多年生单子叶植物的越冬休眠研究进展

doi:10.16420/j.issn.0513-353x.2021-1064

摘要/Abstract

摘要：

多年生单子叶植物具有特殊的地下芽、地面芽和地下器官，其越冬休眠特性明显有别于双子叶植物。汇总了环境、激素、碳水化合物调控多年生单子叶植物越冬休眠的机制，比较了多年生单、双子叶植物越冬休眠调控机制及研究进展，并结合研究现状，对未来多年生单子叶植物越冬休眠调控机制的研究方向提出建议。

关键词: 多年生单子叶植物, 越冬休眠, 需冷量, 脱落酸, MADS-box

Abstract:

Perennial monocots have special underground buds，ground buds，and underground organs. Their dormancy characteristics are obviously different from those of eudicots with aboveground buds. This review article summarized the regulation mechanisms of winter dormancy from environmental，hormonal and carbohydrate in monocots. The regulation mechanisms and research progress of winter dormancy in perennial monocots and eudicots were compared. Combined with the current research progress，suggestions for future directions on the study of winter dormancy regulation in perennial monocots were provided.

Key words: perennial monocot, winter dormancy, chilling requirement, abscisic acid, MADS-box

中图分类号:

S6
Q 945

许曈, 邵灵梅, 王小斌, 张润龙, 张凯靖, 夏宜平, 张佳平, 李丹青. 多年生单子叶植物的越冬休眠研究进展[J]. 园艺学报, 2022, 49(12): 2703-2721.

XU Tong, SHAO Lingmei, WANG Xiaobin, ZHANG Runlong, ZHANG Kaijing, XIA Yiping, ZHANG Jiaping, LI Danqing. Research Progress on Winter Dormancy of Perennial Monocots[J]. Acta Horticulturae Sinica, 2022, 49(12): 2703-2721.

图/表 5

图1 多年生单子叶植物越冬休眠研究论文数量、内容和通信作者所属国家

Fig. 1 Number and content of papers and country of corresponding author on winter dormancy of perennial monocots

表1 参与多年生单子叶植物越冬休眠调控的ABA合成与信号转导基因

Table 1 Genes involved in ABA biosynthesis and signal transduction for the regulation of winter dormancy in perennial monocots

基因名称 Gene name	基因全称 Full name of gene	基本功能 Gene function	研究物种 Research species	验证方法 Verification method	参考文献 Reference
NCED	9-cis-epoxycarotenoid dioxygenases	ABA生物合成 ABA biosynthesis	唐菖蒲 Gladiolus hybridus	基因沉默 Gene silencing	Wu et al.，2019a
CYP707A1	Cytochrome P450，Family 707，Subfamily A 1	参与ABA分解代谢途径 Involved in ABA catabolic pathway	滇黄精 Polygonatum kingianum	转录组分析 Transcriptome analysis	Wang et al.，2019
ABI5	Abscisic acid-insensitive 5	ABA信号调节因子 ABA signaling regulator	唐菖蒲 Gladiolus hybridus	基因沉默 Gene silencing	Wu et al.，2015
ABF	ABA-responsive element binding factors	ABA响应因子 ABA response factor	柳枝稷 Panicum virgatum	转录组分析 Transcriptome analysis	Palmer et al.，2017
PP2C	Protein phosphatase 2C	ABA信号转导因子 Regulators of ABA signal transduction	柳枝稷 Panicum virgatum	转录组分析 Transcriptome analysis	Palmer et al.，2017
PP2C1	Protein phosphatase 2C 1	ABA信号转导因子 Regulators of ABA signal transduction	唐菖蒲 Gladiolus hybridus	基因沉默 Gene silencing	Wu et al.，2019b
PYL	Pyrabactin-like	ABA受体 ABA receptors	柳枝稷 Panicum virgatum	转录组分析 Transcriptome analysis	Palmer et al.，2017
NAC83	NAC domain containing protein 83	促进ABA信号转导和合成 Positively regulates ABA signaling and biosynthesis	唐菖蒲 Gladiolus hybridus	基因沉默 Gene silencing	Wu et al.，2019b
SnRK	Snf1-related protein kinase	ABA信号转导因子 Regulators of ABA signal transduction	柳枝稷 Panicum virgatum	转录组分析 Transcriptome analysis	Palmer et al.，2017

表1 参与多年生单子叶植物越冬休眠调控的ABA合成与信号转导基因

Table 1 Genes involved in ABA biosynthesis and signal transduction for the regulation of winter dormancy in perennial monocots

