西藏高原高寒植物长鞭红景天生殖特征成功及影响因子

为了探索高寒植物长鞭红景天(Rhodiola fastigiata)濒危的生殖生态学机理,寻找保护和恢复该物种的有效途径,对其繁育系统、开花物候特征、生殖过程(开花、结实、种子产量和发育)及其影响生殖成功的主要因子进行调查分析。结果表明:长鞭红景天包括两性植株和雌性植株两种类型,其中两性植株仅执行雄性功能,种群繁殖为异交,属风媒传粉方式。种群花期出现在1a中降雨和气温较高的季节,6月下旬开始开花,7月下旬结束,时间30d左右。单株开花动态呈现单峰和双峰曲线两种模式,两性植株花期(9.29d)大于雌性植株花期(7.06d),并先于雌性植株开放,但雌性植株开花的高峰期比两性花早。从花蕾到开花,雌性单株有16%—25%的花蕾损失;开花到结实,单株36%—57%的花败育;至种子成熟,共有84%—94%的胚珠损失,仅6%—16%的胚珠发育成种子。雌性单株平均种子产量估计值约为285粒,说明长鞭红景天生殖潜力较大。人工条件下种子发芽率为93.6%,可见种子质量不是导致其濒危的原因,花粉限制和种子产量可能是影响生殖成功的主要因素。     

濒危植物金花茶开花物候和生殖构件特征

对桂林雁山迁地保护的金花茶(Camellia nitidissima)开花物候和生殖构件特征进行了研究.结果表明:(1)单花开放过程可分为蕾松、初开、盛开、落花4个阶段,单花寿命约2～8 d;(2)个体花期长,始花时间变异大,单株花期平均为37 d,始花时间可相差2个多月,种群始花时间为1月上旬,3月上旬达到开花高峰期,花期持续近3个月;(3)生殖构件的数量与个体径级和树冠层次有一定关系,地径大于6 cm的个体,其单株花数、生殖枝数、生殖枝比率显著高于地径小于4 cm和4～6 cm个体,而每生殖枝花数则无显著差异;不同树冠层次之间,平均生殖枝数和生殖枝比率,以中、下层显著高于上层,而每生殖枝花数以上层最高,花数在不同冠层之间无显著差异;(4)花芽期败育率为16.33%,花期败育率为63.05%,果期败育率为44.09%,败育率在不同径级个体和不同冠层之间有一定差异;败育率高,结实率低,种子产量少,是金花茶濒危的一个主要原因;(5)开花数目与花期长度呈正相关,个体始花时间与花期长度、座果率之间存在显著的负相关关系.     

两种互逆资源梯度影响下刺叶锦鸡儿的种子选择性败育格局

种子的选择性败育在被子植物中普遍存在,开展结籽格局及其影响因素的研究有助于深入了解种子的形成机制及多样化的生殖对策。以刺叶锦鸡儿Caragana acanthophylla为材料,通过传粉过程、胚珠发育动态和资源分配状况的研究,以探讨种子的选择性败育格局及相关影响因素。结果显示：(1)刺叶锦鸡儿具高度自交不亲和性,为泛化的传粉系统,蜂类是主要的传粉者。自然状态的结实率为(86.00±4.96)%,不存在传粉限制,但受精具明显的时间效应。(2)刺叶锦鸡儿单花期4—5d,每朵花有(14.00±0.14)粒胚珠。胚珠的发育从荚果顶部开始,其中,开花后第3天荚果顶部胚珠开始膨大;第11天绝大多数胚珠出现了膨大,此时,在荚果基部的胚珠开始败育,随后荚果顶端受精后的胚珠也出现败育,最终仅荚果中部形成2—3粒种子。其结籽格局为成熟荚果中部胚珠,而败育荚果顶部和基部胚珠的选择性败育类型。(3)受微地形影响,居群内开花植株具斑块分布,对较少开花植株通过添加水肥进行资源调控后,结籽率显著提高,说明在种子形成过程中存在资源限制。综上所述,受精顺序和资源限制两个互逆的资源梯度决定了刺叶锦鸡儿的结籽格局。其中,...     

