绣球茜的二型花柱及其传粉生物学初步研究

首次对绣球茜(Dunnia sinensis Tutch)的花药和柱头高度、花粉和柱头形态,以及人工辅助授粉的结实情况等进行了研究.结果表明:绣球茜在同一居群中同时具有长花柱(Pin 型)和短花柱(Thrum)两种花型,长花柱植株的柱头和雄蕊高分别为9.61±0.36 mm和6.79±0.38 mm,短花柱则为6.11±0.56 mm和8.96±0.59 mm,表现出互补式雌雄异位的花部特征.通过人工辅助授粉,两种花型植株的自交和同型植株间异交都不结实,仅在型间异交方能正常结实,说明绣球茜中存在极强的自交不亲和和型内不亲和现象.在花粉与柱头形态方面:长柱型花的单花花粉量(38555.6±6258.4)显著地小于短柱型花(52145.45±8924.5)(t=4.009,P＜0.01),长柱花和短柱型花的胚珠数分别为38.73±1.85和39.1±2.25(t=-4.13,P＞0.01);长柱型花花粉粒的大小(极径21.27±0.70 μm)显著地小于短柱花(21.92±0.79μm)类型(t=3.890,P＜0.01),而柱头乳突大小长柱类型要大于短柱类型,表现出明显的二型性.研究结果表明绣球茜是一种典型的二型花柱植物.此外,对其传粉昆虫及昆虫访花行为的观察表明,蝶类(青风蝶属Graphium)和蜂类(熊蜂Bombus sp.和木蜂Xylocopa magnifica Smith)为绣球茜的主要访花昆虫和传粉昆虫,它们能够在两种不同花型植株之间有效地传粉.     

二型花柱植物金荞麦的繁殖生态学研究

通过野外实验观察和人工控制套袋等方法,从金养麦的花部基本特征、开花动态、花粉胚珠比(P/O)、花粉活力、柱头可授性、杂交指数、套袋实验及访花昆虫等方面对其繁殖生态学进行了研究.结果显示:(1)金荞麦在同一居群中同时具有长柱型(L-型)和短柱型(S-型)两种花型,短花柱花直径(7.25±0.11 mm)显著大于长花柱花直径(6.79±0.11 mm),长花柱花的柱头和花药高分别为3.39±0.04 mm和1.80±0.02 mm,短花柱花的柱头和花药高则分别为1.89±0.04 mm和3.19±0.06 rnm,表现出互补式雌雄异位的花部特征.(2)金荞麦8～10月开花,单花序的花期为15～23 d,单花的花期为1～3 d,开花进程中,内轮雄蕊先散粉,外轮雄蕊后散粉;S-型花的花粉活力高于L型花的花粉活力,但其单花花粉量低于L-型花,显示了两种花型的自我调控机制.(3)套袋实验表明,金荞麦自花自交不亲和,但具有型内和型间亲和性,L型和S-型花的P/O值分别为810±40.48和526.5±42.24,表明其繁育系统为兼性异交,需要传粉者,但L-型花更倾向异交.(4)金荞麦的主要访花昆虫为膜翅目(Hymenoptera)、双翅目(Diptera)、鳞翅目(Lepidoptera)、半翅目(Hemiptera)和蜻蜒目(Odonata) 19个科37种昆虫,其中,两种花型拥有12种共同的访花昆虫,L-型花访花昆虫14种,S-型花访花昆虫35种,表明S-型花对昆虫具有更强的吸引力.研究表明,金荞麦是典型的二型花柱植物,虽然具有严格的自交不亲和性,但显示一定的型内亲和性.     

