Pollen and resource limitations to lifetime seed production in a wild population of the endangered plant Disanthus cercidifollus var.Iongipes H.T.Chang (Hamamelidaceae)

Disanthus cercidifolius Maxim.var.longipes H.T.Chang,a plant species that only occurs in a few counties in Hunan,Jiangxi and Zhejiang Provinces and with a relatively small number of individuals,is ranked as a second Class endangered species for conservation in China.We have studied the effect of pollen and resources available to female reproduction,and the reproductive mechanism of "excess flowers with low fruit set" in Disanthus cercidifolius Maxim.var.longipes H.T.Chang was discussed.Results are as follows:Pollen from different sources has significant effects on fruit set and seed set of Disanthus cercidifolius Maxim.var.longipes H.T.Chang.The pollen source rather than pollen numbers significantly affected reproduction of this species.In wild populations,producing one fruit needs about 54.8 flowers,and one satiation seed needs about 6.60 flowers or 83.19 ovules.After fertilizing,which was propitious to flower development,the abortion rate of flower buds was decreasing,but the flowering rate was increasing.The fruit set and seed set was also significantly increasing,while abortion rate of fruit was significantly decreasing.With the increasing percentages of cutting leaves,the fruit set decreased,but the abortion rate of fruit shows no significant differentiation among treatments.After cutting branches that were puny,broken and insectinfested branches,the flower number seemed to be decreasing,but the fruit set and seed set all increased significantly.After removing some flowers,the fruit set was calculated with respect to the number of flowers remaining after the treatment increased with increasing of percentages of flower removal,whereas fruit set calculated with respect to the initial number of flowers remained constant,and the mean weights of per fi'uit and per seed all decreased significantly.Sufficient spatial or temporal heterogeneities in nutrient levels might allow limitation of seed set by resources and pollen in a natural population,while supplying resources may indirectly affect pollination by increasing attraction of the flowers to pollinators.There were very low fruit and seed sets in natural populations ofDisanthus cercidifolius Maxim.var.longipes H.T.Chang.Different factors may have interacted to effect a low fruit set.A joint adoption of the "selection abortion hypothesis","ovary reserve hypothesis" and "male function hypothesis" seems to be the most likely explanation for the reproductive strategy of"excess flowers with few fruit sets" in Disanthus cercidifolius Maxim.var.longipes H.T.Chang.     

Pollen and resource limitations to lifetime seed production in a wild population of the endangered plant Disanthus cercidifolius var. longipes H. T. Chang (Hamamelidaceae)

Disanthus cercidifolius Maxim. var. longipes H. T. Chang, a plant species that only occurs in a few counties in Hunan, Jiangxi and Zhejiang Provinces and with a relatively small number of individuals, is ranked as a second Class endangered species for conservation in China. We have studied the effect of pollen and resources available to female reproduction, and the reproductive mechanism of “excess flowers with low fruit set” in Disanthus cercidifolius Maxim. var. longipes H. T. Chang was discussed. Results are as follows Pollen from different sources has significant effects on fruit set and seed set of Disanthus cercidifolius Maxim. var. longipes H. T. Chang. The pollen source rather than pollen numbers significantly affected reproduction of this species. In wild populations, producing one fruit needs about 54.8 flowers, and one satiation seed needs about 6.60 flowers or 83.19 ovules. After fertilizing, which was propitious to flower development, the abortion rate of flower buds was decreasing, but the flowering rate was increasing. The fruit set and seed set was also significantly increasing, while abortion rate of fruit was significantly decreasing. With the increasing percentages of cutting leaves, the fruit set decreased, but the abortion rate of fruit shows no significant differentiation among treatments. After cutting branches that were puny, broken and insectin-fested branches, the flower number seemed to be decreasing, but the fruit set and seed set all increased significantly. After removing some flowers, the fruit set was calculated with respect to the number of flowers remaining after the treatment increased with increasing of percentages of flower removal, whereas fruit set calculated with respect to the initial number of flowers remained constant, and the mean weights of per fruit and per seed all decreased significantly. Sufficient spatial or temporal heterogeneities in nutrient levels might allow limitation of seed set by resources and pollen in a natural population, while supplying resources may indirectly affect pollination by increasing attraction of the flowers to pollinators. There were very low fruit and seed sets in natural populations of Disanthus cercidifolius Maxim. var. longipes H. T. Chang. Different factors may have interacted to effect a low fruit set. A joint adoption of the “selection abortion hypothesis”, “ovary reserve hypothesis” and “male function hypothesis” seems to be the most likely explanation for the reproductive strategy of “excess flowers with few fruit sets” in Disanthus cercidifolius Maxim. var. longipes H. T. Chang. __________ Translated from Acta Ecologica Sinica, 2006, 26(2): 496–502 [译自: 生态学报]     

