簇枝补血草花的二态性及繁育系统研究

白花丹科被列为异型花柱较多的科,该文以补血草属簇枝补血草为材料,通过对野外居群的花部特征、花期物候和繁育系统等研究,探讨花部形态多态性及其对生殖的影响。簇枝补血草具有两种类型植株,具玉米状柱头和A型花粉的植株称为C型植株;具乳头状柱头和B型花粉的植株称为P型植株,且A/B型花粉的大小、外壁纹饰不同。单花发育过程中两型植株的花萼、雌蕊大小差异明显,随着花的成熟差异逐渐消失。两型植株都为雌蕊高于雄蕊,柱头探出式雌雄异位;C型植株的花期较P型植株长约2 d。一天当中,A型花粉活力高峰期为13:00,B型花粉在11:00和15:00都表现为高峰期;玉米状柱头可授性最强时期在13:00～15:00,乳头状柱头则在15:00。两型花粉可育性在盛花期最高,终花期时B型花粉可育性明显高于A型。荧光染色观察发现花粉萌发和花粉管的生长速度都为:异型授粉＞同型授粉,B型花粉在乳头状柱头上不能完成受精。此外,人工授粉实验也表明异型授粉结实率显著高于同型,P型植株经同型授粉结实现象并不明显,存在少量无融合生殖。研究表明簇枝补血草花部具二态性,繁育系统为兼性异交,其中乳头状柱头对A型花粉亲和,对B型花粉不亲和,但玉米状柱头却对A/B型花粉都亲和,这打破了该科的异型自交不亲和的例证。     

Floral development in the androdioecious tree Tapiscia sinensis: Implications for the evolution to androdioecy

The evolutionary pathway between hermaphroditism and dioecy (females and males in a single population) draws widespread interests, and androdioecy (bisexuals and males in a single population) is rarely achieved as an intermediate state between the two breeding systems. Flower bud differentiations in the pistils of hermaphrodites and the pistillodes of males in androdioecious Tapiscia sinensis Oliv. are investigated by routine paraffin section technology, light microscopy, and scanning electron microscopy. A phylogenetic approach is used to analyze the origin of androdioecy. In T. sinensis, hermaphroditic flowers (HF) and male flowers (MF) experienced a similar development pattern in early flower bud differentiation, including the initiation of tepals and stamens. However, the carpel differentiation of MF and HF proceed in different patterns. In HF, the central zone bulges out and produces a ring meristem on which two to three carpel primordia emerge, which eventually developed into a normal pistil with a stigma, a style, and an ovary. However, in most MF, vestigial pistils are stem‐like (type I), and very few have an empty ovary (type II) or a sterile ovule (type III). Moreover, the evolution of sexual systems within the Huerteales indicates that hermaphroditism is the primitive character of T. sinensis. Tapiscia sinensis shows different degrees of reduction between male flowers and bisexual ones in the evolution to dioecy. Functional androdioecy originated from a hermaphroditic ancestor in T. sinensis and, as an intermediate sexual system, involves evolution from hermaphrodites to dioecy.     

黑河中游和烟台海滨中国柽柳的传粉生态学研究

中国柽柳(Tamarix chinensis)是干旱及半干旱地区的重要建群物种.为便于其有效利用和引种栽培,对黑河中游和烟台海滨的中国柽柳的开花动态、花粉活力、柱头可授性、访花者种类、花粉-胚珠比(P/O)、杂交指数(OCI)及座果率进行观测,结果表明:黑河中游和烟台海滨柽柳的花期和柱头可授性相似,花序花期为2-5天,开花当天柱头可授活性最大,之后逐渐下降,第4天柱头无可授性;花粉活力和访花者种类差别较大,黑河柽柳开花第1天花粉活力为(62±4.31)％,最高为(82.12±7.23)％,4天后花粉完全失活.烟台柽柳的花粉活力在开花后第1天最高为(89.13±7.23)％,第4天花粉活力下降为(11±1.12)％.黑河中游和烟台海滨柽柳的访花者均以膜翅目昆虫为主,黑河柽柳以黑带斑沙蜂(Bembix melanura)、卡切叶蜂[Megechile (Amegachile) kagiana]和黑带食蚜蝇(Episyrphus balteatus)等为主要访花者;烟台柽柳以姬蜂(Ichneumon sp.)、大灰食蚜蝇(Metasyrphus corollae)和灰地种蝇(Delia platura)等为主要访花者.二者P/O值、OCI及座果率差异不显著,黑河和烟台柽柳的P/O值分别为373.6±71.9和382.1±86.2;最高座果率分别为(89.72±7.76)％和(93.17±8.21)％;OCI均为3.上述研究结果显示,黑河中游和烟台海滨柽柳均为兼性异交的繁育系统.     