梅花个体内花柱长度的变异及其对繁殖成功的影响

植物个体内花型的变异影响繁殖成功, 雌性繁殖性状的变异可能影响雌性的繁殖成功, 也可能作为花粉的受体影响雄性的繁殖成功。然而, 植物个体内不同花柱长度的花产生的花粉是否影响植物的繁殖成功却少有研究。梅(Armeniaca mume)是原产我国的重要木本花卉和经济果树, 我们的野外观察发现在同一植株内, 不同花的花柱长度有变异, 存在长柱型、短柱型和雄花型(雌蕊败育)三种花型, 是比较雌性繁殖性状的变异对两性繁殖成功的影响的理想材料。本文主要测量了不同花型的花部特征, 统计花期, 并开展体外花粉萌发以及人工控制授粉实验。结果表明: 长柱型的花冠直径、雌蕊长、单花花粉数、花粉体积显著大于短柱型和雄花型。长柱型的单花期以及雌花期显著长于短柱型。长柱型、短柱型以及雄花型花粉在活体柱头上的萌发率没有显著性差异, 雄花型的花粉管长度显著高于长柱型和短柱型。长柱型为母本的花粉萌发率以及花粉管的长度要显著高于短柱型。长柱型、短柱型、雄花型花粉授粉与自然对照处理的坐果率没有显著性差异, 而长柱型为母本的坐果率要显著高于短柱型为母本的坐果率。这些结果表明野生梅花的长柱型为母本有利于花粉的萌发和花粉管的伸长, 有高的坐果率; 但其作为花粉供体的雄性功能与其他花型没有差异。     

连翘花的结构与繁育系统研究

通过定株观察、解剖和人工套袋交叉授粉试验对连翘花的生长发育和繁殖系统进行研究.结果表明:(1)连翘具有2种避免自交的方式,雌雄异位和雌雄异熟.其雌雄异位表现为长雄蕊短雌蕊花型和短雄蕊长雌蕊花型;雌雄异熟表现为长雄蕊短雌蕊花型为雄蕊先于雌蕊成熟,短雄蕊长雌蕊花型表现为雌蕊先于雄蕊成熟.(2)连翘花P/O值测定和户外套袋交叉授粉试验显示,P/O值为2 000±300;不同类型花的异花授粉结果率在50.64%~80.32%,其中短型花的花粉授到长型花柱头的结果率最高,达80.32%,而长型花和短型花的同型花授粉结果率分别为2.92%和34.15%,表明连翘的异花授粉结果率高于自花授粉,以异交为主,其繁育系统属兼性异交.研究结果认为,连翘雌雄异位和雌雄异熟是其自然结果率低下的主要原因,为进一步探讨连翘在木犀科中的系统进化提供了生殖生态学的依据.     

黑果枸杞两种花型的花部综合征与传粉特性

黑果枸杞(Lycium ruthenicum)是中国西北地区极端环境中分布的国家二级保护植物, 该物种在新疆南部的自然种群中出现了同型花柱类型(同位花)和柱头探出式雌雄异位类型(异位花)个体, 并且遭遇沙尘暴频繁的种群中异位花个体出现频率减少。该研究对喀什市自然种群中黑果枸杞两种不同花型植株的花部综合征和传粉特性进行比较研究, 以期探讨该物种不同花型植株在南疆早春极端环境中的花部特征的可塑性及其适应性机制。结果表明: 同位花雌雄蕊高度间无显著差异, 而异位花雌蕊高度显著高于雄蕊; 同位花花冠直径、花冠筒长、胚珠数均高于异位花, 而异位花雌雄蕊空间距离、花粉数及花粉胚珠比均比同位花高。黑果枸杞同位花个体比例(68%)高于异位花个体(32%), 种群水平及个体水平同位花花期((117.00 ± 2.25) d, (101.65 ± 1.98) d)比异位花((26.00 ± 1.00) d, (18.75 ± 1.00) d)长, 而单花水平上异位花单花寿命((4.50 ± 0.14) d)比同位花((3.13 ± 0.11) d)长。两种类型花在花早期(紫色)分泌的花蜜量均高于花后期(白色)。在紫色花阶段(花开放早期), 同位花上的主要传粉者意大利蜜蜂、熊蜂和食蚜蝇的访花频率和停留时间均高于异位花; 而白色花阶段(花开放后期)意大利蜜蜂、熊蜂在异位花上的访花频率比同位花高。在不同花色阶段, 同位花柱头花粉落置数、花粉移出率、花粉传递效率均比异位花高, 并且同位花自然坐果率及结籽率均比异位花高。在新疆南部的沙尘暴极端环境下, 同位花通过较高的自交亲和性保障繁殖, 而异交为主的异位花提高了异交率。异位花与同位花在花部综合征和花报酬上的差异, 是影响其繁殖成功的主要因素。     