濒危植物凹叶厚朴的花部综合特征和繁育系统

为了解凹叶厚朴(Magnolia officinalis subsp.biloba)濒危的生殖生物学原因,通过野外观测,运用杂交指数、花粉-胚珠比、花粉活力和柱头可授性观测、人工授粉和套袋实验等方法,对凹叶厚朴种群的开花生物学、繁育系统进行了研究。结果表明:凹叶厚朴单花花期一般为4～5d,单花花期根据形态和散粉时间可分为4个时期:蕾期、初开期、盛开期和凋零期;种群于4—5月开花,一般开花历时30d左右,但不同地点、年份间有所差异;种群无开花高峰,单株平均开花数量为171.4～188.8朵,平均开花振幅为5～6朵·株-1·d-1;凹叶厚朴杂交指数为4,花粉胚珠比为5728.86,结合人工套袋和授粉实验结果可确定,繁育系统为异交,传粉过程需要传粉者;花粉活力较高(94%),且花粉活力可维持至散粉后的第6天。散粉后,柱头可授性显著下降,花粉高活力期与柱头可授期有5～6h的重叠;自然状况下结果率和单果出种百分率低,异株异花授粉可明显提高结果率和单果出种百分率,二者相对自然状况分别提高14.25%和43.66%。表明自然状态下传粉效率低、柱头最佳可授期短、同株自花授粉败育严重是其濒危的主要生殖生物学原因。     

大血藤生殖生物学研究

通过野外观测、实验室分析以及人工套袋授粉等方法,研究了大血藤的繁育系统、开花习性、传粉方式以及花粉-胚珠比等生殖生物学特征.结果表明,大血藤的花为单性花,雌雄同株.雄花单花花期为7~10 d,雌花单花花期为2~3 d,种群开花历时14~17 d,散粉高峰期为4月11~17日,属风媒传粉植物.大血藤花粉-胚珠比为600±20,属兼性异交类型,人工套袋实验进一步证明,大血藤的交配系统为兼性异交类型,但以异交为主.     

濒危植物长柄双花木自然种群生殖构件的时空动态

根据野外观测数据,研究了濒危植物长柄双花木(Disanthus cercidifolius var.longipes)在不同生境条件下自然种群开花物候变异以及生殖构件的时空动态.结果表明,不同群落类型中长柄双花木开花物候存在一定差异,在纯林中个体的平均花期最长,达75d,而个体间开花同步性最低,仅为0.717;竹林中的个体花期最短,只有47d,而同步指数最高,达0.968.生殖枝数量及花序数均随个体年龄增长而增加,在年龄为30～35年时达最高,此后又呈下降趋势;在海拔为810m左右处,个体生殖枝数及着生的花序数均达最大值,且冠层间差异明显.纯林中个体着生的生殖枝数及每生殖枝上着生的花序数均最高,分别为411.39和7.857;竹林中个体每生殖枝着生的花序数显著低于其它群落.年龄小的个体,各时期生殖构件败育率均高于年龄大的个体;另外,生殖构件的败育与海拔高度间无相关性,而与构件发育时间、分布的树冠层次以及所处的群落类型相关.影响生殖构件败育的主要因子是光照条件.     

