植物生态学报 ›› 2010, Vol. 34 ›› Issue (7): 867-875.DOI: 10.3773/j.issn.1005-264x.2010.07.012
所属专题: 生物多样性
收稿日期:
2009-12-11
接受日期:
2010-02-26
出版日期:
2010-12-11
发布日期:
2010-07-01
通讯作者:
任明迅
作者简介:
* E-mail: renmingxun@yahoo.com.cnReceived:
2009-12-11
Accepted:
2010-02-26
Online:
2010-12-11
Published:
2010-07-01
Contact:
REN Ming-Xun
摘要:
雄蕊运动指雄蕊在自身能量支持下发生的主动运动, 不包括雄蕊在访花者触碰下造成的被动位移。该文总结了雄蕊的应激运动、快速猛烈弹射、缓慢运动以及级联运动等4种主要类型, 分析了这些运动类型的系统分布及繁殖适应意义等方面的研究进展。雄蕊的应激运动由访花者或其他外力诱发, 可能起到促进散粉和实现繁殖保障的作用; 雄蕊快速猛烈的弹射运动可将花粉猛然撒向空中或访花者身上, 促进了花粉的风媒或虫媒散布; 缓慢运动的雄蕊可能通过在不同花期改变雄蕊的空间位置和雌雄异位程度来调节繁殖策略, 或主动将雄蕊花药移至特定部位(如柱头表面)实现自交; 雄蕊逐一、依次发生的级联运动较为复杂, 主要分布在刺莲花科、梅花草科、旱金莲科和芸香科中, 目前还缺乏实验研究; 但根据“花粉呈现理论”以及其他类型的雄蕊运动研究结果, 雄蕊的级联运动可以将花粉分批呈现给不同的传粉者, 通过不同传粉者的分别传粉来提高花粉的输出; 而且可避免已散粉雄蕊对即将散粉雄蕊的干扰, 同时可能也降低了雌雄功能干扰和(或)花内自交。在芸香(Ruta graveolens)中, 级联运动之后的雄蕊还会在花末期再同时向花中央运动; 这种多向、多次运动方式是目前发现的最复杂的雄蕊运动类型。雄蕊运动领域值得今后开展进一步实验研究的方向主要有: 1)雄蕊运动尤其是级联运动对雌雄功能干扰(性别间干扰)、雄蕊与雄蕊的“性别内干扰”等植物繁殖格局的影响; 2)雄蕊运动与雌雄异熟、雌雄异位等花部特征的相互作用; 3)雄蕊运动复杂类型的生理与发育机制。
任明迅. 两性花的雄蕊运动: 多样性和适应意义. 植物生态学报, 2010, 34(7): 867-875. DOI: 10.3773/j.issn.1005-264x.2010.07.012
REN Ming-Xun. Stamen movements in hermaphroditic flowers: diversity and adaptive significance. Chinese Journal of Plant Ecology, 2010, 34(7): 867-875. DOI: 10.3773/j.issn.1005-264x.2010.07.012
图1 加拿大山茱萸(Cornus canadensis)雄蕊的快速猛烈弹射现象(改自Edwards et al. , 2005)。 箭头所示为花药顶端速率最大的时候(花药在此时才会释放花粉)。A, 雄蕊弹射过程中花丝顶端与花药顶端的运动轨迹(三角形: 花丝顶端; 圆点: 花药顶端), 横轴与纵轴分别表示花水平与垂直方向上的位移。B, 花药顶端(上图)与花丝顶端(下图)的运动速率;x、y分别表示水平方向与垂直方向上的速率。
Fig. 1 Stamen quick and explosive movement in Cornus canadensis (revised from Edwards et al., 2005). The arrows indicate the maximum speed of anther tip (the time pollen grains released from anthers). A, The track of filament tip and anther tip during stamen explosive movement (triangle for filament tip; dot for anther tip). B, Speed of anther tip (above) and filament tip (bottom), and x and y show the horizontal and vertical direction, respectively.
