新疆野生三叶草弹花机制及繁殖特性

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王玉祥, 张博. 新疆野生三叶草弹花机制及繁殖特性. 草业科学, 2016,33(4):662-668
Wang Yu-xiang, Zhang Bo. Research on tripping phenomenon and propagation characteristics of wild Trifolium in Xinjiang . Pratacultural Science,2016,33(4): 662-668 复制到剪切板

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《草业科学》编辑部

新疆野生三叶草弹花机制及繁殖特性

王玉祥, 张博

新疆草地资源与生态重点实验室,新疆农业大学草业研究所,新疆乌鲁木齐 830052

张博(1963-),男,新疆五家渠人,教授,硕士,主要从事牧草遗传育种研究。E-mail:[email protected]

第一作者:王玉祥(1980-),男,安徽亳州人,讲师,硕士,主要从事牧草遗传育种和种子生产研究。E-mail:[email protected]

收稿日期: 2015-12-08

基金: 新疆维吾尔自治区重点实验室开放课题“新疆野生白三叶的形态和生理指标与抗寒性的关系研究”(XJDX0209201304)

摘要

为了解新疆三叶草( Trifolium)弹花现象和繁殖特性,本研究对三叶草开花习性、雄蕊散粉时间及不同交配方式对结实性的影响进行研究。结果表明:1)单花从现蕾至萎蔫持续时间为50~52 h,花粉展示属于龙骨瓣闭合不严;2)雌蕊柱头在翼瓣生长至旗瓣长1/2处时开裂,雄蕊在翼瓣完全打开时开始散粉;3)自主自交中昌吉三叶草结实率最高,为98.61%;同株异花授粉中乌鲁木齐南山山地草甸的三叶草结实率最高,为100%;自由授粉中呼图壁林带的三叶草结实率最低,为96.30%。本研究结果可为进一步开展三叶草的繁育系统和生殖生物学的研究提供参考依据。

关键词: 三叶草; 开花习性; 散粉特性; 繁殖

中图分类号:S541⁺.203.8 文献标志码:A 文章编号:1001-0629(2016)4-0662-07 doi: 10.11829/j.issn.1001-0629.2015-0480

Research on tripping phenomenon and propagation characteristics of wild Trifolium in Xinjiang

Wang Yu-xiang, Zhang Bo

Key Laboratony of Grassland Resource and Ecology of Xingjiang, Pratacultural Institute of Xinjiang Agricultural University, Urumqi 830052, China

Corresponding author: Zhang Bo E-mail:[email protected]

Abstract

To understand tripping phenomenon and reproductive characteristics of Xinjiang Trifolium, flowering habit, pollen spreading time and the influence of different mating pattern on reproductive characteristic were studied. The results as followed: 1) flower from budding to wilting maintained 50~52 h, pollen display belongs to keel valvular insufficiency; 2) the pistil stigma crazed when wing petal grow to 1/2 of flag petal; 3) the podding rate of Trifolium from Changji desert steppe in the autonomous selfing was highest (98.61%); the podding rate of Trifolium from Urumqi Nanshan mountain meadow under homologous strain cross-pollination was the highest (100%) ; the podding rate of Hutubi Trifolium under forest belt with free pollination was the lowest (96.30%). The results of this study can provide some references for further study on breeding system and reproductive biology of Trifolium repens.

Keyword: Trifolium; flowering habits; pollination features; propagation

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蝶形花亚科(Papilionideae)植物具有4种式样的花粉展示方式, 分别是毛刷机制、活塞机制、弹花机制和瓣膜机制^{[1, 2]}。弹花机制又称为爆炸机制, 是指昆虫作用于翼瓣和龙骨瓣, 当龙骨瓣破裂时其内部的张力被释放, 同时使花粉瞬间“ 喷出” , “ 溅到” 昆虫身上(主要是头部和腹部)。Leppik^[3]提出弹花机制是蝶形花植物在进化中形成的最经济和最有效的花粉释放机制, Atluril等^[4]认为花粉释放是借助外力作用使花器内累积的能量突然释放出来, 在使花柱暴露的同时也把花粉释放出来, 因此, 花粉的爆炸释放本质上是能量的释放。另外, 有研究发现还存在鸟媒爆炸机制, 当鸟落在花上时, 由于鸟的重量对花的作用, 使花腔内受力, 花就会瞬间爆炸, 将花粉释放到鸟的前额上^[5]。如, 苜蓿属(Medicago)植物一旦受到外界压力就会出现将花药柱头紧紧包裹着的龙骨瓣弹开, 花药和柱头弹向并紧贴翼瓣, 不再回到原来的位置; 而三叶草属(Trifolium)植物的花粉展示式样为龙骨瓣闭合不严, 拜访者采用外力促使龙骨瓣打开, 在拜访过程内使柱头和花药露出, 在拜访者离开的时候, 柱头和花药又被隐藏起来。

从适应异花授粉的方式来看, 有花植物的习性可分为三大类:雌雄异位、雌雄异熟和自交不亲和系统^{[6, 7]}。自交是指同一朵花或同一个植物体内的传粉、受精; 而异交是指同一种植物中的不同个体间的传粉、受精, 这种交配方式能够促进遗传多样性的形成, 是形成繁育系统中传粉多样性的主要动力^{[8, 9, 10]}。三叶草为异花授粉植物, 大部分不能自花授粉, 自交不亲和是结实率低的重要原因。目前, 国内对植物交配系统的研究大都集中于濒危稀有植物中, 而有关三叶草的繁育特性的研究相对较少。新疆野生三叶草物种资源丰富, 由于其独特的地理位置和自然风貌, 三叶草在长期逸生蔓延和自然选择的影响下形成了不同类型的表型特征, 不同群落间具有明显的表型差异, 并具有较强的抗逆性, 是优良的种质资源。有关新疆野生三叶草的研究不多, 仅有王玉祥等^{[11, 12]}、潘奥等^[13]对新疆三叶草形态特征的研究, 而未见关于新疆野生三叶草开花习性和繁殖特性的报道。基于此, 新疆农业大学通过长期的研究比较, 从野生群落中筛选出5个[白三叶(Trifolium repens)4个, 草莓三叶草(T. fragiferum)1个]综合性状较好的材料, 开展三叶草开花习性、雄蕊散粉时间以及不同交配方式对结实性影响的研究, 探寻新疆野生三叶草的弹花机制和繁殖特性, 以期为进一步开展新疆野生三叶草的交配系统、繁殖特性、纯合基因型等研究提供参考依据。