濒危植物长柄双花木自然种群结实的花粉和资源限制

长柄双花木（Disanthus cercidifolius Maxim．vsr．longipes H．T．Chang）只零星分布于湖南、江西和浙江等省的少数县,已处于濒危状态。通过对其野外自然种群试验,研究了花粉和资源的有效性对雌性生殖的影响,并进一步探讨了长柄双花木“花多果少”的生殖机制。结果表明：不同来源的花粉对长柄双花木结果率及结籽率均产生显著影响。花粉来源而不是花粉数量对长柄双花木结实存在显著影响。自然条件下,长柄双花木“花一果转化率”约为54馏,“胚珠-种子转化率”约为83．19。施肥处理有利于花芽发育,可使植株花芽败育率明显下降,开花比例升高;同时座果率和结籽率均显著提高,而果实败育率则显著降低。随剪除叶片量的增加,结果率明显降低,但果实败育率在处理间无显著差异;剪除弱小枝条及病虫枝后,个体总开花数虽有所下降,但是结果率和结籽率均显著提高。去除花后,长柄双花木平均单果重、单粒种子重均显著降低。营养水平的时空异质性使自然种群结籽率受到资源和花粉的限制;补充资源可以通过增强花对传粉者的吸引而间接影响传粉。长柄双花木自然种群的坐果率极低,而且存在严重的“大小年”结果现象,不同的因子可能对其低水平坐果率产生相互作用。选择性败育、子房供应和雄性功能假说似乎是长柄双花木“花多果少”的生殖策略的最合理解释。     

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。     

大花百子莲的开花物候与生殖特性

连续2年对引种植物大花百子莲在中国上海市人工种群的开花物候、结实情况进行观测记录,从不同水平分析了大花百子莲的开花物候对其生殖的影响。结果显示:（1）大花百子莲在引种地的开花时间为6月初至7月末;种群花期历时40 d左右,单株花期在14～31 d波动,单花寿命主要为2～3 d;2年的开花振幅平均在6朵·株^-1·d^-1左右,花朵开放百分率值在3.69%~4.87%左右,平均开花式样为11~17朵。（2）大花百子莲种群内2007年比2008年的各项开花物候指标均提前6～9 d;种群开花进程为渐进式的双峰曲线;种群花期重叠时间较长,2年的平均开花同步指数分别为0.663、0.695;植株个体相对开花强度主要分布频度为50%～70%。（3）开花物候指数与座果率的相关分析表明,始花时间与开花数目、座果数、花期长度呈显著负相关关系,开花数目与座果数、花期长度以及花期长度与座果数均呈显著正相关关系,同一花序内单花所发育成果实的数量与单花开放时间呈极显著负相关关系。研究表明,大花百子莲始花时间早的植株比始花时间晚的植株花期更长,开花数目多的植株比开花数目少的植株花期更长,花期长的植株比花期短的植株座果率更高;而且在同一花序上,越早开放的单花其结实的可能性越大。作为引种植物,大花百子莲开花物候年度间具有相似性,说明其在引种地可以适应环境压力,形成稳定的开花时间与开花模式;较长的花期重叠和大量的开花数目可以吸引较多的传粉者访问,为其生殖成功奠定了良好基础。     

双岛湾大叶藻的有性生殖特征

对威海双岛湾大叶藻(Zostera marina L.)种群的有性生殖特性进行了研究。结果表明:该水域中,大叶藻种群于3月下旬或4月上旬进入生殖枝始现期,5月上旬进入始花期,6月下旬进入种子散落高峰期,7月上旬进入生殖枝死亡始期,7月下旬生殖期结束,其有性生殖物候进程与水温密切相关,15℃是其开花适宜水温;生殖枝的侧枝数和扇状聚伞花序数在进入始花期时达到最大,分别为4.4和4.9个·株-1,佛焰苞数和生殖枝高度均在种子散落始期最大,分别为18.6个·株-1和96.2 cm,种子败育率较高,平均为40%,成熟种子产量为44.8粒·株-1;生殖枝密度在始花期时达到最大,为64株·m-2,生殖枝分化率平均仅有12.6%,有性生殖期间营养枝密度和种群总密度的下降幅度超过40%;生殖枝生物量在初果期达到最大(27.9 g·m-2),从始花期至种子散落高峰期平均种群生物量分配格局为地下部38.4%,营养枝52.4%,生殖枝9.2%,生殖分配较低,成熟种子产量为1859粒·m-2。     

Pollen and Resource Limitation in Veratrum nigrum L. （Liliaceae）, an Andromonoecious Herb

