3种雌雄异熟樟科植物开花动态的比较研究

为了解樟科植物雌雄异熟的繁育系统特点,对3种樟科植物阴香(Cinnamomum burmannii)、紫楠(Phoebe sheareri)和浙江楠(Phoebe chekiangensis)雌雄异熟花的开花动态进行了比较研究.结果表明,3种植物雌性功能期的开始时间、雌性功能期和雄性功能期的时间分配有差异.3种植物的主...     

二重雌雄异型异熟物种色木槭3种表型花的细胞形态学研究

二重雌雄异型异熟,即雌雄功能按雄-雌-雄顺序分别表达,是有花植物中较为少见的异熟机制.槭属(Acer)大部分物种具有二重雌雄异型异熟的开花物候,且第一期雄花与第二期在形态上差别较大.为深入探讨花发育模式对花形态和开花物候的影响,本文应用石蜡切片技术和光镜技术,对色木槭(Acer mono)各个发育时期的3种类型花(雄花...     

元宝枫花芽分化及花开放过程研究

为探明元宝枫花性别、交配系统及花芽分化过程,通过石蜡切片和连续解剖观察,对元宝枫花开放及花芽分化过程进行了研究。结果表明：(1)元宝枫花性别可分为雌能花和雄能花两种,雌能花花药不开裂,只表现雌性功能,而雄能花分Ⅰ型和Ⅱ型两种,花药均可产生成熟花粉并开裂散粉,只表现雄性功能。(2)元宝枫是较为少见的二重雌雄异型异熟树种,且交配系统有雄先型和雌先型两种;雄先型小花开放顺序为：雄能花→雌能花→雄能花,雌先型小花开放顺序为雌能花→雄能花→雄能花。(3)元宝枫花芽的形态分化期开始于7月上旬,花序和小花分化期为8月上旬,雄蕊和雌蕊分化开始于8月下旬,雌配子体发育晚于雄配子体。(4)元宝枫花性别分化的关键期为第二年的3月下旬至4月上旬。     

雌全同株植物三脉紫菀花期偏雄的个体大小依赖的性别分配

经典的虫媒传粉植物个体大小依赖的性别分配模型通常预期:分配给雌性功能的资源比例将随着个体大小的增大而增加;但一些研究表明,花期个体大小依赖的性别分配模式表现出随个体大小增大而偏雄的趋势.我们以植株高度衡量个体大小,从花和花序两个水平上研究了雌花、两性花同株植物三脉紫菀(Aster ageratoides)花期个体大小依赖的性别分配策略.随着植株高度的增大,植株产生的头状花序数量增加,表明三脉紫菀投入到繁殖的资源不是固定不变的,而是随个体大小增大而增加的.在花和花序水平上,繁殖资源在雌雄性别功能之间的分配均表现为随个体大小的增大而更偏雄的模式,即花粉/胚珠比增加,产生花粉的两性花占两性花和雌花总花数的比例升高.这些结果与花期个体越大、性别分配越偏雄的预期一致.花期更偏雄的性别分配可能有助于植物在花期通过输出花粉提高雄性适合度,从而实现个体适合度的最大化.     