同域分布柽柳属两种植物的传粉生物学研究

以同域分布于新疆南部的早春开花植物短穗柽柳(Tamarix laxa Willd)和紫杆柽柳(T.androssowii Litw)为材料,对它们的开花式样、花部特征、花报酬特点、传粉特性及交配方式进行比较研究。结果表明,2种植物居群和个体水平的花期及单花寿命间存在显著差异;紫杆柽柳的居群花期比短穗柽柳短,而短穗柽柳个体花期和单花寿命比紫杆柽柳长。2种柽柳的花瓣、雌雄蕊长度、花粉胚珠数及花粉/胚珠比值(P/O)间也存在显著差异。二者的花瓣、雌雄蕊均能分泌香味,访花频率与花蜜量均存在正相关。主要传粉者在短穗柽柳上的访花频率和沉积花粉效率高于紫杆柽柳,且存在显著差异。2种植物通过自花和异花授粉均可结实,属于混合交配系统;二者的自交不亲和指数及花粉限制程度间存在显著差异;紫杆柽柳的自交亲和性高于短穗柽柳,而短穗柽柳的花粉限制指数高于紫杆柽柳。因此,2种柽柳的开花式样、花部特征及花报酬的差异,是它们在传粉者受限的早春环境中为吸引更多的访花昆虫、提高雌性繁殖率所形成的繁殖对策。     

入侵植物酢浆草在中国的花型分布特点及自花授粉

通过野外调查和馆藏标本(IBSC)的查阅,对广泛分布于中国的入侵植物酢浆草Oxalis corniculata的花型、开花物候、访花昆虫和花粉胚珠比(P/O)进行了观测.通过人工辅助授粉实验检测了其繁育系统;使用扫描电子显微镜(SEM)观察了自花花粉在柱头上的萌发情况并使用荧光显微镜(FM)观察了花粉管在花柱中的生长情况.结果表明:酢浆草在中国有3种花型,半等长花柱型、准等长花柱型和长花柱型,大部分的居群只有半等长花柱型,分布于人为干扰较大的生境(如路边);云南安宁松树林下的自然居群和采自广东的两份标本为长花柱型花:少部分居群内兼有半等长花柱型和准等长花柱型植株.酢浆草花期从2月到11月.其中3_9月为开花高峰期:单花花期2天,花开当天傍晚花瓣会合拢,第二天早上花瓣再次打开,到花开第二天傍晚花瓣萎蔫.花蕾开放前花药已开裂,柱头有活性,大部分花粉散落到柱头上,且花粉已开始萌发;花粉胚珠比为83.3±33.08.有少量花蜜分泌.唯一的访花昆虫为一种小灰蝶Pseudozizeeria sp.,但并不是有效的传粉者.开花前套袋处理结实率(88.12±3.38)与自然结实率(88.03±2.48)基本没有差异:荧光显微镜观察证明自花花粉的花粉管可在花开第一天生长到达胚珠,说明酢浆草半等长花柱植株是自花授粉.     

艳丽耳草的二型花柱及异型自交不亲和系统

异型花柱是一种受遗传因素控制的花型多态性现象,包括二型花柱和三型花柱两种类型.本文以茜草科艳丽耳草(Hedyotis pulcherrima)为实验材料,通过对其野外居群的花型、花部形态及花粉特征等观察,发现艳丽耳草野外居群同时存在长花柱型花和短花柱型花,长/短花柱型花的数量比例为1∶1.两型花具有精确的交互式雌雄异位特征,并且该特征与花冠长度相关性显著.长/短花柱型花的柱头裂片长度、花粉大小及淀粉含量等具有二型性.花粉体外培养时花粉萌发率及花粉管生长速率无显著性差异.人工授粉后,艳丽耳草长/短花柱型花型间异交花粉管生长形态正常,授粉24 h后花粉管均已进入子房.而长/短花柱型花在自交及型内异交下均表现为不亲和,花粉管生长停止于柱头,花粉管顶端累积胼胝质并膨大.艳丽耳草没有无融合生殖现象,型间人工辅助异交授粉结实率为100％,显著高于自然结实率.本研究结果表明,艳丽耳草是典型的二型花柱植物,并具有异型自交不亲和系统.     