Pollen limitation and resource limitation were invoked to account for the pattern that flowering plants produce more flowers and ovules than fruits and seeds. This study aimed to determine their relative importance in Veratrum nigrum, a self-compatible, perennial, andromonoecious herb. In order to determine whether female production was limited by pollen grains on stigmas or by available resources, we performed supplemental hand pollination in three populations, male-flower-bud removal in three other populations, and emasculation of hermaphroditic flowers in still another population, resulting in a total of seven populations experimentally manipulated. Across the three populations, supplemental hand pollination did not significantly increase fruit set, seed number per fruit, and total seed production per individual, nor did emasculation of hermaphroditic flowers. Taken together, our results suggest that pollen grains deposited on stigmas were abundant enough to fertilize all the ovules. Male-flower-bud removal significantly increased the mean size of hermaphroditic flowers in all three populations. Female reproductive success was increased in one population, but not in the other two populations possibly due to heavy flower/seed predation. We concluded that the female reproductive success of V. nigrum was not limited by pollen grains but by available resources, which is consistent with Bateman＇s principle. Furthermore, the female reproduction increase of male-flower-bud removal individuals might suggest a trade-off between male and female sexual functions.     

Accessory costs of seed production

Accessory costs of reproduction are those that are necessary to mature a seed, but that do not involve the direct cost of provisioning the seed itself. This study aims to quantify accessory costs in a range of species, and test whether they decrease as a proportion of total reproductive expenditure with increasing seed mass, as might be expected if economies of scale came into play at larger seed sizes. We also test whether accessory costs varied with growth form, pollination mode, and dispersal mode, with the expectation that biotic pollination and dispersal modes should incur greater costs. Reproductive allocation (dry biomass) over one season, was calculated for 14 diclinous angiosperm species. Accessory costs averaged 73% of total reproductive allocation, with the majority spent on packaging and dispersal. Total accessory costs, packaging and dispersal costs, and costs incurred prior to pollination were proportional to direct costs of reproduction in major axis regressions. However, larger seeded species incurred significantly greater costs associated with aborted seeds and fruits, and matured a smaller proportion of ovules. This is consistent with larger seeded species being more selective of the ovules/embryos matured than small-seeded species. Total accessory costs, and proportion of ovules aborted, were also significantly greater for biotically dispersed species, but only due to an association with larger seed masses. Costs associated with abortions were lower for biotically pollinated species, due to a general trend of more ovules per ovary, resulting in greater cost sharing. This study demonstrates that expenditure on items other than seeds accounts for the majority of reproductive allocation in flowering plants. Yet, far more literature exists on seed mass variation than on investment in accessory structures. We found a proportional relationship between accessory costs and seed mass that warrants further investigation within the context of selection on margin returns on investment.     

峨眉拟单性木兰的开花生物学特性与繁育系统

为阐明峨眉拟单性木兰(Parakmeria omeiensis)在自然条件结实率低的原因和确定最佳人工授粉时期,该文通过观察峨眉拟单性木兰的开花动态,采用杂交指数估算、花粉胚珠比、花粉活力及柱头活性检测、人工授粉试验等方法对其繁育系统进行了研究。结果表明:(1)植物园保育的峨眉拟单性木兰花期在4月底到5月中下旬,持续17～23 d,雄株始花期比雌株早3～4 d,但两者花期可遇。(2)两性花经检测雄蕊败育,实为功能上的雌性,部分雄株个体的雄花上残留1～2个心皮,其性别分化是通过雌、雄蕊选择性败育形成的,为隐性雌雄异株(cryptic dioecy)。(3)雄花、两性花开放经历佛焰苞开裂、花被片开裂、展开、闭合、二次开放、凋落6个阶段,历时4 d。(4)雄花初次展开时花粉活力最高,达92.8%,开花2 d后活力显著下降;两性花柱头在花被片展开期可授性最强,盛开后柱头部分可授。(5)杂交指数为5,P/O比为2.14×10~4。(6)套袋试验表明,峨眉拟单性木兰不能进行自花传粉,人工异花授粉的结实率和出种数显著高于自然授粉,且不存在无融合生殖。这说明峨眉拟单性木兰繁育系统为专性异交,传粉过程需要传粉媒介,自然条件下结实率低,主要是受传粉昆虫和柱头可授期短的限制。     