西藏极小种群野生植物密叶红豆杉种群的性比及雌雄空间格局

雌雄异株植物种群具有性比和雌雄个体空间分布格局, 这对个体成功繁殖、种群生存潜力、天然更新能力和遗传多样性的维持都是重要的, 对于珍稀濒危雌雄异株植物种群尤其如此。密叶红豆杉(Taxus fuana)为国家I级重点保护植物, 是首批列入《全国极小种群野生植物拯救保护工程规划》的120种极小种群野生植物之一, 具有重要的生态和经济价值。但目前关于密叶红豆杉种群生态学方面的研究, 尤其是性比结构与空间分布格局在国内外鲜有报道。本文对西藏吉隆地区的6个天然密叶红豆杉种群(吉普、多甫、朗久、吉隆、开热和唐蕃)进行了实地调查, 研究其性比结构及其空间分布格局。结果表明, 调查的6个种群中总计雄株1,651株, 雌株1,231株, 仅吉普(雄/雌 = 1.89)与吉隆(雄/雌 = 1.39)两个种群有显著的性别偏倚现象且显著偏雄性(P < 0.001)。6个种群雌雄植株间的空间相关性不强, 整体趋于相互独立。不同径级间, 吉隆和开热种群的性比格局相似, 均在小径级上性比显著偏雄性, 而吉普种群则在中等径级上显著偏雄性。综上, 不同密叶红豆杉种群的大小、性比、雌雄个体的大小级结构以及空间分布格局等均表现出不同。因此, 需要结合各个种群本身的发展动态、受干扰的类型以及各区域环境因子的差异进行有针对性的保护。     

多年生龙胆属植物个体大小与花期资源分配研究

于各物种花中前期对青藏高原东部高寒草甸6种多年生龙胆属植物花期的繁殖分配和性分配进行分析,结果表明:(1)多年生龙胆属植物的植株个体越大,繁殖投入越高,繁殖分配越低;(2)随着植物个体的增大,对雌性、雄性和吸引结构的投入都在增加,这可保证资源的充分利用,不会因为单一部分的增加而造成资源的浪费;(3)6种龙胆属植物中,有4种其性分配结果与性别分配(SDS)的理论预测一致,即大个体更偏向雌性器官的资源投入,但麻花艽(Gentiana atraminea)和达乌里秦艽(Gentiana dahurica)的性分配与个体大小则没有表现出负相关,可能与其本身具有的雌雄异熟———雄性先熟特点有关;(4)资源在雌雄功能间的分配没有表现出权衡关系,可能是由于植物必须在许多不同生活史性状之间进行资源分配,而不是两两之间非此即彼.     

浑善达克沙地黄柳雌雄群体年龄结构及动态特征

植物种群年龄结构和动态对于揭示其生活史特征和发展趋势具有重要意义,但针对雌雄异株植物两性群体的研究却鲜有报道。黄柳是浑善达克沙地常见的一种雌雄异株植物,也是该地区最重要的固沙先锋植物,该研究通过对浑善达克沙地黄柳天然种群的实地调查,采用年龄结构图、静态生命表、存活曲线、生存分析以及时间序列分析等方法对黄柳雌雄群体现状和未来变化趋势进行研究。结果表明:(1)浑善达克沙地黄柳雌雄群体在年龄结构及发展趋势方面均存在差异,雌株群体的年龄结构表现为稳定型,雄株群体的年龄结构表现为衰退型。(2)浑善达克沙地黄柳种群中雌株个体数量约占总数的70%,雄株约占30%,性比偏雌现象明显。(3)动态变化指数表明,研究区黄柳雌株群体呈现"前期缺乏、中后期稳定"的变化特征,雄株群体呈现"前期缺乏、中后期波动"的变化特征。(4)研究区黄柳雌雄群体的存活曲线都趋近Deevey-Ⅱ型,且与生存分析结果共同显示雌株群体比雄株群体进入衰退期的时间早。(5)时间序列分析表明,随着时间的推移研究区黄柳的老龄个体数量逐渐增多,幼龄个体缺乏是导致雌雄群体衰退的重要原因;建议通过人工扦插补充雌雄幼龄个体以及平茬复壮中、老龄个体的方法以保证该地区黄柳天然种群的正常繁衍。     

蒙古沙冬青花序内性分配的变化、传粉者运动与繁殖成功

性分配理论主要研究繁殖资源在雌雄功能间的最优分配,从雌雄功能的角度考虑其个体适合度.对花序内不同部位花的雌性与雄性资源分配变化的研究,对于我们理解植物采取哪种繁殖对策保障繁殖成功具有重要意义.本文对生长在中国科学院吐鲁番沙漠植物园内的蒙古沙冬青(Ammopiptanthus mongolicus)连续开花花序内不同部位...     

