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。     

青篱柴通过促进亲和花粉生长而提高传粉精确性

异型花柱植物的适应意义在于提高亲和花粉的传递, 从而促进异交, 但是有不少研究发现柱头上落置了大量不亲和花粉。目前, 很少有人研究柱头是否有利于亲和花粉的萌发和花粉管伸长, 以及通过去雄处理排除花内和植株内不亲和花粉干扰从而验证二型花柱植物是否可以促进异交。本研究以亚麻科青篱柴属(Tirpitzia)的二型花柱植物青篱柴(T. sinensis)为研究对象, 分居群调查了青篱柴长、短柱型的植株数量, 测量不同柱型的花粉与柱头大小以及观察其表面纹饰, 测定不同开花时间长、短柱型的柱头活性与花粉活力, 统计自然状态下柱头上落置的不同表型花粉比例, 进一步统计花内、株内去雄和自然对照下柱头上落置的不同表型花粉比例, 计算型间、型内和混合授粉花粉管长度和花粉萌发率, 人工授粉检测其是否型内异交不亲和与自交不亲和。结果表明, 自然居群中长、短柱型青篱柴植株数量没有显著性差异; 短柱型青篱柴花粉体积显著大于长柱型, 且二者花粉表面纹饰不同, 但不同柱型的柱头表面积与表面纹饰无差异; 青篱柴开花第一天的花粉活力显著高于开花第二天, 而柱头活性在开花前两天无显著性差异; 自然状态下柱头上落置的不亲和花粉比例显著高于亲和花粉; 在长柱型青篱柴中, 花内和株内去雄显著提高了柱头上亲和花粉的落置比例; 柱头上授型内花粉, 其花粉萌发率和花粉管的长度都显著低于授型间花粉; 授粉结果表明青篱柴为型内异交不亲和与自交不亲和。二型花柱植物通过促进亲和花粉的萌发和花粉管的伸长而有利于型间授粉, 从而显著提高其传粉精确性。     

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.     

七叶树花粉活力和柱头可授性变化的研究

采用花粉离体萌发法研究不同蔗糖浓度、硼酸浓度和不同温度对七叶树花粉萌发的影响及花粉活力测定,用联苯胺-过氧化氢法测定柱头可授性.结果表明:七叶树花粉萌发的最佳培养基是12％蔗糖+30 mg/L硼酸,花粉萌发的最适温度为25℃.雄花在开花当天花粉活力最高达75.69％,并在开花当天的上午10:00时,花粉活力最强,10:00～16:00花粉活力保持较高活力,是最佳授粉时段.两性花的柱头可授期持续时间较长,为8～9 d,开花3d达到最强,开花1～4 d柱头可授性保持较高水平,为授粉的最佳时间段.因此,从七叶树的花部特征、花粉活力与柱头可授性及花粉萌发的条件看,在长期的自然选择下七叶树在花部结构和开花生理上都是相配合的,以保障生殖成功.     

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。     

琼花生殖器官结构及传粉昆虫的观察

为了探讨影响琼花Viburnum macrocephalum f.keteleeri有性生殖的因素,对其生殖生物学和传粉生物学进行了研究。研究内容主要包括琼花花部特征、生殖器官解剖结构、花粉活力、花粉胚珠比(P/O)、繁育系统、传粉形式、传粉昆虫和花粉管生长路径等。结果表明:(1)琼花聚伞花序由大型不孕花和小型可孕花组成,可孕花雌雄蕊发育正常,雄蕊5枚,雌蕊1枚,干型柱头,单子房,倒生胚珠;不孕花的雌雄蕊在发育早期正常,而在花期时退化。雄蕊退化表现为雄蕊消失、花丝缩短或消失或花药大小不一;而雌蕊退化表现为雌蕊缩小或柱头开裂;有时雌雄蕊也存在瓣化现象。(2)自然条件下,单花花粉活性在散粉4–5d后明显下降,居群花粉活力在4月25日–28日开始显著下降。(3)花粉胚珠比P/O值为12800–18700,属专性异交型;繁育系统为异株异花授粉,属虫媒传粉植物。(4)可孕花大量散粉时间为9:00–16:00,昆虫访花高峰时段为11:00–15:00,访花昆虫中以蝶类和蜂类为主,蝶类访花频率最高。(5)落置柱头的花粉萌发率较高,花粉粒授粉后1h左右开始萌发,花粉管从柱头乳突细胞的间隙穿入花柱,沿花柱中央引导组织生长,18h左右进入子房,20h左右从珠孔进入胚囊。讨论了琼花的不孕花形成、花部结构适应、花型演化趋势、低结实率和花粉管生长特点。     

