开花时间对巫山淫羊藿花蜜分泌和结实的影响

运用单因素方差分析研究了巫山淫羊藿(Epimedium wushanense)开花时间对花蜜分泌、有效传粉者活动及结实性的影响.结果表明:(1)开花时间对巫山淫羊藿花蜜常备量和含糖量的影响极其显著.3个种群早期花蜜常备量较高,而花蜜含糖量在晚期较高.(2)开花时间对有效传粉者行为也具显著影响.一天中,有效传粉者的访花活动主要集中在中午;在整个花期进程中,晚期有效访花频率明显大于早期.(3)开花时间对巫山淫羊藿结实率、果实种子数及果实种子败育率的影响均显著,其中早期和中期的结实率和果实种子数大于晚期;而果实种子败育率则在晚期较高.因此,人工栽培巫山淫羊藿应注意花期调控,并选择早期果实进行实验或育种,其效果更好.     

淫羊藿开花的时空效应对其结实的影响

运用非参数检验和单因素方差分析,研究了淫羊藿开花的花序水平的位置效应与空间效应对结实的影响.结果表明:(1)开花时间效应对内萼片长、内萼片宽、花药长、距宽、距长的影响显著,其参数均是早期和中期大于晚期;花部空间对内萼片长和内萼片宽的影响显著,而对花药长、距长和距宽的影响不显著.(2)开花时期对淫羊藿结实率无显著差异,但早期和中期的结实率要远远高于晚期;时间效应对果实形态参数的影响极显著,且大小均为早期大于中期和晚期;早期和中期的果实重量和种子数要显著高于晚期,而在晚期的果实种子数败育率较高.(3)在开花的空间效应中,基部和中部之间的结实率无显著差异,但两者显著高于顶部的结实率;基部的果实形态参数要高于中部和顶部;从果实重量和种子数来看,基部和中部要大于顶部,但顶部的果实败育率高于基部和中部.因此,对于淫羊藿的实际繁殖或药用,应采摘早期和中期开花的基部果实,其效果会更好.     

巫山淫羊藿开花物候与生殖特征

研究巫山淫羊藿(Epimedium wushanense)3个种群的开花物候进程和生殖特征,分析了不同种群中的花部特征、开花物候指数和结实率差异,以及开花物候指数对巫山淫羊藿生殖成功的影响。结果表明:巫山淫羊藿花期为3月初到4月中下旬,种群开花一般历时22～27d;单株花期(花序)约12～17d;单花花期一般为3～4d。3个种群的个体水平的开花振幅曲线呈单峰型曲线,均具有较高的开花同步指数,表现出一种集中开花式样。相关分析表明:坐果数与开花高峰期、终花期、花期持续时间和花蕾数均呈极显著正相关,但坐果数与始花期无相关关系。开花物候变异系数分析表明:3个种群中花期持续时间、单花持续时间、终花期、开花高峰期和坐果数差异极显著;总花数、开花振幅和结实率差异显著;开花同步指数和始花期无显著差异性。3个种群中除同步性指数外,其余指数的变异系数均有差异,此外,巫山淫羊藿中结实率受到距的长短的影响。表明巫山淫羊藿的开花物候和生殖特征受到微环境的影响,同时生殖特征还受到花部特征距的影响。     

不同大花类群淫羊藿属植物的花部特征及生殖特征研究

淫羊藿是我国重要的草本植物,在中药、功能性食品和园林观赏等领域都具有重要用途,受地域差异性影响,淫羊藿属植物的植株形态、花部特征、开花物候和生殖特征各不相同。本研究对四川不同产地野生种群大花类淫羊藿（巫山淫羊藿、粗毛淫羊藿和宝兴淫羊藿）的开花动态、花部特征、访花昆虫与繁殖系统进行了对比研究。结果表明：（1）3种淫羊藿属植物的开花期较为集中,巫山淫羊藿和粗毛淫羊藿的花期为3月下旬～4月下旬,单花花期均为3～4 d;宝兴淫羊藿的花期为4月中旬～5月中旬,单花花期5～6 d;（2）膜翅目和双翅目昆虫为3种淫羊藿属植物的有效访花者,有效访花者访花特性与花部特征之间具有明显的相关性;（3）3种淫羊藿属植物的花序、花和果实的数量受所处环境影响较大,自然结实率：巫山淫羊藿>宝兴淫羊藿>粗毛淫羊藿;温度和光照是最主要的影响因子。     

