自交与异交象牙参属植物的蜜腺结构及其生物学意义

蜜腺是有花植物与传粉昆虫构建互惠关系的关键花部结构,解析不同繁殖特性物种间蜜腺结构的差异可为理解有花植物繁殖特性的演化提供理论依据。以传粉系统高度特化、异交实现有性生殖的早花象牙参（Roscoea cautleoides）和缺乏有效传粉者、主动自交实现有性生殖的无柄象牙参（R. schneideriana）为材料,通过野外测量2个物种蜜腺特性、超景深显微镜观察和石蜡切片染色,探究早花象牙参与无柄象牙参在蜜腺分布位置、外部形态及内部结构的异同。结果表明：2个物种均具有结构蜜腺,从外部形态来看,早花象牙参蜜腺体积较大、分泌糖浓度较高的花蜜,而无柄象牙参蜜腺体积较小、无花蜜分泌;从内部结构来看,早花象牙参蜜腺结构特化、各个组成部分有明显的区分,而无柄象牙参蜜腺出现一定程度的退化。该研究结果为揭示早花象牙参与无柄象牙参在花部特征、传粉机制及繁育系统的差异以及无柄象牙参蜜腺无花蜜分泌的可能机理提供了重要的形态学依据,也为深入地探讨植物与传粉动物间的协同进化关系以及理解蜜腺多样性的起源与维持机制奠定了科学基础。     

濒危植物翅果油树种群传粉生物学研究

对山西省南部平陆、乡宁和翼城3个翅果油树(Elaeagnus mollisDiels)种群分别进行了传粉生物学观察.结果表明:(1)翅果油树花期一般为8~12 d,单花花期3~4 d,结实率仅为0.42%;(2)访花者多属于膜翅目、鞘翅目、双翅目、鳞翅目昆虫,蜂类是有效的传粉者;(3)翅果油树依靠芳香气味、花蜜、花粉和适度紧缩的生殖枝吸引访花昆虫;(4)昆虫的访花活动主要集中在11:00~16:00,访花活动易受气象因素影响;(5)自然条件下,翅果油树平均每日每花被访问的频次为(1.93±1.64)次,人工放蜂可显著增加传粉强度;(6)交配系统属居群内的异花传粉,仍属于广义的自花传粉范畴.传粉昆虫少,传粉效率低是翅果油树致濒的重要因素之一.     

干热河谷麻疯树访花昆虫及主要传粉昆虫

膏桐为一种重要的油料作物,其种子可以提炼生物柴油,生物柴油在我国将成长为有一定规模的产业。为了增加膏桐结籽率,提高其产量,给膏桐规模化栽培方面提供传粉生物学的科学依据,于2006年对云南元江坝区半栽培膏桐(Jatropha curcas L.)居群的访花昆虫及主要传粉昆虫种类进行调查。膏桐开花示样醒目,具花香与蜜腺,花形态没有特化,这些特征使得它提供的酬物适合于不同的昆虫采食,其访花昆虫种类较丰富,共有35种访花昆虫,分属5目20科。综合访花昆虫的传粉数量,及传粉质量包括访花行为、访花频率与日活动规律等确定主要传粉昆虫,确定大头金蝇Chrysomya(Compsomyia)megacephala Fabricius、中华蜜蜂Apiscerana Fabricius、迁粉蝶(淡色型)Catopsilia pomona f.Crocale为该居群的主要传粉昆虫。3种传粉昆虫具有不同的访花习性和日活动规律。雌雄迁粉蝶日活动规律相似。访花者的组成受气象因素、开花示样及生境条件的影响。最后,探讨膏桐与访花昆虫的关系,以及主要传粉者的组成随时空变异而改变的规律。     

