持壁蜂种群成为急需解决的重要问题之一。【方法】以苹果园和周围生境为研究区域,在四类不同生境类型中(苹果园中心区与边缘区,苹果周围生境中心区和边缘区),共选取18个点,采用粘虫板(白板和黄板)和蜂巢(芦苇管和塑料巢箱)两种不同调查方式监测凹唇壁蜂及其天敌叉唇寡毛土蜂种群数量动态,重点分析苹果园周围生境对凹唇壁蜂种群的维持作用。【结果】粘虫板调查结果表明:整体上凹唇壁蜂的数量在苹果园周围生境(65%±17%)高于苹果园生境(35%±12%)。芦苇巢管调查表明:凹唇壁蜂筑巢数量在苹果园周围生境(69%±26%)高于苹果园生境(31%±9%);塑料巢管调查表明,凹唇壁蜂筑巢数量在苹果园周围生境(48%±13%)与苹果园生境(52%±13%)基本相同。苹果园的周围生境可以为释放在果园内的壁蜂提供稳定觅食筑巢条件,起到有效维持凹唇壁蜂种群的作用。【结论】苹果园景观中的凹唇壁蜂种群受生境类型的影响,果园周围的生境可以为凹唇壁蜂等授粉昆虫提供栖息空间与食物资源,具有重要价值。在苹果园实际生产与管理时,应考虑在其周围生境中保护与种植更多的开花植物,为以凹唇壁蜂为代表的授粉昆虫提供适宜的栖息环境和补充食物,维持其种群。     

两种生境间大花百合不同性别表型开花及传粉特征的差异

植物的花期物候与花部综合特征均体现了其对生存环境的适应, 是与周围生物及非生物环境长期适应进化产生的结果。大花百合(Lilium concolor var. megalanthum)野外种群中具有雄性植株和两性植株两种不同性别表型。该研究以人为干扰较弱的孤山屯湿地以及人为干扰较强的金川湿地的大花百合为材料, 通过对两种生境间不同性别表型植株的花期物候、花部综合特征以及主要传粉昆虫的访花行为进行比较研究, 探究大花百合的开花及传粉特征在两种生境间的差异。结果表明: 孤山屯湿地内大花百合种群的始花期、盛花期、末花期均早于金川湿地。孤山屯湿地内大花百合的雄性植株在初开时花粉活力高于金川湿地, 而两种生境间的大花百合两性植株的花粉活力总体变化趋势一致。孤山屯湿地内大花百合雄性植株初开时的花蜜含量较高, 而后缓慢下降; 而在金川湿地内则是先增长, 48 h后开始下降。孤山屯湿地内大花百合两性植株的花蜜含量在开花24 h后开始下降, 金川湿地内则是48 h后才开始下降。孤山屯湿地内大花百合无论是雄性植株还是两性植株, 糖含量达到峰值的时间均比金川湿地晚。两种生境中大花百合两性植株的柱头可授性均保持较高水平。两种生境分布的大花百合主要传粉昆虫为中华蜜蜂(Apis cerana)、蓝灰蝶(Everes argiades)和老豹蛱蝶(Argyronome laodice)。孤山屯湿地内中华蜜蜂及蓝灰蝶对大花百合的访花频率均显著低于金川湿地, 但老豹蛱蝶的访花频率显著高于金川湿地。3种访花昆虫中, 中华蜜蜂的访花效率最高, 蓝灰蝶与老豹蛱蝶起到补充授粉的作用。两种生境间大花百合花期物候和花部综合特征的变化与当地小气候和传粉昆虫数量有关, 这种变化是大花百合经过长期适应而产生的。     

5种地黄花部综合特征与繁育系统的初步研究

生殖隔离是物种分化与延续的基础,地黄属植物是否存在生殖隔离仍不清楚。通过实地观测地黄属5个物种(天目地黄、湖北地黄、裂叶地黄、地黄和茄叶地黄)自然种群的花部形态、开花动态、花蜜体积、花蜜的糖浓度及其日变化、杂交指数、花粉-胚珠比等指标,结合人工授粉套袋,对地黄属植物的花部综合特征和繁育系统进行研究。结果显示:(1)地黄属植物的花为两性花,单花花期5～7 d,种群花期40～60 d;5种地黄在花序类型、花形态、花粉量、花粉-胚珠比等方面都存在差异,其中茄叶地黄的花粉完全败育。(2)5种地黄属植物的繁育系统属于异交,部分自交亲和,需要传粉者,不具有无融合生殖特性,地黄与茄叶地黄存在强烈营养繁殖。(3)5种地黄属植物的花朵基部有蜜腺可以产生糖浓度较高的花蜜,这也支持传粉者的存在,但除地黄确定为蜜蜂传粉外,其它地黄植物在野外并未观测到有效的传粉者,可能与地黄的散生特性、生境相互隔离以及传粉观测期气象条件恶劣有关。研究认为,5种地黄属植物较长的花期能有效弥补传粉者访问频率低的不足;除遗传屏障外,推测地黄属5个物种花展示方式的差异可能吸引不同的传粉者,加之地理、生态因素的作用导致了5种地黄属植物繁育系统的分化。     