1 材料与方法

1.1 试验材料

试验材料为新疆野生三叶草, 由新疆草地资源与生态重点实验室提供(表1), 种植在新疆农业大学呼图壁草地生态站。

表1 材料信息 Table 1 Information of material

1.2 试验地概况

试验地设在乌鲁木齐以西80 km的新疆农业大学呼图壁草地生态试验站, 该站地处准葛尔盆地南缘, 属于中亚型干旱荒漠气候。土壤主要是盐化草甸土, 盐分含量较高, pH> 8.5^[14]。

1.3 研究内容及方法

试验以新疆野生三叶草为材料, 开展三叶草开花习性、雄蕊散粉特性以及同株同花(自花)授粉、同株异花授粉和自由(开放)授粉对结实性的影响研究。

开花习性观测:在单花现蕾期, 每个材料随机选择50朵单花, 从单花现蕾至单花萎蔫, 从06:00时至20:00时、每隔1 h记录1次单花的发育状态。

雄蕊散粉时期观测:由于E材料开花相对较其它材料早, 因而以其为标样开展雄蕊散粉特性研究。在单花现蕾期, 随机标定20个花序, 从单花露白开始至翼瓣完全展开, 每隔2 h采集10朵发育一致的单花进行镜检, 同时记录单花的发育状态。

单花的选择:在三叶草花期, 随机选取长势一致的花序, 剔除不合格单花、保留发育良好且生长一致的单花4~5朵, 每个处理随机取40~50个花序、200朵单花进行试验, 处理后套袋隔离。其中, 每个处理的200朵单花随机分为5组, 每40朵单花作为一个重复进行处理, 以便于统计分析。

同株同花授粉方法是用食指和拇指轻轻挤压即将绽放的单花的龙骨瓣, 让雌蕊和雄蕊充分接触, 达到自花授粉的目的; 同株异花授粉方法是先轻轻剥开单花的龙骨瓣, 用镊子、棉花棒等去除雄蕊, 然后摘取同株异花的3~5朵成熟单花的雄蕊混合后放到雌蕊上, 让雌雄蕊充分接触, 以保证授粉效率; 自由授粉方法是单花不去雄, 先轻轻剥开龙骨瓣, 直接摘取异株的3~5朵单花的雄蕊混合后放到雌蕊上, 让雌雄蕊充分接触, 然后闭合龙骨瓣。

授粉时间:06:30-10:00进行授粉工作, 为保证试验处理和结果的准确性, 授粉的工具均为一次性用品。

种子成熟后进行单独收获, 拷种。统计结种子荚果数、单荚种子数、粒长、粒宽和千粒重等。其中, 每个处理随机选取30~50粒种子, 用游标卡尺(± 0.02 mm)测量粒长和粒宽; 同时用万分之一电子天平测量千粒重, 每个处理重复5次, 每次50~100粒种子。

结实率=含种子荚果数/处理小花数× 100%。

1.4 数据处理

利用SPSS 17.0软件对试验数据进行方差分析。

2 结果与分析

2.1 开花习性与弹花机制

三叶草的开花习性观测结果表明(表2), 三叶草的单花从现蕾至萎蔫持续时间一般为50~52 h。其中, 单花从现蕾至旗瓣露白需要2 h, 旗瓣露白至旗瓣完全伸出萼片需要4~6 h, 从旗瓣伸出花萼到单花萎蔫需要44~46 h, 一般单花集中在12:00-16:00时开放。其中, A、B、D、E材料的单花集中在12:00-14:00时开放, 持续2 h; C材料的单花集中在12:00-16:00时开放, 持续4 h。原始生境不同, 三叶草的开花习性也不尽相同, C材料花瓣露白至完全伸出花萼及单花现蕾至萎蔫持续时间均最长, 这可能是因为C材料来自荒漠草地, 因其生长环境恶劣, 为保证基因的延续, 产生的一种适应环境的生殖策略; A和E材料的单花展示时间一致, 可能是因为其生长环境是河谷和草甸的原因, 二者温湿度相对平和, 有助于其生长发育, 仍有待于进一步验证。

表2 小花展示时间 Table 2 Display time of flora

豆科植物单花花粉的展示式样从空间和时间上可分为4种:1)龙骨瓣不闭合或闭合不严; 2)龙骨瓣具喙, 上部闭合; 3)具次级花粉展现, 即花粉被雌蕊的上部带到龙骨瓣的外面; 4)爆破式花粉展现。试验通过对昆虫访花过程和三叶草弹花授粉过程进行观测发现, 昆虫停留在龙骨瓣上时, 龙骨瓣没有像苜蓿的花朵一样弹开, 而是先在昆虫体重的压力作用下向下凹向柱头, 此时昆虫再利用前足轻轻挑开龙骨瓣, 取食花蜜, 同时进行传粉受精, 当昆虫取食花蜜结束, 整个传粉受精过程结束, 昆虫离开后龙骨瓣恢复原状。即三叶草在花粉展示过程中, 龙骨瓣并未完全打开, 龙骨瓣始终处于半闭合状态, 昆虫利用龙骨瓣之间的间隙为花完成传粉受精过程。因此, 三叶草花粉展示属于龙骨瓣闭合不严, 即昆虫授粉时龙骨瓣保持半闭合状态。

2.2 雄蕊散粉时期

在开展植物自交和杂交的试验中, 去雄是否彻底是一个关键步骤, 如果去雄不彻底将会严重影响试验的准确性和可靠性, 在雄蕊散粉前进行去雄, 既能够降低工作量, 又能够提高授粉效率; 如果在雄蕊散粉后开展自交, 将增加试验的难度, 对试验的结果具有一定的影响。因此, 自交试验开展前, 试验以开花相对较早的E材料为研究对象, 对三叶草雄蕊的散粉时期进行观测表明(图1), 在旗瓣完全打开时, 雌雄蕊均未开裂散粉; 当翼瓣生长至旗瓣长1/2处时, 雌蕊柱头已经开裂, 雄蕊仍旧未散粉; 当翼瓣生长至完全打开时, 雄蕊的花粉开始散开。因此, 三叶草最佳的去雄时期是从翼瓣开始伸长至生长到旗瓣长1/2处时, 这期间雌蕊柱头已经开裂, 雄蕊花粉粒逐渐发育成熟并即将散粉。