濒危植物长柄双花木开花物候与生殖特性

研究了长柄双花木开花过程中花部表型的变化。连续4a对其野生种群、1a对人工种群的开花物候进行了观察,并运用相对开花强度和同步性等开花物候指数分析了开花物候对其生殖的影响。结果如下：长柄双花木开花时间为9月上、中旬至11月中、下旬;单花花期一般为6～7d,单花依其形态和散粉特征可以分为4个时期：散粉前期、散粉初期、散粉盛期和凋谢期。个体开花持续时间49～55d,种群花期历时63～71d。种群内不同年度间开花物候指数没有显著差异,而种群间则存在显著差异,野生种群开花进程为渐进式单峰曲线。人工种群则为“钟”形曲线,二者均属于“集中开花模式”。长柄双花木具有2个相对开花强度的分异趋势,这种分异趋势具有进化意义。开花物候指数与生殖间的相关分析表明,始花时间与开花数量、座果率及花期长度之间均具显著负相关关系,而开花数量与花期长度之间则呈显著正相关,但均为线性相关。长柄双花木开花物候在种群间的差异和种群内年度间的相似性说明,其开花时间可能是由与其相关的复杂的微生境特征和(或)由其遗传因子决定的,同时也反映了种群间的遗传分异和种群内个体间的遗传一致性。作为一种濒危物种,长柄双花木在这种环境的选择压力之下,形成了“大量、集中开放的花”的开花模式,吸引到更多的传粉者的访问,从而达到生殖成功。     

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。     

Isolation and characterization of nine polymorphic microsatellite loci in the endangered shrub Disanthus cercidifolius var. longipes (Hamamelidaceae)

Nine polymorphic microsatellite loci were isolated and characterized from an AC-enriched genomic library of Disanthus cercidifolius var. longipes. Microsatellite polymorphism was investigated using 24 individuals from one natural population. The observed number of alleles per locus ranged from two to four. Observed and expected heterozygosities ranged from 0.17 to 0.92 and from 0.16 to 0.72, respectively. These polymorphic microsatellite loci provide useful tools for the ongoing population genetic studies of D. cercidifolius var. longipes.     

华北驼绒藜自然种群结实的花粉和资源限制

运用人工授粉、补充无机营养、去叶处理以及疏花处理研究了华北驼绒藜自然种群花粉和资源有效性对结实的影响,并进一步探讨了该植物的选择性败育现象和繁殖对策。结果表明:花粉来源而不是花粉数量对华北驼绒藜结实存在显著影响;同枝授粉结籽率和结实率均低于自然授粉,异株授粉则显著高于同枝授粉和自然授粉;异株授粉种子千粒重显著高于自然授粉和同株异枝授粉,极显著高于同枝授粉。华北驼绒藜结实存在资源限制,补充施肥提高了单枝开花数和结籽率。随摘除叶片数量的增加,结籽率明显降低,摘除叶片处理与对照之间均形成显著差异,摘除1/2叶、3/4叶以及去全叶处理使种子千粒重显著低于对照。人工疏花实验表明华北驼绒藜存在选择性败育现象。华北驼绒藜可根据花粉和资源的可利用性来调整性分配、授粉方式和结实以获得最大适合度,具有复杂的繁殖对策,表现出较好的环境适应性。     

长柄双花木光合功能对光强的适应

在光强较高的纯林和阔叶林内,长柄双花木净光合速率呈“双峰曲线”,并伴随有胞间CO2的下降和气孔限制值的增大;随着日间光强的增大,叶片PQ还原程度和非光化学猝灭（NPQ）增加,ΦPSⅡ则下降,但后者午后随着光强的下降可得到完全恢复;叶片吸收光能分配于光化学反应的比例随着光强的增大而迅速减少;而天线色素热耗散部分及反应中心过剩的光能则显著增加,在午后光强减弱后两者均可恢复到早晨的初始水平。在低照度光的竹林内,净光合速率呈现“单峰曲线”,叶片PQ还原程度、NPQ以及ΦPSⅡ均无明显变化,叶片吸收光能也主要用于光化学反应,热耗散及过剩光能比例均较低。     

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.     