核桃揪雌雄异熟性的初步研究

对核桃揪开花特性进行了观察。结果表明：核桃揪多数植株为雌雄异熟,有的植株上雌花先开（雌先型）,有的植株上雄花先开（雄先型）。每一植株上雌雄异熟的次序是相当稳定的。并且多数能为雌先型植株提供授粉机会的雄先型群体,其雌花期与雌先型群体的雄花期也能较好的吻合。但是少数花期不同的同一类型（即：雌先型与雌先型或雄先型与雄先型）植株之间也有授粉机会。在坐果率、产量、核果的整齐度、放叶早晚、落叶期、形态差异的调查分析中未发现异熟性与这些性状上有必然的联系。     

柠条长角象雌雄个体的快速鉴别方法

【目的】柠条长角象Trigonorhinussp.是内蒙古地区新发现为害中间锦鸡儿Caraganaliouana Zhao Y. Chang&Yakovlev的蛀干害虫。为便于种群调查和分辨雌雄成虫,本文对柠条长角象雌雄性别的形态差异进行研究。【方法】通过测量柠条长角象成虫体长、体宽和第6腹板以及腹部末端和鞘翅末端是否完全覆盖腹部腹板等形态特征,进行雌雄成虫个体判别。【结果】采用独立样本t检验分析并比较发现,雌雄两性成虫体长（t=﹣0.294,P=0.771）、体宽（t=0.981,P=0.337）和体宽与体长之比（t=1.352,P=0.191）差异不显著。通过对柠条长角象雌雄成虫个体的体征观察发现,差异主要表现在：1）雌虫腹部末端于中央圆弧处凹入呈二裂形,雄虫腹部末端钝圆;2）雌性的第6腹板长度（0.48±0.01）mm明显较第5腹板长,而雄性（0.30±0.01）mm显著短,且雌虫第6腹板所占体长比例（0.16±0.01）mm大于雄虫所占体长比例（0.11±0.02）mm;3）雌虫鞘翅完全覆盖腹部末端,雄虫鞘翅未完全覆盖腹部末端。【结论】仅依据柠条长角象体型大小鉴别雌雄成...     

Influence of local floral density and sex ratio on pollen receipt and seed output: empirical and experimental results in dichogamous Alstroemeria aurea (Alstroemeriaceae)

Local density and sexual composition are two aspects of floral neighborhoods thought to influence pollination and seed output of recipient plants. I characterized the floral neighborhood of 436 flowering ramets of Alstroemeria aurea, a southern Andean perennial, distributed among three sites. On each ramet, I measured total pollen receipt and seed output. The long-lived, bumblebee-pollinated flowers of A. aurea are synchronously protandrous with a given ramet being either all male or all female and thus incapable of self or geitonogamous pollination at the ramet level. Even though each ramet changes sex over time, A. aurea forms floral neighborhoods that remain stable with respect to density and sex ratio during the span of a focal ramet female phase. Contrary to expectation, under field conditions neither local density nor sexual identity explained significant amounts of variation in pollen receipt. Density of neighboring flowering ramets marginally affected pollen receipt in two of the three populations but in opposite directions. Despite the absence of strong effects of neighborhood sexual composition on pollen receipt, the sexual identity of neighbors affected seed output which suggests effects on the quality of pollination due to changes in patterns of pollen flow. I also compared pollen loads on the stigmas of artificially isolated ramets (6 m) with those on experimental focal ramets surrounded by six close neighbors (20 cm) that were either all male or all female. Here, pollen receipt by focal ramets in all-male neighborhoods was 1.3 times greater than in isolated ramets, and 3.8 times greater than in ramets in all-female neighborhoods. In these artificial neighborhoods, stigmatic pollen deposition increased significantly over time. In nature, rates of bumblebee visits were higher in female-biased (early-flowering) than in male-biased (late-flowering) co-occurring floral patches. Thus, spatio-temporal shifts in visitation frequencies associated with the sexual composition of floral neighborhoods might compensate for spatial variability in pollen availability within populations and explain the discrepancies between empirical and experimental results. Received: 11 June 1996 / Accepted: 24 March 1997     