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。     

欧报春二型花柱的繁育特性研究

二型花柱植物通常具有自交不亲和性,在不同植物中所表现的可育性不同。欧报春(Primula vulgaris)是典型的二型花柱植物,为了探究欧报春的繁育特性,通过温室栽培,对欧报春的长花柱和短花柱的花部特征、花粉和花柱形态、花粉活力、柱头可授性、花粉/胚珠比、杂交指数、杂交亲和性、花粉管观察进行研究。结果表明,(1)长花柱和短花柱的花冠直径、花冠筒长度、柱头高度、花药高度和花筒中部直径均表现出两型性;(2)花粉和柱头观察发现花粉极轴长、花粉赤道轴长、花柱直径、柱头乳突细胞和花粉数量均有差异;(3)长花柱的花粉/胚珠比为384.20,短花柱为369.70,属于兼性异交类型;长花柱和短花柱的花粉活力和柱头可授性能在较长时间内维持较高活力;(4)长花柱杂交指数值为5,短花柱为4,表明繁育系统类型为异交,部分自交亲和,需要传粉者;6种授粉组合均能结实,异型花间授粉的结实数明显高于同型自花授粉和异株同型授粉;短花柱为母本的异型花间授粉组合亲和性优于其他组合;欧报春存在自交不亲和性,长花柱的自交不亲和性低于短花柱。     

Is sexual organ reciprocity related to legitimate pollen deposition in distylous Pulmonaria (Boraginaceae)?

Heterostyly, i.e. the reciprocal positioning of anthers and stigmas, is a floral polymorphism that is thought to promote disassortative (i.e. between‐morph) pollination and to maintain genetic diversity within populations. Recent research, however, has shown several cases of heterostylous plant species in which the reciprocal positioning of the sexual organs varies, which may affect the likelihood of ‘legitimate’ pollination between compatible morphs, and hence morph fitness and ultimately the stability of this floral polymorphism. In this study, we investigated floral morphology and stigmatic pollen deposition patterns in nine Pulmonaria species (Boraginaceae) across Europe to test the hypothesis that sexual organ reciprocity is related to legitimate pollen deposition. We used a recently developed adaptive accuracy concept to assess the level of reciprocity and to relate inaccuracies in positioning of sexual organs to legitimate pollen deposition measured on stigmas in the field. The nine investigated Pulmonaria species showed substantial deviations from perfect reciprocity, with total inaccuracies varying between 2.9 and 20.3% relative to the squared mean of all anther and stigma heights in the population. In almost all cases, most of the inaccuracy was generated by the high‐level organs. Total pollen deposition was always higher on stigmas of long‐styled individuals, but stigmas of short‐styled individuals received significantly more legitimate pollen than stigmas of long‐styled individuals (71.23% and 38.75% respectively). We also found a weak but significant relationship between measures of inaccuracy and the level of disassortative pollination. Under these conditions of pollen flow, the distylous mating system is predicted to persist in only eleven (27%) of all sampled populations. Overall, these results indicate that deviations from perfect reciprocal herkogamy are common in Pulmonaria and have an impact on legitimate pollen deposition.     

High herkogamy but low reciprocity characterizes isoplethic populations of Jasminum malabaricum,a species with stigma‐height dimorphism