Genetic Variability and Population Structure of Disanthus cercidifolius subsp. longipes (Hamamelidaceae) Based on AFLP Analysis

Disanthus cercidifolius subsp. longipes is an endangered species in China. Genetic diversity and structure analysis of this species was investigated using amplified fragments length polymorphism (AFLP) fingerprinting. Nei''s gene diversity ranged from 0.1290 to 0.1394. The AMOVA indicated that 75.06% of variation was distributed within populations, while the between-group component 5.04% was smaller than the between populations-within-group component 19.90%. Significant genetic differentiation was detected between populations. Genetic and geographical distances were not correlated. PCA and genetic structure analysis showed that populations from East China were together with those of the Nanling Range. These patterns of genetic diversity and levels of genetic variation may be the result of D. c. subsp. longipes restricted to several isolated habitats and “excess flowers production, but little fruit set”. It is necessary to protect all existing populations of D. c. subsp. longipes in order to preserve as much genetic variation as possible.     

濒危植物长柄双花木(Disanthus cercidifolius var. longipes)叶功能性状随生长发育阶段的变化

对珍稀濒危植物长柄双花木(Disanthus cercidifolius var. longipes H. T. Chang)不同生长发育阶段的叶厚度(LT)、叶面积(LA)、叶含水量(LWC)、比叶面积(SLA)、叶片全氮含量(LNC)和叶片全磷含量(LPC)等叶功能性状及性状症候群的变化进行了研究。结果显示,不同发育阶段长柄双花木的叶功能性状存在显著差异,其中LT、LA随植株发育呈递增趋势,而LWC、SLA、LNC和LPC则呈递减趋势。主成分分析显示,随着长柄双花木的不断发育,性状症候群朝着LT、LA增大而LWC、SLA、LNC和LPC减小的方向变化。表明该物种的经济策略随着发育进程由快收益向慢收益转变。     

濒危植物长柄双花木自然种群年龄结构及其生态对策

通过对濒危植物长柄双花木自然种群的年龄结构、图解生命表以及生殖价分析等途径,研究了其种群结构动态及生态对策,结果表明:长柄双花木种群为衰退型种群,虽然种群中也存在一定数量的幼龄个体,但个体死亡率较高。长柄双花木种群表现出r～K对策的混合生态适应策略。该种群数量在下降过程中还存在波动,仍具有一定的实现生殖和恢复的可能,但依靠有性生殖途径实现恢复的能力有限。     

濒危植物猪血木（Euryodendron excelsum H. T. Chang）自然种群结构及动态

猪血木(Euryodendron excelsum H. T. Chang)是中国特有的山茶科单型属猪血木属的珍稀濒危植物,目前只在广东省阳春县八甲镇有分布且种群数量不足200株.通过样地调查和数据统计,绘制了猪血木种群的高度结构和径级结构图,在此基础上编制了种群的特定时间生命表,分析了存活曲线、死亡率曲线和寿命期望等重要参数,并运用时间序列模型预测种群数量动态.结果表明:猪血木种群幼年个体丰富,中老年个体相对较少,受环境因素和人为干扰的影响,种群在第Ⅱ级出现死亡高峰,且只有少量幼年个体能进入成年阶段生长,个体平均生存能力的期望在第Ⅳ级最大,种群存活曲线属于Deevey-Ⅲ型,猪血木种群目前仍表现为稳定型种群.时间序列预测分析表明,猪血木种群具备一定的恢复潜能,故现有植株和生境斑块的保护是保持猪血木种群自然更新和进行种群恢复的关键因素.     