不同生境与花部特征对巫山淫羊藿结实特性的影响

对不同生境中巫山淫羊藿结实特性进行比较研究。结果表明:(1)三个种群在繁殖时期中不同花部特征的植株繁殖投资差异不大,如每株分枝数、每株花序数和每株开花数差异均不显著;(2)三个种群中每株结实数、每株结实率、每花序结实数在不同花部特征中差异显著,PLS和SAA要显著大于PESS;(3)三种花部特征在不同生境中单株分枝数和单株花序数均无显著差异。在不同花部特征中种群1开花数量相较少,因而在结实方面均值均要小于种群2和种群3;(4)不同花部特征的果实饱满种子数、败育数和败育率均具有显著差异,而种子数差异不大,但在种群间均无显著的差异。说明种子的多少更多地受到花部特征的影响。总之,巫山淫羊藿的结实特性受到生境和花部特征的双重影响,其中花型比生境对巫山淫羊藿结实特性的影响更为明显。     

不同栽培方式对巫山淫羊藿生长的影响

测定和比较了3种栽培方式(栽培密度、栽培根茎长度和栽培深度)下巫山淫羊藿叶长、株高、分枝数和花蕾数等生理生长指标,探寻巫山淫羊藿最适生长条件和生长投资策略,为药用淫羊藿人工栽培提供理论基础和实验依据。结果表明:在每平方米18000株的栽培密度下,植株各项生理指标均显著优于其它密度处理;15cm根茎长度处理下的巫山淫羊藿分枝数明显高于其它组;5cm深度处理有利于巫山淫羊藿叶生长,20cm深度处理有利于巫山淫羊藿茎生长。进一步分析表明:栽培密度和栽培根茎长度对巫山淫羊藿营养生长影响较大;栽培深度对植株的营养生长和生殖生长影响都较大。     

淫羊藿种子产量与生境的关系

以四川省南充市金城山的淫羊藿为研究对象,对小檗科淫羊藿产果数量,种子生产和种子质量与生境的关系进行了研究。研究表明:不同生境的淫羊藿植株高度和花序长度不同,单株结果率也不同,单果产种量差异较小,出种量一般在1~8粒之间,但每个生境单果主要出种量不同。对千粒重和不同果枝结果率单因素方差分析表明,不同生境的淫羊藿种子千粒重有显著差异,不同生境的淫羊藿植株的主次果枝结果率也有显著差异。不同生境中的淫羊藿果实的虫食率也不同。淫羊藿种子的低种子数,小粒种子,而不同生境的结实率虽有差异,但是总体偏低,是导致淫羊藿种子难以收集的主要内因,也导致种群受干扰后不易恢复。     

金城山柔毛淫羊藿的开花物候与生殖特性

采用实地观测的方法,对南充市金城山三个不同生境中柔毛淫羊藿的开花物候特征及其生殖特性进行了研究。结果表明:柔毛淫羊藿花期为3月上旬至4月上旬,其种群、个体、花序和单花的花期分别历时30~41、22~34、9~18和4~8 d。三个生境中柔毛淫羊藿种群的开花物候进程基本相同,均呈单峰集中开花式样,因而能够吸引更多的传粉者访问而达到生殖成功。开花物候指数与坐果数之间的相关分析结果表明,坐果数与始花日期存在显著的负相关关系,与花期持续时间和开花数均呈极显著的正相关关系。揭示了药用植物柔毛淫羊藿的开花物候特征与生殖特性。     

朝鲜淫羊藿（Epimedium koreanum Nakai）不同生境种群生物量与更新潜力

朝鲜淫羊藿（Epimedium koreanum Nakai）为重要药用植物资源,由于种子繁殖问题较大,而尚未实现真正的人工栽培,野生资源在长期过度利用中已遭到一定的破坏。研究结果表明：朝鲜淫羊藿的有性生殖能力较弱,平均开花和结果率分别不超过8%和6%,最低开花和结果率均在次生杂木林下,分别仅有2%和0.8%。朝鲜淫羊藿主要以无性繁殖维持种群更新和增长,其中,2龄根茎无性繁殖活力最高,其产生的潜伏芽和1龄根茎数量平均占80%以上;3龄和4龄根茎产生的潜伏芽和1龄根茎数量显著减少,而4龄根茎的潜伏芽100%不能发育成1龄根茎;5龄以后的根茎不再有无性繁殖活力。朝鲜淫羊藿无性繁殖潜伏芽数量具有较大冗余性,并随着根茎龄级的增高和生境条件的变差其冗余性也增大。另外,朝鲜淫羊藿的开花率、结果率和根茎产生潜伏芽及1龄根茎数量,均随生境条件的不适宜性增强和根茎龄级增大而变异系数明显增大,说明生境条件变差和生理功能衰退导致其种群增殖的不确定性增加,而有序性降低。总之,朝鲜淫羊藿是极易遭到利用威胁的易危物种,应注意对野生资源利用强度和方式的研究与控制。[编者按]     