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	图1 雌雄蕊发育过程Fig.1 Development of female and stamen

2.3 不同交配方式对结实性的影响

不同交配方式对三叶草的结实率、单荚种子数、种子特征有影响(表3)。其中, 同株同花自交试验中, B材料结实率最高, 为98.61%, 且与其它处理差异极显著(P< 0.01), A材料次之, E材料结实率为0, 说明B材料同株同花自交亲和性高, E材料自交严格不亲和; 同株异花授粉试验中, 材料A、B、C、E结实率均为100%, 且极显著高于同株同花授粉(P< 0.01), 说明同株异花授粉能够提高结实率; 在自由授粉条件下, 5个材料的结实率均大于96%, 材料间差异不显著(P> 0.05)。材料A、B、C、E的自由授粉结实率均高于同株同花, 但低于同株异花的授粉结实率, 且A、C、E的结实率在3种处理之间的差异极显著(P< 0.01); 只有材料D的自由授粉结实率均高于同株同花和同株异花, 且差异极显著(P< 0.01)。不同材料, 交配方式不同, 结实率差异较大, 尤其是自由授粉的结实率低于同株异花的授粉结实率, 这可能与材料自身的遗传基因有关, 也可能与异株的单花数量、花粉数量等有关, 有待进一步深入研究。通过对试验数据进一步分析表明, 不同交配方式对三叶草的结实率影响较大, 同株异花授粉和开放授粉的结实率均高于同株同花授粉, 即异交和杂交能够提高三叶草结实率。

表3 不同交配方式对结实率和种子特征的影响 Table 3 Effects of different mating types on seed setting rate and seed characteristics

材料编号 Material code	特征 Characteristics	同株同花授粉 Self pollination	同株异花授粉 Geitonogamy	自由授粉 Open pollination
A	结实率Sit pod rate/%	25.00± 1.42dD	100.00± 0.00aA	98.75± 0.57bB
	单荚种子数Seed number per pod	2.33± 0.42bB	2.35± 0.21bB	1.60± 0.42cC
	粒长Grain length/mm	1.13± 0.03cC	1.10± 0.67cC	1.26± 0.28bB
	粒宽Grain width/mm	0.90± 0.14cC	1.01± 0.01cC	1.00± 0.03cC
	千粒重1 000 Grain weight/g	4.60± 0.29dD	5.60± 0.49bB	5.50± 0.85bB
B	结实率Sit pod rate/%	98.61± 2.76bB	100.00± 0.00aA	98.68± 0.71bB
	单荚种子数Seed number per pod	1.45± 0.26cC	1.79± 0.14Cc	1.20± 0.14dD
	粒长Grain length/mm	1.53± 0.08aA	1.57± 0.07aA	1.61± 0.01cC
	粒宽Grain width/mm	1.10± 0.18abAB	1.20± 0.16aA	1.24± 0.57aA
	千粒重1 000 Grain weight/g	11.93± 0.42aA	12.56± 0.14aA	13.50± 0.78aA
C	结实率Sit pod rate/%	4.01± 0.43eE	100.00± 0.00aA	98.04± 0.57bB
	单荚种子数Seed number per pod	1.00± 0.71eE	1.87± 0.23cC	2.40± 0.14bB
	粒长Grain length/mm	1.02± 0.04cC	1.10± 0.69cC	1.15± 0.06cC
	粒宽Grain width/mm	0.78± 0.05Dd	0.96± 0.07cC	0.94± 0.03cC
	千粒重1 000 Grain weight/g	3.00± 0.28eE	5.60± 0.42bB	4.50± 0.14dD
D	结实率Sit pod rate/%	1.41± 0.26eE	77.78± 1.27cC	96.30± 0.85bB
	单荚种子数Seed number per pod	1.00± 0.28eE	0.29± 0.28fF	3.80± 0.42aA
	粒长Grain length/mm	1.14± 0.03cC	1.12± 0.03cC	1.14± 0.04cC
	粒宽Grain width/mm	0.82± 0.05cdCD	0.94± 0.08cC	0.91± 0.01cC
	千粒重1 000 Grain weight/g	5.00± 0.42cC	4.30± 0.21dD	4.90± 0.21cC
E	结实率Sit pod rate/%	0.00± 0.00fF	100.00± 0.00aA	98.11± 0.85bB
	单荚种子数Seed number per pod	0.00± 0.00gG	2.34± 0.14bB	1.80± 0.50cC
	粒长Grain length/mm	0.00± 0.00dD	1.23± 0.01bB	1.13± 0.01cC
	粒宽Grain width/mm	0.00± 0.00eE	1.03± 0.03cC	0.95± 0.08cC
	千粒重1 000 Grain weight/g	0.00± 0.00fF	5.70± 0.35bB	4.90± 0.49cC

Note: Different capital and lower case letters within the same parameter mean significant difference at 0.01 and 0.05 level, respectively.

注:同一指标不同大小写字母表示差异极显著(P< 0.01)和显著(P< 0.05)。

表3 不同交配方式对结实率和种子特征的影响 Table 3 Effects of different mating types on seed setting rate and seed characteristics

同株同花授粉试验中, 单荚种子数最多是材料A, 为2.33粒· 荚^-1, 材料B次之, 为1.45粒· 荚^-1, 材料C和D均为1粒· 荚^-1, 材料E无种子; 在同株异花授粉试验中, 材料E单荚种子仅次材料A(2.35粒· 荚^-1), 为2.34粒· 荚^-1, 材料D最少, 为0.29粒· 荚^-1; 而在自由授粉条件下, 单荚种子数较多的是材料D, 为3.8粒· 荚^-1, 材料C次之, 为2.4粒· 荚^-1, 材料B最少, 为1.2粒· 荚^-1。材料A、B、E同株异花授粉的单荚种子数均高于同株同花和自由授粉, 而C、D自由授粉的单荚种子数均高于同株同花和同株异花, 且差异极显著(P< 0.01)。

