铁皮石斛不同授粉方式对蒴果发育及种子萌发的影响

为研究不同授粉方式对铁皮石斛蒴果发育以及无菌播种种子萌发的影响,选用健康的铁皮石斛植株花朵进行了自花授粉、异花授粉和自然授粉;选用不同种龄的种子在离体培养条件下进行萌发实验。研究结果表明:铁皮石斛自花授粉和异花授粉均能结实,异花授粉的结实率较高,达64.8%;铁皮石斛在开花的第2~3天授粉的结实率最高;120~210 d种龄的种子萌发率较高,均为90%以上。选择人工异花授粉和适宜种龄的种子离体培养,不仅可培育优良品种,还可缩短繁殖周期,为铁皮石斛遗传改良提供依据。     

对植物白花授粉和异花授粉概念的探析

自花授粉和异花授粉是2种截然不同的授粉方式。通过比较和研究国内外有代表性的生物学、植物学、遗传学、育种学和种子学等相关学科教材及专著对植物授粉方式的论述,发现至今对植物自花授粉和异花投粉的概念不仅缺乏一个统一的认识,而且还存在着明显矛盾之处。作者分别从单花、单株、品种、基因型、遗传特性和天然异交率的角度,在对自花授粉与异花授粉的不同之处进行剖析的基础上,对自花授粉和异花授粉的概念作出了新定义,并对自花授粉和异花授粉的具体概念作出了新界定。     

濒危植物安徽羽叶报春两种花型的繁育特性及其适应进化

二型花柱的维持机制和自然选择压力多年来一直是生态学和进化学研究领域的热点之一。通过实验室栽培和野外观察统计相结合的方法,对安徽羽叶报春两种花型(长柱花和短柱花)的形态特征、花粉活力、柱头可授性、花粉/胚珠比、自然授粉及结籽能力、自交亲和性等繁育特性进行了比较研究。结果表明:长、短柱花的花冠直径和裂片宽无明显差异,而花冠筒、雌蕊和雄蕊高、花粉数目及大小、P/O比均有显著差异。在自然条件下,长柱花所接受的总花粉数要明显高于短柱花的总花粉数,但所接受的异型花花粉数和平均每果结籽数两者无显著差异。长柱花和短柱花的花粉和柱头活力相似,均能在较长时间内维持较高活力,仅在开花末期显著下降。两种花型的花在自花授粉、同型异花授粉、异型花授粉条件下均能结籽,但异型花授粉的结籽数均明显高于自花授粉和同型异花授粉结籽数。在长柱花各种授粉方式中,花粉萌发率无明显差异,但异型花花粉管的生长速度明显比同型异花花粉和自花花粉的快,而在短柱花柱头上表现为异型花授粉的萌发率最高,但只要萌发后在花柱中的生长速度无显明差异。此外,综合上述结果,对二种花型花部综合征的维持机制及自然选择压力进行了探讨。     

大花百子莲的结实和结籽格局及种子产量影响因素分析

在2007和2008年对大花百子莲(Agapanthus praecox subsp.orientalis‘Big Blue’)在群体、果序和单果水平上的结实和结籽格局进行了观测,并依据每花序单花数、单花胚珠数和单粒种子质量推算其单株潜在种子产量;此外,采用人工补充授粉方法研究了不同授粉方式对大花百子莲结实和结籽的影响,并对其种子产量的影响因素进行了分析。结果表明:从始花期、盛花期至末花期,随开花时间的延迟,大花百子莲植株的每花序单花数和座果率均逐渐降低,其中始花期开花的植株每花序单花数和座果率均最高。从果序基部、中部至顶端,座果率和结籽率均逐渐减小且差异显著。在果序基部和中部,果实内种子败育率沿果实基部至顶端依次降低,而单粒种子质量则依次增加;且果序基部的果实各部位的种子败育率均低于果序中部的果实,而单粒种子质量则差异不大。大花百子莲的单株潜在种子产量约为43.18 g,而其单株实际种子产量约为2.87 g,远低于其潜在种子产量。自然授粉的大花百子莲每柱头花粉数为27.5粒,而自花授粉、同株异花授粉和异株授粉的每柱头花粉数均超过100粒;与自然授粉、自花授粉和同株异花授粉植株相比,异株授粉植株的座果率和结籽率均显著增加。研究结果显示:大花百子莲的结实和结籽格局呈现非随机分布规律,在引种地存在有性繁殖障碍;养分和授粉状况是制约大花百子莲种子产量的重要因素。     