珍稀濒危植物瑶山苣苔开花生物学及繁育系统研究

对仅分布于广西金秀老山自然保护区的国家Ⅰ级保护植物瑶山苣苔进行了开花生物学和繁育系统研究.结果显示:(1)瑶山苣苔花期从8月底至11月初,开花无固定时间.单花开放过程可分为萌动、露白、盛开、凋落4个阶段,且花朵具明显的增大再生长现象.(2)单花、花序、单株水平上的开花物候都表现出开花不同步性,单花花期约5~14 d,单花序花期约11~20 d,单株花期约8~20 d.花粉活力在散粉后6 d内相对较高,但花粉在野外通常于散粉3 d内被昆虫啃食完.(3)柱头在花药散粉时明显高于花药,便于接受异花花粉,柱头在散粉后第4天具可授性,可持续5~6 d.(4)瑶山苣苔杂交指数(OCI)为5,花粉-胚珠比(P/O)为379.64±145.61,繁育系统为兼性异交,自交亲和,传粉过程需要传粉媒介.研究表明,不稳定的传粉环境可能是该物种至濒的主要生殖生物学原因,自发自交"是其在开花期间对不稳定传粉环境的一种适应.     

Growth and Reproduction of Double-Ended Pipefish, Syngnathoides biaculeatus, in Moreton Bay, Queensland, Australia

Life-history characteristics of the double-ended pipefish, Syngnathoides biaculeatus (Bloch), were investigated to determine growth rate, degree of sexual dimorphism, size at maturity, and reproductive biology. Growth rates of wild juveniles and adults calculated from monthly progression of length-frequency modes ranged from 0.8mmd^–1 (fish lengths 120–145mm standard length (SL)) in summer to 0.2mmd^–1 in winter (185–200mm SL). Growth of laboratory-reared juveniles up to 63d old was greater, ranging from 0.8 to 2.3mmd^-1. The von Bertalanffy growth constant K was estimated at 0.0076d^{- 1}, or 2.8year^–1. Morphological differentiation between the sexes based upon abdominal pattern was possible for fish larger than 120mm SL, with females possessing a zigzag pattern on the abdomen. The association between this pattern and sex was confirmed by histological gonad analysis. Males were significantly longer than females during four of seven seasons examined, and a 1:1 sex ratio was determined for all seasons except autumn when the ratio was female biased. The breeding season was marked by the appearance of pregnant males between October and April, and during courtship both species exhibited increased pigmentation. The minimum paternal size at maturity was 185mm, the maximum length recorded 260mm. Clutch size ranged between 60 and 200 eggs, with a mean of 153. Ovaries had a sequential pattern of egg development, resulting in egg batches that approximated the number of eggs carried by brooding males. Additionally, all eggs in a brood were at the same developmental stage. This suggests that one female provides all of the eggs for one male per breeding event in a monogamous mating system.     

不同香水月季染色体倍性鉴定及其繁育系统分析

【目的】香水月季（Rosa odorata）是一类重要的蔷薇属植物种质资源,具有较高的赏价值及利用价值,为了促进香水月季种质资源的开发和利用,对不同香水月季染色体倍性、花部综合特征和繁育系统特性进行研究。【方法】以9种香水月季为材料,采用流式细胞仪、杂交指数检测等对不同香水月季的染色体倍性、花粉活力、柱头可授性、花部特征等进行观察及测定,并对繁育系统进行分析。【结果】9种香水月季均为二倍体,其花为两性花,花色明亮,多重瓣或半重瓣,其中4种香水月季雌雄蕊异熟,同时,9种香水月季变种的花粉活力普遍偏低,花粉活力最高仅为35.47%,所有香水月季变种杂交指数（OCI）均≥4,其花粉量-胚珠比(P/O）值数值范围为6 709~75 521。【结论】结合杂交指数（OCI）和花粉量-胚珠比（P/O）值指示9种香水月季的繁育系统类型均以异交为主,且其繁育系统受花粉粒数、花瓣数等多个因素影响。     

