木荷主要传粉昆虫的传粉行为

木荷(Schima superba)是我国珍贵的优质阔叶用材和高效生物防火树种,属于异花授粉植物,其种子生产必须依靠传粉昆虫。为了明确木荷主要传粉昆虫的组成及优势传粉昆虫的传粉效率,2015—2016年对浙江省兰溪市木荷种子园主要传粉昆虫的访花行为、携粉部位及数量、访花频率、花粉移出和柱头花粉沉降数目等进行了观测。结果表明,中华蜜蜂(Apis cerana)、白星花金龟(Protaetia brevitarsis)和棉花弧丽金龟(Popillia mutans)是木荷的主要传粉昆虫,中华蜜蜂和棉花弧丽金龟的访花高峰均发生在10:00—11:00,白星花金龟无明显的访花高峰。3种传粉昆虫体表均携带花粉,白星花金龟的携粉量显著高于棉花弧丽金龟(P0.001)和中华蜜蜂(P0.001),棉花弧丽金龟的携粉量显著高于中华蜜蜂(P0.001)。体表不同部位携粉量不同,胸部腹板和腹部腹板的携粉量高于其他部位,3种传粉昆虫的主要传粉部位均为胸腹部。中华蜜蜂每次访花的花粉移出数、柱头花粉沉降数及传粉效率均低于白星花金龟和棉花弧丽金龟,但不显著,而中华蜜蜂在访花频率上占有明显优势,分别为白星花金龟的29倍和棉花弧丽金龟的8.3倍;单花停留时间也显著低于白星花金龟和棉花弧丽金龟。综合各项特征表明,中华蜜蜂是木荷最有效的传粉者。     

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.     

琼花生殖器官结构及传粉昆虫的观察

为了探讨影响琼花Viburnum macrocephalum f.keteleeri有性生殖的因素,对其生殖生物学和传粉生物学进行了研究。研究内容主要包括琼花花部特征、生殖器官解剖结构、花粉活力、花粉胚珠比(P/O)、繁育系统、传粉形式、传粉昆虫和花粉管生长路径等。结果表明:(1)琼花聚伞花序由大型不孕花和小型可孕花组成,可孕花雌雄蕊发育正常,雄蕊5枚,雌蕊1枚,干型柱头,单子房,倒生胚珠;不孕花的雌雄蕊在发育早期正常,而在花期时退化。雄蕊退化表现为雄蕊消失、花丝缩短或消失或花药大小不一;而雌蕊退化表现为雌蕊缩小或柱头开裂;有时雌雄蕊也存在瓣化现象。(2)自然条件下,单花花粉活性在散粉4–5d后明显下降,居群花粉活力在4月25日–28日开始显著下降。(3)花粉胚珠比P/O值为12800–18700,属专性异交型;繁育系统为异株异花授粉,属虫媒传粉植物。(4)可孕花大量散粉时间为9:00–16:00,昆虫访花高峰时段为11:00–15:00,访花昆虫中以蝶类和蜂类为主,蝶类访花频率最高。(5)落置柱头的花粉萌发率较高,花粉粒授粉后1h左右开始萌发,花粉管从柱头乳突细胞的间隙穿入花柱,沿花柱中央引导组织生长,18h左右进入子房,20h左右从珠孔进入胚囊。讨论了琼花的不孕花形成、花部结构适应、花型演化趋势、低结实率和花粉管生长特点。     