山羊臭虎耳草的延迟自动自花授粉机制

通过对山羊臭虎耳草6个花形态及其发育阶段、授粉过程的连续观察和对花粉活性、柱头可授性的连续检测,报道了一种新的延迟自动自花授粉机制,即山羊臭虎耳草花丝运动引导的延迟自动自花授粉。利用电子显微镜研究了花粉囊的破裂方式和柱头结构,同时利用苏丹Ⅲ研究了花粉的组织化学性质。结果显示:(1)第四生长阶段,当花丝引导大部分花粉囊运动至柱头上方时,花粉和柱头的可授性才均达到峰值[可授性分别为(94±6)%和(96±13)%],这种运动消除了第三生长阶段花粉活性较高[(90±8)%]而柱头可授性较低[(6±17)%]的异花授粉特征,即消除了花粉与柱头的时空隔离;自动自花授粉发生于异花授粉特征消除之后,因此山羊臭虎耳草具有典型的延迟自动自交行为。(2)第四发育阶段柱头空腔的大量柱状突起和花粉鞘的油脂性特征,增加了花粉和柱头的接触表面,并且使花粉更好地互相堆积粘附在柱头上,提高授粉效率。(3)对山羊臭虎耳草的繁育系统研究表明,强制自花授粉和人工自花授粉产种量无明显差异,2007年和2008年的自交系数高达0.92和0.91,说明山羊臭虎耳草具有很好的自交能力并且不存在花粉数量限制;人工异花授粉的产种率显著高于人工自花授粉,说明自花花粉的质量显著低于异花花粉,虎耳草延迟自动自花授粉存在生殖衰退,生殖保障系数2007年为0.92,2008年为0.91,表明在极端环境下自动自花授粉为该植物提供了繁殖保障,但极端环境下,没有观察到有效的授粉昆虫。研究表明,作为高山冰缘植物的山羊臭虎耳草,保留了开花植物异花授粉的特征,在极端环境下利用延迟自动自花授粉机制完成授粉,保障生殖。     

前自交型小红花寄生种子特性及寄生关系的建立

寄生是生物有机体之间一种常见的互作关系,其有趣的生理、生态和进化问题日益受到关注与研究。以小红花寄生(Scurrula parasitica var. graciliflora)为研究对象,研究了该植物的开花物候、繁育系统、种子萌发特性及寄生关系的建立。结果表明:小红花寄生的开花高峰期为4月中下旬至5月下旬。花冠筒闭合期间与花冠筒开裂24 h后的花粉活力均 90%,且两者之间没有显著差异(P0.05),但花冠筒开裂48 h之后的花粉活力≤70%。柱头可授性在花冠筒闭合期间显著高于花冠开裂之后(P0.05)。自然授粉的对照组和花蕾期套袋的实验组的座果率(分别是82.5%和83.1%)显著高于花开放之后的异花授粉的实验组的座果率(33.8%)(P0.05),而去除雄蕊的花则未能结实。小红花寄生的种子对脱水敏感,严重脱水显著降低种子萌发率。适宜的种子萌发温度为25℃和30/20℃。仅在桂花幼树的茎顶叶腋处获得了6.25%的寄生成功率,而其他部位的人工寄生均不成功。综合分析表明,两性花的小红花寄生依赖于前自交型的自花授粉;种子萌发表现出明显的温度偏好性,但对光照没有依赖性。人工寄生成功率显示,种子依附的寄主部位直接影响小红花寄生的幼苗与寄主寄生关系的建立。     