多叶斑叶兰繁育系统与传粉生物学研究

为探讨多叶斑叶兰(Goodyera folisa)的繁育系统与传粉生物学特征,对其开花物候、花粉活力与柱头可授性、人工授粉、花的挥发性成分以及昆虫传粉行为进行了研究。结果表明,多叶斑叶兰的唇瓣黄色,萼片白色或白色带红褐色;单花花期为(9.4±0.8)d;花粉在开花后第1天具备活力,柱头在开花后第2天具备可授性,花粉活力和柱头可授性都在开花后第5天达到峰值。去雄套袋和不去雄套袋都不能结实,人工自花授粉、同株异花授粉和异株异花授粉的结实率分别为93.3%、95.0%和96.7%,自然结实率为43.3%。花朵的主要挥发性成分为1-辛烯-3-醇、3-辛醇和N,N-二甲基甲酰胺。多叶斑叶兰传粉者为中华蜜蜂(Apis cerana)。多叶斑叶兰具有自交亲和能力,但在自然界不具有主动自交现象,必须依赖中华蜜蜂传粉,花色及花香气味为吸引传粉者的主要因素。     

草乌传粉过程中的广告效应与回报物质研究

虫媒传粉植物与其传粉者之间的相互作用被认为是被子植物花多样性的一个重要选择压力。这种相互作用体现在植物对传粉者的吸引以及传粉者行为对花粉的转运两个方面。本项研究通过去除不同的花部器官研究了草乌(Aconitum kusnezoffii)对其传粉者的吸引, 并结合传粉者的访问行为和草乌花的生物学特性探讨了传粉过程与交配系统的适应。红光熊蜂(Bombus ignites)是草乌的有效传粉者。去除花萼片显著降低了红光熊蜂的访问频率, 但去除特化成蜜腺叶的花瓣并没有显著改变红光熊蜂的访问频率, 这表明草乌吸引红光熊蜂的主要结构是由5枚萼片组成的花部外观形态, 而非花瓣。花蜜是草乌提供给红光熊蜂的回报物质, 糖浓度为39.23%, 组氨酸浓度为0.25 μg/μL。草乌花较大、单花花期长、雌雄异熟、花粉寿命长, 是一个自交亲和但需传粉者传粉完成繁殖过程的异交物种。草乌花序是无限花序, 当上部的花处于雄性阶段时, 下部的花正好处于雌性阶段。而红光熊蜂在草乌花序上的访问顺序通常自下而上, 带来异交花粉为下部的花进行异花授精, 同时又带走上部花的花粉, 这就很好地促进了草乌的异交。草乌雄蕊自外向内逐渐成熟, 是一种有效的限制传粉者单次访问浪费大量花粉的花粉装配策略, 能提高植物通过花粉散布获得的雄性适合度。     

中国特有种——沙芥繁育系统和传粉生物学研究

运用定位观察、杂交指数、花粉-胚珠比、套袋试验和重力玻片法等方法,对沙芥自然居群和人工栽培种群的开花日动态、繁育系统和传粉习性进行了研究.结果显示:沙芥在自然居群和人工栽培种群开花日高峰分别在19:00和15:00～18:00.沙芥繁育系统为异花授粉植物.沙芥传粉媒介中风的飘移能力非常微弱;沙芥自然居群的访花昆虫有19种,分属于6个目;人工栽培种群的访花昆虫有11种,分属于6个目.意大利蜜蜂、熊蜂、黑带食蚜蝇和拟蜂食蚜蝇是两个居群共有的主要传粉昆虫,在自然居群熊蜂每天的访花频率呈双峰曲线,其中访花最高峰与开花日高峰基本重叠.     

