植物生态学报 ›› 2023, Vol. 47 ›› Issue (12): 1611-1628.DOI: 10.17521/cjpe.2022.0336
• 综述 • 下一篇
收稿日期:
2022-08-18
接受日期:
2023-02-20
出版日期:
2023-12-20
发布日期:
2023-03-01
通讯作者:
*(liuhuiliang@ms.xjb.ac.cn)
基金资助:
Dilixiadanmu TASHENMAIMAITI1,2, LIU Hui-Liang1,3,4,*()
Received:
2022-08-18
Accepted:
2023-02-20
Online:
2023-12-20
Published:
2023-03-01
Contact:
*(liuhuiliang@ms.xjb.ac.cn)
Supported by:
摘要:
荒漠一年生植物在严峻的环境条件下形成其特殊的萌发时间格局, 即种子异时萌发: 在特定条件下, 同胞种子群形态一致的部分种子可以即时萌发, 而另一部分仍保留在种子库中的现象。目前对荒漠一年生植物的异时萌发研究多集中在种子萌发特性、萌发后性状及繁殖分配等方面。该文系统总结了国内外种子异时萌发性的研究工作, 主要内容包括: 1)种子异时萌发的概念; 2)具有异时萌发性的一年生植物种类及分布; 3)异时萌发对植物生活史类型及模式的影响; 4)异时萌发的植株在表型响应上的不同, 如存活率、物候特征、繁殖产量、生物量分配等; 5)异时萌发的生物学和环境因素。在综述文献的基础上, 对今后的研究进行了展望, 这有助于从更深层面理解荒漠一年生植物的异时萌发特性及其适应机制。
迪力夏旦木·塔什买买提, 刘会良. 荒漠一年生植物异时萌发研究进展. 植物生态学报, 2023, 47(12): 1611-1628. DOI: 10.17521/cjpe.2022.0336
Dilixiadanmu TASHENMAIMAITI, LIU Hui-Liang. Review of research on germination heterochrony of desert annual plants. Chinese Journal of Plant Ecology, 2023, 47(12): 1611-1628. DOI: 10.17521/cjpe.2022.0336
图1 异时萌发的十字花科(A)、菊科(B)和紫草科(C)一年生植物分布图(以属分类)。黄色部分为2001-2020年全球的干旱半干旱区, 划分依据为FAO提出的干旱指数。所有物种分布数据来源于GBIF (https://www.gbif.org/)。
Fig. 1 Distribution map of heterochronous germinating annuals of Brassicaceae (A), Asteraceae (B) and Boraginaceae (C) (classified by genus). The yellow section shows the global arid and semi-arid region for 2001-2020. The classification is based on the FAO proposed aridity index. All species distribution data were obtained from GBIF (https://www.gbif.org/).
图2 异时萌发的一年生植物分布图(以科分类)。黄色部分为2001-2020年全球的干旱半干旱区, 划分依据为FAO提出的干旱指数。所有物种分布数据来源于GBIF (https://www.gbif.org/)。
Fig. 2 Distribution map of heterochronous germinating annuals (classified by family). The yellow section shows the global arid and semi-arid region for 2001-2020. The classification is based on the FAO proposed aridity index. All species distribution data were obtained from GBIF (https://www.gbif.org/).
图3 一年生植物的两年生活史周期及共存生活史模式。改自Han等(2016)。A, 冬季一年生植物, 秋季萌发, 春季繁殖, 以一年为生活史周期, 幼苗需越冬。B, 春季一年生植物, 春季萌发, 夏季繁殖, 以一年为生活史周期。C, 两年生活史周期内共存, 在两年内的同一种生活史模式, 不同的萌发时间使得春萌和秋萌幼苗共存。D, 两种生活史模式混合共存, 同一物种内存在两种独立的生活史模式, 都以一年为生活史周期, 因此在一个种群内春秋萌植株共存。
Fig. 3 Two-year life history cycle and coexistence life history pattern of annual plants. Adapted from Han et al. (2016). A, Winter annuals, which germinate in autumn, reproduce in spring, have a life history cycle of one year, seedlings need to be overwintered. B, Spring annuals, which germinate in spring, reproduce in autumn, have a life history cycle of one year. C, Coexistence within a two-year life history cycle. The same life history pattern over two years. Different germination times allow coexistence of spring- and autumn-germinating seedlings. D, Mixed coexistence of two life history patterns. Two independent life history patterns exist within the same species, both with a one-year life history cycle. Therefore, spring- and autumn- germinating plants coexist within a population. A, autumn; S, spring; SU, summer; W, winter.