霍山石斛种苗繁殖与栽培研究

为探讨霍山石斛(Dendrobium huoshanense C. Z. Tang et S. J. Cheng)的种苗繁殖与栽培技术,对霍山石斛的人工辅助授粉方式、无菌播种和试管苗移栽技术进行了研究。结果表明,人工辅助授粉时,栽培大棚内的异花授粉结果率可达65%以上,而自花授粉结果率不到4%,异花授粉的果实发育更好,但二者果实成熟所需时间基本一致。在无菌培养条件下,试管开花植株的异花授粉结果率可达到18.18%,并且果实成熟所需时间比栽培大棚的短67 d。不论是栽培大棚和无菌培养的异花授粉的成熟种子,还是在4℃下贮存2.5年的种子,无菌播种的萌发率均超过80%,比自花授粉的种子萌发率高20%左右。培养基中添加土豆、香蕉和苹果汁等添加物,对霍山石斛的壮苗均有明显的促进作用,以添加香蕉匀浆的效果最好。试管苗移栽的适宜基质为分层基质:下部3/10树皮+上部7/10木屑,其移栽成活率(78%)和萌芽数较高(每丛2.7个芽),这可能与该基质良好的保水性和透气性等有关。     

线叶忍冬开花动态特征与繁育系统研究

为了解线叶忍冬(Lonicera alberti)的开花特性及繁育系统,本研究通过野外调查,记录其花部特征、开花进程及访花昆虫特性,采用TTC、MTT、联苯胺-α萘酚和醋酸洋红四种染色方法测定花粉活力,采用联苯胺—过氧化氢法和MTT两种方法测定其柱头可授性,运用花粉/胚珠比率、杂交指数、人工授粉试验等方法检测线叶忍冬的繁育系统。结果表明:线叶忍冬花朵有花冠裂片逐一展开和花冠裂片同时展开两种开放方式;花粉在散粉后6 h活力达到最高值,柱头可授性在花朵开放后不断增强;膜翅目蜜蜂科意蜂(Apis mellifera L.)在传粉过程中起重要作用;P/O值为583.43±99.9;杂交指数为4;人工授粉套袋试验中人工异株异花授粉结实率为79.17%,自花授粉情况下结实率为9.52%。其繁育系统为混合交配系统,异花传粉需要传粉者。     

关于四棱草的二型花的研究

陈介(1964)在四棱草属(Schnabelia)中发现具有二型花,即开花授粉型的花(chasmogamous)和闭花授粉型的花(cleistogamous)。他根据当时所掌握的材料,认为二型花"似乎是异株"。笔者在采自安徽祁门的四棱草(S.oligophylla)上发现,二型花无疑是生在同一个体上,但这两种花的花期和花的着生位置是不同的。开花授粉型花的花期早,持续时间短,为一次性开花;闭花授粉型的花是不断地从叶腋发出的,花期持续时间长。     

不同蜂为设施辣椒授粉的授粉效果比较

蜜蜂授粉对开花植物的结实具有重要作用,但目前的研究主要集中在蜜蜂授粉对异花授粉作物的授粉效果上,而对自花授粉植物的授粉效果鲜有报道。本研究以设施农业中主要的自花授粉作物—辣椒为研究对象,研究了意大利蜜蜂(意蜂)和小峰熊蜂(熊蜂)为设施辣椒授粉时的出巢访花活动规律及其授粉效果,并初步估算了其可能带来的经济效益。对两种蜂为辣椒授粉过程中的出巢访花活动规律研究表明:意蜂的出巢和工作起点温度均显著高于熊蜂,表明熊蜂每天可比意蜂多工作1.0-1.5 h。对不同蜂的授粉效果的研究表明:两种蜂授粉后的结实率差异不显著,但均显著高于自花授粉的结实率;三种授粉方式的坐果时间长短依次为自花授粉意蜂授粉熊蜂授粉;花后30 d三种授粉方式的单果重和果实长度差异显著,单果重依次为43.04、57.23和67.58 g,对应果实长度依次为17.41、19.14和20.28㎝。两种蜂授粉后的果实籽粒数(121.35和127.25粒)显著高于人工授粉后的105.40粒。研究表明意蜂和熊蜂可使单位面积自花授粉的设施辣椒增产6.62%和9.79%,具有较好的增产效果,相应的,辣椒对两种蜂的依赖度D值分别为0.06和0.09,采用国际通用的生物经济的方法初步推算出这两种授粉方式在本世纪前十年每年均可使我国辣椒的经济效益得到可观的增加。     