Reproductive biology and pollinators in two enantiostylous Qualea species (Vochysiaceae) in the Brazilian Cerrado

• Enantiostyly is a floral polymorphism in which two floral forms in the same species differ in deflection of the stigma to right or left position. In monomorphic enantiostylous plants, flowers of the two morphs occur within the same individual, usually in the same proportion. In self‐compatible species the function of monomorphic enantiostyly is proposed to increase outcrossing rates and offer a reproductive advantage under pollination limitation. Enantiostylous species are usually self‐compatible and show heteranthery, with poricide anthers and pollen as pollinator reward; however, there are families, such as Vochysiaceae, that have different characteristics.
• We analysed the reproductive system and pollination biology of Qualea parviflora and Q. multiflora, two enantiostylous species from the Brazilian Cerrado that have specific morphological and physiological traits. For this, we characterized flower traits, performed hand pollinations and studied floral visitors.
• We found no differences between morphs in the proportion of flowers, nectar produced or its concentration, pollen quantity and fruit set. Both species were self‐incompatible and quite generalist regarding floral visitors.
• Enantiostyly in self‐incompatible plants seems to confer a reproductive advantage by reducing self‐interference resulting from stigma clogging. This novel result helps to expand our knowledge on this complex floral polymorphism and opens new avenues for future research on this topic.

     

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。     

Reproductive biology of Pachira aquatica Aubl. (Malvaceae: Bombacoideae): a tropical tree pollinated by bats,sphingid moths and honey bees

We investigated the reproductive biology, including the floral biology, pollination biology, breeding system and reproductive success, of Pachira aquatica, a native and dominant tropical tree of fresh water wetlands, throughout the coastal plain of the Gulf of Mexico. The flowers present nocturnal anthesis, copious nectar production and sugar concentration (range 18–23%) suitable for nocturnal visitors such as bats and sphingid moths. The main nocturnal visitors were bats and sphingid moths while bees were the main diurnal visitors. There were no differences in legitimate visitation rates among bats, moths and honey bees. Bats and honey bees fed mainly on pollen while moths fed on nectar, suggesting resource partitioning. Eight species of bats carried pollen but Leptonycteris yerbabuenae is probably the most effective pollinator due to its higher pollen loads. The sphingid moths Manduca rustica, Cocytius duponchel and Eumorpha satellitia were recorded visiting flowers. Hand pollination experiments indicated a predominant outcrossing breeding system. Open pollination experiments resulted in a null fruit set, indicating pollen limitation; however, mean reproductive success, according to a seasonal census, was 17 ± 3%; these contrasting results could be explained by the seasonal availability of pollinators. We conclude that P. aquatica is an outcrossing species with a pollination system originally specialized for bats and sphingid moths, which could be driven to a multimodal pollination system due to the introduction of honey bees to tropical America.     

Reproductive Ecology of Rhynchanthus beesianus W. W. Smith （Zingiberaceae）in South Yunnan, China： A Ginger with Bird Pollination Syndrome

Rhynchanthus beesianus W. W. Smith （Zlnglberaceae） Is an eplphytlc tropical ginger with a very conspicuous floral display, but almost no fruit set under field conditions. The reproductive ecology encompassing phenology, floral biology, and pollination and breeding systems was Investigated In an evergreen broadleaved forest In Yunnan Province, Southwest China. The flowers possess a typical bird pollination syndrome, but no effective pollinators were observed during 138 h of observation. Female Black-breasted Sunbird （Aethopyga saturata） and bumblebees visited R. beesianus regularly, but they all played roles as nectar robbers. No fruit was found In the bagging treatment, and fruit set following manual self-pollination （（57.55 ± 4.08）%） was comparable with cross-pollination （（64.32 ± 4.42）%）, suggesting that R. beeslanus is self-compatible but spontaneous self-pollination In this species does not occur. Seed set of open-pollination （（26.42 ± 3.11）%） was significantly lower than manual self-pollination （（73.41± 4.16）%） and cross-pollination （（75.56 ± 4.52）%）, confirming that R. beeslanus was dependent on animals for fertilization and suffered a serious pollinator-limitation.     