珍稀濒危植物蒙古扁桃花生物学特性

蒙古扁桃的植株可分为长花丝植株、短花丝植株和中花丝植株。群落花期约50d,单花花期约8d,分为露粉、微开、盛开、凋谢4个时期。过氧化物法测定4个时期花粉均具活力,可保持30d左右,联苯胺-过氧化氢测定柱头可授性,花粉活力与柱头可授性重叠,长花丝植株为8d左右,而中花丝约为5d。蒙古扁桃花一般在9:00开始泌蜜,11:00分泌量达到高峰,之后产蜜量减少直至停止,日泌蜜和散粉集中在10:00～14:00。蒙古扁桃开花受环境的影响。     

异型花柱植物喀什补血草的传粉者功能群与花粉转移效率

异型花柱是受遗传控制的花柱多态现象, 被达尔文认为是植物通过在传粉者体表不同部位滞落花粉以促进型间花粉准确传递的一种适应。该现象虽已受到广泛关注, 但在一些花型变异较大且不稳定的传粉系统中, 不同传粉者对各花型繁殖所产生的影响仍知之甚少。该研究以分布于新疆天山南坡的一个有同长花柱共存的异型花柱植物喀什补血草(Limonium kaschgaricum)种群为研究对象, 对其花型构成及频率、传粉者及花粉转移效率等进行了调查分析。结果表明: 1)种群中除了存在雌/雄蕊长度交互对应的长(L)/短(S)花柱型花外, 还有雌/雄蕊同长的花(H型), 且各花型花的花冠口直径、花冠筒长及花粉量等参数间无差异, 但花粉纹饰和柱头乳突细胞形态具二型性。其中, H型花的花粉和柱头形态与L型花(或S型花)的一致。2)花型内和自花授粉均不亲和; 型间授粉时, 花粉和柱头形态不同的花型间亲和, 反之不亲和。3)种群内存在长/短吻两类传粉昆虫。在以短吻传粉者为主的盛花初、中期, L和H型花柱头上的异型花粉数均显著高于S型花的, 且L和S型花高位性器官间的异型花粉传递效率高于低位性器官间的; 而在以长吻传粉者为主的盛花后期, L和S型花的柱头间异型花粉数无显著差异, 且高/低位性器官间具有相同的异型花粉转移效率; 与传粉者出现时期相对应的、在花期不同阶段开放花的结实率也明显不同。4)长/短吻昆虫具明显不同的传粉功能, 短吻昆虫只能对L和H型花进行有效传粉, 且访花频率和型间花粉转移效率较低, 为低效传粉者; 而长吻昆虫对各花型均能有效传粉, 具高的访花频率和型间花粉转移效率, 为高效传粉者。因为长吻昆虫的阶段性出现所形成的不稳定传粉系统, 使低效的短吻昆虫可能会成为种群中花型变异的驱动力, 并使S型花受到更大的选择压力。H型花克服了柱头缩入的弊端, 可能会成为不稳定传粉系统下的一个替代花型而持续存在。     

珍稀濒危植物陕西羽叶报春的开花及传粉生物学研究

陕西羽叶报春为中国珍稀濒危植物,研究其开花物候、传粉者及访花行为、花粉活力、柱头可授性、繁育系统,以明确陕西羽叶报春的传粉特性及制濒机制。以勉县陕西羽叶报春野生居群为研究对象,通过野外观察和人工授粉实验首次对陕西羽叶报春的传粉生物学进行研究。结果表明:(1)陕西羽叶报春种群花期为2-4月中旬,单花花期一般为14~17 d;单花开花进程中花部形态有6个明显发育变化:全部被萼片包裹、花苞伸出并为深粉色、花朵打开为淡粉色、花冠为白色、花朵枯萎、花冠脱落花萼宿存。(2)陕西羽叶报春为异花授粉植物,蜜蜂属为其主要传粉者。(3)柱头在花粉未开裂前就具有可授性,可授性逐渐增强;花药在花瓣打开后的第3天才开始开裂、散粉,在第4天活力最高,为98.18%,此后逐渐减弱。(4)繁育系统检测显示,长、短柱型花的花粉/胚珠比分别为902.26和831.48,杂交指数为4;套袋实验显示,陕西羽叶报春异型交配可育,同型交配不可育。由此表明,陕西羽叶报春的繁育系统为异交型,自交不亲和,需要传粉者。综上所述,陕西羽叶报春传粉者种类少、自交不亲和、胚珠败育现象可能是导致陕西羽叶报春濒危的重要原因。     

南瓜雌蕊与自花及远缘花粉的相互作用

南瓜柱头表面经去垢剂、蛋白酶及Con A处理后花粉不能萌发或花粉管生长受阻,Con A能专一地与柱头表面结合。柱头块加入培养液可促进花粉萌发。不同的远缘花粉授粉后在雌蕊不同部位受阻。在成熟南瓜雌蕊提取液中检测到血凝活性,凝集素可能参与雌蕊对远缘花粉的抑制。     