不同交配方式对三叶草种子粒长、粒宽和千粒重有影响。以材料A为例, 自由授粉的种子粒长较长, 同株异花授粉的粒长较短; 同株异花授粉的种子粒宽相对较宽, 同株同花授粉的粒宽较窄; 同株异花授粉的种子千粒重较大, 为5.60 g, 自由授粉次之, 同株同花的最小。试验数据表明, 同种材料, 交配方式不同, 种子粒长、粒宽和千粒重数值不同, 差异的显著性也不同。

通过对试验数据进一步分析表明, 三叶草的结实率、单荚种子粒数、种子特性等不仅受材料本身遗传特性的影响, 还与其交配方式有关, 交配方式不同, 花粉来源、数量、质量不同, 授粉效率不同, 对花粉粒的萌发、花粉管和子房发育等均产生影响, 从而影响种子的数量和质量。

3 讨论与结论

植物不同的开花习性对植物的繁衍、基因的延续有着重要的影响^{[15, 16]}。本研究通过对新疆三叶草的开花习性观测发现, 单个小花从现蕾至萎蔫持续时间为50~52 h, 单花的绽放时间主要集中在12:00-16:00, 且授粉时龙骨瓣始终处于半闭合状态, 授粉结束后闭合, 这与苜蓿属植物完全不同。苜蓿属植物受到外界压力后龙骨瓣弹开不能复原, 花药和柱头弹向并紧贴翼瓣暴露在空气中^[6]。

授粉是种子植物有性生殖的一个重要环节, 人工去雄时要确保植物花粉未散粉, 即植物未受精^{[17, 18]}。新疆野生三叶草雌蕊柱头在翼瓣生长至旗瓣长1/2处时开裂, 雄蕊在翼瓣完全打开时开始散粉, 三叶草柱头开裂时期在散粉, 因而在翼瓣展开前去雄为宜。

自交不亲和性是大多数高等植物防止近亲繁殖的一种遗传屏障, 自交亲和能有效确定植物基因的纯度。自交不亲和性作为一种抵制近亲繁殖、保持种群遗传多样性的重要机制普遍存在于植物(特别是被子植物)中^{[19, 20, 21, 22]}。植物的自交不亲和性使植物处于高度的杂合状态, 最大程度地保持物种活力, 对其生存和繁衍及其有利。自交不亲和性植物都具有强度不同的不亲和性, 其花期自交坐果率或结实率一般介于0~5%^{[23, 24, 25]}。而新疆野生三叶草自交结实率差异较大, 有的材料高度自交不亲和, 而有的材料亲和度为100%, 这可能与三叶草的基因或者生境有一定的关系。通过对新疆野生三叶草的弹花机制和繁殖特性的研究, 能为三叶草的杂种优势的保持、物种的进化、品质的改良提供有利条件。同时, 通过自交亲和度的研究, 能够依据材料自身特点保持后代基因型的纯合性, 对纯和基因的研究有着重要的意义。

本研究对新疆三叶草的开花习性、散粉特性和繁殖特性等进行了初步探索, 结果能够为进一步开展三叶草研究提供参考依据。但研究仅仅得到了结果, 而没有对促成这种结果的具体原因进行进一步的研究探索, 如交配方式不同, 结实率、单荚种子数、种子大小等不同, 仅仅是材料本身遗传特性造成的吗, 这与其生长的环境、授粉亲本的大小、所授花粉的数量和质量等有内在联系吗, 目前都是未知数, 有待于进一步探究, 这也是以后研究的侧重点。

The authors have declared that no competing interests exist.

参考文献

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2014

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黄利春, 金樑, 李晶, 张晓强, 杨阳, 王晓娟. 蝶形花亚科植物花部适应机制与传粉系统. 生态学报, 2014, 34(19): 5360-5368.

Huang L

, Jin

, Li

, Zhang X

, Yang

, Wang X

Floral morphology and its relationship with pollination systems in Papilionoideae.

Acta Ecologica Sinica, 2014, 34(19): 5360-5368. (in Chinese)

The formation of the diverse pollination systems in Papilionoideae is greatly dependent on the relationships between species richness and their functional groups of pollinators. The floral morphology in Papilionoideae plants is complex. The flowers of most taxa of Papilionoideae have a full complement of organs: pentamerous alternating whorls of sepals, petals, two stamen whorls and a single carpel. The floret has a median sagittal sepal, a bigger vexillum on the outside, and two wings close to the vexillum that parcel two keels of contention. The ten stamens are usually united into two groups (mainly 9+1, few 5+5), called diadelphous stamens. In this paper, we reviewed floral syndrome, functional groups of pollinators, rewards and pollination system in Papilionoideae. Particularly, floral morphology and chemical compositions and their relationships with evolution of pollination systems and mechanism of pollen release were discussed. Pollination system in Papilionoideae plants is mainly bee pollination, with avian pollination, squirrel pollination and bat pollination also present. The pollination system is usually an evolutionary outcome of selection and adaptation between floral morphology and pollinators. The mechanism of pollen release is one of the most important characters that exemplify such interactions between Papilionoideae plants and their pollinators. There are four pollen release mechanisms in Papilionoideae, explosive mechanism, piston mechanism, valvular mechanism and brush mechanism. The selection and adaptation between floral syndrome and the functional groups of pollinators established the pollination system, in which the coordination and adaptation among the components are essential for successful pollination.

蝶形花亚科（Papilionoideae）植物种类丰富、繁育系统多样，与其传粉者关系密切，主要表现在花部适应与传粉系统的形成。蝶形花形态复杂，为完全花，花萼、花瓣5基数，花萼呈箭头状，相邻花瓣螺旋轮生，旗瓣较大位于最外边，两片翼瓣紧贴旗瓣着生并包裹两片合生的龙骨瓣，龙骨瓣内包裹着雌、雄蕊，雌蕊位于正中央，雄蕊轮生，构成二体雄蕊（多数9+1，少数5+5）。对蝶形花亚科植物的花部特征、传粉功能群、酬物与传粉系统构建进行了回顾，重点论述了蝶形花形态和化学组成与传粉系统的进化关系以及花粉呈现机制。其中，泛化的传粉系统以蜂媒传粉为主，同时存在鸟媒传粉、蝙蝠传粉和松鼠传粉等方式，花部结构和传粉者的相互选择和相互适应推动了传粉系统的演化。花粉释放是体现蝶形花植物与其传粉功能群相互作用的重要方面，二者协同进化形成了以下4种花粉呈现机制：弹花机制、活塞机制、瓣膜机制和毛刷机制。蝶形花的花部特征与传粉功能群的相互作用形成相应的传粉系统，而传粉系统是执行传粉功能的重要集合体，各个构件的相互协调和适应保证了蝶形花亚科植物传粉过程的顺利完成。

... 蝶形花亚科(Papilionideae)植物具有4种式样的花粉展示方式,分别是毛刷机制、活塞机制、弹花机制和瓣膜机制^[1,2] ...