基因名称 Gene name	基因全称 Full name of gene	基本功能 Gene function	研究物种 Research species	验证方法 Verification method	参考文献 Reference
NCED	9-cis-epoxycarotenoid dioxygenases	ABA生物合成 ABA biosynthesis	唐菖蒲 Gladiolus hybridus	基因沉默 Gene silencing	Wu et al.，2019a
CYP707A1	Cytochrome P450，Family 707，Subfamily A 1	参与ABA分解代谢途径 Involved in ABA catabolic pathway	滇黄精 Polygonatum kingianum	转录组分析 Transcriptome analysis	Wang et al.，2019
ABI5	Abscisic acid-insensitive 5	ABA信号调节因子 ABA signaling regulator	唐菖蒲 Gladiolus hybridus	基因沉默 Gene silencing	Wu et al.，2015
ABF	ABA-responsive element binding factors	ABA响应因子 ABA response factor	柳枝稷 Panicum virgatum	转录组分析 Transcriptome analysis	Palmer et al.，2017
PP2C	Protein phosphatase 2C	ABA信号转导因子 Regulators of ABA signal transduction	柳枝稷 Panicum virgatum	转录组分析 Transcriptome analysis	Palmer et al.，2017
PP2C1	Protein phosphatase 2C 1	ABA信号转导因子 Regulators of ABA signal transduction	唐菖蒲 Gladiolus hybridus	基因沉默 Gene silencing	Wu et al.，2019b
PYL	Pyrabactin-like	ABA受体 ABA receptors	柳枝稷 Panicum virgatum	转录组分析 Transcriptome analysis	Palmer et al.，2017
NAC83	NAC domain containing protein 83	促进ABA信号转导和合成 Positively regulates ABA signaling and biosynthesis	唐菖蒲 Gladiolus hybridus	基因沉默 Gene silencing	Wu et al.，2019b
SnRK	Snf1-related protein kinase	ABA信号转导因子 Regulators of ABA signal transduction	柳枝稷 Panicum virgatum	转录组分析 Transcriptome analysis	Palmer et al.，2017

图2 代表性多年生单、双子叶植物中DAM/SVP基因的系统进化分析杨树、葡萄、猕猴桃等基因组信息来源于Phytozome v13，鸢尾、蝴蝶花、百合、换锦花SVP蛋白序列来源于本实验室测序的转录组数据，唐菖蒲SVP蛋白序列来由NCBI中PRJNA491310数据拼接获得（Wu et al.，2019b）。选取拟南芥AtSVP作为对象，采用Blastp、Pfam搜索各物种中与其序列一致性较高的蛋白序列，运用mafft软件进行氨基酸序列比对，使用MEGA7.0以邻接法构建SVP蛋白的系统进化树。利用MEME在线软件进行蛋白保守结构域分析，得到16个保守基序，分别命名为Motif1 ~ Motif16。 Prupe：桃;Actinidia：美味猕猴桃;VIT：葡萄;Loh：东方百合;Ls：换锦花;IJ：蝴蝶花;GlaUn：唐菖蒲;IT：鸢尾;Aco：菠萝;GSMUA：香蕉;scaffold：无油樟;Nycol：蓝星睡莲;KAF：侏儒卢旺达睡莲。

Fig. 2 Phylogenetic analysis of DAM/SVP genes in representative perennial monocots and eudicots The genomic information of Populus trichocarpa，Vitis vinifera，Actinidia chinensis and other species in this evolutionary tree comes from Phytozome v13. The SVP protein sequence of Iris tectorum，I. japonica，Lilium oriental hybrid，and Lycoris sprengeri are derived from the transcriptome data sequenced by our laboratory，and the SVP protein sequence of Gladiolus gandavensis is derived from the data of PRJNA491310 in NCBI（Wu et al.，2019b）. The Arabidopsis thaliana AtSVP protein was selected as the query，Blastp and pfam was used to identify homologs of SVP in the above species. Mafft software was used for amino acid sequence comparison，and then the phylogenetic tree of SVP protein was constructed using MEGA7.0 by the neighbor-joining method. Using MEME online software to analyze the protein conserved domains，16 conserved motifs were obtained，and named as Motif1-Motif16. Prupe：Prunus persica;Actinidia：Actinidia deliciosa;VIT：Vitis vinifera;Loh：Lilium oriental hybrid;Ls：Lycoris sprengeri;IJ：Iris japonica;GlaUn：Gladiolus × gandavensis;IT：Iris tectorum;Aco：Ananas comosus; GSMUA：Musa nana;scaffold：Amborella trichopoda;Nycol：Nymphaea colorata;KAF：Nymphaea thermarum.

图3 DAM/SVP基因和ABA介导的多年生单、双子叶植物越冬休眠分子调控模型虚线：间接调控或不确定途径;橙色：多年生单子叶植物越冬休眠分子调控途径;绿色：多年生双子叶植物越冬休眠分子调控途径。

Fig. 3 DAM/SVP gene and ABA-mediated molecular regulation model of winter dormancy in perennial monocots and eudicots Dashed line：Indirect regulation or indeterminate pathway;Orange：Molecular regulation pathway of winter dormancy in perennial monocots;Green：Molecular regulation pathway of winter dormancy in perennial eudicots.

图4 多年生单、双子叶植物越冬休眠调控机制研究的比较

Fig. 4 Comparison of researches on the regulation mechanisms of winter dormancy in perennial monocots and eudicots

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