传粉生物学研究获新成果

对大多数的开花植物来说,需要将一朵花里面的花粉搬运到另一朵花的柱头上,才有可能完成雌雄交配,并产生后代,而搬运花粉的任务,多数情况下需要动物来完成。在动物协助植物传粉的同时,多数植物为动物提供花粉、花蜜等作为报酬。     

露蕊乌头(毛茛科)不同海拔居群的传粉生态学

对青藏高原东北部两个不同海拔地点（互助,2460m和海北,3200m）的露蕊乌头Aconitum gymnandrum的传粉生态学进行了比较研究。研究结果表明高海拔居群的单花花期、雄性期持续时间、雌性期持续时间比低海拔长。露蕊乌头不存在无融合生殖现象,尽管高度自交亲和,但产生种子必须依赖传粉媒介。熊蜂是露蕊乌头的主要传粉昆虫,两个居群均存在传粉限制。熊蜂取食花蜜为主,在同一个花序上的访问顺序以由下向上为主,但在互助居群和海北居群有3．9％和2．7％的访花是由上向下进行的。统计发现有37．7％和29．3％的访问行为发生在同一植株内;因此,同株异花传粉造成的自交在露蕊乌头中仍然不能避免。低海拔居群的熊蜂种类和访花频率均高于高海拔居群,但自然状态下两个居群的结实率并没有显著差异。高海拔露蕊乌头居群可能具有较长的柱头持续时间,从而补偿了由于访花频率低带来的结实率降低。     

慈姑花的开放式样及其花粉流

对一野生慈姑（ＳａｇｉｔａｒｉａｔｒｉｆｏｌｉａＬ．）居群的性表达状态及其花粉流散布规律进行了观测。慈姑雌雄同株,单株雄花数是雌花数的３．６倍。雌雄花单花期均为１ｄ,但二者开放式样不同。尽管雌花的数目较少,但在１ｄ内开放较多的花,其花的开放速率比雄花快。在居群水平上表现出雌雄花比例随时间大幅度波动。在传粉者进行的２６１２次访花活动中,传粉者表现出偏“雌性”访问,对每朵雌花的访问是雄花的１．４倍。用番红染色法标记慈姑的花粉运动表明,花的不同开放式样影响花粉流的散布。当雄花数占高比例时,花粉主要在邻近的个体间散布;而雄花的比例较低、雌花相对较多时,花粉远距离散布的强度加大。     

Pollination biology of Albizia lebbeck (L.) Benth. (Fabaceae: Mimosoideae) with reference to insect floral visitors

Indian siris, Albizia lebbeck (L.) Benth. (Fabaceae: Mimosoideae) has significant importance to human beings for its multipurpose use. Insects play a crucial role in the pollination biology of flowering plants. In the current study, we studied the pollination biology of A. lebbeck with special reference to insect floral visitors. The effectiveness of floral visitors was investigated in term of visitation frequency, visitation rate and pollen load during 2012 and 2013. In the second experiment, effect of pollinators on yield of A. lebbeck was studied in open and cage pollination experiments. Floral visitor fauna of A. lebbeck included eight-bees, two wasps, two flies, and two butterflies species. Among them, Apis dorsata, Apis florea, Amegilla cingulata, and Nomia oxybeloides had maximum abundance ranging from 349–492, 339–428, 291–342 and 235–255 numbers of individuals, respectively during two flowering seasons. A. dorsata had the highest visitation frequency (6.44 ± 0.49–8.78 ± 0.48 visits/flower/5min) followed by Amegilla cingulata (6.03 ± 0.43–7.99 ± 0.33 visits/flower/5min) and A. florea (3.61 ± 0.31–4.44 ± 0.18 visits/flower/5min). A. dorsata, N. oxybeloides, and Amegilla cingulata had the highest visitation rates (18.904 ± 1.53–11.43 ± 1.17 flower visited/min) and pollen load (15333 ± 336.22–19243 ± 648.45 pollen grains). The open pollinated flowers had significantly higher capsule weight (4.97 ± 0.21 g), seed weight (1.04 ± 0.05 g), seed numbers per pod (9.80 ± 0.34) and seed germination percentage (84.0 ± 1.78%) as compared to caged flowers. The results suggested bees especially A. dorsata, N. oxybeloides and Amegilla cingulata could be effective pollinators of A. lebbeck.     