2007

0.0

... 蝶形花亚科(Papilionideae)植物具有4种式样的花粉展示方式,分别是毛刷机制、活塞机制、弹花机制和瓣膜机制^[1,2] ...

1966

0.0

... Leppik^[3]提出弹花机制是蝶形花植物在进化中形成的最经济和最有效的花粉释放机制,Atluril等^[4]认为花粉释放是借助外力作用使花器内累积的能量突然释放出来,在使花柱暴露的同时也把花粉释放出来,因此,花粉的爆炸释放本质上是能量的释放 ...

2004

0.0

2008

0.0

... 另外,有研究发现还存在鸟媒爆炸机制,当鸟落在花上时,由于鸟的重量对花的作用,使花腔内受力,花就会瞬间爆炸,将花粉释放到鸟的前额上^[5] ...

2008

0.0

... 从适应异花授粉的方式来看,有花植物的习性可分为三大类:雌雄异位、雌雄异熟和自交不亲和系统^[6,7] ...

... 苜蓿属植物受到外界压力后龙骨瓣弹开不能复原,花药和柱头弹向并紧贴翼瓣暴露在空气中^[6] ...

2006

0.0

... 从适应异花授粉的方式来看,有花植物的习性可分为三大类:雌雄异位、雌雄异熟和自交不亲和系统^[6,7] ...

2013

0.0

侯真珍, 何爽, 王风雷, 张爱勤. 珍稀克隆植物北极花有性繁殖动态及影响因素. 生态学杂志, 2013, 32(12): 3167-3172.

Hou Z

, He

, Wang F

, Zhang A Q. Dynamics of rare clonal plant Linnaea borealis L. sexual reproduction and related affecting factors. Chinese Journal of Ecology, 2013, 32(12)3167-3172. (in Chinese)

Sexual reproduction is a crucial process of clonal plant life history, and to evaluate the plant sexual reproduction effort is of significance in judging the fate of species. This paper studied the sexual reproduction dynamics of  Linnaea borealis  and related affecting factors from the aspects of clonal architecture, ramets, and population.  L. borealis had a clonal architecture of “guerilla”, with a quite fast clonal expansion and an obvious patchy distribution. At ramet level, small ramets had a higher reproductive effort than larger ramets. There were significant differences in the number ratio of assimilation shoots to flowering shoots and the mean fruitsetting rate between smaller ramets (L L >80 cm, 7.51±1.3, and 27.3%±6.52%, respectively) ( P P L. borealis  patches with the same densities, the patches with more companion species had fewer sexual branch number than the patches with lesser companion species ( P L. borealis  mainly through affecting the number of flowering shoots per unit area.

( 1 College of Life Science and Technology, Xinjiang University, Urumqi 830046, China;  2 Ecological Research Center, Kanas Management Committee, Buergin 836600, Xinjiang, China)

有性繁殖是克隆植物生活史中至关重要的环节，评估植物有性繁殖的程度对判断物种的命运有重要的参考价值。从北极花（ Linnaea borealis L.）克隆构型、分株及居群水平上对其有性繁殖动态及其影响因素进行研究。结果表明：北极花为典型的游击型克隆构型，具有较为快速的克隆扩展能力和明显的斑块状分布。个体水平上，在分株形成的早期，对有性繁殖的投入较大。在较小分株（ L L >80 cm，7.51±1.3，27.3%±6.52%）均呈极显著差异（ P P P

... 而异交是指同一种植物中的不同个体间的传粉、受精,这种交配方式能够促进遗传多样性的形成,是形成繁育系统中传粉多样性的主要动力^[8,9,10] ...

2009

0.0

吕林有, 魏臻武, 赵艳, 耿小丽, 刘高军, 武自念. 苜蓿自交亲和性、授粉方式及后代性状分离的研究. 草业科学, 2009, 26(4): 33-36.

Lyu Y

, Wei Z

, Zhao

, Geng X

, Liu G

, Wu Z

A study of self-compatibility and pollinations and separation of offspring traits of alfalfa.

Pratacultural Science, 2009, 26(4): 33-36. (in Chinese)

Eight alfalfa cultivars from different sources were used to study the characteristics of selfcompatibility and pollination approaches. The results showed that the selfcompatibility of alfalfa among the cultivars was different, and the selfcompatibility indexes of all materials were ranged from 0 to 1.39. In which, the selfcompatibility index of 2 cultivars were over 1.00, 5 cultivars was from 1.00 to 0, and 1 cultivar was 0. The phenomenon of gradual death in batches was found in the S1 growth course, and the final survival rate was only 34.80% and 32.30%. the seed rate of selfcousins is the highest in 4 selfpollinations, so the selfpollination of selfcousins was appropriate to popularize and apply in alfalfa selfpollinate.

对8份不同苜蓿Medicago sativa材料的自交亲和性及授粉方式的研究表明：参试苜蓿材料的自交亲和性存在较大差异，自交亲和指数为0～1.39，其中有2个自交亲和材料(自交亲和指数＞1.00)，5个自交不亲和材料(1.00≥自交亲和指数＞0)和1个极端自交不亲和材料；2个自交亲和材料自交所得S1具有出苗后分批逐渐死亡的自交衰退现象，最终叶片多叶型和枝条高且分枝多型成活率分别仅为34.80%和32.30%；在4种自交授粉方式中兄妹自交结实率最高，适宜在苜蓿自交授粉中推广应用。

... 而异交是指同一种植物中的不同个体间的传粉、受精,这种交配方式能够促进遗传多样性的形成,是形成繁育系统中传粉多样性的主要动力^[8,9,10] ...

2002

0.0

张卫东, 刘公社, 刘杰, 刘凡, 齐冬梅, 李芳芳, 董贵俊, 赵晶晶. 羊草自交不亲和性初步研究. 草地学报, 2002, 10(4): 287-292.

Zhang W

, Liu G

, Liu

, Qi D

, Li F

, Dong G

, Zhao J

A Preliminary study on self-incompatibility of Leymus chinensis.