异型花柱植物喀什补血草的传粉者功能群与花粉转移效率

异型花柱是受遗传控制的花柱多态现象, 被达尔文认为是植物通过在传粉者体表不同部位滞落花粉以促进型间花粉准确传递的一种适应。该现象虽已受到广泛关注, 但在一些花型变异较大且不稳定的传粉系统中, 不同传粉者对各花型繁殖所产生的影响仍知之甚少。该研究以分布于新疆天山南坡的一个有同长花柱共存的异型花柱植物喀什补血草(Limonium kaschgaricum)种群为研究对象, 对其花型构成及频率、传粉者及花粉转移效率等进行了调查分析。结果表明: 1)种群中除了存在雌/雄蕊长度交互对应的长(L)/短(S)花柱型花外, 还有雌/雄蕊同长的花(H型), 且各花型花的花冠口直径、花冠筒长及花粉量等参数间无差异, 但花粉纹饰和柱头乳突细胞形态具二型性。其中, H型花的花粉和柱头形态与L型花(或S型花)的一致。2)花型内和自花授粉均不亲和; 型间授粉时, 花粉和柱头形态不同的花型间亲和, 反之不亲和。3)种群内存在长/短吻两类传粉昆虫。在以短吻传粉者为主的盛花初、中期, L和H型花柱头上的异型花粉数均显著高于S型花的, 且L和S型花高位性器官间的异型花粉传递效率高于低位性器官间的; 而在以长吻传粉者为主的盛花后期, L和S型花的柱头间异型花粉数无显著差异, 且高/低位性器官间具有相同的异型花粉转移效率; 与传粉者出现时期相对应的、在花期不同阶段开放花的结实率也明显不同。4)长/短吻昆虫具明显不同的传粉功能, 短吻昆虫只能对L和H型花进行有效传粉, 且访花频率和型间花粉转移效率较低, 为低效传粉者; 而长吻昆虫对各花型均能有效传粉, 具高的访花频率和型间花粉转移效率, 为高效传粉者。因为长吻昆虫的阶段性出现所形成的不稳定传粉系统, 使低效的短吻昆虫可能会成为种群中花型变异的驱动力, 并使S型花受到更大的选择压力。H型花克服了柱头缩入的弊端, 可能会成为不稳定传粉系统下的一个替代花型而持续存在。     

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。     

濒危植物伯乐树传粉生物学特性研究

通过野外观察和室内测定等方式,对伯乐树的花部结构、访花昆虫及访花行为进行了观察,并利用石蜡切片法对开花期的雌蕊花柱及柱头进行显微观察,有关其传粉生物学特性的研究结果表明:伯乐树自然开花正常,其有效胚珠的比率为23.3%。伯乐树为虫媒植物,传粉者主要依赖黄胸木蜂,伯乐树的授粉方式可能属于居群内的花粉交流,石蜡切片显示柱头与花柱道相通,花柱道长,空隙很大,在一定程度上影响本物种的传粉。     

珍稀濒危植物小花木兰传粉生物学研究

基于定株观测,对小花木兰（Magnolia sieboldii）野生种群和人工栽培种的开花物候进行了研究。结果表明,小花木兰花期为5月中下旬至6月中下旬,花期20d左右,单花花期一般为5～7d,盛花期为6月1～5日;开花过程中花部表型变化明显,花粉在花开时不成熟,但雌蕊已经成熟,花开后雌蕊柄伸长,约20h后下柱头依然湿润,上柱头已萎缩变褐色,再过约40h下柱头也萎缩失去活力．此时花粉成熟,花药裂开;花的柱头接受到的花粉量由下往上依次减少。检测花粉活力、萌发率发现,小花木兰花粉在散发6d后基本失去活力和萌发能力。重力玻片法观测表明,小花木兰为虫媒传粉;花柱的授粉率不高,为65％;柱头上平均花粉量为3．5个;自然状态下结实率低,为13．5％,人工栽培为20．1％;外轮花被片全部去掉,结果率下降至11．3％;去掉雄蕊,结实率下降至10．7％;再辅以人工授粉,结果率上升至25．0％。小花木兰的访花昆虫种类较少,访花频率低,其中蜜蜂和一种蚜科（Aphididae）小昆虫对其传粉影响较大。同花期灯台附（Cornus controversa）等植物对其传粉产生昆虫竞争作用。环境因素如光照、温度、阴雨、海拔等均对其传粉产生一定影响。     

二型花柱植物金荞麦的繁殖生态学研究

通过野外实验观察和人工控制套袋等方法,从金养麦的花部基本特征、开花动态、花粉胚珠比(P/O)、花粉活力、柱头可授性、杂交指数、套袋实验及访花昆虫等方面对其繁殖生态学进行了研究.结果显示:(1)金荞麦在同一居群中同时具有长柱型(L-型)和短柱型(S-型)两种花型,短花柱花直径(7.25±0.11 mm)显著大于长花柱花直径(6.79±0.11 mm),长花柱花的柱头和花药高分别为3.39±0.04 mm和1.80±0.02 mm,短花柱花的柱头和花药高则分别为1.89±0.04 mm和3.19±0.06 rnm,表现出互补式雌雄异位的花部特征.(2)金荞麦8～10月开花,单花序的花期为15～23 d,单花的花期为1～3 d,开花进程中,内轮雄蕊先散粉,外轮雄蕊后散粉;S-型花的花粉活力高于L型花的花粉活力,但其单花花粉量低于L-型花,显示了两种花型的自我调控机制.(3)套袋实验表明,金荞麦自花自交不亲和,但具有型内和型间亲和性,L型和S-型花的P/O值分别为810±40.48和526.5±42.24,表明其繁育系统为兼性异交,需要传粉者,但L-型花更倾向异交.(4)金荞麦的主要访花昆虫为膜翅目(Hymenoptera)、双翅目(Diptera)、鳞翅目(Lepidoptera)、半翅目(Hemiptera)和蜻蜒目(Odonata) 19个科37种昆虫,其中,两种花型拥有12种共同的访花昆虫,L-型花访花昆虫14种,S-型花访花昆虫35种,表明S-型花对昆虫具有更强的吸引力.研究表明,金荞麦是典型的二型花柱植物,虽然具有严格的自交不亲和性,但显示一定的型内亲和性.     