阿魏传粉昆虫及其访花行为的初步研究

对新疆阿魏访花昆虫种类、昆虫访花频次、昆虫访花时间及昆虫访花行为进行了初步研究。在盛花期观察中,(1)发现有5目、23科、45种昆虫进行了访花,其中传粉昆虫4科7种,均属蜜蜂总科;(2)各类昆虫访花频次在不同观察时间变化较大,传粉蜜蜂仅出现在其中一个观察日中;(3)不同种类昆虫在花上的停留时间差异很大,传粉昆虫停留时间很短,鞘翅目昆虫停留时间最长。进入始果期后,昆虫的访花行为是围绕花盘的蜜腺展开的。     

芭蕉科花蜜腺形态比较: 兼论其与传粉者的关系

对狭义芭蕉科3个属的代表性种芭蕉(Musa basjoo)、象腿蕉(Ensete glaucum)和地涌金莲(Musella lasiocarpa)的花蜜腺形态进行了比较研究。结果表明它们的蜜腺属于隔膜蜜腺。雌花的蜜腺着生于子房的上部, 胚珠的上方; 雄花蜜腺占据了整个败育子房的位置。蜜腺结构由许多腔道组成, 这些腔道在横切面上呈现出复杂的发散式迷宫状结构。这3种植物花蜜腺的栅栏状表皮细胞、维管束和蜜腺开口方式相似, 而从纵切面和横切面上观察其结构存在一些差异。PAS反应显示象腿蕉泌蜜组织中淀粉粒含量高于其他两个种; 芭蕉和象腿蕉的蜜腺腔里有许多纤维状物质存在。3种植物的传粉综合征多样化, 花序和花的特征(如花序下垂或直立、苞片的颜色、泌蜜量和泌蜜时间等)和传粉样式之间有密切关系。它们的蜜腺结构和传粉者行为之间没有明显的相关性, 但是胶质或水质的花蜜对传粉者的取食方式有一定影响。     

芭蕉科花蜜腺形态比较：兼论其与传粉者的关系（英文）

对狭义芭蕉科3个属的代表性种芭蕉(Musa basjoo)、象腿蕉(Ensete glaucum)和地涌金莲(Musella lasiocarpa)的花蜜腺形态进行了比较研究。结果表明它们的蜜腺属于隔膜蜜腺。雌花的蜜腺着生于子房的上部,胚珠的上方;雄花蜜腺占据了整个败育子房的位置。蜜腺结构由许多腔道组成,这些腔道在横切面上呈现出复杂的发散式迷宫状结构。这3种植物花蜜腺的栅栏状表皮细胞、维管束和蜜腺开口方式相似,而从纵切面和横切面上观察其结构存在一些差异。PAS反应显示象腿蕉泌蜜组织中淀粉粒含量高于其他两个种;芭蕉和象腿蕉的蜜腺腔里有许多纤维状物质存在。3种植物的传粉综合征多样化,花序和花的特征(如花序下垂或直立、苞片的颜色、泌蜜量和泌蜜时间等)和传粉样式之间有密切关系。它们的蜜腺结构和传粉者行为之间没有明显的相关性,但是胶质或水质的花蜜对传粉者的取食方式有一定影响。     

海南三七（姜科）的食源性欺骗传粉

有花植物为繁殖成功,进化出各种各样的花部特征来吸引传粉者,如为传粉者提供花蜜、花粉、栖息地等,然而在33科146属的被子植物中也存在着不提供任何报酬而欺骗昆虫为其传粉的现象。这种欺骗性传粉模式主要出现在高度进化的具有多样化传粉模式的兰科植物中。报道了在姜科植物中首次发现的食源性欺骗传粉模式。对姜科山柰属海南三七进行连续2年的传粉生物学观察和研究发现,海南三七的花在早上5:30～6:00之间开放,下午17:00～18:00左右闭合萎蔫,持续大约11～12h。开花过程中花粉活性与柱头可受性均保持较高水平(>90%)。花粉/胚珠比率(P/O)为82.20±47.89(n=20)。木蜂是其主要的访花和传粉昆虫,访花目的是吸取花蜜。海南三七虽有细长线形的蜜腺,但并不分泌花蜜作为传粉昆虫访花的报酬,采用食源性欺骗的方式欺骗木蜂为其传粉。繁育系统的研究表明广西弄化的海南三七居群主要是通过根茎进行无性繁殖。     

