菠菜AT-hook基因家族鉴定与表达谱分析

doi:10.11983/CBB24117

植物学报 ›› 2025, Vol. 60 ›› Issue (3): 377-392.DOI: 10.11983/CBB24117 cstr: 32102.14.CBB24117

菠菜AT-hook基因家族鉴定与表达谱分析

杨莉, 曲茜彤, 陈子航, 邹婷婷, 王全华, 王小丽^*()

上海师范大学生命科学学院/植物种质资源开发协同创新中心, 上海 200234

收稿日期:2024-07-30 接受日期:2024-12-26 出版日期:2025-05-10 发布日期:2024-12-27
通讯作者: *王小丽, 上海师范大学生命科学学院副教授, 硕士生导师。主要从事园艺植物营养品质调控、遗传鉴定及新品种选育等工作。目前研究团队以菠菜为主要研究对象, 收集了世界各地菠菜种质资源千余份, 完成菠菜资源的深度转录组测序和全基因组测序工作, 利用生理生化、遗传学和分子生物学等手段对其重要农艺性状进行研究。E-mail: wangxl@shnu.edu.cn
基金资助:
上海市科技兴农项目(2022-02-08-00-12-F01116)

Identification of the Spinach AT-hook Gene Family and Analysis of Expression Profiles

Yang Li, Qu Xitong, Chen Zihang, Zou Tingting, Wang Quanhua, Wang Xiaoli^*()

Development and Collaborative Innovation Center of Plant Germplasm Resources, College of Life Sciences, Shanghai Normal University, Shanghai 200234, China

Received:2024-07-30 Accepted:2024-12-26 Online:2025-05-10 Published:2024-12-27
Contact: *E-mail: wangxl@shnu.edu.cn

摘要/Abstract

摘要： AHL (AT-hook)蛋白家族在植物生长发育、器官构建、逆境胁迫及植物激素信号应答中发挥关键作用。为揭示菠菜(Spinacia oleracea) AHL基因家族的生物学功能, 在基因组范围内对SoAHL家族成员进行鉴定和分析, 并探究外源水杨酸对其表达水平的影响。结果表明, SoAHL家族包含19个成员, 在6条染色体间不均匀分布, 系统发育树将其分为2个亚族, I、II亚族分别包含10、9个SoAHL成员; 不同亚族间PPC和AT-hook保守基序序列组成有差异; SoAHL主要定位在细胞核、细胞质和线粒体; SoAHL I亚族成员无内含子, II亚族成员含4-5个内含子; SoAHL启动子上游分布有不同数量的植物激素和非生物胁迫响应相关顺式作用元件。SoAHL成员在根、叶片及叶柄中均有表达, 大多数在根中表达水平较高。水杨酸处理条件下, SOV6g041850.1和SOV2g038950.1受水杨酸显著诱导; SOV2g031340.1和SOV4g018880.1主要在地上部表达, 与叶酸含量的组织分布特征相似。瞬时过表达SOV4g018880.1能提高菠菜叶酸含量1.75倍。研究结果可为进一步解析SoAHL的功能奠定基础。

关键词: 菠菜, AT-hook基因家族, 生物信息学, 水杨酸

Abstract: INTRODUCTION: The AT-hook motif nuclear localized (AHL) gene family is a highly conserved transcription factors involved in plant growth, development, and stress responses, but their roles in spinach are still unknown. RATIONALE:To reveal the basic characteristics of the AHL family in spinach, members of the spinach SoAHLfamily were identified at the whole-genome level, and their physicochemical properties, gene structure, conserved motifs, promoter elements, and salicylic acid-responsive expression profiles were analyzed in this study.RESULTS: The results revealed 19 SoAHL family members in the spinach genome, which were unevenly distributed across six chromosomes. These SoAHL members can be classified into three branches, with 10 members in subfamily I and 9 members in subfamily II. The sequence composition of PPC and AT-hook conserved motifs varies among subfamilies; most of the SoAHL genes are located in the nucleus, cytoplasm, and mitochondrion. Members of subfamily I of SoAHL have no introns, whereas members of subfamily II contain 4-5 introns. The varying numbers of cis-acting elements relate to phytohormones and abiotic stress responses were distributed upstream of the promoters of the SoAHL members. The SoAHL genes can be expressed in roots, leaves, and petioles, with most genes expressed at relatively high levels in roots. The expression of two SoAHL genes (SOV6g041850.1 and SOV2g038950.1) was significantly induced by salicylic acid treatment. The expression profiles and salicylic acid-induced expression levels of SOV2g031340.1 and SOV4g018880.1 were highly correlated with the folic acid content, which may play a role in the spinach response to the salicylic acid signaling pathway. The transient overexpression of SOV4g018880.1 increased the folate content of spinach leaves by 1.75 times.CONCLUSION: The results from the sequence characteristics, expression profiles and exogenous salicylic acid treatment revealed that the SoAHLs had potential functional diversity and that specific members may have positive effects on spinach folate accumulation. Our results will lay the foundation for further resolving the function of spinach AT-hook genes.