The Effects of Inflorescence Size and Flower Position on Biomass and Temporal Sex Allocation in <Emphasis Type="Italic">Lobelia sessiliflora</Emphasis>

To test the prediction of sex allocation theory that plants or flowers high in resource status emphasize the female function, we explored the variation in both biomass (the number of pollen grains and ovules) and temporal (male and female durations) sex allocation among and within plants of protandrous Lobelia sessilifolia in relation to plant size and flower position within plants. Among plants, the mean number of pollen grains and ovules per flower of a plant increased with plant size, whereas the mean P/O ratio (number of pollen grains/number of ovules ratio) decreased with plant size. The mean male duration, the mean female duration, and the mean ratio of male duration/flower longevity per flower of a plant were not correlated with plant size. Thus, large plants emphasized female function in terms of biomass sex allocation, which is consistent with the prediction of size-dependent sex allocation theory. The results for temporal sex allocation, however were inconsistent with the theory. Within plants, the mean number of pollen grains and ovules per flower at each position decreased from lower to upper flowers (early to late blooming flowers) and that of the P/O ratio increased from lower to upper flowers. The mean male duration and the mean female duration per flower decreased from lower to upper flowers, whereas the mean ratio of male duration/flower longevity increased from lower to upper flowers. The population sex ratio changed from male-biased to female-biased. Thus, later blooming flowers emphasized the male function in terms of both biomass and temporal sex allocation, consistent with the sex allocation theory, regarding the change in the population sex ratio.     

Effects of floral sexual investment and dichogamy on floral longevity

Aims Floral longevity, the duration that a flower remains open and functional, varies greatly among species. Variation in floral longevity has been considered to be optimal strategy for resource allocation under different ecological conditions, mainly determined by the rates of pollination and cost of flower maintenance. However, it is unclear whether an intrinsic factor, floral sexual investment, constrains evolution of floral longevity. The theoretical model also predicts that dichogamy favors long-lived flowers, but empirical studies to test this prediction remain unexplored.Methods To examine the effect of floral sexual investment on floral longevity, we measured flower size together with pollen and ovule production in 37 sympatric flowering plants in a natural community. The duration of the female and male phase in 21 protandrous species and floral longevity of the other 16 adichogamous species were documented in the field.Important findings Floral longevity varied from 1 day to 15 days, while pollen number per flower varied from 643 to 710880 and ovule number per flower from 1 to 426 in the 37 species. Flower size was correlated with pollen production as well as ovule production. Floral longevity was positively related to pollen production but not to ovule production. Consistent with the prediction that dichogamy favors long-lived flowers, we found the floral longevity of protandrous species was significantly longer than that of adichogamous species. In the protandrous species, pollen production per flower was observed to be positively related to male duration, while ovule production was not related to female duration. Our analyses of variation in floral longevity and sexual investment among different species suggest that the floral sexual investment could be an intrinsic factor contributing to the selected floral longevity, particularly the male phase, and that high pollen production could potentially increase pollen removal, i.e. male productive success.     