• Studies of floral polymorphisms have focused on heterostyly, while stigma‐height dimorphism has received considerably less attention. Few studies have examined the reproductive biology of species with stigma‐height dimorphism to understand how factors influencing mate availability and pollen transfer are related to morph ratios in populations.
• Floral morphological traits, especially herkogamy and reciprocity, pollinator visitation, breeding system and spatiotemporal mate availability, are known to affect inter‐morph pollination and morph ratios in species with stigma‐height dimorphism. In this study, we investigated the presence of stigma‐height dimorphism and estimated morph ratios in four naturally occurring populations of Jasminum malabaricum. We quantified morph‐ and population‐specific differences in the abovementioned factors in these populations to understand the observed morph ratios.
• The positions of anthers and stigmas were characteristic of stigma‐height dimorphism, the first report of this polymorphism in the genus. All study populations were isoplethic, implying equal fitness of both morphs. Herkogamy was higher in the short‐styled morph, while reciprocity was higher between the long‐styled stigma and short‐styled anthers. Long‐ and short‐tongued pollinators were common floral visitors, and we observed no differences between morphs in spatiotemporal mate availability or pollinator visitation. Neither morph exhibited self‐ or heteromorphic incompatibility.
• The short‐styled stigma had lower reciprocity but likely receives sufficient inter‐morph pollen from long‐tongued pollinators, and also by avoiding self‐pollination due to higher herkogamy. These results highlight the importance of sufficient effective pollinators and floral morphological features, particularly herkogamy, in maintaining isoplethy in species with stigma‐height dimorphism.

     

Ovarian self‐incompatibility in Narcissus papyraceus (Amaryllidaceae) is the result of a pre‐zygotic response

Ovarian self‐incompatibility, including pre‐ and post‐zygotic reactions, is a complex mechanism for which we still lack many details relating to its function and significance. The joint presence of ovarian self‐incompatibility with style polymorphism is a rare combination that is found in the genus Narcissus. Usually, style polymorphic species have heteromorphic (diallelic and linked to style length locus) incompatibility, which prevents fertilization between individuals of the same morph, thereby helping to maintain equal proportions of floral morphs in populations. However, when present, self‐incompatibility in Narcissus is not linked to style polymorphism and cross‐fertilization within each morph is possible. Hence, self‐incompatibility in Narcissus is of particular interest when attempting to unravel the nature of the rejection reaction and aiming to assess possible cryptic differences in the fertilization process in intra‐ and inter‐morph crosses, which might ultimately explain the wide variation of morph‐ratio in the field. We examined the breeding system of Narcissus papyraceus, a style‐dimorphic species that has biased morph ratios in most of its populations. We studied pollen‐tube growth in the pistil and ovule fate after experimentally controlled hand pollinations. The growth of pollen tubes in self‐ and intra‐ and inter‐morph crosses was similar up to the point of micropyle penetration in both morphs but, subsequently, a pre‐zygotic failure appeared to affect male and female gametophytes in selfed pistils. A high proportion of ovules from self‐pollinated flowers showed signs of collapse and self‐pollen tubes were blocked or behaved abnormally before entering the embryo sac. Self‐incompatibility was stronger in the long‐styled morph than in the short‐styled morph. We did not find any conclusive sign of differential functioning between intra‐ and inter‐morph cross‐pollinations in any morph. These results enable us to rule out the possible effects of pollen–pistil interactions in N. papyraceus as a cause of morph‐ratio biases and confirm the exceptional nature of the self‐incompatibility mechanism in this polymorphic species. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 629–643.     

Adaptive significances of sexual system in andromonoecious Capparis spinosa (Capparaceae)