珍稀濒危植物猬实的开花特性与传粉生物学研究

对迁地保护的珍稀濒危植物猬实的开花动态、花粉活力、柱头可授性、访花昆虫种类、访花频率、花粉与胚珠比(P/O)、异交指数(OCI)进行观测.结果表明：猬实的单花期约为7 d,种群花期约为14 d,花冠展开后2~3 h,花粉活力达到最高,约为90％,3 d后花粉基本不具活力.在整个单花期,柱头一直具可授性,花冠展开后第2天可授性最强.花粉与胚珠比(P/O)为(398．1±63．7),异交指数(OCI)为3,表明猬实是以异交为主,自交亲和的繁育系统,有时需要传粉者完成传粉过程.猬实的访花昆虫有10余种,以膜翅目蜜蜂科和隧蜂科昆虫为主,另外还有少量双翅目食蚜蝇科昆虫,偶见鳞翅目的柑橘凤蝶和长喙天蛾,其中以蜜蜂科蜜蜂属昆虫传粉效率最高.迁地保护的猬实未见结籽,其原因还有待于进一步研究.     

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。     

Removal and insertion of pollinia in flowers of Oxypetalum (Asclepiadaceae) in southeastern Brazil

Pollinarium removal and pollinium insertion of seven Oxypetalum species (O. alpinum var. alpinum, O. appendiculatum, O. banksii subsp. banksii, O. jacobinae, O. mexiae, O. pachyglossum and O. subriparium) were recorded in Vi?osa. Minas Gerais. They presented a tendency of one or two pollinarium removals and one pollinium insertion (single insertion), except O. appendiculatum. In this species, mainly, two pollinia of the same pollinarium were inserted per stigmatic chamber (double insertion), resulting exceptionally in 6-10 inserted pollinia in a flower, an unusual occurrence among the Asclepiadaceae. No association between removal and insertion was found, e.g., O. subriparium and O. banksii subsp. banksii had the highest pollinarium removal (1.78 and 1.45, respectively) and one of the lowest pollinium insertions (0.02 in both species), per flower. Oxypetalum mexiae showed the lowest pollinarium removal and pollinium insertion per flower (0.09 and 0.01, respectively) among the studied species and other Asclepiadaceae. Oxypetalum subriparium, O. banksii subsp. banksii and O. mexiae might be having reproductive limitations. Pollinarium removal and pollinium insertion per flower of the studied species varied from site to site, similarly to what was recorded for other Asclepiadaceae.     

Effects of nutrient contents and defense compounds on herbivory in reproductive organs and leaves of Iris gracilipes

Optimal defense theory (ODT) states that the plant tissue with the highest value to fitness will receive the most protection compared with other plant parts. ODT can be applied to the differences in defenses among floral organs, although most studies have concentrated on the comparison between leaves and flowers. Using Iris gracilipes, we investigated whether ODT is supported when primary and accessory floral organs and leaves are distinguished. We found that anthers and perianths tended to be attacked more severely than ovaries and leaves in the bud and flower stages and that anthers contained the highest nitrogen and phosphorus concentrations. Although ovaries were also found to contain high nitrogen and phosphorus concentrations, they were less severely attacked by herbivores than anthers, perhaps because ovaries contained the highest condensed tannins concentrations among the floral organs except for perianths in the flower stage. Thus, noting that the number of ovules is very much smaller than that of pollen grains, we concluded that ovaries are the most intensively protected, consistent with the prediction of ODT as applied to floral organs. ODT is applicable to the difference in defense allocation among floral organs.     

