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。     

芍药的传粉生物学研究

对内蒙古赤峰市高格斯台罕乌拉自然保护区2个自然居群和内蒙古农校芍药园内2个栽培品种的芍药进行了连续2年的观察和研究表明,不同居群、不同品种的芍药花期相同,均为23~27d。芍药不存在无融合生殖,有微弱的自交性,不存在闭花授粉的现象。是以昆虫为媒介以异交为主要传粉方式的物种。不同花期的结实率之间没有显著差异(F=1.501)。芍药的结实率较低,野生居群平均只有23.27%的胚珠发育成种子。两个野生居群的传粉强度和传粉效率小于栽培品种单瓣花,结实率也低于栽培品种单瓣花。三个实验样地不同居群间的人工授粉结实率基本一致,且均高于其他组合。     

濒危植物丰都车前的繁育系统与传粉生物学研究

丰都车前为三峡库区特有珍稀濒危植物,目前仅在重庆市江津支坪镇涂家村附近大中坝上有一小种群分布。以野生种群和移栽种群为对象,通过野外观察和人工授粉实验对丰都车前传粉生物学及繁育系统进行研究。结果表明:(1)丰都车前单花花期约4 d,花序花期约8 d,种群花期约3个月;单花开花进程可分为6个时期:花蕾期、雌蕊伸长期、雄蕊初露期、花瓣展开期、花瓣反折期、枯萎凋谢期。(2)花瓣展开前,柱头先伸出花冠,达最高可受性,之后花瓣展开雄蕊才露出,而此时大部分柱头已枯萎失活丧失可受性,只有少部分具可受性。表现出雌雄异熟的特征,一定程度避免了自交的发生。(3)花粉活力始于雄蕊初露期,花瓣展开、花药开裂时花粉活力最高,至枯萎凋谢前均具有活力。花粉活力最高期避开了柱头可受性最佳期,仅有少部分重叠。(4)繁育系统检测显示,花粉/胚珠比为15373.29,杂交指数为3。结合套袋实验表明,丰都车前可能同时具有风媒和虫媒传粉机制,传粉昆虫为中华蜜蜂、食蚜蝇,但数量偏少,不存在无融合生殖,异交为主,部分自交亲和,有时需要传粉者。(5)开花时期避开了其分布地洪水淹没期,能够顺利完成开花授粉及座果。综上所述,丰都车前的花部特征、开花物候和繁育系统为适应分布地夏季洪水水淹胁迫提供了一定的生殖保障,促进了该种群的续存。     

黄花牛耳朵(苦苣苔科)的传粉生物学

通过野外观察和繁育系统的实验,对黄花牛耳朵(Chirita lutea Yan Liu et Y.G.wei)的传粉生物学进行了研究.结果表明,黄花牛耳朵的花期从7月初至8月底,单株花期约35～47 d,单花化期约6～10 d,花的开放无固定的时间.在花期内花粉活性约80%,柱头可授性约75%～90%.花粉/胚珠比率(P/O)为1 215.73±266.13.柱头在花药散粉时已生长至花筒口部,明显高于花药,便于接受异花花粉.黄花牛耳朵不存在无融合生殖,高度白交亲和,但较难发生自动的白花授粉,产生种子主要依靠传粉媒介.自然授粉的结实率明显低于人工授粉的结实率,存在传粉限制,蜜蜂(Apidae sp.)、方头泥蜂(Crabro sp.)、无垫蜂(Ameglla sp.)是主要的传粉者.     

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.     

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。     

珍稀濒危植物陕西羽叶报春的开花及传粉生物学研究

陕西羽叶报春为中国珍稀濒危植物,研究其开花物候、传粉者及访花行为、花粉活力、柱头可授性、繁育系统,以明确陕西羽叶报春的传粉特性及制濒机制。以勉县陕西羽叶报春野生居群为研究对象,通过野外观察和人工授粉实验首次对陕西羽叶报春的传粉生物学进行研究。结果表明:(1)陕西羽叶报春种群花期为2-4月中旬,单花花期一般为14~17 d;单花开花进程中花部形态有6个明显发育变化:全部被萼片包裹、花苞伸出并为深粉色、花朵打开为淡粉色、花冠为白色、花朵枯萎、花冠脱落花萼宿存。(2)陕西羽叶报春为异花授粉植物,蜜蜂属为其主要传粉者。(3)柱头在花粉未开裂前就具有可授性,可授性逐渐增强;花药在花瓣打开后的第3天才开始开裂、散粉,在第4天活力最高,为98.18%,此后逐渐减弱。(4)繁育系统检测显示,长、短柱型花的花粉/胚珠比分别为902.26和831.48,杂交指数为4;套袋实验显示,陕西羽叶报春异型交配可育,同型交配不可育。由此表明,陕西羽叶报春的繁育系统为异交型,自交不亲和,需要传粉者。综上所述,陕西羽叶报春传粉者种类少、自交不亲和、胚珠败育现象可能是导致陕西羽叶报春濒危的重要原因。     