图4 种子休眠状态变化的年周期示意图。改自Baskin和Baskin (2014a)。
Fig. 4 Schematic diagram of the annual cycle variation of seed dormancy. Adapted from Baskin & Baskin (2014a).
图5 种子休眠比例及类型与果实类型的关系。红色散点数据表示各果实类型所占的百分比。MD, 形态休眠; MPD, 形态生理休眠; ND, 不休眠; PD, 生理休眠; PD+PY, 生理+物理休眠; PY, 物理休眠; PY+MD, 物理+形态休眠。1, 蒺藜科; 2, 车前科; 3, 罂粟科; 4, 牻牛儿苗科; 5, 石竹科; 6, 大戟科; 7, 桔梗科; 8, 虎耳草科; 9, 报春花科; 10, 田基麻科; 11, 花荵科; 12, 粟米草科; 13, 马齿苋科; 14, 十字花科; 15, 毛茛科; 16, 菊科; 17, 蓼科; 18, 蔷薇科; 19, 紫草科; 20, 唇形科; 21, 莎草科; 22, 禾本科; 23, 藜科; 24, 苋科; 25, 豆科; 26, 伞形科。
Fig. 5 Proportion and kinds of seed dormancy in relation to fruit type. The red scatter plot data indicate the percentage of each fruit type. ND, nondormancy; PY, physical dormancy; PD, physiological dormancy; MD, morphological dormancy; MPD, morphophysiological dormancy; PD+PY, physiological and physical dormancy; PY+MD, physical and morphophysiological dormancy. 1, Zygophyllaceae; 2, Plantaginaceae; 3, Papaveraceae; 4, Geraniaceae; 5, Caryophyllaceae; 6, Euphorbiaceae; 7, Campanulaceae; 8, Saxifragaceae; 9, Primulaceae; 10, Hydroleaceae; 11, Polemoniaceae; 12, Molluginaceae; 13, Portulacaceae; 14, Brassicaceae; 15, Ranunculaceae; 16, Asteraceae; 17, Polygonaceae;18, Rosaceae; 19, Boraginaceae; 20, Lamiaceae; 21, Cyperaceae; 22, Poaceae;23, Chenopodiaceae;24, Amaranthaceae; 25, Fabaceae; 26, Apiaceae.
图6 脱落酸(ABA)-赤霉素(GA)网络对萌发时间的调控示意图。改自Abley等(2021)。箭头代表有效促进, 钝箭头代表有效抑制。该模型表现为相互抑制回路和相互激活回路, 分别表现在: GA与复合物相互抑制; ABA与复合物相互促进。复合物中的DELLA蛋白是GA信号转导的负调控因子, 转录因子ABI4、ABI5是ABA信号转导的正调控因子, ABI4、ABI5以及DELLAs蛋白相互作用形成转录复合物, 抑制萌发。转录复合物通过作用于激素的生物合成或分解代谢来影响GA和ABA水平, 精密调控种子的萌发。
Fig. 6 Schematic representation of the regulation of germination time by the abscisic acid (ABA)-gibberellic acid (GA) network. Adapted from Abley et al. (2021). The arrow represents effective facilitation and the blunt arrow represents effective inhibition. The model behaves as a mutual inhibition circuit and a mutual activation circuit, respectively: GA inhibits integrator and vice versa; ABA promotes integrator and vice versa. DELLA protein is a negative regulator of GA signaling. Transcription factors ABI4 and ABI5 are positive regulators of ABA signal transduction. ABI4, ABI5, and DELLA proteins interact to form a transcriptional complex that inhibits germination. The transcriptional complex affects GA and ABA levels by acting on hormone biosynthesis or catabolism to precisely regulate seed germination.
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