Acta Agrestia Sinica, 2002, 10(4): 287-292. (in Chinese)

The selfing and outcrossing seeding capability of wildrye ( Leymus chinensis )were investigated under the natural environments. The viability of pollens was estimated by the way of FDA fluorescent staining and the developmental characters of pistals and stamina at time and space were observed. The results showed that the selfing setting percentage was far less than the outcrossing setting percentage in the fields. The mean value of viable pollens was above exist temporal and spatial isolation between stigmas and anthers. In addition, thioredoxsin h gene was considered to be related to self-incompatibility in poaceae plants. Using the primers designed from it, the anticipated fragment was detected in the genome of Leymus chinensis by PCR amplification. So, the authors considered that no or much lower selfing seeding percentage may lie in the self-incompatible mechanism under the genetic control in Leymus chinensis ,and the thioredoxsin h gene may be related to its self-incompatibility. However, this is only an initial conclusion and much evidence is needed to prove this in cytology and molecular biology.

1. Key Laboratory of Photosynthesis and Environmental Molecular Physiology, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China; 2. Beijing Vegetable Research Center, Beijing Academy of Agriculture and Forestry Sciences, Beijing 100089, China

在自然条件下进行羊草自交、异交结实性实验。采用FDA染色法检测羊草小孢子活力,观测雌、雄蕊发育的时空特点。结果表明,在草原上羊草异交结实率远大于其自交结实率。在成熟花药中有生活力的花粉平均达95.2%以上。在发育时间顺序和空间结构上,雌、雄蕊并不妨碍自体授粉。另外,利用禾本科植物自交不亲和性(SI)有关的硫氧还蛋白 h 基因设计的引物在羊草DNA中检测到预期片断。因此,笔者认为,羊草自交不结实或结实率低的主要原因可能在于基因控制下的自交不亲和反应机制,硫氧还蛋白 h 基因可能与羊草的自交不亲和性有关。这仅为初步结论,仍需在细胞学和分子生物学方面取得更多的依据。

... 而异交是指同一种植物中的不同个体间的传粉、受精,这种交配方式能够促进遗传多样性的形成,是形成繁育系统中传粉多样性的主要动力^[8,9,10] ...

2014

0.0

... 有关新疆野生三叶草的研究不多,仅有王玉祥等^[11,12]、潘奥等^[13]对新疆三叶草形态特征的研究,而未见关于新疆野生三叶草开花习性和繁殖特性的报道 ...

2012

0.0

王玉祥, 张博. 新疆野生白三叶表型性状变异研究. 草地学报, 2012, 20(6): 1163-1169.

Wang Y

, Zhang

Study on phenotypic variation of Trifolium repens in Xinjiang.

Acta Agrestia Sinica, 2012, 20(6): 1163-1169. (in Chinese)

Phenotypic variation was revealed by observing and measuring the height, leaf shape, leaf color, flower color, stem length, internode length and 1000-grain weight of thirteen wild Trifolium repens species collected from Xinjiang province. Phenotypic variation among and within populations was analyzed by using variance analysis and correlation analysis. Results showed that there were abundant genetic diversities and significant differences among the morphological characters of different populations. Hierarchical cluster analysis divided these thirteen populations into three groups: Ⅰ. Hutubi-M; Ⅱ. Hutubi-J, Hutubi-K, Hutubi-L, Urumqi Nanshan-D and Changji-F; and Ⅲ. Urumqi Nanshan-A, Urumqi Nanshan-B, Urumqi Nanshan-C, Urumqi Nanshan-E, Nileke-G, Tekesi-H and Nalati-I. These findings provide basic data for further study on biological characteristics and genetic breeding.

Key Laboratory of Grassland Resource and Ecology of Xinjiang, Xinjiang Agricultural University, Urumqi, Xinjiang 830052, China

以新疆13份白三叶( Trifolium repens )为材料,通过观测其株高、叶形、叶色、花色、茎长、茎节间长和千粒重等特征,研究新疆野生白三叶表型性状差异。结果表明:白三叶的各个指标在群体间存在显著差异,具有丰富的遗传多样性。通过聚类分析可将13个群体分为3类,呼图壁M单独聚为1类;呼图壁J、呼图壁K、呼图壁L、乌市南山D和昌吉F聚为1类;其余7个聚为1类。本研究结果能够为开展野生白三叶的生物学特性和遗传育种研究提供基础数据。

2013

0.0

潘奥, 王玉祥, 李陈建, 张博. 新疆白三叶的形态特征及坪用性. 草业科学, 2013, 30(12): 1960-1967.

Pan

, Wang Y

, Li C

, Zhang

Research on morphological characteristics and lawn-using characters of Trifolium repens from Xinjiang.

Pratacultural Science, 2013, 30(12): 1960-1967. (in Chinese)

There are abundant resources of Trifolium repens which have stronger stress tolerance in Xinjiang Province. The present study analyzed various morphological and turf indices including branch number, leaf characteristics and flower number of T.repens to evaluate their lawn characters. The results indicate that T.repens have lawn characters such as strong stress tolerance, early blooming, bright-colored flowers, persistence of florescence and green period and ornamentals. The germplasm of T.repens from HuTuBi performed as a better ornamental grass with turning green early, lower grass layer (17.1~21.3 cm), longer flowering period (150~154 d) and longer green period (221~225 d). The germplasm of T.repens from Heaven Pool(E) and ChangJi(F) had higher grass layer (24.3 and 26.1 cm, respectively) which are suitable for forage. And the germplasm of T. repens from TeKeSi (H) performed as abetter nectar plant with great intensity leaf and flowers (0.335and 0.216 cm 2 , respectively).

Key Laboratory of Grassland Resource and Ecology of Xinjiang, Xinjiang Agricultural University, Urumqi 830052, China

新疆野生白三叶( Trifolium repens )资源丰富且抗逆性强,试验通过分析白三叶的分枝数、叶片特征、花序数等形态指标及坪用性指标,对白三叶的利用价值进行分析研究。结果表明,供试的材料具有抗性强,开花早,花色鲜艳,花期、绿期持续时间长和观赏性强等特点。其中,呼图壁的材料均具有返青早、草层低矮(17.1～21.3 cm)、花期(150～154 d)和绿期(221～225 d)较长等优点,适宜作为观赏性草坪草;天池的材料E(24.30 cm)和昌吉的材料F(26.10 cm)草层较高,适宜作为饲用牧草;特克斯的材料H叶量大(0.335个·cm -2 )且花(0.216个·cm -2 )多,适宜作为蜜源植物。

2015

0.0

王玉祥, 李陈建, 陈述明, 朱进忠. 施氮肥时间对苜蓿结荚率及种子产量的影响. 草业科学, 2015, 32(2): 231-235.