濒危植物大别山五针松母树林花粉传播规律

大别山五针松是典型的风媒传粉植物,花粉流是种群基因流的重要形式。用重力玻片法研究大别山五针松母树林的花粉飞散特征,分析了时间和空间对花粉飞散的影响,结合林分密度和气象因子探讨花粉飞散的特点。结果表明:大别山五针松从5月中旬开始传粉,散粉期为14d,盛粉期为3d,种植密度低的下坡花粉密度比密度大的上坡的要高。日散粉最大值出现在10:00～14:00,植株中上部花粉密度较高,距花粉源50m处花粉密度最高。建议对种植密度大的区域移栽以促进花芽生成,以保证结实率与提高种子品质。     

Pollen viability and limitation of seed production in a population of the circumpolar cushion plant, Silene acaulis (Caryophyllaceae)

Pollen viability among genders and limitation of female seed production in a natural trioecious population of the circumpolar cushion plant Silene acaulis was examined. Pollen viability was estimated by an in vitro pollen germination experiment. Both male and hermaphrodite flowers displayed large variation in pollen viability (0–53% in hermaphrodite and 0–54% in male flowers). There was a significant difference between genders in pollen viability: male plants had on average higher pollen viability than hermaphrodite plants. Resource and pollen limitation of seed production was studied by an experiment consisting of three treatments; (I) hand-pollination and removal of all other flowers on the cushion, (II) hand-pollination without removal of other flowers, and (III) open pollination without removal of flowers. Hand-pollination increased seed production, whereas removal of flowers had no effect on seed production. Abortion of pollinated ovules during seed development and seed mass did not differ among treatments. To control for effect of fruit number on seed production, data from naturally pollinated individuals was used. There was a positive correlation between both total number of seeds and fruit number, mean seed number per fruit and fruit number, respectively. These results indicate that seed production of 5. acaulis is mainly limited by pollen availability whereas resource competition between fruits is not important as a limiting factor. The possible role of male quality differences between genders and pollen limitation of seed production for maintenance of trioecious reproductive systems is discussed.     

Distribution, stand characteristics and habitat of a critically endangered plant Euryodendron excelsum H. T. Chang (Theaceae): implications for conservation

In the present study, the distribution, population size and structure, habitat and stand disturbance of a critically endangered plant Euryodendron excelsum H. T. Chang were examined using field investigations. A total of 179 individual plants, including 23 adult trees, were found. The population was distributed in 10 highly isolated and fragmented patches. Half of the 10 patches had less than 10 individual plants and two patches had only one remnant adult; there were no adult plants in two of the patches. Although the overall population structure trends to a reversed 'J' shape, two fragile stages of E. excelsum limit its natural recruitment and regeneration: seed germination and seedling growth toward adulthood. The major threats for the survival of this species are its specific small population size and the high frequency of anthropogenic destruction. Based on the results, some strategies were put forward for the conservation of E. excelsum populations.     

Conservation of genetic diversity in the endangered plant Eriogonum ovalifolium var. vineum (Polygonaceae)

The purpose of his research was to describethe organization of genetic variation in thefederally endangered plant taxon Eriogonumovalifolium var. vineum using allozymes. Such information can help prioritize sites andmanagement choices for capturing andmaintaining genetic variation and can reducethe number of populations necessary to committo conservation, thus reducing costs andconflicts with competing land uses. Information on genetic diversity patterns alsoprovides insight into evolutionary anddemographic history of a taxon which canprovide means to assess future risk of erosionof diversity. Similar to other Eriogonumovalifolium varieties, E. ovalifoliumvar. vineum is diverse atboth the taxon (proportion of polymorphic loci[P] = 0.55 and alleles per locus [A] =5.45 [SD = 2.5]) and population (P = 0.56 [SD =0.11] and A = 2.68 [SD = 0.35]) levels. Gene diversity (H _e) withinpopulations averaged 0.19 (SD = 0.03). Wefound some evidence for limited clonalreproduction within populations. Populationswere moderately differentiated from one another(_S = 0.14) and showed moderatedeviations from Hardy-Weinberg equilibriumwithin populations (f = 0.14 and F =0.19). Mean Nei's genetic distance (D)among all pairs of populations was 0.02. Populations with high levels of multiplemeasures of genetic diversity, high levels ofdifferentiation, and low levels of apparentinbreeding are suggested as conservationpriorities.     