Pollinator response to female and male floral display in a monoecious species and its implications for the evolution of floral dimorphism

Pollinator-mediated selection has been hypothesized as one cause of size dimorphism between female and male flowers. Flower number, ignored in studies of floral dimorphism, may interact with flower size to affect pollinator selectivity. In the present study, we explored pollinator response, and estimated pollen receipt and removal, in experimental populations of monoecious Sagittaria trifolia, in which plants were manipulated to display three, six, nine or 12 female or male flowers per plant. In this species, female flowers are smaller but have a more compressed flowering period than males, creating larger female floral displays. Overall, pollinators preferred to visit male rather than female displays of the same size. Both first visit per foraging bout and visitation rates to female displays increased with display size. However, large male displays did not show increased attractiveness to pollinators. A predicted relationship that pollen removal, rather than pollen receipt, is limited by pollinator visitation was confirmed in the experimental populations. The results suggest that the lack of selection on large male displays may affect the evolution of floral dimorphism in this species.     

The floral biology of Curcumorpha longiflora (Zingiberaceae): a ginger with two-day flowers

Curcumorpha longiflora is a perennial chasmophyte (rock fissure plant) ginger that usually grows in crevices of calcareous rocks and forms patches on the understory of limestone monsoon rainforests. The pollination ecology of C. longiflora was studied by monitoring phenology and flowering behavior, observing pollinator activity (frequency and behavior of visitors), and the quantity and quality of pollination services. We also investigated the germination of pollen grains and growth of pollen tubes after different pollination treatments to detect its breeding system. Based on the results: (1) for the first time in Zingiberaceae a new protandrous mechanism was found with a two-day flowering to avoid autogamy in this species; (2) under field conditions, all individuals of C. longiflora usually produced only one flower every other day to keep geitonogamy to a minimum; (3) germination of pollen grains and growth rates of pollen tubes under different pollination treatments were the same 4 h later after pollination, suggesting that C. longiflora is completely self-compatible; (4) among the limited visitors, Bombus sp. and Apis florae were effective pollinators, but they were active at different times and at different stages of the flower, probably receiving different rewards.     

Floral antagonists counteract pollinator‐mediated selection on attractiveness traits in the hummingbird‐pollinated Collaea cipoensis (Fabaceae)

Pollinator‐mediated selection toward larger and abundant flowers is common in naturally pollen‐limited populations. However, floral antagonists may counteract this effect, maintaining smaller‐ and few‐flowered individuals within populations. We quantified pollinator and antagonist visit rates and determined a multiplicative female fitness component from attacked and non‐attacked flowers of the Brazilian hummingbird‐pollinated shrub Collaea cipoensis to determine the selective effects of pollinators and floral antagonists on flower size and number. We predicted that floral antagonists reduce the female fitness component and thus exert negative selective pressures on flower size and number, counteracting the positive effects of pollinators. Pollinators, mainly hummingbirds, comprised 4% of total floral visitation, whereas antagonist ants and bees accounted for 90% of visitation. Nectar‐robbers involved about 99% of floral antagonist visit rates, whereas florivores comprised the remaining 1%. Larger and abundant flowers increased both pollinator and antagonist visit rates and the female fitness component significantly decreased in flowers attacked by nectar‐robbers and florivores in comparison to non‐attacked flowers. We detected that pollinators favored larger‐ and many‐flowered individuals, whereas floral antagonists exerted negative selection on flower size and number. This study confirms that floral antagonists reduce female plant fitness and this pattern directly exerts negative selective pressures on flower size and number, counteracting pollinator‐mediated selection on floral attractiveness traits.     

Flowering phenology of Campanula on Mt Olynipos, Greece

We studied the fowering phenology of the genus Campanula (represented by nine species) along the elevation gradient of Mt Olympos (Greece) in order to assess whether there are elevation patterns at the genus level and whether these relate to patterns previously observed along such gradients at the community level. The traits examined were time and duration of the flowering period, flower life span, and duration of male and female flower phases. Floral attributes, such as number of flowers per plant, flower biomass, flower size, were also studied in order to examine whether they change with elevation or influence flower phenology. Flowering of Campanula species started in mid-May and ended in late September. The average duration of flowering of the genus was ca 27 d and the average floral longevity 4 d. In all but one species, the female phase lasted longer than the male. Campanula versicolor differed remarkably from all others in flower phenology and other floral trails. Nearly all Campanula populations studied had right or positively skewed flowering distributions indicating that flowering begins more abruptly than it ends. At the genus level, the time of flowering increased with elevation by 2 3d for every 100 m. Floral longevity also increased with elevation, by 0,2 d for every 100 m. Neither duration of flowering nor duration of the flower phases showed any consistent change with elevation. The same is true of the non–phenological floral traits examined. No trade–off between duration of flowering and flower life span or between structure and maintenance of flowers was apparent. The pattern of increasing floral longevity along the elevation gradient suggests a mechanism of compensation for reduecd pollinator availability.     

