植物生态学报 ›› 2009, Vol. 33 ›› Issue (1): 222-236.DOI: 10.3773/j.issn.1005-264x.2009.01.024
• 综述 • 上一篇
收稿日期:
2008-05-21
接受日期:
2008-07-08
出版日期:
2009-05-21
发布日期:
2009-01-30
作者简介:
E-mail: renmingxun@yahoo.com.cn
基金资助:
Received:
2008-05-21
Accepted:
2008-07-08
Online:
2009-05-21
Published:
2009-01-30
摘要:
对花内雄蕊存在显著分化的现象进行了分析与归纳, 总结了花内雄蕊分化的各种主要形式及其繁殖适应意义。“花内雄蕊分化”是指花内雄蕊与雄蕊之间存在显著分化的现象, 这一概念可以把二强雄蕊、四强雄蕊和异型雄蕊等以往单独进行研究的相关性状结合起来, 并明确区分了几种新的花内雄蕊分化形式, 以期更准确全面地认识这些相关性状的适应意义与进化。该文将花内雄蕊分化区别为花丝的分化、花药的分化、雄蕊合生的分化、雄蕊运动的分化、退化雄蕊5大类。花丝的分化主要是花丝长度的分化, 如四强雄蕊、二强雄蕊和单强雄蕊; 花药分化主要指花药颜色、花药与花粉粒大小和花药开裂时间等的分化; 雄蕊合生的分化主要体现为花内部分雄蕊合生而部分雄蕊离生; 雄蕊运动的分化指的是花内雄蕊在运动时间或方式上存在差异, 造成雄蕊处于不同的成熟阶段和位于不同的空间位置; 退化雄蕊则是花内部分雄蕊失去了生产花粉的繁殖功能, 通常也发生了花药形态上的巨大改变。异型雄蕊不仅存在花丝和花药的形态分化, 还存在着明显的功能分化, 是分化程度很高的一类特殊的花内雄蕊分化形式。一些特殊的繁育系统, 如异长花柱和镜像花柱等在种内不同个体上存在着不同形式的花内雄蕊分化。花内雄蕊分化在花内造成了多个不同的花药位置, 在很大程度上影响了雌雄异位程度, 对植物自交与异交水平、花内雌雄功能干扰等有着潜在作用; 花内雄蕊分化形成的多个不同空间位置的雄蕊还增加了对多种访花者的吸引与适应潜力, 有可能影响到访花者的类型与访花行为, 得以适应多种传粉者。此外, 花内雄蕊分化可将花粉逐渐分批次分发给访花者, 提高花粉散布效率, 可看成是“花粉呈现理论”所指的花粉装配与分发机制之一。现有的实验研究发现, 花内雄蕊分化能够吸引传粉者、保护正常花药和花粉、促进花粉散发(降低花粉竞争)、实行延迟自交和降低花内雌雄功能干扰等。花内雄蕊分化还缺少系统研究, 有些雄蕊分化形式如单强雄蕊和雄蕊运动的分化还没有针对性的实验揭示其适应意义, 鸭跖草科和某些豆科植物的雄蕊三型分化等现象也缺少进化适应意义的研究。花内雄蕊分化对植物雌性和雄性适合度可能不同的影响、如何与访花者相互作用、如何与其它花部特征一起影响植物繁殖过程等, 可能是这一领域值得今后优先研究的课题。
任明迅. 花内雄蕊分化及其适应意义. 植物生态学报, 2009, 33(1): 222-236. DOI: 10.3773/j.issn.1005-264x.2009.01.024
REN Ming-Xun. INTRAFLORAL STAMEN DIFFERENTIATIONS AND THEIR ADAPTIVE SIGNIFICANCES. Chinese Journal of Plant Ecology, 2009, 33(1): 222-236. DOI: 10.3773/j.issn.1005-264x.2009.01.024
分化形式 Types of stamen differentiations | 主要的系统分布 Taxonomic distributions | 适应意义 Adaptive significances | 主要文献 Main references | ||
---|---|---|---|---|---|
1 花丝长度的分化 Differentiations in filament length | |||||
1.1 单强雄蕊 Monodynamous | 金虎尾科 Malpighiaceae 风筝果属 Hiptage | 未知 Unknown | |||
1.2 二强雄蕊 Didynamous | 唇形科 Labiatae 玄参科 Scrophulariaceae 紫葳科 Bignoniaceae 苦苣苔科 Gesneriaceae 爵床科 Acanthaceae | 不同长度雄蕊的散粉时间不同, 提高了花粉输出效率, 且常与雌雄异熟一起限制了花内自交 Differentiated stamens disperse pollen in different time, and often accompany with dichogamy to prevent selfing | Foerste, 1888 | ||
1.3 四强雄蕊 Tetradynamous | 十字花科 Cruciferae | 长雄蕊接触传粉者散粉, 短雄蕊增加访花者访花时间和通过自交实现繁殖保障 Longer stamens disperse pollen, shorter stamens attract pollinators and promote delayed autogamy | |||
1.4 多强雄蕊 Polydynamous | 杜鹃科 Ericaceae 茄科 Solanaceae 野牡丹科 Melastomataceae 豆科云实属 Caesalpinia | 长雄蕊接触传粉者散粉, 短雄蕊吸引传粉者和延迟自交 Long stamens disperse pollen, short stamens attract pollinators and for delayed autogamy | et al., 2001 | ||
1.5 三型花柱 Tristyly | 雨久花科 Pontederiaceae 酢浆草科 Oxalidaceae 千屈菜科 Lythraceae | 不同长度花药接触传粉者不同部位, 给其它花型对应长度柱头授粉; 且能吸引不同类型的传粉者提高花粉输出 Differentiated stamens contact the different parts of pollinator’s body and pollinate reciprocal stigma between floral morphs; Differentiated stamens can also enhance pollen output via contacting different pollinators | |||