梯度洗脱法同时快速分离测定不同品种不同部位淫羊藿中7种异戊烯基黄酮类成分的研究

建立一种同时快速分离测定不同品种、不同部位淫羊藿中双藿苷A、淫羊藿属苷A、朝藿定C、淫羊藿苷、淫羊藿属苷C、意卡瑞苷A和去多甲基羟基淫羊藿素7种异戊烯基黄酮类成分的梯度洗脱方法,其中双藿苷A、意卡瑞苷A和去多甲基羟基淫羊藿素为首次报道其含量测定。实验比较了三种不同的梯度洗脱条件和三种不同的提取分离条件,选出了实验的最佳条件:采用RP-HPLC法,以美国WATERS SUNFIRETM-C18为色谱柱,乙腈-水为流动相,梯度洗脱(乙腈:0 min,20%;10 min,40%;15 min,45%;20 min,60%),流速为1 mL/min,检测波长270 nm,在此条件下,30 m in内所有组分均得到了良好的分离。实验同时考察比较了巫山淫羊藿植株不同药材部位和其它十种不同品种淫羊藿中7种异戊烯基黄酮类成分的分布。结果表明:不同品种淫羊藿中黄酮类成分的含量差异很大,但大部分品种以朝藿定C为主;同一品种巫山淫羊藿不同部位中则双藿苷A和朝藿定C的含量较大(在巫山淫羊藿根中,两者的含量分别为现行质量控制指标淫羊藿苷的20倍和30倍),可见巫山淫羊藿显著不同于其它品种的淫羊藿,朝藿定C在其药用效果中可能比淫羊藿苷扮演着更重要的角色尤其是在根部作为药用时,这也为民间将巫山淫羊藿根作为小黄连使用提供了一定的实验支持。结论:对于品种较多,分布资源较广的淫羊藿药材,作为药用时,应注意品种和药用部位,考察药材的内在质量,不能一概而论。实验所得到的方法能被用做淫羊藿植株中二次代谢产物分布的快速考察、筛选符合要求的淫羊藿原药材和淫羊藿制备药物中的质量考察。     

The movement patterns of carpenter beesXylocopa micans and bumblebeesBombus pennsylvanicus onPontederia cordata inflorescences

The movement patterns of carpenter bees (Xylocopa micans) and bumblebees (Bombus pennsylvanicus) foraging for nectar on vertical inflorescences ofPontederia cordata were studied near Miami, Florida. The floral biology ofP. cordata is unique in several ways: (a) many short-lived flowers per inflorescence, (b) constant nectar production throughout the life span of each flower, and (c) abscence of vertical patterning of nectar and age of flowers. Inflorescences ranged between 3.5 and 15.8 cm long and had between 9 and 55 open flowers. Both carpenter bees and bumblebees arrived mostly on the bottom third of the inflorescence and left after visiting flowers on the top third of the inflorescence. The departure position from the inflorescence was higher up than observed in studies of other insect pollinators foraging on other speces of plants. This pattern of departure probably occurs in the absence of a vertical gradient of nectar or floral morphology.     

Fruit size decline from the margin to the center of capitula is the result of resource competition and architectural constraints

Plants produce repeated structures, such as leaves, flowers, and fruits, which differ in size and shape. One example of this is fruit size, which is commonly observed to decrease from proximal to distal positions within an inflorescence. The resource limitation hypothesis proposes that because proximal fruits usually develop first, they have temporal priority on access to resources over distal fruits. The non-mutually exclusive architectural effects hypothesis suggests that these position effects in fruit size may also be due to inherent architectural variation along infructescence axes. We separated out the effects of resource competition and inflorescence architecture by removing the outer or the inner flowers within capitula of Tragopogon porrifolius. We also studied if fruit position influenced germination and seedling performance in order to assess fitness consequences of position effects. Inner fruits were significantly heavier when outer flowers were removed. However, outer fruits did not significantly increase when inner flowers were removed, suggesting later fruits were limited by the development of early fruits. Our findings also suggest that architectural constraints restricted the size of inner fruits in comparison with outer ones. We found that both resource competition and inflorescence architecture affected the fruit size of T. porrifolius, even though this species does not have linear, indeterminate inflorescences. We advance the hypothesis that, when such effects on fitness occur, resource competition-mediated position effects could turn, in evolutionary time, into architectural position effects.     