EFFECT OF DENSITY ON SECONDARY SEX CHARACTERISTICS AND SEX RATIO IN SILENE ALBA (CARYOPHYLLACEAE)

A field survey of plant and flower sex ratio and secondary sex characteristics was made in Silene alba. Female-biased plant sex ratios were found, as seems typical for the species. Sex ratio distribution correlated with a gradient of soil moisture (with the more moist area having a more female-biased ratio) and with changes in the density of Silene (intermediate and higher density areas having greater female bias). The floral sex ratio was significantly female-biased only at the site that was most female-biased in terms of plant sex ratio. Otherwise the population of flowers was significantly male-biased. Male and female plants harvested from the field differed in secondary sexual characteristics. Males had more flowers and invested proportionately more biomass in leaf, but less in root, stem and reproductive tissue than did females. Although both males and females were larger in terms of total dry weight at the moist site, males produced more flowers at the driest (high density) site. Here the female bias in plant sex ratio was intermediate, but the floral sex ratio was significantly male-biased. A glasshouse experiment was performed in which plants were grown at four densities. Density significantly influenced plant survivorship and the probability of flowering, and increased female bias in the pots, but it did not affect patterns of biomass allocation in flowering plants. Patterns of male and female biomass allocation did not differ in the experiment, except in terms of reproductive allocation (greater in females) and allocation to leaf, greater in males, but only at the lowest density. This work urges caution in interpreting differences between males and females in the field as secondary sex characteristics, since we find such properties to be overlapping under experimental conditions. It supports the idea that males and females of a species may sustain different reproductive output under differing conditions.     

Temporal variation in phenotypic gender and expected functional gender within and among individuals in an annual plant

Background and Aims

Adaptive explanations for variation in sex allocation centre on variation in resource status and variation in the mating environment. The latter can occur when dichogamy causes siring opportunity to vary across the flowering season. In this study, it is hypothesized that the widespread tendency towards declining fruit-set from first to last flowers on plants can similarly lead to a varying mating environment by causing a temporal shift in the quality (not quantity) of siring opportunities.

Methods

A numerical model was developed to examine the effects of declining fruit-set on the expected male versus female reproductive success (functional gender) of first and last flowers on plants, and of early- and late-flowering plants. Within- and among-plant temporal variation in pollen production, ovule production and fruit-set in 70 Brassica rapa plants was then characterized to determine if trends in male and female investment mirror expected trends in functional gender.

Key Results

Under a wide range of model conditions, functional femaleness decreased sharply in the last flowers on plants, and increased from early- to late-flowering plants in the population. In B. rapa, pollen production decreased more rapidly than ovule production from first to last flowers, leading to a within-plant increase in phenotypic femaleness. Among plants, ovule production decreased from early- to late-flowering plants, causing a temporal decrease in phenotypic femaleness.

Conclusions

The numerical model confirmed that declining fruit-set can drive temporal variation in functional gender, especially among plants. The discrepancy between observed trends in phenotypic gender in B. rapa and expected functional gender predicted by the numerical model does not rule out the possibility that male reproductive success decreases with later flowering onset. If so, plants may experience selection for early flowering through male fitness.     

Influence of inflorescence size on sexual expression and female reproductive success in a monoecious species

Sex allocation theory forecasts that larger plant size may modify the balance in fitness gain in both genders, leading to uneven optimal male and female allocation. This reasoning can be applied to flowers and inflorescences, because the increase in flower or inflorescence size can differentially benefit different gender functions, and thus favour preferential allocation to specific floral structures. We investigated how inflorescence size influenced sexual expression and female reproductive success in the monoecious Tussilago farfara, by measuring patterns of biomass, and N and P allocation. Inflorescences of T.?farfara showed broad variation in sex expression and, according to expectations, allocation to different sexual structures showed an allometric pattern. Unexpectedly, two studied populations had a contrasting pattern of sex allocation with an increase in inflorescence size. In a shaded site, larger inflorescences were female-biased and had disproportionately more allocation to attraction structures; while in an open site, larger inflorescences were male-biased. Female reproductive success was higher in larger, showier inflorescences. Surprisingly, male flowers positively influenced female reproductive success. These allometric patterns were not easily interpretable as a result of pollen limitation when na?vely assuming an unequivocal relationship between structure and function for the inflorescence structures. In this and other Asteraceae, where inflorescences are the pollination unit, both male and female flowers can play a role in pollinator attraction.     