雀榕及其传粉昆虫传粉生态研究

雀榕（Ficus virens)花是单性的,雌雄同株,每一个隐头花序里同时具有雌花和雄花,雌花有长柱花和短柱花（瘿花）的分化,但二者均能结实和形成虫瘿,在生理上仍未分离,在这些花序中生长着4种小蜂总科的昆虫,其中榕小蜂科的2个种是雀榕的传粉省,尤以雀榕小蜂（Blastophagea sp.)为雀榕的主要传粉者,是共生体系的真正的互惠共生伙伴,冠缝榕小蜂（B.coronata)则是次要的传粉者也是主要传粉者的竞争者,食榕小蜂（Sycophila sp.)和刻腹小蜂（Ormyrus sp.)是上述2种榕小蜂的寄生者,但是刻腹小蜂的雄蜂也能参与出飞孔开掘,首次报道了我国雌雄同株的榕属植物和传粉昆虫的季节性环境胁迫下的共生关系,并探讨了共生体系维持的对策。     

柴黄姜的传粉生物学特性

对产自湖北武当山的柴黄姜（Dioscorea nipponica subsp．rosthornii Prain et Burkill）的花部特征和访花昆虫种类及访花行为、访花频率和携粉量进行了研究,并对柴黄姜的传粉方式、结实和种子萌发状况进行了检测。结果表明,柴黄姜具有虫媒花的典型花部特征,共观测到访花昆虫4目34种,其中传粉昆虫6种,隧蜂属（Halictus sp．）和地蜂属（Andrena sp．）的2种昆虫是柴黄姜的主要传粉昆虫。访花昆虫对雄株的访花高峰时段为11：00至14：00,对雌花没有明显的访花高峰时段。在自然条件下,柴黄姜的座果率为18．86％,结实率为7．12％,种子萌发率为64．76％。在武当山,柴黄姜种群扩展仍以有性繁殖为主,无性繁殖为重要补充。     

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。     

Specialized pollination in the African milkweed Xysmalobium orbiculare: a key role for floral scent in the attraction of spider-hunting wasps

Specialized pollination by prey-hunting wasps is poorly documented in rewarding plants. Furthermore, the mechanisms of achieving specialization are not clear since flowers typically produce exposed nectar and have no morphological adaptations (such as long spurs) to exclude non-pollinating visitors. We investigated the pollination of Xysmalobium orbiculare and explored the functional roles of floral scent and nectar in attracting pollinators and deterring nectar robbers. Floral visitor observations showed that this milkweed is visited almost exclusively by pompilid wasps in the genus Hemipepsis. These wasps were the only insects to carry pollinia, and a cage experiment confirmed their effectiveness in removing and inserting pollinia on flowers. Hand-pollinations showed that plants are genetically self-incompatible and thus reliant on pollinators for seed set. Palatability experiments with honeybees showed that nectar is distasteful to non-pollinating insects and is therefore likely to play a functional role in deterring nectar thieves. Choice experiments in the field showed that the wasp pollinators are attracted primarily by floral scent rather than visual cues. Analysis of spectral reflectance of flowers revealed that flowers are dull colored and are unlikely to stand out from the background vegetation. We conclude that X. orbiculare is specialized for pollination by spider-hunting wasps in the genus Hemipepsis and utilizes floral scent to selectively attract its pollinators and unpalatable nectar to deter non-pollinating visitors.     