Variation in floral sex allocation, female success, and seed predation within racemiform synflorescence in the gynomonoecious Ligularia virgaurea (Asteraceae)

Studies of diclinous species have showed that floral sex allocation and female reproductive success were quite variable within inflorescences. However, little attention has been paid to gynomonoecious species, in which individuals produce both female and bisexual flowers. The purpose of this study was to investigate the variations in reproductive patterns at different capitulum positions within racemiform synflorescence in Ligularia virgaurea, and to determine selective mechanisms of variations in reproductive patterns. We conducted observational and experimental studies in natural populations of the gynomonoecious composite L. virgaurea. Floral sex allocation, seed production and pre-dispersal seed predation were quantified in the field. The results showed several patterns of variation from top to bottom capitula, including an increase in bisexual flowers and flower number per capitulum, but a decrease in seed set and size. Removing earlier capitula during bud stage did not change floral sex allocation in later capitula. And no effect was found on seed set under supplemental pollination. Thus, although it has been reported many times in previous studies, the variation of floral sex allocation in L. virgaurea may not result from architectural effect or mating environment, and the variation of seed production could not be fully explained by pollination success. Additionally, our results showed that L. virgaurea was susceptible to high levels of bisexual biased predation, which was greater for top capitula. We therefore suggest that these variations may help to enhance reproductive success of L. virgaurea in the face of bisexual-biased seed predation.     

Pollination ecology of Isoglossa woodii,a long‐lived,synchronously monocarpic herb from coastal forests in South Africa

Synchronous monocarpy in long‐lived plants is often associated with pollination by wind, in part because infrequent mass flowering may satiate pollinators. Selfing in synchronous monocarps may provide reproductive assurance but conflict with the benefits of outcrossing, a key evolutionary driver of synchrony. We predicted that animal‐pollinated species with synchronous flowering would have unspecialised flowers and attract abundant generalised pollinators, but predictions for selfing and outcrossing frequencies were not obvious. We examined the pollination biology of Isoglossa woodii (Acanthaceae), an insect‐pollinated, monocarpic herb that flowers synchronously at 4–7‐year intervals. The most frequent visitor to I. woodii flowers was the African honeybee, Apis mellifera adansonii. Hand‐pollination failed to enhance seed production, indicating that the pollinators were not saturated. No seed was set in the absence of pollinators. Seed set was similar among selfed and outcrossed flowers, demonstrating a geitonogamous mixed‐mating strategy with no direct evidence of preferential outcrossing. Flowers contained four ovules, but most fruits only developed one seed, raising the possibility that preferential outcrossing occurs by post‐pollination processes. We argue that a number of the theoretical concerns about geitonogamous selfing as a form of reproductive assurance do not apply to a long‐lived synchronous monocarp such as I. woodii.     

Gender variation of sequential inflorescences in a monoecious plant Sagittaria trifolia (Alismataceae)

In protogynous plants, female flowers of early blooming plants are at a reproductive disadvantage because they cannot set fruit due to the lack of available pollen. To study this phenomenon, gender expression of the monoecious herb Sagittaria trifolia was investigated over the entire flowering season in two field and two cultivated populations in Hubei and Hunan Provinces, China. In racemes of S. trifolia, flowers open sequentially from bottom to top, with female flowers opening first followed by male flowers. This creates a temporal separation of sexes in the species. Under field conditions small plants are often male, with production of both male and female flowers increasing with plant size. Femaleness increased among sequential inflorescences since female flower production increased whereas male flower production did not. Seed production was greater in large inflorescences because they contain more female flowers, and the number of ovules increased in female flowers at basal positions within the raceme. A consistent pattern of high seed set was observed in flowers from both field and cultivated populations. About 1 % of unfertilized ovules resulted from no pollination and 2 % of the seeds produced were only partly developed due to resource limitation. In the first inflorescence of the six experimental populations, 6.7-40.0 % of individuals produced only male flowers, and female flowers of 1.9-6.5 % individuals were aborted. The occurrence of male flowers in early blooming inflorescences could be an adaptive strategy to conserve resources and enhance pollination of female flowers in protogynous S. trifolia.     