山姜属植物花柱卷曲性传粉机制的研究

暂性雌雄异株是一种在两性花或雌雄同株植物个体上表现出的雄蕊和雌蕊成熟时间不同而形成的暂时性雄性阶段和雌性阶段不重叠的现象。这种现象减少了自交亲和的雌雄同株和两性花种类发生自交的可能性。作系统研究了花柱郑曲性传粉机制在西双版纳姜科山姜属（Alpinia Roxb.）植物中的普遍性,这一机制使得植物通过完全的暂性异株达到了异交的目的。这类植物的种群中具有两种独特的表现型,其差异央开花行为的不同：柱头下垂型个体在刚开花时其柱头向上反卷。位于已开裂的花药上方,到中午开始向下运动,下午则位于花药下方;柱头上举型个体的柱头在开花初期向花冠内卷曲,位于未开裂的花药和唇瓣之间中午开始向上运动,当柱头上举到花药上方后,花药开裂,花偻开始散发。两种个体的开花行为是同步的,通过两种柱头反向运动授粉方式（不同时间的自花和异花授粉）具有相同的结实率,但套代的花序几科不结实,两种表型在自然种群中的比例为1：1。     

野生翠雀传粉生物学特性研究

对从北京郊区引种的野生翠雀(Delphiniun grandiflorum L.)的开花习性进行了观察,对其花粉活力、柱头可授性,花粉/胚珠比(P/O),杂交指数(OCI)进行了检测,并进行了人工授粉实验。结果表明:(1)翠雀的花序为总状花序,具3~15朵花,花梗长1.5~3.8cm,单花期10d,花序花期15~18d。(2)翠雀花粉活力日变化呈先升高后降低的趋势;柱头在开花当天至开花后4d无可授性,在开花后的5d、6d有可授性。(3)翠雀的花粉胚珠比(P/O)为2 159.14,杂交指数OCI为5;人工授粉实验表明,翠雀在自然条件下能够结实,结实率为100%;存在自主自花授粉现象,部分自交亲和,结实受传粉者限制;不存在无融合生殖现象。(4)翠雀主要的访花昆虫有四条无垫蜂和短腹管蚜蝇。研究认为,翠雀的繁育系统为兼性异交或专性异交,部分自交亲和,需要传粉者。     

深山含笑传粉生物学研究

为明确深山含笑(Michelia maudiae)的传粉生物学特性,该文以西华师范大学校园内的深山含笑为研究材料,采取野外观察法记录了深山含笑的开花动态、访花者及访花频率;用游标卡尺法,测定了深山含笑的异交指数(OCI);用醋酸洋红法,测定了花粉胚珠比(P/O);用套袋和人工授粉法,测定了深山含笑的繁育系统。结果表明:(1)深山含笑为早春开花植物,其种群花期为2~3月,持续30d左右,单花花期5~6d;在单花开放初期,最内层的花瓣未绽开,是适应潮湿或多雨的环境而保护花药与柱头免受雨水的冲刷,提高雌雄性适合度,确保繁殖成功的一种策略;(2)深山含笑的有效传粉者为蜜蜂,访花时间集中在晴朗天气的中午;(3)深山含笑的异交指数(OCI)等于5,花粉胚珠比(P/O)为2 933±50;(4)深山含笑为雄蕊先熟,柱头可授性在开花第2~3天达到最高;(5)套袋和人工授粉表明深山含笑为兼性自交和异交授粉,其中异交授粉的座果率和结籽率显著高于自交授粉。早春开花的深山含笑,其传粉者为单一的蜜蜂,繁育系统为异交,部分自交亲和,结果为引种栽培、良种选育等提供依据更好地利用和保护此资源奠定基础。     

Study on the Flexistyly Pollination Mechanism in Alpinia Plants (Zingiberaceae)

Temporal dioecy, a flowering mechanism of synchronous alternation of pistillate and staminate phases in hermaphroditic (bisexual) flowers or monoecious individuals, decreases the probability of self-fertilization in self-compatible (SC) species. The authors report the prevalence of flexistyly in native species of Alpinia Roxb. in Xishuangbanna, Southwest China, resulted in dimorphism with two temporal dioecious floral phenotypes that facilitates out-crossing. Populations of ginger plants Alpinia have two specific phenotypes that differ in flowering behavior: (1)“cataflexistyle”individuals with the stigma held erect beyond dehiscent anther when anthesis begins in the morning and becoming decurved under the anther at noon; (2)“hyperflexistyle” individuals with the receptive stigma decurved under the indehiscent anther first and moving into a reflexed superior position above the anther as it begins to shed pollen at midday. The stigmatic movements in the two floral phenotypes, which occur in a ratio of 1∶1 in natural populations, are synchronous and pollination is effected only between floral forms. Field experiments indicate that species of Alpinia are self-compatible and insect dependent fertilization plants.     

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.     