Oxypetalum banksii subsp. banksii: a taxon of Asclepiadaceae with an extragynoecial compitum

 The floral morphology of seven Oxypetalum species and, in particular, the spatial relationship between the five stigmatic chambers and two separate ovaries of their flowers with respect to transmission of the pollen tube are studied. In all species, except O. banksii subsp. banksii, floral morphology is similar to that in other Asclepiadeae, and the flowers pollinated with one pollinium develop only one follicle, which means compitum absence. In O. banksii subsp. banksii flowers, the secretory interstaminal tissue lines the inner walls of the stigmatic chambers as in the other species studied, but it also reaches the upper part of the inner surface of the filament tube, where it surrounds the styles, an unprecedented feature for Asclepiadaceae. This tissue secretes nectar and mucilage; the latter acts as transmitting medium for the growth of pollen tubes from pollinia inserted and hydrated in stigmatic chambers (“hyperstigmas”). Mucilage also functions as an extragynoecial compitum: in flowers pollinated with one pollinium both carpels develop into a follicle. Received August 28, 2001; accepted April 9, 2002 Published online: October 14, 2002 Addresses of the authors: Milene Faria Vieira (e-mail: mfvieira@mail.ufv.br), Departamento de Biologia Vegetal, Universidade Federal de Vi?osa, 36571-000, Vi?osa, Minas Gerais, Brazil. George John Shepherd, Departamento de Botanica, Instituto de Biologia, Universidade Estadual de Campinas, C.P. 6109, 13083-970, Campinas, S?o Paulo, Brazil.     

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.     

THE INTRAFLORAL PHENOLOGY OF STREPTANTHUS TORTUOSUS (BRASSICACEAE)

The sequence of floral events during anthesis was examined in Streptanthus tortuosus to determine the relationship between the male and female floral phases. The flowers are strongly protandrous. In the staminate phase, the anthers mature sequentially over a 3–4-day period. Because pollinators quickly remove pollen from the anthers, sequential anther maturation prolongs the male phase relative to what it would be if anthers did not mature sequentially. Pollen applied to the stigma during the staminate phase does not adhere readily and does not germinate. The length of the pistillate phase depends on pollinator activity, as pollination accelerates the abscission of floral parts. Unpollinated flowers remain pistillate for 3–4 days, during which time stigmatic receptivity declines gradually. In the field, 72% to 80% of flowers are staminate at any time, indicating that the staminate phase is three times longer than the pistillate phase when pollinators have access to the flowers. The consequences of the relative length of the floral phases and the schedule of stigmatic receptivity are discussed in terms of outcrossing mechanism, floral longevity, and sexual selection models.     

铁皮石斛花粉块结构及发育过程的解剖学特征研究北大核心CSCD

兰科植物的有性生殖特殊,每朵花只有1个花药,且花粉有聚集成块发育的特征。为了揭示铁皮石斛花粉块的发育特征,该研究以野生铁皮石斛不同时期的花药为材料,采用半薄切片和植物组织化学方法对其发育过程进行解剖学观察分析,并对成熟花粉块进行离体培养,观察花粉管的萌发状况。结果表明:(1)铁皮石斛花药壁由1层表皮细胞,2层药室内壁细胞,1层中层细胞和1层绒毡层细胞组成。开花时,绒毡层细胞退化,中层细胞没有退化,药室内壁细胞则形成纤维状细胞壁;药室中的小孢子母细胞没有明显的胼胝质壁结构。(2)小孢子发生属同时型,减数分裂后四分体小孢子不分散,以四合花粉状态发育,并进一步连接形成花粉块。(3)在小孢子发育中,孢粉素覆盖在整个花粉块表面形成花粉外壁,但花粉块内部的花粉没有花粉外壁结构;在花粉块表面的花粉外壁上未见花粉萌发孔。(4)在花粉离体萌发实验中,具有花粉外壁的花粉块表面花粉未见萌发,仅由花粉块内部的花粉萌发出花粉管。     

Reduced visitation to buzz‐pollinated Cyanella hyacinthoides in the presence of other pollen sources in the hyperdiverse Cape Floristic Region

