姜科、闭鞘姜科植物繁育系统与传粉生物学的研究进展

综述了姜科Zingiberaceae植物繁育系统和传粉生物学方面的研究及其进展。姜科有50余属1500多种,是一个泛热带分布的、动物传粉的单子叶植物大科。姜科植物中存在着雌花两性花异株、雄花两性花同株、花柱卷曲性、雄性先熟、自交不亲和等多种性表达方式和花部机制。姜科植物重要的传粉动物包括各种蜂类、天蛾、蝴蝶、鸟类等,不同的传粉动物对应不同的花部特征。在相对有限的研究中,姜科植物展现出了一些独特的传粉和繁育机制,在豆蔻属Amomum、山姜属Alpinia等植物中发现的花柱卷曲性被认为是植物界中一种独特的促进异交的行为机制,在对黄花大苞姜Caulokaempferia coenobialis的研究中发现了植物界中一种全新的花粉滑动自花传粉机制,这些新的研究和发现丰富了我们对姜科植物传粉和繁育系统多样性的认识。本文提出了今后姜科植物繁殖生物学研究的建议:对更多未知姜科植物类群的传粉和繁育系统进行研究;从系统发育的角度开展姜科不同类群植物的传粉和繁育系统的演化研究;对花柱卷曲性这一独特的性多态现象开展全面深入的研究。     

Reproductive biology of Alpinia blepharocalyx (Zingiberaceae): another example of flexistyly

The floral biology and breeding system of Alpinia blepharocalyx were studied in Yunnan province, southwest China. Our results indicate that populations of A. blepharocalyx have two floral morphs, which differ in flowering behaviour: the cataflexistylous morph in which the stigma is held erect above the dehiscent anther when anthesis begins in the morning and becomes curved under the anther at afternoon, and the anaflexistylous morph in which the receptive stigma is curved under the indehiscent anther first and moves into a reflexed superior position above the anther as it begins to shed pollen in the afternoon; the flowering period of both floral morphs is 12 hours; the stylar movements in the two floral morphs are synchronous, and they have similar traits to those found in other Alpinia species previously reported. Also, on average, cataflexistylous flowers are larger than anaflexistylous flowers, especially the labellum and corolla tube length; moreover, the P/O ratio of the two floral morphs is significantly different: the cataflexistylous morph has more pollen grains and fewer ovules than the anaflexistylous morph; the evolutionary significance of this comparison is discussed. Cataflexistylous flowers were observed to produce a lower concentration of nectar than that of anaflexistylous flowers before 11:00 a.m., but they then have higher nectar concentration; cataflexistylous flowers always have more nectar than anaflexistylous flowers. Flowers of A. blepharocalyx were visited by fewer insects at a lower frequency, including honey bees (Apis cerana cerana) and two species of carpenter bees (Xylocopa spp.). No significant differences were found in the fruit set among the hand-pollination treatments regardless of selfing (geitonogamy and autogamy) or crossing (intermorph and intramorph); but all hand-pollination treatments exhibited much higher fruit set than the controls; meanwhile, no fruit set occurred in the unpollinated bagged plants. Pollen tube growth was examined using fluorescence microscopy following self, intramorph, and intermorph hand pollinations at different times of the day; the pollen tubes of both floral morphs have the same growth rate and the capability to reach the ovary. Both the results of pollen tube growth and fruit sets under different pollination conditions show that A. blepharocalyx is self-compatible and dependent upon insects for pollination. The distinctive flexistyly mechanism of A. blepharocalyx is likely a floral mechanism that promotes effective intermorph pollen transfer.     

山地五月茶的蝇类传粉研究

观测了山地五月茶Antidesma montanum的开花物候、开花动态、访花者种类和访花行为,并对其繁育系统、花粉组织化学、花粉胚珠比（P／O）、花粉活力进行了检测。结果表明：山地五月茶是雌雄异株植物;其总状花序花期可长达7天;雄花／雌花花序数目为140．33±27．79／208．33±33．65（n=6）,雌雄花的颜色为很淡的黄绿色小型花;雄花单花花期为2天,花药开裂当天有活力,花粉为非淀粉型,花粉胚珠比为3333．33±607．18;单花花蜜量可达0．34±0．03μL,花蜜含糖量3．69％±0．30％。花的结构和开花式样适合蝇类传粉。芳香的气味是吸引蝇类的直接物质：花蜜是传粉者的报酬。主要的传粉昆虫为双翅目Diptera丽蝇科Calliphoridae的Chrysomya megacephala、Chrysomyasp．、寄蝇科Tachinidae的Drino sp．和蝇科Muscidae的Spilogona sp．、Mitroplatia sp．。套网不授粉的处理不结实,表明山地五月茶不存在无融合生殖,人工辅助授粉的坐果率（39．1％）略高于自然坐果率（36．7％）,二者无显著差异,表明其坐果率主要受自身资源分配限制。还讨论了蝇类传粉与雌雄异株性系统以及蝇类传粉与热带林中小型黄绿色花植物的相关性。     