Are mismatches the norm? Timing of flowering,fruiting, dispersal and germination and their fitness effects in Frangula alnus (Rhamnaceae)

The close morphological and temporal links between phases of plant growth and reproduction call for integrated studies incorporating several reproductive phases from flowering to recruitment, and associated plant‐animal interactions. Phenological strategies, as well as plastic phenological response to climate change, incorporate complex interactions between developmental constraints, pollination and seed dispersal. Relationships between reproductive phenology and components of fitness were studied for two years in the north‐temperate, self‐incompatible, insect‐pollinated, and bird‐dispersed shrub Frangula alnus (Rhamnaceae). Fruit set, dispersal, germination and juvenile survival, as well as seed mass and juvenile size were measured in relation to flowering, fruiting and germination time. The results suggest that effects of flowering and fruiting time prevailed in subsequent phases, to some extent as far as to the juvenile phase, but effects of timing were complex and had partly opposing effects on different fitness components. Early flowers had higher fruit‐set and experiments indicated that synchronous peak flowering increased fruit‐set, but later flowers had higher seed mass. Peak fruiting was not associated with peak dispersal. Late fruits derived from late flowers promoted dispersal. Juvenile recruitment was enhanced by increasing seed size. We conclude that the phenology of flowering and fruiting in F. alnus comprises several features, each with different and sometimes counteracting effects on fitness components. From a general perspective, this result implies that we should not expect to find finely tuned matches in timing specifically between flowering and pollinators, and fruiting and seed dispersing birds.     

Variation of pollen limitation in the early flowering Mediterranean geophyte Narcissus assoanus (Amaryllidaceae)

A recent literature review indicates that pollen limitation of female fertility is a common feature of flowering plants. Despite the ecological and evolutionary significance of pollen limitation, most studies have only examined fertility in a single population at one time. Here we investigate pollen limitation of fruit and seed set in five populations of Narcissus assoanus, a self- sterile, insect-pollinated geophyte, over 2–3 years in southern France. In common with many early spring flowering plants, pollinator visitation to N. assoanus is often infrequent. Supplemental hand-pollination of flowers with outcross pollen significantly increased overall fruit and seed set by 11% and 19%, respectively. Four of the five populations experienced some pollen limitation during the study. For a given year, there was significant variation in pollen limitation among populations. Two of the populations were pollen limited in one year but not in other years in which they were studied. Seed:ovule ratios for open- and hand-pollinated flowers averaged 0.29 and 0.33, respectively. While hand pollination significantly increased the seed:ovule ratio, the low value obtained indicates that the majority of ovules in flowers do not mature seeds despite hand pollination. The role of genetic and environmental factors governing low seed:ovule ratios in N. assoanus is discussed. Received: 28 December 1999 / Accepted: 6 April 2000     

赤霉素诱导小桐子产生两性花

小桐子（Jatropha curcas）种子含油率高,油脂组成适合于生产生物燃油,是制备航空生物燃油和生物柴油的理想原料。小桐子是一种雌雄同株植物,雌雄花着生于同一花序,其花序中雌花比例很低,可能是其种子产量低的主要原因之一。本文研究用不同浓度的赤霉素外源喷施处理小桐子花序芽对其花和种子发育的影响。结果表明：外源赤霉素处理能够诱导小桐子产生两性花,且处理浓度越高,两性花数量越多;随着两性花数量的增加,雌花数量相应越少,但雌花与两性花数量之和在各处理和对照之间差异不显著,这表明赤霉素诱导出的两性花可能来源于雌花。另外,高浓度(500~1500mg·L-1)的赤霉素处理会导致小桐子的种子不能正常发育, 表现为每个果实中的种子数量、大小、单粒种子重量、每个果序的种子重量以及种子的含油量都显著减少。这些结果有助于深入理解赤霉素在小桐子花器官形成及种子发育过程中所发挥的生理作用,为从分子水平上对小桐子进行遗传改良、提高其种子产量奠定基础。