Pollinator limitation and position dependent fruit set in the high Andean orchidMyrosmodes cochleare (Orchidaceae)

Myrosmodes cochleare is a terrestrial orchid restricted to high elevations of the northern Andes. It is self-compatible but not autogamous. Flowering begins at the apex of the inflorescence, but flower size increases steadily towards the base. Fruit set of selfed flowers was found to increase significantly from the apex to the base of the inflorescence. Open-pollinated fruit set was significantly lower than hand-pollinated fruit set overall, but the highest levels occurred in the middle of the inflorescence. Eumenid wasps and a calliphorid fly were observed to pollinate the flowers, but visits were rare and occurred only during infrequent warm periods.     

毛翠雀花花序内的性分配和繁殖成功

两性花植物花序内不同位置的性分配和繁殖成功一般存在差异,通常认为资源竞争、结构效应和交配环境是形成这种差异的主要原因。为了研究雄性和雌性繁殖资源在花序内不同位置间的最优分配问题,该文以青藏高原高寒草甸典型高山植物毛翠雀花为材料,通过比较花序内不同位置的花部特征和种子性状,对其花序内的性分配模式和雌性繁殖成功进行研究,并通过观察传粉者运动特点以及人工去花和补授花粉实验,探讨花序内资源竞争和交配环境对繁殖资源分配的影响。结果表明:(1)不同位置间的雄蕊数、雄蕊鲜重/雌蕊鲜重、花粉数及花粉胚珠比从花序基部到上部显著增加,而雌蕊鲜重和胚珠数逐渐减少,表现出上部花偏雄的性分配;上部花的结籽率显著低于基部花和中部花,不同位置间的发育种子数/果实和发育种子重/果实随着花位置的升高而显著降低,说明基部花具有更佳的雌性繁殖成效。(2)去花处理后,剩余果实的单个种子重/果实显著增加,但发育种子数/果实没有显著增加;而给上部花人工补授异花花粉后,位置间结籽率的差异消失,说明传粉限制而非资源竞争导致了花序内位置依赖的种子生产模式。(3)毛翠雀花雄性先熟的开花特征,以及传粉者苏氏熊蜂从花序基部到上部的定向访花行为,导致了花序内交配环境的变化。综上结果表明,毛翠雀花花序内的性分配和繁殖成功差异是对交配环境适应的结果,对其在高山环境中实现雌雄适合度最优化具有重要意义。     

Variation in fruit- and seed set among and within inflorescences of Melampyrum roseum var. japonicum (Scrophulariaceae)

Abstract Variations in fruit set and seed set among and within inflorescences of the annual herb, Melampyrum roseum var. japonicum, were studied. Under natural conditions, although the mean fruit set was slightly different among inflorescences, the mean seedset was not significantly different among inflorescences within the plants. In constrast, within the inflorescences, the flowers located at a lower position of the inflorescence and which opened earlier showed higher fruit set than those at a higher position and which opened later. However, the seed set of matured capsules were not significantly different from each other, regardless of the position of flowers within the inflorescences. Patterns of the fruit- and seed set under open pollination indicated that variation in seed reproduction of M. roseum is due to variation in fruit production. The results of clipping experiments of flowers revealed that there was no functional limitation in seed production among flowers located at various positions within the inflorescence. It seemed that the variation in the fruit set within the inflorescences of M. roseum was not attributable to 'architectural effects'. Reduction of the number of flowers within the inflorescences resulted in an increase of fruit set and seed weight, indicating that the flowers in an inflorescence compete for resources. This phenomenon supports the 'resource competition hypothesis', and variation in fruit set within the inflorescence is attributable to competition among flowers within the inflorescence for limited resources. Consequently, it was concluded that, under natural conditions, the early blooming flowers located at lower positions of the inflorescences obtain more resources and produce more fruits than the late blooming flowers located at higher positions in M. roseum .     