Ypsilorchis and Ypsilorchidinae,a new genus and a new subtribe of Orchidaceae

A new orchid genus, Ypsilorchis Z. J. Liu, S. C. Chen & L. J. Chen, is established based on Ypsilor-chis fissipetala (Finet) Z. J. Liu, S. C. Chen & L. J. Chen (basionym: Liparis fissipetala Finet). The new genus differs from Liparis and its allies by having two granular-waxy pollinia each with a somewhat elastic caudicle, deeply bilobed petals and strongly crisped-margined leaves with an apical awn to 1 mm long. These features are an indication of its distant relation to the members of the subtribe Liparidinae, and thus a new subtribe, Ypsilor-chidinae Z. J. Liu, S. C. Chen & L. J. Chen, is proposed.     

A new species of Selaginella from Hainan Island, China, with a key to the Hainan species

Selaginella hainanensis X.C.Zhang & Noot. sp. nov. , a new species from Hainan Island, South China, is described and illustrated. Diagnostic notes and a key to the species distributed in Hainan are also provided.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 323–327.     

An example of flexistyly in a wild cardamom species (Amomum maximum (Zingiberaceae))

We studied the floral and pollination biology of Amomum maximum. Our observations suggest that flexistyly is present in this species, and stigmatic behavior is similar as reported for Alpinia species. Two phenotypes occur in a ratio of 1:1 in natural populations. No fruit set occurred in the unpollinated bagged plants. Significant differences in seed set between selfing and crossing manipulations demonstrate that inbreeding depression or partial self-incompatibility is apparent in this species. Despite the resemblance in floral behavior, there are still some differences between the previous studies and ours: with special floral design, the body size of legitimate visitors ranged wider in Amomum maximum than in Alpinia species; the presence of anther appendage is suggested to prevent competing and delayed selfing. This is the first report of flexistyly in a wild cardamom species, which is the second largest group in ginger family (Zingiberaceae).     

Reproductive biology and pollination ecology of Triplaris gardneriana (Polygonaceae): a case of ambophily in the Brazilian Chaco

• Triplaris gardneriana (Polygonaceae) is a dioecious pioneer tree reported as insect‐pollinated, despite possessing traits related to anemophily. Here, we analyse the possible roles of insects and wind on the pollination of this species to establish whether the species is ambophilous.
• We carried out observations of floral biology, as well as on the frequency and behaviour of pollinators visiting flowers in a population of T. gardneriana in the Chaco vegetation of Brazil. We conducted experimental pollinations to determine the maternal fertility of female plants and whether they were pollen‐limited, and we also conducted aerobiological experiments to provide evidence of how environmental factors influence atmospheric pollen dispersal.
• The population comprised an area of approximately 152.000 m² and was composed of 603 female and 426 male plants (sex ratio = 0.59:0.41). We observed 48 species of insects visiting flowers of T. gardneriana, of which the bees Scaptotrigona depilis and Apis mellifera scutellata were the most effective pollinators. We recorded pollen grains dispersed by wind on 74% of the glass slides placed on females, located at different distances (1–10 m) from male plants.
• Airborne pollen concentration was negatively correlated with relative humidity and positively correlated with temperature. Our observations and experimental results provide the first evidence that T. gardneriana is an ambophilous species, with pollen dispersal resulting from both animal and wind pollination. This mixed pollination strategy may be adaptive in T. gardneriana providing reproductive assurance during colonisation of sites with different biotic and abiotic conditions.

     

A preliminary study on pollination biology of Omphalogramma souliei Franch. (Primulaceae), a species endemic to China

The pollination biology of Omphalogramma souliei Franch., a species endemic to southwestern China, was investigated. Floral phenology, flower visitors, pollen/ovule ratio, attractants and rewards to the visitors were observed, measured, and recorded. Bagging experiments to exclude pollinators were carried out in the wild for two years. Our results revealed five important aspects of the reproductive biology of Omphalogramma souliei. 1) The pollen-ovule ratio was 1748±233. The breeding system was self-compatible, with facultative xenogamy. 2) The pollination syndrome is entomophily, and this species could not be pollinated by wind if the pollinators were unavailable. 3) Six insect species were observed visiting the flowers of Omphalogramma souliei in the wild, of them, three species of hymenoptera, Lasioglossum sp., Heriades parvula Cockerell and Micrapis florae Fabricius, are the principal floral visitors and effective pollinators. 4) The visual attractants to the visitors are floral color and shape, the large yellow anthers, and the rewards for visitors are pollen and nectar. 5) Cleistogamy may also occur, since the anthers of some flowers dehisced before opening of the corolla. The results of floral biology and pollination characters suggest that xenogamy predominated and autogamy played an assistant role in the evolution of reproduction and breeding system of O. souliei.     