The sexual system of the perennial shrub Capparis spinosa L. (Capparaceae), which is distributed in arid deserts of northern Xinjiang, was investigated. The main results can be summarized as follows. (1) The species is andromonoecious, i.e. individuals possess both male and perfect flowers. Stamens of two floral morphs are normal and can be divided into long and short ones in each flower. The perfect flowers have well developed pistils, but male flowers have aborted pistils and only function as males. (2) There were very significant differ-ences among the populations in daily ratio of the two floral morphs, number of long and short stamens of perfect flowers and length of filaments and anthers of short stamens in male flowers (P<0.01), but no significant differ-ences occurred in biomass of floral organ (P>0.05). (3) Anthesis was nocturnal and lasted 15–16 h. Both male and perfect flowers opened about 18:00 at dusk. The number of two floral morphs produced on each individual was indeterminate every day during flowering, which would make the individual temporally androdioecious. However, the total number of male flowers was more than that of perfect flowers within the population every day. (4) The P/O ratios of perfect flowers in three populations were 1.57×104, 1.65×104 and 1.71×104. There was no significant difference in pollen numbers (P>0.05) within population between male and perfect flowers or between long and short stamens, and also no significant difference among the populations in pollen numbers of male and perfect flowers, and in ovule numbers and P/O ratios of perfect flowers (P>0.05). (5) Dynamic curves of pollen viability of long and short stamens in the two floral morphs were similar in three populations. Pollen longevity of both long and short stamens was about 18–20 h, and duration of stigmatic receptivity was about 16–18 h. (6) Floral visitors were hymenopterous and lepidopterous insects. There was a total of seven species of floral visitors in the three populations. The activities of them were greatly affected by the climate of the desert environment. (7) Perfect flowers were not apomictic and could produce fruits after self-pollination (autogamy and geitonogamy) and cross-pollination; thus they had a mixed mating system. These results suggest that the sexual system of andro-monoecy may reflect an environment-heredity interaction in C. spinosa. The male flowers increase the number of pollen grains, the P/O ratio and enhance male fitness of individuals. They also increase the floral display and consequently effect cross pollination by attracting more pollinators that bring cross-pollen to the stigmas of per-fect flowers, thus increase outcrossing rate and female fitness of individuals, which insure reproductive success of C. spinosa in the extreme desert environment.     

Pollen and ovule production, floral nectary structure, and nectar secretion dynamics in tristylous Lythrum salicaria L.

Lythrum salicaria L. (Lythraceae) is tristylous, each plant forming one of three floral morphs that differ in reciprocal placement of the stigma and two sets of anthers. Several reproductive traits were compared quantitatively among these morphs. Although mean pollen viability (??93%) and total pollen per mature, indehiscent anther within a staminal level (mean CV?=?11%) were constant, the patterns of mean pollen production per anther were complex, being significantly lowest in long stamens (1,490 grains) of the short-styled morph, but highest in intermediate stamens (3,590) of the long-styled morph. Overall, pollen production was greatest (38,200) in long-styled flowers and least (22,000) in short-styled ones. On the contrary, ovule quantities per ovary (mean 107) were similar among floral morphs; thus, pollen-to-ovule ratios spanned 192 (short-styled morph) to 364 (long-styled morph), relatively low values for a strictly xenogamous species. Each morph had a recessed annular nectary of similar dimensions encircling the ovary base, with equal numbers of modified stomata distributed uniformly on the nectary surface. Most stomata were solitary (94%), whereas 5% occurred in pairs and 1% of stomatal units had just one guard cell. During nectar secretion, about 16% of pores were closed plus 28% of pores were fully occluded. Similarly, nectar volumes and solute concentrations, peak rates of nectar secretion (mean 72?C79???g sugar?h^?1) at early afternoon, and the nectar??s sucrose prevalence [S/(G?+?F) ??4.3] were not significantly different among morphs. Based on these similarities in nectary structure and nectar-secretion dynamics, traits rarely studied in tristylous species, the preferential visitation of any particular floral morph of L. salicaria during nectar foraging by insects is unlikely. Indeed, lack of discrimination among morphs by potential pollinators may be a key tenet of successful sexual reproduction in tristylous species.     