Pollen and stigma size changes during the transition from tristyly to distyly in Oxalis alpina (Oxalidaceae)

• Pollen and stigma size have the potential to influence male fitness of hermaphroditic plants, particularly in species presenting floral polymorphisms characterised by marked differences in these traits among floral morphs. In this study, we take advantage of the evolutionary transition from tristyly to distyly experienced by Oxalis alpina (Oxalidaceae), and examined whether modifications in the ancillary traits (pollen and stigma size) respond to allometric changes in other floral traits. Also, we tested whether these modifications are in accordance with what would be expected under the hypothesis that novel competitive scenarios (as in distylous‐derived reproductive system) exert morph‐ and whorl‐specific selective pressures to match the available stigmas.
• We measure pollen and stigma size in five populations of O. alpina representing the tristyly–distyly transition.
• A general reduction in pollen and stigma size occurred along the tristyly–distyly transition, and pollen size from the two anther levels within each morph converged to a similar size that was characterised by whorl‐specific changes (increases or decreases) in pollen size of different anthers in each floral type.
• Overall, results from this study show that the evolution of distyly in this species is characterised not only by changes in sexual organ position and flower size, but also by morph‐specific changes in pollen and stigma size. This evidence supports the importance of selection on pollen and stigma size, which increase fitness of remaining morphs following the evolution of distyly, and raises questions to explore on the functional value of pollen size in heterostylous systems under pollen competition.

     

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。     

Reproduction in Seagrasses: Nature of the Pollen and Receptive Surface of the Stigma in the Hydrocharitaceae

This paper describes the characteristics of the pollen and thereceptive surfaces of the stigmas in the three marine angiospermsincluded in the Hydrocharitaceae. The pollen in Enhalus acoroidesand Thalassia hemprichii is spherical and has an ornamentedexine. An exine layer is not found in Halophila stipulacea wherereniform pollen grains are contained within transparent moniliformtubes. Cytochemical tests show that the pollen wall in the threespecies contains acidic and neutral polysaccharides and acidhydrolase acitivity is detected in the intine of H. stipulaceaand T. hemprichii. In Thalassia, one of the intine enzymes,acid phosphatase, is unambiguously associated with cytoplasmicinclusions. Flowering in Thalassia is coincident with the spring tides andthe pollen is released as a mass suspended in a thecal slimewhich contains approximately 5 per cent by weight carbohydrate,the principal mono-saccharide being mannose. Electrophoreticanalysis of the pollen-free slime shows a single glycoproteincomponent. The stigmas of the three seagrasses are papillate and of the‘dry’ type possessing a continuous protein-aceouspellicle subtended by a cuticle. The stigma pellicle exhibitscytochemically detectable esterase activity and binds the lectinconcanavalin A. Acid phosphatase activity is localized beneaththe cuticle at the tips of the stigma papillae. The discoveries show that the characteristics of the pollenand stigmas in the seagrasses are comparable with those foundin terrestrial flowering plants. The similarity in enzymaticproperites of the pollen wall and stigma pellicle suggests that,intriguingly, a similar mechanism of cuticle erosion might wellfollow compatible pollination both on land and in the sea. Enhalus acoroides, Thalassia hemprichii, Halophila stipulacea, Halophila decipiens, seagrasses pollen wall, stigma surface, hydrolytic enzymes     

Abnormality in Male Organ Development and Fertility of Photoperiod sensitive Genic Male Sterile Rice Plants Under Short Day Condition

Ten photoperiod-sensitive genic male sterile rice (PGMSR) lines were studied to observe the development of male organs of their short day (SD) plants. The results were shown as follows: 1 ) The self seed-setting rate and the percentage of fertile pollen of these PGMSR lines were much lower than those of the controls. 2) All the male sterile line plants had variant barriers in anther dehiscing and pollen scattering. In addition, some of their stigmas were devoid of pollen, and some self pollen grains on their stignas germinated weakly. 3) Female fertility of those plants was believed to be normal through observation in their embryo sac development and cross seed-setting rate. The authors deduced that the male sterile gene expression becomes the major factor for low rate of self seed-setting of PGMSR plants under SD condition.     