Floral traits of mammal‐pollinated Mucuna macrocarpa (Fabaceae): Implications for generalist‐like pollination systems

Floral traits are adapted by plants to attract pollinators. Some of those plants that have different pollinators in different regions adapt to each pollinator in each region to maximize their pollination success. Mucuna macrocarpa (Fabaceae) limits the pollinators using its floral structure and is pollinated by different mammals in different regions. Here, we examine the relationships between floral traits of M. macrocarpa and the external morphology of mammalian pollinators in different regions of its distribution. Field surveys were conducted on Kyushu and Okinawajima Island in Japan, and in Taiwan, where the main pollinators are the Japanese macaque Macaca fuscata, Ryukyu flying fox Pteropus dasymallus, and red‐bellied squirrel Callosciurus erythraeus, respectively. We measured the floral shapes, nectar secretion patterns, sugar components, and external morphology of the pollinators. Results showed that floral shape was slightly different among regions and that flower sizes were not correlated with the external morphology of the pollinators. Volume and sugar rate of nectar were not significantly different among the three regions and did not change throughout the day in any of the regions. However, nectar concentration was higher in Kyushu than in the other two regions. These results suggest that the floral traits of M. macrocarpa are not adapted to each pollinator in each region. Although this plant limits the number of pollinators using its flower structure, it has not adapted to specific mammals and may attract several species of mammals. Such generalist‐like pollination system might have evolved in the Old World.     

The relative strength of different floral visitors driving floral evolution within a <Emphasis Type="Italic">Primula secundiflora</Emphasis> population

Floral visitor assemblages within plant populations are usually composed of different visitors, and the relative abundance of these visitors also varies. Therefore, identifying the relative strength of these floral visitors driving floral evolution within the population is an important step in predicting the evolutionary trajectory of floral traits. Using supplemental hand pollination and nectar-robbing exclusion treatments, we experimentally identified the relative strengths of legitimate pollinators (that visit flowers through the corolla tube entrance) and nectar robbers (that visit flowers by biting a hole in the corolla tube or using an existing hole) driving floral evolution within the Primula secundiflora population. We also estimated legitimate pollinator- and nectar robber-mediated selection separately for pin and thrum flowers. Both legitimate pollinators and nectar robbers mediated selection on pollination efficiency traits in P. secundiflora population. Legitimate pollinators mediated selection for wider corolla tubes, whereas nectar robbers mediated selection for longer corolla tubes. In addition, nectar robber-mediated selection on corolla tube length marginally varied between the pin and thrum flowers. Nectar robber mediated selection for longer corolla tube length in the pin flowers not in the thrum flowers. These results indicate that legitimate pollinators and nectar robbers within a population can drive differential evolutionary trajectories of floral traits.     

Pollinators and nectar robbers cause directional selection for large spur circle in Impatiens oxyanthera (Balsaminaceae)

Nectar spurs have an important role in floral evolution and plant–pollinator coadaptation. The flowers of some species possess spurs curving into a circle. However, it is unclear whether spur circle diameter is under direct selection pressure from different sources, such as pollinators and nectar robbers. In this study, we quantified selection on some floral traits, such as spur circle diameter in Impatiens oxyanthera (Balsaminaceae) using phenotypic selection analysis and compared the relative importance of pollinators and nectar robbers as selective agents using mediation analysis. The study showed that pollinators caused significant selection on corolla length, spur curvature and spur circle diameter while nectar robbers only imposed strong selection on spur circle diameter. Pollinators favored flowers with large corolla, curly spurs and large spur circle while nectar robbers preferred flowers with small spur circle. More pollinator visits resulted in higher female reproductive success, while robbery reduced female fitness. Conflicting selection on spur traits from pollinators and nectar robbers was not found. Mediation analysis showed that selection on floral traits through nectar robbing was stronger than selection through pollination. The results suggested that pollinators and nectar robbers jointly mediated the directional selection for large spur circle, and nectar robbers caused stronger selection than pollinators on floral traits.     