Temporal floral sex allocation in protogynous Aquilegia yabeana contrasts with protandrous species: support for the mating environment hypothesis

We tested one of the predictions of Brunet and Charlesworth (1995) that relative floral sex allocation will vary temporally with the mating environment and that the form of dichogamy (protandry vs. protogyny) will select for the pattern of variation in male versus female resource allocation. In many hermaphroditic plant species, allocation to female function (ovule number) decreases from early to late flowers within inflorescences as a result of resource limitation or ontogenetic changes. This pattern may obscure the effects of the mating environment and dichogamy on selection for allocation patterns in protandrous species (male allocation increases regardless). By examining a protogynous species the alternative pattern of temporal variation in resource allocation is predicted, namely that allocation to male function should decrease (or female allocation increase) throughout the flowering sequence. This pattern was observed in protogynous Aquilegia yabeana (Ranunculaceae), in which ovule number per flower remained constant whereas pollen number decreased in sequentially blooming flowers. These observations support the temporal sex allocation hypothesis of Brunet and Charlesworth (1995).     

The effect of flower position on variation and covariation in floral traits in a wild hermaphrodite plant

Background  

Floral traits within plants can vary with flower position or flowering time. Within an inflorescence, sexual allocation of early produced basal flowers is often female-biased while later produced distal flowers are male-biased. Such temporal adjustment of floral resource has been considered one of the potential advantages of modularity (regarding a flower as a module) in hermaphrodites. However, flowers are under constraints of independent evolution of a given trait. To understand flower diversification within inflorescences, here we examine variation and covariation in floral traits within racemes at the individual and the maternal family level respectively in an alpine herb Aconitum gymnandrum (Ranunculaceae).     

Relationship between floral longevity and sex allocation among flowers within inflorescences in Aquilegia buergeriana var. oxysepala (Ranunculaceae)

Understanding the fitness of plants with inflorescences requires examining variation in sex allocation among flowers within inflorescences. We examined whether differences in the duration of the male and female phases of flowering lead to variation in sex allocation and reproductive success among flowers within inflorescences. In 2002 and 2003, we quantified floral longevity, floral sex allocation, and reproductive success between the first and the second flowers within inflorescences in a protandrous species, Aquilegia buergeriana var. oxysepala. Floral longevity was greater in the first flowers than in the second ones in both years. The male phase lasted longer, and the initial number of pollen grains and the number of pollen grains removed were greater in the first flowers than in the second ones in both years. Within first flowers, the number of pollen grains removed was greater in flowers that had longer male phases, thus duration of the male phase may positively affect male reproductive success in the first flowers. The female phase lasted longer and the number of ovules was greater in the first flowers than in the second only in 2002. However, seed production per flower and female phase duration in both years were not significantly related. The variation in the number of pollen grains among flowers in this species may be caused by the variation in male phase duration.     

Gender plasticity in Sagittaria sagittifolia (Alismataceae), a monoecious aquatic species

 Aquatic plants are well known for their high degree of phenotypic plasticity in vegetative structures, particularly leaves. Less well understood is the extent to which their sexuality can be modified by environmental conditions. Here we investigate gender plasticity in the European clonal monoecious aquatic Sagittaria sagittifolia (Alismataceae) to determine how floral sex ratios may vary with plant size and inflorescence order. We sampled two populations from aquatic habitats in East Anglia, U.K. and measured a range of plant attributes including ramet size and the number of female and male flowers per inflorescence. The two populations exhibited similar patterns of phenotypic gender, despite contrasting patterns of total allocation to female and male flower number. Plants produced male-biased floral sex ratios but female flower number increased from the first to the second inflorescence whereas male flower number decreased. Size-dependent gender modification occurred in both populations, but the patterns of allocation to female flower production differed between the two populations. Our results are consistent with the view that monoecy is a sexual strategy that enables plants to adjust female and male allocation in response to changing environmental conditions. Received September 16, 2002; accepted October 23, 2002 Published online: March 20, 2003