Pollen source and resource limitation to fruit production in the rare species Eremosparton songoricum (Fabaceae)

Eremosparton songoricum (Litv.) Vass. is a rare, central Asian desert species which shows lower fruit set and seed set (<16%) than most hermaphroditic species. We hypothesized that fruit production was limited by pollen and resources. To evaluate potential fruit abortion due to pollen limitation, supplemental hand‐pollination was undertaken, the mating system was investigated and the foraging behavior of pollinators was recorded. To investigate possibile resource limitation, flowers and young pods were artificially removed and fertilization were manipulated. The results showed that under natural pollination, the number of pollen deposited on the stigma greatly exceed the number of ovules per ovary. Mating system experiments showed that the species is self‐compatible, but depended on pollinators to set seeds. Supplemental outcross pollination increased fruit set significantly. The most frequent effective pollinator Megachile terminate Morawitz, was observed pollinating many flowers of the same individual plant (74.5±1.3%). These results suggested that fruit production is affected by insufficient outcross pollen rather than by pollen quantity. Removal of 2/3 of the flowers and young pods led to significantly higher fruit set, as did addition of fertilizers (N–P–K: 0.025–0.05–0.013 g, N–P–K: 0.05–0.1–0.025 g) showing that reducing resource acceptors and increasing inorganic resources both helps to improve fruit set. We therefore conclude that reproductive success of E. songoricum is limited by both outcross pollen and available nutrients.     

Insect pollinators improve seed production in globe artichoke (Cynara cardunculus var. scolymus)

The role of many insect species in crop pollination has been widely studied. The use of entomophilous pollination, commonly adopted for other crops, may be of key importance in order to improve seed yields in seed-propagated globe artichoke (Cynara cardunculus var. scolymus) cultivars. In this regard, in a 2-year field experiment, the role of two honeybees (Apis mellifera ligustica and A. mellifera siciliana) and one bumblebee (Bombus terrestris) was evaluated on the seed production of two globe artichoke cultivars in caged-protected environments. Overall, the number and weight of seeds plant⁻¹ and head⁻¹, 1,000 seed weight and fruit setting were higher in caged plants with imposed pollination than open field plants. The pollination efficiency was both insect and cultivar dependent. Both honeybee species performed better on the Mediterranean cultivar ‘Violetto di Sicilia’, while the bumblebee performed better on the Brazilian cultivar ‘NP4’. These results could be very useful to modernise the agronomic management of globe artichoke and reduce the costs of cultivation. In addition, such knowledge can be used to improve the seed production in globe artichoke seed-propagated cultivars and cardoon. This will benefit the bioenergy, nutraceutical, cosmetic and ornamental applications.     

Inbreeding depression influences lifetime breeding success in a wild population of red deer (Cervus elaphus)

Evolutionary and conservation biologists have a long-standing interest in the consequences of inbreeding. It is generally recognized that inbred individuals may experience reduced fitness or inbreeding depression. By the same token, relatively outbred individuals can have greater than average fitness, i.e. heterosis. However, nearly all of the empirical evidence for inbreeding depression comes from laboratory or domestic species. Inbreeding depression and heterosis are difficult to detect in natural populations due to the difficulties in establishing pedigrees. An alternative method is to correlate heterozygosity, which is measured using genetic markers, with a trait related to fitness. The typically studied traits, such as juvenile survival and growth rates, either cover only early life or are weakly correlated with lifetime breeding success (LBS). In this paper we show that heterozygosity is positively associated with male and female adult LBS in a wild population of red deer (Cervus elaphus) on the Isle of Rum, Scotland. To the authors' knowledge, this is the first time that inbreeding depression and/or heterosis have been detected for a trait highly correlated with overall fitness in both sexes in a wild population.     

Effects of seed release timing on plant life-history and seed production in a population of a desert annual,shape Blepharis sindica (Acanthaceae)

Growth, phenology, survivorship, and seed production were observed in a population of a desert annual, Blepharis sindica, with reference to the variation in the timing of seedling emergence. The population consisted of several cohorts induced by rain-cued seed release within a growing season. The fate of 100 individuals of six cohorts was monitored throughout the growing season. Earlier-established cohorts had significantly larger plant sizes and higher reproductive outputs than later cohorts. The time and duration of each phenological stage varied among the cohorts, and they were also influenced by plant size. Mortalities at the seedling stage, vegetative stage, and reproductive stage increased with the delay of seed release. Seed release was concentrated in the early growing season. Fecundity was highest in the earliest cohort and decreased monotonically in later cohorts. The results suggested that even in temporally varying environments, the superiority of early emergent plants was evident. The seed release patterns in temporally fluctuating desert environments are discussed as a compromise between 'diversified bet-hedging' and an optimal timing for maximizing the reproductive success in a growing season.