火把花的繁殖生物学研究

火把花（Colquhounia coccinea?）表现出典型的鸟媒综合征,因此具有作为引鸟景观植物的开发潜力。以自然、人工生境的火把花居群为研究对象,通过观察和试验对其开花物候、花部综合特征、访花动物及其行为、繁育系统、种子萌发特性进行研究,以明确火把花的繁殖特性及对不同访花动物的吸引潜力。结果显示,火把花的整体花期持续约3个月,单花花期为9.6±0.6 d;花蜜较丰富且稀薄,具有较短的花冠管和己糖为主的花蜜糖组成;访花动物主要是中华蜜蜂和多种食蜜鸟类,尤其是短喙的泛化鸟类,且在非自然生境中仍然能吸引鸟类访花;完全自交亲和但需要传粉者才能完成授粉,不存在花粉限制;不同授粉处理种子的发芽能力无显著差异;中华蜜蜂能有效传粉,鸟类的传粉作用需进一步验证。综上所述,火把花可供观赏的时间很长,具有明显的吸引鸟类访花的能力,容易通过有性繁殖途径快速获得大量幼苗。     

Autogamy of an endangered species: Loropetalum subcordatum (Hamamelidaceae)

The reproductive biology of Loropetalum subcordatum was studied. The floral phenology, pollen histochemistry, pollen-ovule ratio (P/O), pollen viability and floral visitors were investigated and determined. Assisted pollination experiments were carried out to examine the breeding system of L. subcordatum. Scanning electron microscopy (SEM) and fluorescence microscopy (FM) were employed to check the pollen germination and the growth of pollen tubes. Results were obtained as follows: (1) The flowering period of L. subcordatum was from September to the next February with a peak at September-October; the longevity of a single flower was 4-6 days. (2) L. subcordatum was protogynous, and pollen grains could be found on self-stigmata when the anthers opened 12-24 h after petals unrolling. (3)The pollen viability (MTT test) maintained for ca. 26 hours; the pollen was starchless; the P/O was 8420±720.86 (n=10); no nectar secretion was observed. (4) Thrips (Thrips sp.) were the only floral visitors observed but which seldom moved among inflorescences, thus played limited role in pollination. (5) Fruit sets of untreated bagged (5.30±1.83%) and hand assisted cross pollinated flowers (6.67±1.91%) were not significantly different from that of open flowers (4.79±1.45%). (6) SEM and FM observations proved that pollen germinated on self-stigmata and pollen tubes growed in self-styles. The results indicated that L. subcordatum was facultatively autogamous and without apomixes. The possibility of outcrossing and the protogyny might indicate that the species was originally a crosser. Flowers pollinated in September-October usually start ovule growth in next summer, and set mature fruits in next October (after the next flowering peak), suggesting the occurrence of retard embryo development. The possibility that autogamy in Hamamelidaceae could have been developed from fly pollination was also discussed.     

濒危植物肉苁蓉（Cistanche deserticola）繁育系统研究

在自然居群中通过野外定位观测,运用杂交指数（OCI）;花粉—胚珠比（P/O）;去雄、套袋、人工授粉等方法对濒危植物肉苁蓉（Cistanche deserticola Ma）的花序发育、开花特性、花粉活力、柱头可授性、传粉方式、繁育系统进行了研究,结果表明：肉苁蓉穗状无限花序的生长发育期可明显分为拱土期、出土期、开花期、结实与种子成熟期四个阶段。两性花,单花开放时间2~3d,花序有花期持续20~25d。种群开花历时36~38d。开花时柱头略高出花药,柱头较大,下垂。重力玻片法检测,风媒导致的异株异花传粉完全可以忽略。主要访花昆虫为蜂类。扫描电镜观察虫体,蜂类携带大量花粉,但传粉飞行距离大部分为0。杂交指数为3;花粉—胚珠比为35.45±8.56;结合去雄、套袋和人工授粉试验结果,认为肉苁蓉属于以自交为主的繁育系统。     

Reproductive biology of Boswellia serrata, the source of salai guggul, an important gum-resin

Detailed studies were carried out on the phenology, floral biology, pollination ecology and breeding system of Boswellia serrata Roxb. (Burseraceae) the source of 'salai guggul'. The trees remain leafless during the entire period of flowering and fruiting. The inflorescence is a terminal raceme and produces up to 90 bisexual, actinomorphic flowers. On average a flower produces 10 044 ± 1259 starch-filled pollen grains. About 85% of the fresh pollen grains are viable; the pollen to ovule ratio is 3348 : 1. The stigma is of the wet papillate type. The style is hollow with three flattened stylar canals filled with a secretion product. The stylar canals are bordered by a layer of glandular canal cells. The inner tangential wall of the canal cells shows cellulose thickenings. The ovary is trilocular and bears three ovules, one in each locule. Flowers offer nectar and pollen as rewards to floral visitors. The giant Asian honey bee ( Apis dorsata ) and A. cerana var. indica (Indian honey bee) are the effective pollinators. The species is self-incompatible and the selfed pollen tubes are inhibited soon after their entry into the stigma. Self-pollen tubes develop a characteristic 'isthmus' as a result of enlargement of the tube soon after emergence through the narrow germ pore. Cross-pollinated flowers allowed normal pollen germination and pollen tube growth, and resulted in fruit- and seed-set. Under open pollination fruit-set was only about 10%. Although manual cross-pollinations increased fruit set, it was only up to about 20%. Low fruit set appears to be the result of inadequate cross-pollination and other constraints, presumably limitation of available nutrients.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 73–82.     