Unpredictability of nectar nicotine promotes outcrossing by hummingbirds in Nicotiana attenuata

Many plants use sophisticated strategies to maximize their reproductive success via outcrossing. Nicotiana attenuata flowers produce nectar with nicotine at concentrations that are repellent to hummingbirds, increasing the number of flowers visited per plant. In choice tests using native hummingbirds, we show that these important pollinators learn to tolerate high‐nicotine nectar but prefer low‐nicotine nectar, and show no signs of nicotine addiction. Nectar nicotine concentrations, unlike those of other vegetative tissues, are unpredictably variable among flowers, not only among populations, but also within populations, and even among flowers within an inflorescence. To evaluate whether variations in nectar nicotine concentrations increase outcrossing, polymorphic microsatellite markers, optimized to evaluate paternity in native N. attenuata populations, were used to compare outcrossing in plants silenced for expression of a biosynthetic gene for nicotine production (Napmt1/2) and in control empty vector plants, which were antherectomized and transplanted into native populations. When only exposed to hummingbird pollinators, seeds produced by flowers with nicotine in their nectar had a greater number of genetically different sires, compared to seeds from nicotine‐free flowers. As the variation in nectar nicotine levels among flowers in an inflorescence decreased in N. attenuata plants silenced in various combinations of three Dicer‐like (DCL) proteins, small RNAs are probably involved in the unpredictable variation in nectar nicotine levels within a plant.     

Flower choice by honey bees (Apis mellifera L.): sex-phase of flowers and preferences among nectar and pollen foragers

Bees foraging for nectar should choose different inflorescences from those foraging for both pollen and nectar, if inflorescences consist of differing proportions of male and female flowers, particularly if the sex phases of the flowers differ in nectar content as well as the occurrence of pollen. This study tested this prediction using worker honey bees (Apis mellifera L.) foraging on inflorescences of Lavandula stoechas. Female flowers contained about twice the volume of nectar of male flowers. As one would predict, bees foraging for nectar only chose inflorescences with disproportionately more female flowers: time spent on the inflorescence was correlated with the number of female flowers, but not with the number of male flowers. Inflorescence size was inversely correlated with the number of female flowers, and could be used as a morphological cue by these bees. Also as predicted, workers foraging for both pollen and nectar chose inflorescences with relatively greater numbers of both male and female flowers: time spent on these inflorescences was correlated with the number of male flowers, but not with the number of females flowers. A morphological cue inversely associated with such inflorescences is the size of the bract display. Choice of flowers within inflorescences was also influenced predictably, but preferences appeared to be based upon corolla size rather than directly on sex phase.     

Pollen and nectar as a reward in the basal epidendroid Psilochilus modestus (Orchidaceae: Triphoreae): A study of floral morphology, reproductive biology and pollination strategy

Psilochilus modestus Barb. Rodr. is a basal epidendroid orchid occurring in both the semi-deciduous and Atlantic rain forests of the state of São Paulo, southeastern Brazil. This species presents a perfect flowering synchrony within populations, since all the mature buds of each plant open simultaneously in the morning hours of the same day. These flowers are available only for 1 day and are pollinated by several species of small solitary and social native bees. These bees exploit both pollen and nectar as a reward. The bees collecting pollen promote a higher fruit set and perform mainly self-pollination while those collecting nectar, which are less numerous, are responsible for an increase in cross-pollinations. P. modestus is self-compatible but pollinator dependent. Natural fruit set (open pollination) is low when compared with the numbers obtained under manual self- and cross-pollination. Low fruit set in natural conditions is related to deficient pollen transfer, and pollinator inefficiency seems to be the main factor. Some factors, such as the small amount of nectar produced, the low number of flowers per inflorescence and their availability for 1 day only added to the perfect flowering synchrony seem to be responsible for the increase of cross-pollinations. The offering of both pollen and nectar as a reward can represent a transitional condition in basal Epidendroideae. Based on floral morphology, reward production and pollinator behavior, the relationship of P. modestus with the basal and most derived groups within Orchidaceae is discussed.     

SEX DIFFERENTIAL FLORAL LONGEVITY,NECTAR SECRETION,AND POLLINATOR FORAGING IN A PROTANDROUS SPECIES

Sex differential nectar production, floral longevity and pollinator foraging were examined in Lobelia cardinalis, a self-compatible, protandrous species that is hummingbird pollinated. The staminate phase of the flowers lasts significantly longer and produces significantly more nectar (total sugar) per day than the pistillate phase of the flowers. Additional pollen is presented throughout the staminate phase. Because inflorescences of L. cardinalis mature acropetally, the nectar reward on any given day is greatest at the top of the inflorescence (where staminate phase flowers are located). Hummingbirds appear to be sensitive to this pattern of nectar presentation as they most commonly began foraging in the middle of an inflorescence and proceeded upward. This foraging pattern tends to promote outcrossing and suggests that staminate phase flowers are visited more often than pistillate phase flowers. We conclude that L. cardinalis emphasizes the male function at anthesis. Others have hypothesized that the features of this species are a logical consequence of intrasexual selection, but further research is needed before we place great confidence in a sexual selection interpretation of our data.