1.6 异型雄蕊 Heteranthery | 血皮草科 Haemodoraceae 异蕊草科 Tecophilaeaceae 雨久花科 Pontederiaceae 茄科 Solanaceae 大多数镜像花柱植物 Most enantiostylous plants | 短雄蕊专门吸引传粉者和/或给传粉者提供可食花粉, 长雄蕊专门传粉 Long stamens disperse pollen, short stamens attract pollinators and provide edible pollen for pollinators | |||
2 花药的分化 Differentiations in anthers | |||||
2.1 花药大小与颜色分化 Differentiation in anther color and size | 雨久花科 Pontederiaceae 鸭跖草科 Commelinaceae 异蕊草科 Tecophilaeaceae 豆科 Fabaceae 大部分异型雄蕊植物 Heterantherous plants | 大而鲜艳的花药吸引传粉者, 小而暗淡的花药接触传粉者散粉 Showy anther attract pollinators and the normal anther directly contact the pollinators’ body to disperse pollen more effectively | |||
2.2 花药散粉时间分化 Differentiation in dehiscing time of anthers | 豆科云实属 Caesalpinia 大部分二强雄蕊植物 Most Didynamous plants | 延长对访花者吸引时间, 通过不同传粉者的多次传粉提高异交花粉输出率 Prolonging the attractiveness of flower, and enhancing pollen dispersal via many visits by different pollinators | |||
3 雄蕊合生的分化 Differentiation in stamen fusion | |||||
3.1 部分雄蕊花丝合生 Some fused with filaments | 豆科 Fabaceae 山茶科 Theaceae 大戟科 Euphorbiaceae | 未知 Unknown | |||
3.2 部分雄蕊花药合生 Some fused with anthers | 苦苣苔科 Gesneriaceae 爵床科 Acanthaceae | 未知 Unknown | |||
4 雄蕊运动的分化 Differentiation in stamens movement | |||||
芸香科 Rutaceae 景天科 Crassulaceae 唇形科 Labiatae 玄参科 Scrophulariaceae | 未知 Unknown | ||||
5 退化雄蕊或变态雄蕊 Staminodes | |||||
鸭跖草科 Commelinaceae 兰科 Orchidaceae 姜科 Zingiberaceae | 吸引传粉者和/或提供可食花粉, 保护蜜腺和子房 Attracting pollinators and providing nutrient pollen for pollinators, protecting nectar and ovule |
表1 花内雄蕊分化的常见形式、系统分布及其适应意义
Table 1 Typical types of intrafloral stamen differentiations and their systematic distributions and possible adaptive significances
分化形式 Types of stamen differentiations | 主要的系统分布 Taxonomic distributions | 适应意义 Adaptive significances | 主要文献 Main references | ||
---|---|---|---|---|---|
1 花丝长度的分化 Differentiations in filament length | |||||
1.1 单强雄蕊 Monodynamous | 金虎尾科 Malpighiaceae 风筝果属 Hiptage | 未知 Unknown | |||
1.2 二强雄蕊 Didynamous | 唇形科 Labiatae 玄参科 Scrophulariaceae 紫葳科 Bignoniaceae 苦苣苔科 Gesneriaceae 爵床科 Acanthaceae | 不同长度雄蕊的散粉时间不同, 提高了花粉输出效率, 且常与雌雄异熟一起限制了花内自交 Differentiated stamens disperse pollen in different time, and often accompany with dichogamy to prevent selfing | Foerste, 1888 | ||
1.3 四强雄蕊 Tetradynamous | 十字花科 Cruciferae | 长雄蕊接触传粉者散粉, 短雄蕊增加访花者访花时间和通过自交实现繁殖保障 Longer stamens disperse pollen, shorter stamens attract pollinators and promote delayed autogamy | |||
1.4 多强雄蕊 Polydynamous | 杜鹃科 Ericaceae 茄科 Solanaceae 野牡丹科 Melastomataceae 豆科云实属 Caesalpinia | 长雄蕊接触传粉者散粉, 短雄蕊吸引传粉者和延迟自交 Long stamens disperse pollen, short stamens attract pollinators and for delayed autogamy | et al., 2001 | ||
1.5 三型花柱 Tristyly | 雨久花科 Pontederiaceae 酢浆草科 Oxalidaceae 千屈菜科 Lythraceae | 不同长度花药接触传粉者不同部位, 给其它花型对应长度柱头授粉; 且能吸引不同类型的传粉者提高花粉输出 Differentiated stamens contact the different parts of pollinator’s body and pollinate reciprocal stigma between floral morphs; Differentiated stamens can also enhance pollen output via contacting different pollinators | |||