Wang Y

, Li C

, Chen S

, Zhu J

Effects of nitrogen application time on pod retention rate and seed yield of alfalfa.

Pratacultural Science, 2015, 32(2): 231-235. (in Chinese)

In order to improve pod retention rate and seed yield of alfalfa, effects of nitrogen fertilizer application time on pod retention rate, 1 000seed weight, seed yield of Medicago sativa cv. Xinmu No.4 were studied. The results showed that the pod retention rate, the 1 000seed weight and seed germination rate with the treatment of fertilized at returning green stage were higher than that in the other treatments which was 63.87%, 2.01 g and 85.33%, respectively. However, the pods number per inflorescence and pod retention rate of this treatment were lower than that in the other treatments. The seed yield was highest if fertilized at the budding stage which was 802.16 kg·ha-1 and 2.05 times of control and the seed yield was higher if fertilized at branching stage which was 641.29 kg·ha-1 and 1.64 times of control. These results provided a theoretical basis for rational fertilization of alfalfa seed field.

Key Laboratory of Grassland Resource and Ecology of Xinjiang, Xinjiang Agricultural University, Urumqi 830052, China

试验以新牧4号紫花苜蓿（Medicago sativa cv. Xinmu No.4）为材料，研究在苜蓿不同生育期施氮肥对苜蓿结荚率、千粒重、种子产量等的影响，为提高苜蓿结荚率、增加种子产量提供依据。结果表明，返青期施氮肥处理的苜蓿荚果结荚率、种子千粒重以及种子发芽率均高于其他处理，依次为63.87%、2.01 g和85.33%；但其单序荚果数和结荚率均低于其他处理；在各个处理中，现蕾期施氮肥处理苜蓿种子产量最高(802.16 kg·hm-2)，分枝期次之（641.29 kg·hm-2），分别是对照（390.34 kg·hm-2）的2.05倍和1.64倍。本研究结果能够为苜蓿种子田的合理施肥提供理论依据。

... 5^[14] ...

2012

0.0

杨旭, 杨志玲, 王洁, 檀国印, 何正松. 松濒危植物凹叶厚朴的花部综合特征和繁育系统. 生态学杂志, 2012, 31(3): 551-556.

Yang

, Yang Z

, Wang

, Tan G

, He Z

Floral syndrome and breeding system of endangered species Magnolia officinalis subsp. biloba.

Chinese Journal of Ecology, 2012, 31(3): 551-556. (in Chinese)

... 3 讨论与结论植物不同的开花习性对植物的繁衍、基因的延续有着重要的影响^[15,16] ...

2004

0.0

... 3 讨论与结论植物不同的开花习性对植物的繁衍、基因的延续有着重要的影响^[15,16] ...

2007

0.0

... 授粉是种子植物有性生殖的一个重要环节,人工去雄时要确保植物花粉未散粉,即植物未受精^[17,18] ...

2011

0.0

孙鹏, 李云, 姜金仲, 袁存权, 张立玲. 植物同源四倍体花器官及种子变异研究进展. 植物研究, 2011, 31(2): 249-256.

Sun

, Li

, Jiang J

, Yuan C

, Zhang L

Progress of variations in floral organs and seeds of autotetraploid plants.

Bulletin of Botanical Research, 2011, 31(2): 249-256. (in Chinese)

The phenotypes of floral organs, female and male gametophytes and seeds vary in various degrees because of the genome reduplication for autotetraploid plants. Compared with original diploid plants, the variations focus on the following aspects. The form and inserted site of inflorescences are diverse. The floral organs generally increase in size. The transmitting tissue in style is not well developed. The content of nucleic acid increases whereas the numerical value of RNA/DNA decreases in buds. Parts of egg apparatus, centrocytes and antipodal cells in sac degenerate or grow abnormally of morphology and location, which result in embryo sac abortion. Male gametophytes (pollen) are bigger than original diploid in size, diversiform and infertile. Seeds of autotetraploid plants have an increasing trend of size, weight, content of protein and tolerability to salt, but few of the seeds can germinate. The abnormal development of endosperm negatively affects the supply of nutrition to embryo, so that a lot of embryos degenerate or develop abnormally.

1.College of Biological Sciences and Biotechnology,Beijing Forestry University,National Engineering Laboratory for Tree Breeding,Key Laboratory of Genetics and Breeding in Forest Trees and Ornamental Plants,Ministry of Education,The Tree and Ornamental Plant Breeding and Biotechnology Laboratory of State Forestry Administration,Beijing　100083；2.Guizhou Normal College,Guiyang　550018；3.Chengde Prefecture Forestry Administration,Chengde　067400

由于染色体数目的加倍，植物同源四倍体花器官、雌配子体、雄配子体及种子的表现型均发生了不同程度的变异，与其起源二倍体相比，四倍体的表现型变异主要体现在以下几个方面：花序形态及着生位置呈现多样化；花器官普遍增大，花柱中引导组织欠发达；花芽中核酸含量增加，RNA/DNA降低；雌配子体（胚囊）中卵器、中央细胞、反足细胞形态位置异常或退化，胚囊育性下降；雄配子体（花粉）体积增大、形态多样，且育性下降；种子体积、重量多数呈增加趋势，蛋白含量提高，对NaCl逆境耐受性增强，萌发力下降；种子胚乳发育的异常有不利于种子胚营养供给的趋势，这种趋势会导致胚畸形或退化。

... 授粉是种子植物有性生殖的一个重要环节,人工去雄时要确保植物花粉未散粉,即植物未受精^[17,18] ...

2014

0.0

郭媛, 邵有全, 郭宝贝, 张旭凤. 梨花粉和柱头发育与温度关系研究. 中国生态农业学报, 2014, 22(12): 1446-1452.