Altitude-related shift of relative abundance from insect to sunbird pollination in Elaeagnus umbellata (Elaeagnaceae)

The evolution of floral traits has been thought to be influenced by local, effective pollinators. However, little attention has been paid to the possibility that altitudinal variation in floral traits could be mediated by local pollinator functional groups, particularly a shift from bees to birds. Plant size, floral traits, pollinators and their pollination roles were investigated in the spring-flowering shrub Elaeagnus umbellata (Elaeagnaceae) at three altitudes (1160, 1676, and 2050 m) in Minshan, Sichuan Province, on the northern rim of the Hengduan Mountains, southwest China. Compared to lower altitudes, higher-altitude plants were smaller but the floral tubes were longer, with a larger volume of nectar of lower sugar concentration but with a greater proportion of sucrose. The visitation frequency of bees decreased with altitude, whereas the sunbirds did the opposite. Birds and bees foraged for nectar but not pollen, and birds deposited more pollen grains per visit relative to bees and least were syrphid flies. Excluding birds decreased seed set at high but not at mid- or low altitude. Our study of E. umbellata revealed an association between altitudinal variation in floral traits and a change in the relative abundance of the major pollinators with altitude from majority bees to majority sunbirds. Although abiotic factors also tend to vary with altitude and can affect floral traits, nectar properties of “pro-bird” pollination were observed at high altitude.     

MACROEVOLUTIONARY TESTS OF POLLINATION SYNDROMES: A REPLY TO FENSTER ET AL.

Studies of floral ecology and evolution are often centered on the idea that particular floral trait combinations, or syndromes, represent adaptations for particular pollinators. Despite the conceptual importance of pollination syndromes, few macroevolutionary studies have statistically examined the relationship between pollinators and floral traits. Using 15 species of Iochroma, Smith et al. applied phylogenetically structured correlation analyses to test the relationship between floral variation and pollination system, quantified in terms of the importance of major pollinator groups. This study revealed that pollinator shifts are tied to changes in nectar reward and floral display but are not significantly correlated with changes in corolla length or color, contrary to what might be predicted from classical pollination syndromes. Fenster et al. question these findings because our pollinator importance estimates included recently introduced honey bees. To address this concern, we recalculated importance values excluding honey bees and repeated the analyses. We found the same patterns as in our original study with significant correlations between pollinators and nectar reward and display. We conclude that phylogenetic approaches provide essential tools for testing macroevolutionary predictions of pollination syndromes and, by applying these approaches to other radiations, we can refine our understanding of the role of pollinators in floral diversification.     

The Role of Jasmonates in Floral Nectar Secretion

Plants produce nectar in their flowers as a reward for their pollinators and most of our crops depend on insect pollination, but little is known on the physiological control of nectar secretion. Jasmonates are well-known for their effects on senescence, the development and opening of flowers and on plant defences such as extrafloral nectar. Their role in floral nectar secretion has, however, not been explored so far. We investigated whether jasmonates have an influence on floral nectar secretion in oil-seed rape, Brassica napus. The floral tissues of this plant produced jasmonic acid (JA) endogenously, and JA concentrations peaked shortly before nectar secretion was highest. Exogenous application of JA to flowers induced nectar secretion, which was suppressed by treatment with phenidone, an inhibitor of JA synthesis. This effect could be reversed by additional application of JA. Jasmonoyl-isoleucine and its structural mimic coronalon also increased nectar secretion. Herbivory or addition of JA to the leaves did not have an effect on floral nectar secretion, demonstrating a functional separation of systemic defence signalling from reproductive nectar secretion. Jasmonates, which have been intensively studied in the context of herbivore defences and flower development, have a profound effect on floral nectar secretion and, thus, pollination efficiency in B. napus. Our results link floral nectar secretion to jasmonate signalling and thereby integrate the floral nectar secretion into the complex network of oxylipid-mediated developmental processes of plants.     