报春花属滇海水仙花开花传粉特性研究

通过田间观察,运用杂交指数、花粉及柱头活力、花粉/胚珠比、坐果率等指标,对滇海水仙花人工种群的花部特征、开花动态及授粉特性等进行研究。在人工栽培条件下,滇海水仙花花期为12月至翌年2月,单株花期30～45 d,单花花期为9～14 d,花朵有长花柱花和短花柱花之分。滇海水仙花的传粉昆虫主要是蜜蜂,此外还有蝴蝶和蝇类,其繁育系统为兼性自交。     

蓝花丹的花部形态二态性及自交不亲和特性

花部形态的二态性在异型自交不亲和中的作用一直颇具争议。为了探讨其对自交不亲和的作用,选择代表植物蓝花丹,观测其花部形态,利用激素调控其花柱高度,对调控后的花柱授粉,并分析结实率的变化。结果表明：(1)蓝花丹的花部形态在众多方面存在二态性;(2)IAA,NAA,GA 在适宜的浓度都能调节其花柱的长度;(3)调节后的花柱自然和人工授粉,结实率与空白对照相比无显著差异。实验结果说明蓝花丹的花部形态有着典型的二态性特征,此特征是蓝花丹对花型间互补式雌雄异位的进化适应策略;而雌雄性器官的空间分布并没有通过影响合法花粉的有效传播而影响自交不亲和性;异型花的自交不亲和性更多地还是体现在生理机制上。     

Sexual dimorphism and gynoecium size variation in the andromonoecious shrub Caesalpinia gilliesii

The degree of sexual dimorphism in flowers and inflorescences can be evaluated early in flower development through the study of floral organ size co-variation. In the present work, the gynoecium-androecium size relationship was studied to assess the degree of sexual expression in flowers and inflorescences of the andromonoecious shrub Caesalpinia gilliesii. The co-variation pattern of floral organ sizes was compared between small and large inflorescences, under the hypothesis that inflorescence size reflected differential resource availability. Also, staminate and perfect flowers were collected from three populations and compared on the basis of gynoecium, ovule length, filament length, pollen size and number. The obtained results indicated that staminate and perfect flowers differed only in the gynoecium and ovule length, whereas filament length, pollen size, and number varied across populations. The gynoecium size was smaller and its variability was much higher in staminate than in perfect flowers, as explained by a recent hypothesis about pollinator-mediated gynoecium size selection acting upon perfect flowers. The analysis of the gynoecium-androecium size relationship during flower development, revealed a dissociation of gynoecium growth relative to other floral structures in some buds. Lower gynoecium-androecium regression slopes and smaller gynoecia length characterized smaller inflorescences, thus reflecting the fact that sexual expression was more male-biased. This trend is in agreement with a differential resource-related response at the inflorescence level, however, post-mating resource allocation and the inclusion of other modular levels may also help us to understand the variation in sexual dimorphism in this species.     

木槿与野西瓜苗花特征和繁育系统的比较研究

为比较分析柱头裂片运动的野西瓜苗和不运动的木槿在花特征和繁育系统上的差异,对其花性状、花粉/胚珠比和不同授粉处理的座果率进行了测定。结果表明:(1)木槿与野西瓜苗在花冠大小、花瓣长及底宽、花萼长与宽、雄蕊长、最上轮雄蕊高度、雄蕊柱长及其底径、花柱长度和雌雄异位间均存在十分显著的差异(P<0.01),但二者的最下轮雄蕊高度间无显著性差异(P>0.05)。(2)木槿的花粉/胚珠比为874.90±20.79,繁育系统属兼性异交类型;野西瓜苗的花粉/胚珠比为24.72±0.68,繁育系统属专性自交类型。(3)木槿自然套袋的座果率为0%,人工自交为2.04%,人工杂交为35.85%;野西瓜苗自动自交和人工自交的座果率均为100%,人工杂交为95.30%。木槿与野西瓜苗在花性状、花粉/胚珠比及花部行为等方面形成了与其繁育系统(兼性异交vs.专性自交)相适应的花特征。     