花柱卷曲性异交机制及其进化生态学意义

 有花植物具有纷繁复杂的繁育系统,以避免或促进自交或异交。花柱卷曲性异交机制(Flexistyly)是最近在热带山姜属(Alpinia)植物中发现的一种促进异交的行为机制。具有这一机制的种类其自然种群中的个体根据开花行为的不同分为两种表型：一种上午散发花粉而其柱头向上反卷,远离昆虫拜访的通道;另一种其柱头上午垂向唇瓣,能够接受拜访昆虫的传粉,但自身的花药却不打开。到中午时分,两种表型花通过互为相反的柱头卷曲运动转换性别——前者柱头向下卷曲,后者柱头向上卷曲且花药打开。每朵花的花期为12 h,两种表型在自然种     

Pollen-Stigma Adhesion in Kale Is Not Dependent on the Self-(In)Compatibility Genotype

The adhesion of pollen on the stigmas of flowering plants is a critical step for the success of reproduction in angiosperms, long considered to present some specificity in terms of self-incompatibility. We carried out quantitative measurements of the pollen-stigma adhesion (expressed in Newtons) in kale (Brassica oleracea), using the flotation force of Archimedes exerted by dense sucrose solutions (50%, w/v) to release pollen grains fixed on the surface of stigmas. We demonstrate that pollen adhesion varies with the genotypes of the plants used as partners, but increases with time in all cases for about 30 to 60 min after pollination. There is no correlation with the self- or cross-status of the pollinations, nor with the self-compatible or -incompatible genotypes of the parents. Only late events of pollination, after the germination or arrest of the pollen tube, depend on compatibility type. Biochemical and physiological dissection of pollen-stigma adhesion points to major components of this interaction: among male components, the pollen coating, eliminated by delipidation (or modified by mutation in the case of the cer mutants of the related species Arabidopsis thaliana), plays a major role in adhesion; the genetic background of the pollen parent is also of some importance. On the female side, the developmental stage of the stigma and the protein constituents of the stigmatic pellicle are critical for pollen capture. The SLG and SLR1 proteins are not involved in the initial stages of pollen adhesion on the stigma but one or both may be involved in the later stages.     

新疆荒漠植物耳叶补血草的爆发式开花式样与传粉模式

荒漠环境中干热、大风、浮尘、温度骤变等突发性天气状况,对植物的有性生殖具不利影响.耳叶补血草(Limonium otolepis)是新疆盐生荒漠的主要建群种,对其适应荒漠恶劣环境的生殖策略仍缺乏研究.为了探讨荒漠植物适应环境的开花式样和传粉模式,作者通过野外定点观测和室内电镜扫描等方法对耳叶补血草的开花进程、花部特征、传粉媒介、结实特性等进行了研究.耳叶补血草的花期在6月上旬至7月中旬,花期较长且具明显不同步的单株开花状态.在一天中,单花表现出极为集中的开花、散粉过程:一般早上8:00开始开放,9:00达开放高峰期,11:00以后基本不再有单花开放;开放后的单花持续6-8 h后闭合.每单花平均有花粉752粒,花粉表面有网状纹饰,有花蜜.传粉者主要为蜜蜂科和食蚜蝇科昆虫,访花高峰期在9:00-14:00.人工授粉实验表明耳叶补血草以异交为主,自动自花授粉率低.自然状态下结实率为36％,在花序中基部第一位上单花的结实率(45.7％)明显高于第二位上单花(3.3％).耳叶补血草爆发式开花、在短时间内快速完成传粉过程,可能是躲避荒漠环境突发性天气条件的一种策略;而居群分散的开花时间及较长的花期在应对不稳定生境,分摊生殖风险方面具重要意义.     

Phenotypic Characterization of a Female Sterile Mutant in Rice

A female sterile mutant, derived from a spontaneous mutation, was first discovered In rlce （Oryzs satlvs L. sep. Indlca） restorer llne 202R. Wlth normal fiowerlng, the mutant exhlblts an extremely low seed-sattlng rate. When the mutant Is crossed as a pollen donor, the seeds set normally; whereas when It Is used as a pollen recelver, no seeds are obtalned even wlth mlxed pollen gralns of dlfferent varletles sprlnkled over the atlgmas. The fioret of the mutant, conslatlng of slx stamens and one platll, looks the same as that of the wlld type In the malefemale organs, except that less than 10% of the mutant florets have three atlgmas on the ovary. Although the mutant has a low seed-setting rate, Its pollen fertility Is approximately 87.1%, which Is equal to that of the wild type. In addition, more than 90% of the mature embryo sacs of the mutant have complete Inner structures. At every stage after pollination, the sperm, embryo, and endosperm are not found In the mutant embryo sac, whereas the disintegration of the egg cell that does not accomplish fertilization Is visible. Through observetlons with a fluorescence microscope, we have found that the pollen grains germinate normally, whereas the pollen tube abnormally elongates in the style-transmitting tissue. The mutant pollen tubes display various defects In the style, such as slower elongation, conversed elongation, distorted elongation, swollen tips, or branched tips. As a result, the growth of the pollen tubes ceases In the style, and, therefore, the pollen tubes cannot reach the embryo sac and the process of double fertilization Is blocked. Based on these observations, we conclude that this mutant, designated as fs-202R, Is a novel type of female sterile mutation In rice, which causes the arrest of the elongation of the pollen tube.