Nectar Sugar Production across Floral Phases in the Gynodioecious Protandrous Plant Geranium sylvaticum

Many zoophilous plants attract their pollinators by offering nectar as a reward. In gynodioecious plants (i.e. populations are composed of female and hermaphrodite individuals) nectar production has been repeatedly reported to be larger in hermaphrodite compared to female flowers even though nectar production across the different floral phases in dichogamous plants (i.e. plants with time separation of pollen dispersal and stigma receptivity) has rarely been examined. In this study, sugar production in nectar standing crop and secretion rate were investigated in Geranium sylvaticum, a gynodioecious plant species with protandry (i.e. with hermaphrodite flowers releasing their pollen before the stigma is receptive). We found that flowers from hermaphrodites produced more nectar than female flowers in terms of total nectar sugar content. In addition, differences in nectar production among floral phases were found in hermaphrodite flowers but not in female flowers. In hermaphrodite flowers, maximum sugar content coincided with pollen presentation and declined slightly towards the female phase, indicating nectar reabsorption, whereas in female flowers sugar content did not differ between the floral phases. These differences in floral reward are discussed in relation to visitation patterns by pollinators and seed production in this species.     

Pollination ecology of Agave macroacantha (Agavaceae) in a Mexican tropical desert. I. Floral biology and pollination mechanisms

In a study of sexual reproduction in long-lived semelparous plants, we observed Agave macroacantha in the tropical desert of Tehuacán-Cuicatlán, Mexico, describing duration of flowering, flower phenology, and nectar production patterns. We also performed two manipulative experiments evaluating (a) the seed production efficiency of different crossing systems (selfing, cross-pollination, apomixis, and control), and (b) the effect of different pollinators (diurnal exposure to pollinators, nocturnal exposure, exclusion, and control) on the seeds produced. Flowering occurred from early May to late July and had a mean duration of 29 days in the individual rosettes. The flowers were protandrous; anthesis occurred in the afternoon of the third day after floral opening, and the pistils matured in the afternoon of the fifth day. The stigmas remained receptive from dusk to the following morning. Pollination was mostly allogamous. Nectar was produced principally during the night, from the first stages of floral aperture until the stigmas wilted and flowering ceased. The flowers were visited during the day by hymenoptera, butterflies, and hummingbirds and during the night by bats and moths. Only the nocturnal visitors, however, were successful pollinators. Agave macroacantha is extremely dependent on nocturnal pollinators for its reproductive success.     

钝叶柃不同性别花的花部形态与传粉特征比较

据有关资料记载,柃属(Eurya Thunb.)植物属于雌雄异株植物,目前仅在柃木(Eurya japonica Thunb.)中有过两性花的报道。近年的调查发现,钝叶柃(Eurya obtusifolia H.T.Chang)也有性别变异,存在两性花。在对其性别特征及功能研究的基础上,进一步跟踪了钝叶柃的开花和传粉过程,对不同性别花的花部形态和传粉特征进行了比较分析。在重庆市北碚区选取了4个样地,测定了花部形态、花朵朝向、花粉活力、柱头可授性、花粉对水的耐受性、单花花粉量和单花泌蜜体积等形态和传粉相关指标,并对其开花动态、传粉昆虫种类和昆虫传粉行为进行了仔细观察,还进行了套袋试验。结果发现钝叶柃不同性别花的花部形态和传粉特征既有一些共同点,又存在着明显差异,性别变异株则呈现出雌雄植株间的过渡特征。钝叶柃是一种花期短,花小而多,开花同步性高的植物,雌花和雄花色味相同。该植物的主要传粉方式为虫媒,主要传粉者为两种蜜蜂科昆虫,但风媒也在其传粉中起着一定作用。该植物不同性别花的主要差异有:1)雄花呈灯笼状,花瓣不反卷,雌花辐射状,花瓣反卷;2)雄花较雌花大;3)雄花倾向于垂直朝下,雌花倾向于斜向下朝向枝条末端;4)雌花寿命较雄花长。钝叶柃在开花和传粉上的许多特征都在一定程度上体现了其对雌雄异株性系统的适应。相应地,传粉者在不同性别植株上的行为也存在一定差异。钝叶柃不同性别花在形态上的差异是其周围各种生物因素及非生物因素共同作用的结果,其中传粉昆虫和雨水的选择可能在这些差异的塑造中起着重要作用。