Floral and pollinator behaviour of flexistylous Bornean ginger, Alpinia nieuwenhuizii (Zingiberaceae)

Flexistyly is a unique floral mechanism involving extreme curving of the style. It was first described in Chinese ginger (Amomum, Zingiberaceae). This is a pioneer report on flexistylous gingers of Malesia, where most species of this family grow. We observed the floral behaviour and flower visitors in Alpinia nieuwenhuizii Val., a Bornean endemic. Although the floral behaviour and effective pollinators (carpenter bees, Xylocopa) were similar between the Bornean species and the previously reported flexistylous Alpinia, the pollinator behaviour between them strikingly differed with regard to the visit frequency of the pollinators showing a bimodal pattern during the day. This was a better match for the floral behaviour of the flexistylous Alpinia. Some gender differentiation observed between the two types of morphs is also discussed in the present study.     

Mating system and stigmatic behaviour during flowering of Alpinia kwangsiensis (Zingiberaceae)

 Little is known about the breeding systems of tropical perennial ginger species. In this paper, we provide information about the breeding system of Alpinia kwangsiensis in Yunnan, Southwest China, specifically self-compatibility, mechanisms promoting outcrossing, and the visitation characteristics of pollinators. Populations of A. kwangsiensis have two specific phenotypes that differ in flowering behaviour: 1) “cataflexistyle” individuals in which the stigma is held erect above the dehiscent anther when anthesis begins in the morning and becomes decurved under the anther at noon and 2) “anaflexistyle” individuals in which the receptive stigma is decurved under the indehiscent anther first and moves into a reflexed superior position above the anther as it begins to shed pollen at mid-day. The stigmatic movements in the two floral phenotypes, which occur in a ratio of 1:1 in natural populations, are synchronous and correlate with the foraging behaviour of floral visitors; pollination is effected only between floral forms. Field experiments indicate that A. kwangsiensis is self-compatible and dependent upon insects for fertilization. This newly reported floral mechanism, which we have named “flexistyly,” adds to the repertoire of devices that have evolved in flowering plants to insure outcrossing. Received November 29, 2001 Accepted January 8, 2002     

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。     

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。     

Adaptive significance of flexistyly in Alpinia blepharocalyx (Zingiberaceae): a hand-pollination experiment

BACKGROUND AND AIMS: Flexistyly is a sexual dimorphism where there are two morphs that differ in the temporal expression of sexual function and also involve reciprocal movement of the stigmatic surface through a vertical axis during the flowering period. The adaptive significance of flexistyly has been interpreted as a floral mechanism for outcrossing, but it may also function to reduce sexual interference in which styles and stigmas impede the pollen export. Here these two explanations of flexistyly were tested in Alpinia blepharocalyx through a hand-pollination experiment. METHODS: Hand-pollinations were performed in two temporal morphs and consisted of two sequential pollination treatments, namely self-pollination in the morning and inter-morph pollination in the afternoon (treatment 1) or conversely inter-morph pollination in the morning and self-pollination in the afternoon (treatment 2), and two simultaneous self- and inter-morph cross-pollination treatments either in the morning (treatment 3) or in the afternoon (treatment 4). Seed paternity was then determined to assess relative success of self- versus cross-pollen using allozyme markers. KEY RESULTS: In the sequential pollination treatments, whether the stigmas of recipients are receptive in the morning is crucial to the success of the pollen deposited. When the cataflexistylous (protandrous) morph served as pollen recipient, early-arriving pollen in the morning can sire only a very small proportion (<15%) of seeds because the stigmas were then unreceptive. However, when the anaflexistylous (protogynous) morph served as pollen recipient, early pollen did gain a large competitive advantage over the late pollen, particularly when cross-pollen arrived first. Simultaneous self- and inter-morph cross-pollination indicated that outcross-pollen is more competitive than self-pollen on receptive stigmas. CONCLUSIONS: Differential maturing of male and female organs in Alpinia blepharocalyx is sufficient for selfing avoidance, obviating the need for style movements. Instead, the upward style curvature of the cataflexistylous morph in the morning and the anaflexistylous morph in the afternoon most likely represents a means of reducing interference with pollen export.     

山姜属中药草豆蔻和益智nrDNA ITS区序列的测定

中药草豆蔻、益智的原植物分别为山姜属草豆蔻（Ａｌｐｉｎｉａ ｈａｉｎａｎｅｎｓｉｓ Ｋ．Ｓｃｈｕｍ．）与益智（Ａ．ｏｘｙｐｈｙｌａ Ｍｉｑ．）。本文采用ＰＣＲ直接测序法,首次测定了它们的核糖体ＤＮＡ ＩＴＳ区序列。结果显示,两者序列长度（ＩＴＳ１＋ＩＴＳ２）分别为４０３ｂｐ与４０４ｂｐ,序列间具有２７个变异位点（包括５．８Ｓ编码区）。本研究为山姜属中药材的ＤＮＡ分子鉴定提供了必要的序列资料。