PLANT DENSITY,CLEISTOGAMY, AND SELF-FERTILIZATION IN NATURAL POPULATIONS OF LITHOSPERMUM CAROLINIENSE

Lithospermum caroliniense has a heteromorphic incompatibility system complemented by self-compatibility via cleistogamy. Plants typically are either solely chasmogamic or solely cleistogamic. The percentage of cleistogamic plants in northern Illinois and Indiana populations varied from zero to 4.7. However, the percentage seed from self-fertilization varied from zero to 51.5, because cleistogamic plants produced 1.5 times as many ovules as chasmogamic plants, and the former type averaged 3.4 seeds per flower in contrast to 0.23 in chasmogamic flowers. Interpopulation differences in cleistogamy and self-fertilization were strongly correlated with plant density, an increase in density being accompanied by increases in the aforementioned features. Most populations had an excess of thrums, but there was no relationship between cleistogamic seed production and thrum:pin ratio.     

Plantago ser. Hispidulae, a taxonomic revision Knud Rahn

Plantago ser. Hispidulae Rahn belongs to subgen. Psyllium (Juss.) Harms sect. Gnaphaloides Barn. 5 species are recognized, they are all narrow–leaved annuals confined to the area west of the Andes. P. limensis Pers. occurs in Peru; P. litorea Phil., P. hispidula Ruiz & Pav., and P. rancagua Steud. in those parts of Chile with drought during the summer, and P. lundborgii Sparre on the island of San Ambrosio. Flowers are cleistogamic with small corolla lobes and small anthers, very rarely chasmogamic with larger corolla lobes and anthers. Experiments demonstrated that pollination is necessary for development of seeds in cleistogamic flowers, and that transfer of pollen from one such flower to another is almost impossible.     

Is plasticity across seasons adaptive in the annual cleistogamous plant Lamium amplexicaule?

Background and aims Many angiosperms exhibit cleistogamy, the production of both cleistogamous flowers (CL), which remain closed and obligately self-pollinated, and chasmogamous flowers (CH), which are potentially open-pollinated. The CH proportion can be plastic. Plasticity is adaptive if environmental changes can be reliably assessed and responded to with an appropriate phenotype and if plastic genotypes have higher fitness in variable environments than non-plastic ones.Methods We studied the plastic response of four natural populations from northern and southern France of an annual cleistogamous plant, Lamium amplexicaule, to predictable seasonal variation. Plants were grown in a semi-controlled environment in spring and in autumn. We assessed the variation in flower number, phenology and cleistogamy-related traits, which were all plastic with respect to season. The CH proportion was higher in spring than in autumn in all four populations.Key Results We showed significant stabilizing selection for cleistogamy traits, with higher optimal CH proportions and more pronounced stabilizing selection in spring than in autumn. Observed CH proportions were close to the predicted optimal CH proportions in each season except in autumn for southern populations, which do not experience the autumnal growing season in nature.Conclusions These results are consistent with adaptive plasticity across seasons of cleistogamy in L. amplexicaule. We propose that adaptive plasticity of cleistogamy could be driven by pollination environment variation, with CL flowers providing reproductive assurance when pollinators are scarce and CH flowers reducing the inbreeding depression in offspring when pollinators are abundant.     

Effects of local densities and abiotic microenvironments on reproductive outputs of a biennial,Lysimachia mauritiana var. rubida

To understand the relative importance of density‐dependent and density‐independent factors on plant fitness, we examined the effects of local densities and surrounding ground surface conditions on the reproductive output of a monocarpic biennial plant, Lysimachia mauritiana var. rubida in a natural population. Observations were conducted during five successive years in open dry habitats on rocky coastal cliffs of the subtropical Bonin (Ogasawara) Islands, Japan. Local density and ground surface conditions, categorized by gravel size, were remarkably heterogeneous in the study population. Therefore, we hypothesized that number of flowers per plant was affected by local density, ground surface condition, and their interaction. During 3 of 5 years, the number of flowers per individual L. mauritiana var. rubida plant was only negatively affected by the local density around the plant. The interaction effects of local density and ground surface conditions were observed in two years. Taken together with the results of previous studies, these results suggest that density effects are dominated survival in the early growing season and reproduction, while the effects of ground conditions are important for seedling recruitment and survival in the late growing season. Thus, interactive effects between density‐dependent and density‐independent factors on the life cycle from germination to reproduction regulate populations of L. mauritiana var. rubida in spatially heterogeneous habitats.     