Comparative pollination biology in two sympatric varieties of Cypripedium macranthos (Orchidaceae) on Rebun Island,Hokkaido, Japan

Cypripedium macranthos sensu lato typically has purple‐pink flowers with no nectar and harvestable pollen. On Rebun Island, Hokkaido, Japan, purple‐pink‐flowered C. macranthos var. macranthos individuals rarely grow among numerous pale‐cream‐flowered C. macranthos var. rebunense plants. In both varieties, flower size is similar, their flowering periods completely overlap, and they share the same pollinator (Bombus pseudobaicalensis). However, in only one of 12 years from 2001 to 2012 did var. macranthos attain an annual fruit‐set ratio (an estimate of pollination success) higher than that of sympatric var. rebunense plants. These findings strongly suggest that in C. macranthos on Rebun Island, flower color results in the differential pollination success, because the pollinator prefers pale‐cream Cypripedium flowers and/or avoids purple‐pink ones, thereby producing pollinator‐mediated selection favoring pale‐cream flowers.     

海南岛青冈属(壳斗科)一新种

描述海南岛昌化岭青冈一新种。     

Hydrophilous pollination and breeding system evolution in seagrasses: a phylogenetic approach to the evolutionary ecology of the Cymodoceaceae

COX, P. A. & HUMPHRIES, C. J. 1992. Hydrophilous pollination and breeding system evolution in seagrasses: a phylogenetic approach to the evolutionary ecology of the Cymodoceaceae. A phylogenetic analysis of seagrasses of the Cymodoceaceae shows the Cymodoceaceae to be monophyletic and Posidoniaceae to be their sister group. Information on the pollination ecologies and breeding systems of the various genera of the Cymodoceaceae were plotted onto the consensus tree obtained for the group. From this analysis, it is suggested that the clade composed of the Zosteraceae, Posidoniaceae and Cymodoceaceae evolved from a freshwater hydrophilous ancestor that developed submarine pollination and filiform pollen in association with invasion of the marine environment. Dioecism and surface pollination appear to have evolved in the progenitor of the Cymodoceaceae, and hence the seagrasses of the Cymodoceaceae are dioecious due to common descent rather than to convergent evolutionary processes in extant genera.     

肋果沙棘(Hippophae neurocarpa S.W.Liu et T.N.He)自然种群的风媒传粉特征

野外调查了青藏高原特有植物肋果沙棘的花期物候、开花式样以及风媒传粉特征,对其花粉生活力、柱头可授性及单花花粉量进行了检测.结果表明,肋果沙棘有许多适应风媒传粉的特征:雌雄异株,花小,无花冠,先叶开放,雄花萼片纵裂形成对流的"风洞",花粉近球形,单花花粉量大;雌花柱头细长捻状,伸出萼筒,授粉面大,密布乳突.花粉生活力可以持续16.5 d,柱头可授性为10～20d.肋果沙棘传粉多发生在昼夜温差大的5月中下旬,在多变的气候条件下,种群花期可持续约12～25d,但在晴朗的天气条件下具有爆发性的集中开花机制,是对恶劣条件的适应.重力玻片法检测表明,离花粉源15m左右远的地点花粉密度最大,135m仍能接收到花粉,夜间接收到的花粉数量也很可观,这些研究结果表明,该植物集中在有利的晴朗天气昼夜进行长距离的风媒散粉,能较好地适应青藏高原的复杂环境.     