图2 芸香(A)和梅花草(B)的雄蕊级联运动。 A1, 芸香花初开。A2, 第1个雄蕊运动至花中央散粉。A3, 第1个雄蕊运动到花外围, 第2个雄蕊运动到花中央。A4, 所有雄蕊都发生了级联运动并位于花外围。A5, 花末期所有雄蕊再次向花中央运动。B1, 梅花草花初开(所有雄蕊花药位于花中央)。B2, 第1个雄蕊在散粉完毕之后运动至花外围。B3, 前一个雄蕊散粉完毕而运动到花外围之后, 下一个雄蕊再次进行散粉、运动的过程。B4, 所有雄蕊已运动至花外围(花药均已掉落)。
Fig. 2 Stamen cascade movement in Ruta graveolens (A) and Parnassia palustris (B). A1, Flower just opened. A2, One stamen moved to the center and dehisced. A3, The dehisced stamen moved to outside and the next one moved to the center. A4, All the stamens moved to the outside after their cascade movements. A5, The stamens moved together to the center at the final phase of blooming. B1, Flower just opened (all the anthers are positioned at the center of flower). B2, One stamen moved to outside after dehiscing at the center. B3, One stamen dehisced and moved to outside after the former one finished moving. B4, All the stamens moved to outside and the anthers fell off.
科 Family | 物种 Species | 雄蕊位置 Position of stamens | 雄蕊形态 Stamen morph | 雌雄异位 Herkogamy | 雌雄异熟Dichogamy | 自交亲和性 Compatibility | 参考文献 Reference |
---|---|---|---|---|---|---|---|
梅花草科Parnassiaceae | 所有种 All species | 花中央-花外侧 Center-outside | 运动前的雄蕊花丝先伸长 Filament will extend before stamen moving | , | 雄先熟 Protandrous | 自交亲合 Self-compatible | Ming-Xun Ren, personal observation |
刺莲花科Loasaceae | 大部分物种, 如刺莲花属和 Nasa属等 Most species of Loasa, Nasa, etc. | 花外侧-花中央- 花外侧 Outside-center- outside | 雄蕊等大同型 All the stamens with same shape and size | 明显 Obvious | 雄先熟 Protandrous | 自交亲合 Self-compatible | |
旱金莲科Tropaeolaceae | 旱金莲 Tropaeolum majus | 花外侧-花中央- 花外侧 Outside-center- outside | 雄蕊等大同型 All the stamens with same shape and size | 明显 Obvious | 雄先熟 Protandrous | 自交亲合 Self-compatible | |
芸香科 Rutaceae | 芸香 Ruta graveolens | 花外侧-花中央- 花外侧 Outside-center- outside | 雄蕊等大同型 All the stamens with same shape and size | 明显 Obvious | 雄先熟 Protandrous | 自交亲合 Self-compatible | Ming-Xun Ren, personal observation |
表1 雄蕊级联运动的主要花部特征
Table 1 Main floral characteristics associated with stamen cascade movement
科 Family | 物种 Species | 雄蕊位置 Position of stamens | 雄蕊形态 Stamen morph | 雌雄异位 Herkogamy | 雌雄异熟Dichogamy | 自交亲和性 Compatibility | 参考文献 Reference |
---|---|---|---|---|---|---|---|
梅花草科Parnassiaceae | 所有种 All species | 花中央-花外侧 Center-outside | 运动前的雄蕊花丝先伸长 Filament will extend before stamen moving | , | 雄先熟 Protandrous | 自交亲合 Self-compatible | Ming-Xun Ren, personal observation |
刺莲花科Loasaceae | 大部分物种, 如刺莲花属和 Nasa属等 Most species of Loasa, Nasa, etc. | 花外侧-花中央- 花外侧 Outside-center- outside | 雄蕊等大同型 All the stamens with same shape and size | 明显 Obvious | 雄先熟 Protandrous | 自交亲合 Self-compatible | |
旱金莲科Tropaeolaceae | 旱金莲 Tropaeolum majus | 花外侧-花中央- 花外侧 Outside-center- outside | 雄蕊等大同型 All the stamens with same shape and size | 明显 Obvious | 雄先熟 Protandrous | 自交亲合 Self-compatible | |
芸香科 Rutaceae | 芸香 Ruta graveolens | 花外侧-花中央- 花外侧 Outside-center- outside | 雄蕊等大同型 All the stamens with same shape and size | 明显 Obvious | 雄先熟 Protandrous | 自交亲合 Self-compatible | Ming-Xun Ren, personal observation |
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