草豆蔻传粉生物学的研究

比较研究了花柱卷曲性植物草豆蔻 Alpinia hainanensis两种表型(上举型和下垂型)的传粉生物学特性。结果表明:草豆蔻的花柱卷曲运动节律与其他已报道的山姜属Alpinia植物基本一致,而下垂型花的花柱卷曲运动明显滞后于上举型花约2 h。草豆蔻的花柱卷曲运动中存在一些不同步的现象,甚至在同一花序上的两朵花的花柱运动节律亦会表现出不一致的现象,但上举型花的花药开裂时间却严格同步,都发生在同类型个体的柱头全部位于花药上方之后进行。草豆蔻上举型花的花蜜分泌量、单花花粉量、花粉/胚珠比率(P/O)均明显比下垂型花多,而两种表型的胚珠数、花蜜糖含量、氨基酸含量无显著差异。在草豆蔻的单花期内不同时间段进行人工授粉,上举型花在其柱头位于花药下方时进行人工授粉,其结实率明显比柱头位于花药上方时人工授粉的处理高,下垂型花则没有显著差异。在自然居群中,草豆蔻的主要传粉者是无垫蜂Ameglla sp.和两种木蜂Xylocopa sp.,但存在传粉者不足而影响结实率的现象。完全套袋、去雄和去雌(去柱头)处理的均不结实,表明草豆蔻中不存在无融合生殖、主动自花授粉和滞后自交的生殖保障现象。而人工自交和异交均具有较高的结实率,表明草豆蔻为自交亲和植物。草豆蔻的繁育系统是具花柱卷曲性运动的异花授粉的交配系统。     

Sex‐specific floral attraction traits in a sequentially hermaphroditic species

• ● Many angiosperms are hermaphroditic and produce bisexual flowers in which male (pollen export) and female (stigma receptivity) functions are separated temporally. This sequential hermaphroditism may be associated with variation in flower size, color, or pattern, all of which may influence pollinator attraction. In this study, we describe variation in these traits across discrete functional sex stages within and between 225 greenhouse‐grown individuals of Clarkia unguiculata (Onagraceae). In addition, to identify the effects of floral phenotype on pollinator attraction in this species, we examine the effects of these floral traits on pollen receipt in ~180 individuals in an experimental field array.
• ● Petal area, ultraviolet (UV)‐absorbing nectar guide area, and blue and green mean petal reflectance differ significantly across the functional sex stages of C. unguiculata. Male‐ and female‐phase flowers display significantly different pollinator attraction traits. Petal and UV nectar guide area increase as flowers progress from male phase to female phase, while blue reflectance and green reflectance peak during anther maturation.
• ● In field arrays of C. unguiculata, female‐phase flowers with large UV nectar guides receive more pollen than those with small nectar guides, and female‐phase flowers with high mean blue reflectance values are more likely to receive pollen than those with low blue reflectance. Female‐phase flowers with green mean reflectance values that differ most from background foliage also receive more pollen than those that are more similar to foliage. These findings indicate that components of flower color and pattern influence pollen receipt, independent of other plant attributes that may covary with floral traits. We discuss these results in the context of hypotheses that have been proposed to explain sex‐specific floral attraction traits, and we suggest future research that could improve our understanding of sexual dimorphism in sequentially hermaphroditic species and the evolution of features that promote outcrossing.

     

香石竹开花结实生物学研究

通过田间观察,运用TTC法、联苯胺-过氧化氢法、杂交指数、花粉-胚珠比和套袋实验等方法,对香石竹的开花状态及繁育系统进行了研究。结果表明:在人工栽培条件下,香石竹种群可周年开花,单株花期35～61d,单花花期9～15d;花粉在开花第1～5天具有活力,柱头在开花第4～12天具可授性,第8～11天可授性最强;杂交指数为5,P/O值为307.84～790.52,结合坐果率结果,判断其繁育系统属于异交,部分自交亲和,需要传粉者。