海南羊山湿地的传粉网络及其季节动态

传粉网络是植物和传粉者之间形成的网状相互作用关系,为理解群落物种多样性形成与维持机制提供了全新的视角。湿地是典型的群落交错区,环境异质性与物种多样性都很高,传粉网络可能比草地和森林等生态系统具有更复杂的结构。该研究针对海南岛海口市南郊的羊山湿地,比较4个样地在旱季(5月)与雨季(8月)的传粉网络及其动态变化,揭示湿地生态系统的传粉网络结构特征以及在干湿季的变化规律。结果表明,羊山湿地传粉网络共有71种开花的植物, 131种传粉者,传粉网络呈现低连接度、高嵌套度、中等网络特化程度的结构特征。在季节动态方面, 4个样地旱季的植物与传粉者种类高于雨季;而传粉网络的连接度、嵌套度与网络特化程度没有明显的季节差异。白花鬼针草和水角等多个物种可同时在雨季和旱季开花,使得植物-传粉者的种间关系虽然存在季节变化,但传粉网络在旱季与雨季间的动态变化不大。总体而言,羊山湿地物种多样性较高,边缘效应较明显,传粉网络结构较稳定。     

深圳香荚兰,首次发现于华南深圳的兰科新种

对兰科新种深圳香荚兰Vanilla shenzhenicaZ．J．Liu＆S．C．Chen作了描述与绘图。此新种为深圳发现的第一个兰科新种,与台湾香荚兰V。somai Hayata有亲缘关系。但是,本新种花序具4花,花较大,不完全开放;唇瓣不裂,紫红色,基部与蕊柱合生长度达3/4,刷状附属物位于唇盘的上部,甚易区别于台湾香荚兰。     

Insect pollination of Musella (Musaceae), a monotypic genus endemic to Yunnan, China

 The pollination biology of Musella lasiocarpa (Franch.) C. Y. Wu ex H. W. Li, a monotypic genus in the banana family (Musaceae) endemic to southwestern China, was investigated. Floral phenology, nectar production, and flower visitors were observed, measured, and recorded. Bagging experiments and comparative breeding system studies were carried out at different study sites and in different seasons. Our results revealed four important aspects of the reproductive biology of Musella. 1) This species blooms year round, but most flowering is concentrated from February to August. The longevity of female flowers is greater than that of male flowers and opening of both types of flowers occurs at anytime during the day, but is concentrated in the morning. 2) Although nectar is produced uniformly while flowers are open, the nectar volume and rate of female flowers are significantly higher than that of male flowers. 3) Insects, such as bumblebees (Bombus eximius and B. montivolans), honeybees (Apis cerana and A. florea), and wasps (Vespa mandarinia) are the primary floral visitors and show a preference for female flowers. 4) Both fruit and seed set are pollinator-limited at both study sites. A comparison of the floral biology and pollination characteristics among the three genera in the banana family suggests that insect pollination has played an important role in the reproductive isolation and breeding system evolution of Musella. Received February 14, 2002; accepted March 26, 2002 Published online: November 22, 2002 Addresses of authors: Aizhong Liu, W. John Kress (correspondence, e-mail: kress.john@nmnh.si.edu), Botany, MRC-166, United States National Herbarium, National Museum of Natural History, Smithsonian Institution, PO Box 37012, Washington DC 20013-7012, USA. Hong Wang, De-Zhu Li (dzl@public.km.yn.cn), Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650204, China.     

广东地区外来种五爪金龙的传粉生物学研究

五爪金龙(Ipomoea cairica)是一种匍匐或攀援生长的入侵杂草。2006–2007年作者在广东地区开展了对该入侵种的传粉生物学研究。五爪金龙在广东全年均可开花, 花序为聚伞花序, 花冠为紫色或蓝紫色喇叭状。开花时间在早晨4: 30–5: 20之间, 下午17: 40闭合, 持续大约12 h。每朵花平均分泌花蜜量为5.82 ± 2.47 μL (n = 10), 花蜜平均含糖量为41.03 ± 6.52%, 氨基酸含量为0.042 mg/mL。开花过程中花粉活性和柱头可受性均保持较高水平(>90%)。花粉/胚珠比率(P/O)为4,778.18 ± 264.58 (n = 10)。木蜂(Xylocopa sp.)是其有效的传粉昆虫, 蝇类和蝶类为随机访问者。开花过程中柱头与花药相接触, 有利于自花授粉。然而, 繁育系统的研究表明五爪金龙是自交不亲和植物, 其种子仅能通过异花授粉获得。     

国家二级保护植物翅果油树传粉生物学的初步研究

对翅果油树进行了传粉生物学的野外观察和实验表明：（1）翅果油树的花具有自花传粉的结构特征,同时具有蜜腺,又具备异花传粉的特征。传粉者主要是一种家养的蜜蜂。（2）翅果油树花形态结构与传粉者的形态和传粉行为非常协调和适应,昆虫的传粉部位主要是头前部和胸部。（3）蜜腺的产蜜量大,开花后1～2天蜜腺分泌量最大。对传粉昆虫竞争中具明显优势。昆虫的访花频率明显高于同花期的其他植物。（4）翅果油树没有融合生殖现象。