Many plant species have floral morphologies that restrict access to floral resources, such as pollen or nectar, and only a subset of floral visitors can perform the handling behaviors required to extract restricted resources. Due to the time and energy required to extract resources from morphologically complex flowers, these plant species potentially compete for pollinators with co‐flowering plants that have more easily accessible resources. A widespread floral mechanism restricting access to pollen is the presence of tubular anthers that open through small pores or slits (poricidal anthers). Some bees have evolved the capacity to remove pollen from poricidal anthers using vibrations, giving rise to the phenomenon of buzz‐pollination. These bee vibrations that are produced for pollen extraction are presumably energetically costly, and to date, few studies have investigated whether buzz‐pollinated flowers may be at a disadvantage when competing for pollinators’ attention with plant species that present unrestricted pollen resources. Here, we studied Cyanella hyacinthoides (Tecophilaeaceae), a geophyte with poricidal anthers in the hyperdiverse Cape Floristic Region of South Africa, to assess how the composition and relative abundance of flowers with easily accessible pollen affect bee visitation to a buzz‐pollinated plant. We found that the number of pollinator species of C. hyacinthoides was not influenced by community composition. However, visitation rates to C. hyacinthoides were reduced when the relative abundances of flowers with more accessible resources were high. Visitation rates were strongly associated with petal color, showing that flower color is important in mediating these interactions. We conclude that buzz‐pollinated plants might be at a competitive disadvantage when many easily accessible pollen sources are available, particularly when competitor species share its floral signals.     

Floral biology and pollen viability of parental lines of onion hybrids

Parental lines (inbred and hybrid male-sterile lines and inbred pollinators) of prospective F, hybrids from the NVRS spring-sown bulb onion breeding programme were examined to study their floral biology and aspects of pollination under glasshouse conditions. Inbreeding depression occurred in both pollinator and male-sterile inbreds in terms of reduced flower numbers per umbel and a greater proportion of defective pistils. Male-sterile and male-fertile lines differed in their stigma and style characteristics and pattern of development. At flower opening, the styles of male-sterile lines were longer and the stigmatic knob became receptive relatively sooner. The inbred male-sterile line II/3 ms appeared to have a reduced receptive area on its stigmatic knob and remained receptive for a relatively shorter period. Individual lines differed for time of flowering, nectar content and its sugar concentration, and in their effectiveness as male and female parents as judged from pollen tube growth down the style in specific pairings. Assessment of pollen viability using the fluorescein diacetate test did not correlate with assessments based on pollen grain germination and subsequent growth down the style, perhaps as a result of differential receptivity in the inbred lines tested. These results are discussed in relation to future hybrid seed production.     

Sexual differences in reproductive characters and pollinator attractiveness in gynodioecious Glechoma longituba (Lamiaceae)

We quantified the differences in floral characters and attractiveness to flower visitors under natural conditions between the sexual types in the gynodioecious plant Glechoma longituba. We also manipulated flowers by altering corolla size or nectar volume, or by removing anthers, to examine the effect of these primary and secondary attractants (i.e. rewards and advertisements) on attractiveness. A change in corolla size and shape reduced visiting frequency and pollen load. Removal of anthers did not affect visiting rates, but significantly reduced pollination rates and stigmatic pollen load. A decrease in the nectar volume of a flower was associated with a reduction in handling time and pollen loads on stigmas. These results show that corolla size is an important advertisement to pollinators (particularly at greater distance), which associate hermaphrodite flowers with a larger corolla and a larger volume of nectar than female flowers. We found that artificial changes in population structure affected the behavior of pollinators as well as the pollination rates of flowers. We suggest that the pattern of distribution of hermaphrodite and female clones in a population may serve to avoid pollen limitation in a female clone or patch. This effect may ensure female reproductive success and allow for the maintenance of female individuals in natural populations of this gynodioecious plant.     

Horizontal orientation of zygomorphic flowers: significance for rain protection and pollen transfer

• Floral traits are recognized to have evolved under selection for abiotic and biotic factors. Complex zygomorphic flowers usually face horizontally. It has been proved that a horizontal orientation facilitates pollinator recognition and pollination efficiency, but its significance in adaptation to abiotic factors remains unknown.
• The floral orientation of Abelia × grandiflora naturally varies around horizontal (with an angle of ?30 to +33° between the floral main axis and the horizontal). We examined whether three different floral orientations affected flower thermal conditions, response to rain and pollination.
• Results showed that floral orientation had no effect on diurnal variations in flower temperature. The anthers of all three flower orientations were wetted by rainfall, but the inclined upward‐facing flowers contained significantly more rainwater. The horizontal flowers received significantly higher visitation by hawkmoths and had a higher stigmatic pollen load. In contrast, the upward flower orientation reduced pollination precision, while downward‐facing flowers had decreased pollinator attraction.
• This study indicates that horizontal flowers may have evolved as a trade‐off between rain protection and pollination. Zygomorphic flowers that deviate from a horizontal orientation may have lower fitness because of flower flooding by rainwater and decreased pollen transfer.