Benzoxazinoids and iridoid glucosides from four Lamium species

A new class of compounds for the plant family Lamiaceae, benzoxazinoids, was found in Lamium galeobdolon. From the aerial parts of the species were isolated the new 2-O-beta-D-glucopyranosyl-6-hydroxy-2H-1,4-benzoxazin-3(4H)-one (6-hydroxy blepharin) together with four known benzoxazinoids, DHBOA-Glc, blepharin, DIBOA, DIBOA-Glc, as well as harpagide, 8-O-acetyl-harpagide and salidroside. Eight known iridoid glucosides, 24-epi-pterosterone and verbascoside were isolated from Lamium amplexicaule, L. purpureum and L. garganicum. The iridoids, 5-deoxylamiol and sesamoside, as well as the phytoecdysone, 24-epi-pterosterone, were found for the first time for the genus Lamium. The phytochemical data are discussed from a systematic and evolutionary point of view.     

Effects of pollen quantity and quality on reproduction and offspring vigor in the rare plant Scorzonera humilis (Asteraceae)

We studied the effects of pollinator exclusion, interparental distance, and supplementary hand pollination on reproduction and progeny vigor in Scorzonera humilis (Asteraceae), a rare plant of fragmented, nutrient-poor grasslands. Caged flowers produced no seeds and selfed flowers only very rarely, indicating that S. humilis is mainly self-incompatible. Seed production, seed mass, and seed germination following between-population crosses were consistently, but not significantly, higher than after within-population crosses. Seed set increased with local density of conspecifics, indicating that the reduced plant density in fragmented populations may reduce plant reproductive success. Seed set was pollen limited in all four populations studied. Supplementary hand-pollination strongly increased the survival of offspring, indicating that either pollinators transferred pollen from related individuals resulting in inbreeding depression in spite of the incompatibility system or that higher pollen loads increased pollen competition and the selectivity among gametes. In one of the populations, adding pollen from a different population strongly increased progeny fitness compared with both natural pollination and pollen supplementation from the same population. The results indicate that S. humilis is sensitive to inbreeding and that pollen limitation can reduce both the number and quality of offspring.     

Studies on the breeding system of Limnocharis flava (Butomaceae)

Limnocharis flava ( Linn. ) Buch. normally produced 2 ～ 13 bisexual flowers in a cincinnus (seorpioid cyme). The anthesis of each flower lasted 5 ～ 13 hours. In natural populations, two species of beetles were observed visiting the flowers in bloom. Several whorls of staminodes outside of the stamens limited the pollen to disperse outward, and made the pollen deposit only on the stigma of the same flower. After the perianth closed, the inner perianth changed into liquid which created a suitable environment for pollen germination. The pollen-ovule ratio, which ranged from 650 to 1120( 930±180) reflects an economical self-pollination system. As the fruits maturied, the inflorescence fell to ground or mud, and the vegetative buds (i.e. vegetative components ), developed at the apex of the inflorescence, grew into new vegetative shoots. One mother plant usually could produce 6 ～ 20 new vegetative shoots in a life cycle. One fruit of L. flava contained 470 ～ 640 seeds, with an average seed-set rate of about 80%. The seeds of Limnocharis flava could disperse a long distance and occupy new suitable habitats to establish new populations. The vegetative buds, however, predominantly could only spread to nearby places and thus enlarge the size of the population. Results from some artificial treatments, such as emasculating and netting, emasculating and bagging, hand-pollinating, netting, and bagging, indicate that L. flava is selfing in general but also cross-compatible. Based on the above results, it is considered that the breeding system of L. flava is predominantly selfing, but mixed with outcrossing, and the likely evolutionary tendencyis from outcrossing to selfing.     