野菊与菊花杂交中花粉活力和柱头可授性及胚胎发育研究

应用石蜡制片、活体压片、光学显微镜及扫描电子显微镜观察等方法,研究了四倍体河南云台山野菊(Dendranthema indicum)与栽培菊花'钟山金山'(D.grandiflorum 'Zhongshanjinshan')种间杂交中父本花粉活力、花粉在柱头萌发、花粉管生长及胚胎发育情况等.结果发现,父本云台山野菊的花粉活力在授粉时为12%左右.人工授粉后的不同时间,在柱头上都观察到正常萌发的花粉粒,且花粉管都能进入柱头,其中,在授粉后0.5 h时,平均每柱头有5.9粒花粉萌发;12 h时,为59.9粒;而24和48 h时,则分别降为47.1和35.7粒.此外,在授粉后8、10、12和15 d时,分别在49.1%、40.8%、39.7%和38.5%子房内观察到正常发育的胚胎,最终杂交结实率为44.8%,而母本自然开放结实率为52.3%.研究表明,授粉前其多数母本雌蕊发育良好、授粉后多数花粉能在柱头正常萌发和花粉管正常生长,在受精后大部分胚胎发育正常是野菊与栽培菊种间杂交较高结实率的重要保证,而授粉前父本较低的花粉活力对杂交结实率影响不大.     

Environmental Effects on Components of Pollen Performance in Faramea occidentalis (L.) A. Rich. (Rubiaceae)1

Studies focusing on gametophytic competition have focused on differences in pollen performance (e.g., pollen germination rate and pollen tube growth rate) among donors in order to examine genetic sources of variation in siring success among them. Donors that produce the fastest growing pollen tubes are expected to fertilize more ovules relative to donors with slow growing pollen tubes under conditions of gametophytic competition. However, the performance of pollen in the field is known to be influenced by environmental conditions in addition to the genotype of donor plants. This field study of Faramea occidentalis was conducted to: (1) determine the effect of environmental conditions during pollination on pollen performance; (2) measure differences among pollen donors in pollen performance; (3) determine if the pollen of different donors responds differently to a variety of environmental conditions surrounding pollinated flowers; and (4) measure differences among pollen recipients in pollen performance. Single-donor crosses were made between four pollen donors and four pollen recipients under a variety of environmental conditions. Pollen performance was then quantified as the growth rate of the fastest pollen tube, the mean pollen tube growth rate, and by a pollen germination index. Pollination environment (the environment surrounding a pollinated flower) and recipient significantly affected all three measures of pollen performance. Pollen donors did not differ overall in pollen performance. However, there was significant among-donor variation in two of the five pollination environment conditions. Future studies of variation in relative siring success may benefit by considering pollination environment in addition to donor and recipient identity.     

Nectarless flowers with deep corolla tubes in Pedicularis: does long pistil length provide an arena for male competition?

Pollinator‐mediated selection does not seem to have a direct influence on the evolution of a long corolla tube in a nectarless flower. We hypothesized that the long pistil length of the nectarless flower with a deep corolla tube provided an opportunity for male competition. Pedicularis siphonantha, a nectarless and partially self‐incompatible lousewort with substantial variation in corolla tube length, was used to test the hypothesis. We tested whether and how corolla tube length affected seed production per capsule and seed germination rate with different pollination treatments. Flowers were hand‐pollinated with pollen from one self donor and one outcross donor and mixed pollen grains consisting of equal amounts from the two donor types, respectively. Additionally, seeds from open‐pollinated flowers with different corolla tube lengths were collected separately for measurement of germination rate. Pollination treatment and corolla tube length significantly affected number of seeds per capsule. Moreover, a significant positive relationship between seeds per capsule and corolla tube length was found when mixed hand pollination was conducted. Seeds of self hand‐pollinated flowers had a lower germination rate than those from outcross‐pollinated flowers. Under open pollination, seeds from flowers with longer corolla tubes tended to have higher germination rate. In P. siphonantha, outcross pollen may have a higher probability of contributing to the next generation when transferred to flowers with longer corolla tubes. The pistil length, therefore, should be seen as a female choice mechanism, which provides an arena for male‐to‐male competition. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 526–532.     