Guo

, Shao Y

, Guo B

, Zhang X

Relationship of pear pollen and stigma development with temperature. Chinese Journal of

Eco-Agriculture, 2014, 22(12): 1446-1452. (in Chinese)

The effect of temperature on pear flower development is a prerequisite for using bees in pear pollination. In this study, the main varieties of Pyrus bretschneideri cv. 'Dangshansuli' and P. bretschneideri cv. 'Yali' served as pollination test materials. Three temperatures (8 ℃, 12 ℃ and 20 ℃) were set to explore the dynamic blossoming and stigma receptivity of 'Dangshansuli', and pollen release, pollen germination, pollen tube growth of 'Yali', and other indicators to lay the theoretical basis for the reasonable use of bee pollination of pears. The results showed that at 8 ℃, the required time for flower opening of 'Dangshansuli' exceeded 144 h. Also the time for stigma receptivity delayed to 112 h and that for pollination persistence shortened. The time for pollen release of 'Yali' extended to 230 h. Pollen germination rate at 8 ℃ was less than that at 20 ℃ ( P P < 0.05). After culturing for 96 h, the number of pollen tubes arriving at the top, middle and base of styles of 'Dangshansuli' was respectively 111, 105 and 92. When temperature was 20 ℃, the required time for the blossoming of 'Dangshansuli' was 44 h. Stigma receptivity was strong and pollination persistence time reached 120 h. The required time for total pollen release of 'Yali' was 64 h and pollen germination rate was high. After culturing for 96 h, the number pollen tubes arriving at the top, middle and base of styles of 'Dangshansuli' was respectively 121, 120 and 119. The temperature of 20 ℃ was more suitable for pear growth and pollination by bees than 12 ℃ and 8 ℃. The present study laid the scientific basis for future studies on biological characteristics of pear blossoming at different temperatures. It also laid the foundation for practical application of honeybee pollination technology in pear farms.

考察环境温度对梨花发育的影响是引进蜜蜂为梨树授粉的前提。本研究以主栽品种'砀山酥梨'及其授粉品种'鸭梨'为试验材料, 设置了8 ℃、12 ℃、20 ℃ 3种温度条件, 考察'砀山酥梨'开花动态、柱头可授性及'鸭梨'散粉、花粉萌发、花粉管生长等指标, 旨在为梨树合理利用蜜蜂授粉提供理论依据。结果表明: 在8 ℃下, '砀山酥梨'花朵全部开放所需时间达到144 h以上, 柱头具有较强可授性出现时间为112 h, 且可授期持续时间短, '鸭梨'全部散粉时间为230 h, 花粉萌发率显著低于20 ℃( P P <0.05), 培养96 h到达花柱上部、中部、基部的花粉管数量分别为111根、105根、92根; 20 ℃时, '砀山酥梨'开花所需时间为44 h, 柱头可授性强, 且可授期持续时间可达120 h, '鸭梨'全部散粉所需时间为64 h, 花粉萌发速度快, 萌发率高, 培养96 h到达花柱上部、中部、基部的花粉管数量分别为121根、120根、119根。20 ℃较12 ℃和8 ℃更适合梨花发育和蜜蜂采集。本试验结果为不同温度下梨花的开花生物学特性研究提供了科学依据, 也为梨树蜜蜂授粉技术的应用奠定了基础。

... 自交不亲和性作为一种抵制近亲繁殖、保持种群遗传多样性的重要机制普遍存在于植物(特别是被子植物)中^{[19,20,21,22]} ...

2015

0.0

... 自交不亲和性作为一种抵制近亲繁殖、保持种群遗传多样性的重要机制普遍存在于植物(特别是被子植物)中^{[19,20,21,22]} ...

2014

0.0

李晶, 金樑, 邓志刚, 杨龙, 王文斌, 王晓娟. 被子植物花粉和雌蕊互作的分子基础. 草业科学, 2014, 31(1): 161-167.

, Jin

, Deng Z

, Yang

, Wang W

, Wang X

Pollen-pistil interaction and its molecular basis angiosperms.

Pratacultural Science, 2014, 31(1): 161-167. (in Chinese)

In angiosperms, a serious of recognition events are involved in pollen-pistil interactions since pollen grains landed on the stigma. Such interaction processes include recognition between pollen and stigma, hydration and germination of pollen grains, pollen tube growth and guidance, sperm release and double fertilization in the embryo sac. In recent years, the genes involved in process of pollen-pistil interactions, pollen tube growth and guidance have been discovered and cloned. In this paper, the advances of adhesion of pollen grains and their hydration, germination and growth have been reviewed. In particular, the machanisms of chemical compounds, such as glycoproteins interactions in pollen adhesion, lipid regulation pathway in pollen hydration and arabinogalactan proteins in polar growth of pollen tube, have been discussed to provide theoretical basis for the reproductive biology in angiosperms.

State Key Laboratory of Grassland Agro-ecosystems, College of Pastoral Agriculture Science and Technology, Lanzhou University, Lanzhou 730020, China

被子植物的花粉自落到柱头上开始, 花粉和雌蕊组织即发生一系列的相互识别作用, 这个过程包括花粉与柱头互作、花粉管极性生长和引导、精细胞在胚囊中释放和完成双受精。近年来, 已相继发现并且克隆了花粉与柱头互作以及花粉管生长和引导相关的基因。本文对花粉粘附、水合、萌发以及花粉管生长和引导的分子机制进行了综述, 重点讨论了花粉粘附的糖蛋白互作、花粉水合的脂类调控途径以及花粉管极性生长的阿拉伯半乳聚糖蛋白等小分子引导机制, 以期为被子植物的繁殖生物学研究提供理论依据。

... 自交不亲和性作为一种抵制近亲繁殖、保持种群遗传多样性的重要机制普遍存在于植物(特别是被子植物)中^{[19,20,21,22]} ...

2012

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... 自交不亲和性作为一种抵制近亲繁殖、保持种群遗传多样性的重要机制普遍存在于植物(特别是被子植物)中^{[19,20,21,22]} ...

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... 自交不亲和性植物都具有强度不同的不亲和性,其花期自交坐果率或结实率一般介于0~5%^[23,24,25] ...

0.0

... 自交不亲和性植物都具有强度不同的不亲和性,其花期自交坐果率或结实率一般介于0~5%^[23,24,25] ...

2012

0.0

... 自交不亲和性植物都具有强度不同的不亲和性,其花期自交坐果率或结实率一般介于0~5%^[23,24,25] ...