1.6 异型雄蕊 Heteranthery | 血皮草科 Haemodoraceae 异蕊草科 Tecophilaeaceae 雨久花科 Pontederiaceae 茄科 Solanaceae 大多数镜像花柱植物 Most enantiostylous plants | 短雄蕊专门吸引传粉者和/或给传粉者提供可食花粉, 长雄蕊专门传粉 Long stamens disperse pollen, short stamens attract pollinators and provide edible pollen for pollinators | |||
2 花药的分化 Differentiations in anthers | |||||
2.1 花药大小与颜色分化 Differentiation in anther color and size | 雨久花科 Pontederiaceae 鸭跖草科 Commelinaceae 异蕊草科 Tecophilaeaceae 豆科 Fabaceae 大部分异型雄蕊植物 Heterantherous plants | 大而鲜艳的花药吸引传粉者, 小而暗淡的花药接触传粉者散粉 Showy anther attract pollinators and the normal anther directly contact the pollinators’ body to disperse pollen more effectively | |||
2.2 花药散粉时间分化 Differentiation in dehiscing time of anthers | 豆科云实属 Caesalpinia 大部分二强雄蕊植物 Most Didynamous plants | 延长对访花者吸引时间, 通过不同传粉者的多次传粉提高异交花粉输出率 Prolonging the attractiveness of flower, and enhancing pollen dispersal via many visits by different pollinators | |||
3 雄蕊合生的分化 Differentiation in stamen fusion | |||||
3.1 部分雄蕊花丝合生 Some fused with filaments | 豆科 Fabaceae 山茶科 Theaceae 大戟科 Euphorbiaceae | 未知 Unknown | |||
3.2 部分雄蕊花药合生 Some fused with anthers | 苦苣苔科 Gesneriaceae 爵床科 Acanthaceae | 未知 Unknown | |||
4 雄蕊运动的分化 Differentiation in stamens movement | |||||
芸香科 Rutaceae 景天科 Crassulaceae 唇形科 Labiatae 玄参科 Scrophulariaceae | 未知 Unknown | ||||
5 退化雄蕊或变态雄蕊 Staminodes | |||||
鸭跖草科 Commelinaceae 兰科 Orchidaceae 姜科 Zingiberaceae | 吸引传粉者和/或提供可食花粉, 保护蜜腺和子房 Attracting pollinators and providing nutrient pollen for pollinators, protecting nectar and ovule |
图1 花内雄蕊分化的6种典型类型 A: 单强雄蕊集中出现于金虎尾科的风筝果属, 其大部分雄蕊都短小, 只有一个雄蕊花丝明显伸长, 使得花药与柱头位置相似 One stamen has a very long filament as compared with most short stamens but shows no difference in anther colors, which can be seen as monodynamous stamens. Monodynamous stamens are mainly found in Hiptage in Malpighiaceae B: 二强雄蕊(唇形科、玄参科等) Didynamous stamens in Labiatae, Scrophulariaceae, etc. C: 十字花科中的四强雄蕊Tetradynamous stamens in Cruciferae D: 雄蕊合生方式和程度的分化 Differentiations in growth pattern of stamens such as some stamens grew with fused filaments while others remain discrete E: 异型雄蕊主要发生在镜像花柱中, 不仅花丝长度、花药大小与颜色、花粉活性等都存在着明显的分化, 还发生了直接参与繁殖、吸引传粉者的功能分化 Heteranthery is mainly found in enantiostyly, in which one stamen has a very long filament and different anther color as compared with all the other short but usually colorful anthers F: 三型花柱中存在3种不同花型, 每种花型内的雄蕊群都分化成在花丝长度、花粉粒大小和亲合性等方面差异显著的两类雄蕊 Three floral morphs in tristyly are found to be reciprocal in positions of anthers and stigmas, each morph has two types of anthers differing in filament length, pollen size and compatibility
Fig. 1 Six typical types of intrafloral stamen differentiations
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