宁夏枸杞的花粉呈现式样及交配系统研究北大核心CSCD

被子植物的花回馈、雌雄蕊时空分离特征和花粉呈现式样等花部特征及传粉者效率会影响雄性适合度和有性繁殖过程。宁夏枸杞(Lycium barbarum)是中国宁夏、新疆、内蒙等干旱和半干旱地区分布的特有种。该研究对新疆喀什地区宁夏枸杞自然种群的花部综合征、花粉呈现式样、花回馈、传粉者行为与交配方式的关系进行观察及统计分析,以探讨其花粉逐步呈现的适应性及其在提高雌雄繁殖过程中的意义。结果表明:(1)宁夏枸杞的单花寿命为(4.07±0.15) d,而雄性持续时间(0.07±0.01 d)比雌性持续时间短(4±0.01 d);花寿命内雌蕊长度比雄蕊长,属柱头探出式异位类型;花粉呈现式样为不完全逐步呈现。(2)花寿命不同阶段花回馈间存在显著差异。(3)意大利蜜蜂(Apis mellifera)、熊蜂(Bombus sp.)、食蚜蝇(Syrphidae sp.)是宁夏枸杞在自然居群的主要访花者,其中熊蜂和食蚜蝇是主要传粉者,但传粉者效率低,属于高移出低沉积类型。(4)宁夏枸杞的花粉胚珠比(2 448.11±448.32)及授粉实验结果均表现出兼性异交特征;自发自交及人工自花授粉花的座果率及结籽率很低,属于自交不亲和类型;自然传粉花的座果率及结籽率比人工去雄异花低,存在其较高的花粉限制(40.71%)。研究发现,宁夏枸杞花部综合征表现出雌雄异位和花粉不完全逐步呈现式样,这是避免雌雄功能及雄蕊各花药间的干扰、减少花粉同步移出及保障其雄性适合度的有效途径,但自然居群的传粉者种类限制、传粉效率低以及自交不亲性是导致宁夏枸杞花粉限制及降低雌性繁殖成功的主要原因。     

Heterostylous morph differences in pollen transfer and deposition patterns in Primula sieboldii on a visitation by a queen bumblebee, measured with a semi-natural experimental system

Abstract Under a semi-natural setting the between-morph pollen exchange patterns were studied in distylous Primula sieboldii flowers by measuring pollen removal from the anthers on a single visit by a Bombus diversus tersatus queen, and stigmatic pollen deposition along the sequence of the visitation of the opposite-morph flowers by the bee. Despite the twofold larger number of pollen grains produced in a single flower of the long-styled morph compared to that of the short-styled morph, no significant difference in pollen removal from a flower was found between the morphs. The stigmas of the long-styled morph received significantly more opposite-morph pollen grains than those of the short-styled morph on a single visit by the bee. Sufficient legitimate pollen grains, surpassing the ovule number, were loaded on the stigmas of 27% and 17% of visited flowers of the long- and short-styled morphs, respectively. The short-styled morph could more efficiently donate pollen to the opposite morph stigmas than the long-styled morph.     

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.     

Both morph‐ and species‐dependent asymmetries affect reproductive barriers between heterostylous species

The interaction between floral traits and reproductive isolation is crucial to explaining the extraordinary diversity of angiosperms. Heterostyly, a complex floral polymorphism that optimizes outcrossing, evolved repeatedly and has been shown to accelerate diversification in primroses, yet its potential influence on isolating mechanisms remains unexplored. Furthermore, the relative contribution of pre‐ versus postmating barriers to reproductive isolation is still debated. No experimental study has yet evaluated the possible effects of heterostyly on pre‐ and postmating reproductive mechanisms. We quantify multiple reproductive barriers between the heterostylous Primula elatior (oxlip) and P. vulgaris (primrose), which readily hybridize when co‐occurring, and test whether traits of heterostyly contribute to reproductive barriers in unique ways. We find that premating isolation is key for both species, while postmating isolation is considerable only for P. vulgaris; ecogeographic isolation is crucial for both species, while phenological, seed developmental, and hybrid sterility barriers are also important in P. vulgaris, implicating sympatrically higher gene flow into P. elatior. We document for the first time that, in addition to the aforementioned species‐dependent asymmetries, morph‐dependent asymmetries affect reproductive barriers between heterostylous species. Indeed, the interspecific decrease of reciprocity between high sexual organs of complementary floral morphs limits interspecific pollen transfer from anthers of short‐styled flowers to stigmas of long‐styled flowers, while higher reciprocity between low sexual organs favors introgression over isolation from anthers of long‐styled flowers to stigmas of short‐styled flowers. Finally, intramorph incompatibility persists across species boundaries, but is weakened in long‐styled flowers of P. elatior, opening a possible backdoor to gene flow through intramorph pollen transfer between species. Therefore, patterns of gene flow across species boundaries are likely affected by floral morph composition of adjacent populations. To summarize, our study highlights the general importance of premating isolation and newly illustrates that both morph‐ and species‐dependent asymmetries shape boundaries between heterostylous species.