Patterns and sources of variation in pollen deposition and pollen tube formation in flowers of the endemic monoecious shrub Cnidoscolus souzae (Euphorbiaceae)

Pollen deposition and pollen tube formation are key components of angiosperm reproduction but intraspecific variation in these has rarely been quantified. Documenting and partitioning (populations, plants and flowers) natural variation in these two aspects of plant reproduction can help uncover spatial mosaics of reproductive success and underlying causes. In this study, we assess variation in pollen deposition and pollen tube formation for the endemic monoecious shrub Cnidoscolus souzae throughout its distribution range in Mexico, and determine how this variation is structured among populations, plants and flowers. We also infer the relative importance of pollen quantity and quality in determining pollination success in this species. While we found no evidence suggesting that pollen receipt limits C. souzae reproduction across 19 populations, we did find extensive variation in pollen load size and pollen tube number per flower. Total variation in pollen receipt and pollen tube number was mostly explained by intra‐individual and among‐population variance. Furthermore, pollen load size had a stronger effect on the number of pollen tubes at the base of the style than pollen germination rate, suggesting that pollen quantity may be more important than quality for pollen tube success in C. souzae. Our results suggest that both small within‐plant flower differences and broad‐scale differences in community attributes can play an important role in determining pollination success. We emphasise the need to evaluate patterns and sources of variation in pollen deposition and pollen tube formation as a first step in understanding the causes of variation in pollination success over broad spatial scales.     

Prolonged clonal growth: escape route or route to extinction?

Many plant species have the capability to reproduce sexually as well as clonally. The balance between clonal reproduction and sexual reproduction varies between different species. It was estimated that 66.5% of all central European flora may form independent but genetically identical daughter plants. Also within species there is great variation in the ratio clonal/sexual reproduction. Clonal reproduction can be considered as an alternative life cycle loop that allows persistence of a species in the absence of the ability to complete the normal life cycle (i.e. seed production, germination and recruitment). Plant populations exhibiting prolonged clonal growth have been referred to as 'remnant populations'. A remnant population in general is defined as "a population capable of persistence during extended time periods despite a negative population growth rate (λ<1) due to longlived life stages and life cycles, including loops, that allow population persistence without completion of the whole life cycle". Here we argue that prolonged and nearly exclusive clonal growth through environmental suppression of sexual reproduction can ultimately lead to local sexual extinction and to monoclonal populations of a species, and that this may imply significant consequences for population viability. Especially obligate or mainly outcrossing clonal plant species may be vulnerable for sexual extinction. We argue that the consequences of reduced sexual recruitment in clonally propagating plants may be understudied and underestimated and that a re-evaluation of current ideas on clonality may be necessary.     

新疆郁金香营养生长、个体大小和开花次序对繁殖分配的影响

植物有性繁殖与资源分配的关系研究对于揭示植物生活史特征及繁育系统进化具有重要意义。新疆郁金香(Tulipa sinkiangensis)是新疆天山北坡荒漠带特有的一种多年生早春短命植物。在自然生境中,该物种仅以有性繁殖产生后代,每株能产生1-8朵花,且不同植株上的花数及果实数以及花序不同位置上的花与果实大小明显不同。本文通过对新疆郁金香有性繁殖与营养生长及植株大小的关系以及花序中不同位置花及果实间的资源分配研究,旨在揭示营养生长、个体大小及开花次序对其繁殖分配的影响。结果表明:在开花和果实成熟阶段,新疆郁金香植株分配给营养器官(鳞茎和地上营养器官)与繁殖器官的资源间均存在极显著的负相关关系(P<0.01),说明其植株的营养生长与生殖生长间存在权衡关系。多花是新疆郁金香的一个稳定性状,其植株上花数目、花生物量、果实生物量和种子数量与植株生物量间均呈极显著的正相关关系(P<0.01),说明新疆郁金香植株的繁殖分配存在大小依赖性。在具2-5朵花的新疆郁金香植株中,花序内各花的生物量、花粉数和胚珠数、结实率、果实生物量、结籽数、结籽率及种子百粒重按其开花顺序依次递减,说明花序内各花和果实的资源分配符合资源竞争假说。植株通过减少晚发育的花或果实获得的资源来保障早发育的花或果实获得较多的资源,从而达到繁殖成功。