Floral nectary,nectar production dynamics,and floral reproductive isolation among closely related species of Pedicularis

Floral nectar is thought to be one of the most important rewards that attract pollinators in Pedicularis;however,few studies have examined variation of nectary structure and/or nectar secretion in the genus,particularly among closely related species. Here we investigated nectary morphology,nectar quality,and nectar production dynamics in flowers of Pedicularis section Cyathophora. We found a conical floral nectary at the base of the ovary in species of the rex-thamnophila clade. Stomata were found on the surface of the nectary,and copious starch grains were detected in the nectary tissues. In contrast,a semi-annular nectary was found in flowers of the species of the superba clade. Only a few starch grains were observed in tissues of the semi-annular nectary,and the nectar sugar concentration in these flowers was much lower than that in the flowers of the rexthamnophila clade. Our results indicate that the floral nectary has experienced considerable morphological,structural,and functional differentiation among closely related species of Pedicularis. This could have affected nectar production,leading to a shift of the pollination mode. Our results also imply that variation of the nectary morphology and nectar production may have played an important role in the speciation of sect. Cyathophora.     

Pollination Ecology and Effect of Nectar Removal in Macleania bullata (Ericaceae)1

The floral syndrome of Macleania bullataYeo (Ericaceae) reflects its adaptation to hummingbird pollination. Its flowers, however, are subject to high levels of nectar robbing. I examined the floral visitor assemblage of M. bullata in a tropical montane wet forest in southwestern Colombia, focusing on the behavior of the visitors. I also tested for the presence of nocturnal pollination and the effects of nectar removal on new nectar production. The principal floral visitors were the nectar robbing hummingbirds Ocreatus underwoodii (19.1% of visits) and Chlorostilbon mellisugus (18.9%). Only two species of long–billed hummingbirds visited the flowers of M. bullata as “legitimate” pollinators: Coeligena torquata (14.7% of visits) and Doryfera ludoviciae (14.3%). The remaining visits constituted nectar robbing by bees, butterflies, and other species of hummingbirds. Nocturnal pollination took place, although fruit set levels were 2.4 times higher when only diurnal pollination was allowed as opposed to exclusively nocturnal pollination. Nectar robbers removed floral nectar without pollinating the flower. Treatments of experimental nectar removal were carried out to examine if flowers synthesize more nectar after nectar removal. Nectar removal increased the total volume of nectar produced by each flower without affecting sugar concentration. Thus, nectar robbing can impose a high cost to the plants by forcing them to replace lost nectar.     

Floral biology of Eriope blanchetii (Lamiaceae) in coastal sand dunes of NE Brazil

Abstract The floral biology of the sand dune shrub Eriope blanchetii (Benth) Harley was investigated in a wild population located within an environment protection area near Salvador, NE Brazil. Inflorescence and flower development were monitored, and the reproductive biology was assessed. Inflorescences are terminal panicles, each bearing from one to 38 small, zygomorphic flowers. The duration of an inflorescence is about 2 months, with few new flowers each day. The rewards to visitors are pollen and nectar, the latter being produced in small quantities at the corolla base, near the stylopodium. Three sequential phases of flower development were identified, based on colour changes, pollen and nectar availability, and stigmatic receptivity. Anthesis takes place throughout the day within each plant, and flowers are strongly protandrous. Flowers are short‐lived, with individual variations of 7–30 h until corolla abscission. This variation is probably associated to a post‐pollinating response, which reduces the length of the female phase and could be advantageous for the economy of nectar production and for increased efficient pollination of the remaining flowers on a plant. The results of the pollination tests (apomixis, spontaneous and manual autogamy, manual cross‐pollination, and control) reinforced the role of pollinators for the reproductive success of E. blanchetii, given that the production of fruits from spontaneous pollination is unlikely to be resulting from protandry and given the morphological floral features. Taken together, the morphological and physiological aspects of E. blanchetii floral biology favour cross‐pollination, and it is likely that most of the fruits in nature are produced by outcrossing.