Sunny-side up: flower heliotropism as a source of parental environmental effects on pollen quality and performance in the snow buttercup, Ranunculus adoneus (Ranunculaceae)

Floral traits affect mating success via their influence on the microenvironment in which sexual reproduction occurs as well as their impact on pollinator attraction. Here we investigate the importance of flower heliotropism as a source of parental environmental effects on pollen quality and performance. Flowers of the snow buttercup, Ranunculus adoneus, closely track the sun's rays. We experimentally restrained flowers to test for effects of heliotropism on pollen quality and performance after pollination. When equivalent amounts of pollen were transferred to recipient pistils, pollen from solar-tracking donor flowers exhibited a 32% advantage in germination compared to pollen from stationary (tethered) donor flowers. By the end of anthesis, pistils of tracking flowers contained 40% more germinating pollen grains and 44% more pollen tubes midway down the style than pistils of stationary ones. Solar tracking had no direct effect on pollen tube growth. The greater amount of germinating pollen in tracking flowers accounted for the treatment effect on pollen tube density. A survey of pollen receipt and pollen germination in naturally tracking flowers indicated that solar tracking primarily affects pollen tube density by promoting pollen germination rather than pollen deposition. We conclude that flower heliotropism, by enhancing the paternal environment for pollen development and the maternal environment for pollen germination, represents a source of positive parental environmental effects on pollen performance in snow buttercups.     

Changes in Florets’ Vertical Direction within Inflorescence Affects Pollinator Behavior,and Fitness in <i>Trifolium repens</i>

Ecological interactions between flowers and pollinators greatly affect the reproductive success. To facilitate these interactions, many flowers are known to display their attractive qualities, such as scent emission, flower rewards and floral vertical direction, in a rhythmic fashion. However, less is known about how plants regulate the relationship between these flower traits to adapt to pollinator visiting behavior and increase reproduction success. Here we investigated the adaptive significance of the flower bending from erect to downward in Trifolium repens. We observed the flowering dynamic characteristics (changes of vertical direction of florets, flowering number, pollen grain numbers, pollen viability and stigma receptivity over time after blossom) and the factors affecting the rate of flower bending in T. repens. Then we altered the vertical direction of florets in inflorescence of different types (upright and downward), and compared the pollinator behaviors and female reproductive success. Our results showed that florets opened sequentially in inflorescence, and then bend downwards slowly after flowering. The bending speed of florets was mainly influenced by pollination, and bending angle increased with the prolongation of flowering time, while the pollen germination rate, stigma receptivity and nectar secretion has a rhythm of “low-high-low” during the whole period with the time going. The visiting frequency of all the four species of pollinators on upward flowers was significantly higher than that of downward flowers, and they especially prefer to visit flowers with a bending angle of 30°–60°, when the flowers was exactly of the highest flower rewards (nectar secretion and number of pollen grains), stigma receptivity and pollen germination rate. The seed set ratio and fruit set ratio of upward flowers were significantly higher than downward flowers, but significantly lower than unmanipulated flowers. Our results indicated that the T. repens could increase female and male fitness by accurate pollination. The most suitable flower angle saves pollinators’ visiting energy and enables them to obtain the highest nectar rewards. This coordination between plants and pollinators maximizes the interests of them, which is a crucial factor in initiating specialized plant-pollinator relationships.     

青篱柴通过促进亲和花粉生长而提高传粉精确性

异型花柱植物的适应意义在于提高亲和花粉的传递, 从而促进异交, 但是有不少研究发现柱头上落置了大量不亲和花粉。目前, 很少有人研究柱头是否有利于亲和花粉的萌发和花粉管伸长, 以及通过去雄处理排除花内和植株内不亲和花粉干扰从而验证二型花柱植物是否可以促进异交。本研究以亚麻科青篱柴属(Tirpitzia)的二型花柱植物青篱柴(T. sinensis)为研究对象, 分居群调查了青篱柴长、短柱型的植株数量, 测量不同柱型的花粉与柱头大小以及观察其表面纹饰, 测定不同开花时间长、短柱型的柱头活性与花粉活力, 统计自然状态下柱头上落置的不同表型花粉比例, 进一步统计花内、株内去雄和自然对照下柱头上落置的不同表型花粉比例, 计算型间、型内和混合授粉花粉管长度和花粉萌发率, 人工授粉检测其是否型内异交不亲和与自交不亲和。结果表明, 自然居群中长、短柱型青篱柴植株数量没有显著性差异; 短柱型青篱柴花粉体积显著大于长柱型, 且二者花粉表面纹饰不同, 但不同柱型的柱头表面积与表面纹饰无差异; 青篱柴开花第一天的花粉活力显著高于开花第二天, 而柱头活性在开花前两天无显著性差异; 自然状态下柱头上落置的不亲和花粉比例显著高于亲和花粉; 在长柱型青篱柴中, 花内和株内去雄显著提高了柱头上亲和花粉的落置比例; 柱头上授型内花粉, 其花粉萌发率和花粉管的长度都显著低于授型间花粉; 授粉结果表明青篱柴为型内异交不亲和与自交不亲和。二型花柱植物通过促进亲和花粉的萌发和花粉管的伸长而有利于型间授粉, 从而显著提高其传粉精确性。