Phenotype (A), total folate content (B) and expression analysis of SoAHL (C) under 50 μmol∙L^-1salicylic acid (SA) treatment for 5 days (D5) and 7 days (D7). CK: Control

Key words: spinach, AT-hook gene family, bioinformatics, salicylic acid

杨莉, 曲茜彤, 陈子航, 邹婷婷, 王全华, 王小丽. 菠菜AT-hook基因家族鉴定与表达谱分析. 植物学报, 2025, 60(3): 377-392.

Yang Li, Qu Xitong, Chen Zihang, Zou Tingting, Wang Quanhua, Wang Xiaoli. Identification of the Spinach AT-hook Gene Family and Analysis of Expression Profiles. Chinese Bulletin of Botany, 2025, 60(3): 377-392.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB24117

https://www.chinbullbotany.com/CN/Y2025/V60/I3/377

图/表 12

表1 引物信息

Table 1 Sequences of the primers used in this study

Gene ID	Forward primer (5′-3′)	Reverse primer (5′-3′)
SOV4g002370.1	GTTCTAACCATCTCCGCAAT	TGAATGGGTGAAATTGGCAT
SOV5g038710.1	GAGCGGCTATGATGGAG	GGATCTGATCTGTCCATGTC
SOV1g029900.1	TAAGATGGAGCAACAACAGG	TCGGGTTCTGATTATCCTCT
SOV6g041850.1	ATATCGGCATCACAACATGG	AGTGGTTTTGGAGTTGAAGT
SOV2g030150.1	ATGGAAGGCCTCTACCACC	GTGGTGAAACTGATGATGCTG
SOV6g047350.1	GACTCTCGTGGCCCATCTGC	GTAAGGTGAATTTGATTGGTGG
SOV2g031340.1	GAGAGGATTAGCCGAGTTAC	CCCGACATTTGATTGGAAAG
SOV3g001110.1	GTAGTTCAGGGGTAACTGTG	TTTTCACCCCTAGCTGTTAC
SOV2g038950.1	GATAACAATGCTCAAAACAATG	GTTGGAATTTGGAAGGATGAG
SOV4g018880.1	CCGAATGAACAAAGCGGAC	GACATTCCATTGGTGGCGG
SOV5g038610.1	TCTGAAAATGACGTCGGTAG	CGGCTTATAAACTGCATTCC
SOV6g041780.1	GTAGTAAGGTCAGATGCTCC	CATACTTCCTTGGTCTACCC
SOV2g029980.1	GGTGACAGTAATAGGAGCAG	TTACTGGAGTAACATCCCCA
SOV5g021850.1	TCAGGACCTGGTAGCTTTTA	AAAGTCATCCCGGAATTAGG
SOV6g040610.1	GGATGGGAGAGAAGGTATGGC	CCAACTGCCCCAGGAATTGT
SOV4g057830.1	ATAACGCCCTGGTGCAACTG	TGTGGCCTCCATCCGAACACAT
SOV4g019010.1	ACGAACAACCCGGACAACTT	ACCGCCATGATTCCATCCAT
SOV6g040700.1	TTGCCTCCGGTCCTGTGGTTATTAT	TCAGGTCTTTGCCAGTGATCTACC
SOV6g047370.1	TCCGGTCCTGTGGTTATTATGGC	TCAGGTCTTTGCCAGTGATCTACC
18S	CCATAAACGATGCCGACCAG	AGCCTTGCGACCATACTCCC

表2 菠菜SoAHL基因家族成员基本信息

Table 2 Information on the SoAHL gene family members in spinach

Gene ID	Genome location	Subcellular localization	Length (bp)	Amino acid (aa)	Molecular weight (kDa)	PI
SOV1g029900.1	Chr. 1: 113135928-113136926	NLS, Chl	999	333	33.84	5.37
SOV4g019010.1	Chr. 4: 52394687-52395604	NLS	918	306	31.47	6.16
SOV4g057830.1	Chr. 4: 186409628-186410449	NLS, Chl	822	274	27.68	6.42
SOV5g038710.1	Chr. 5: 123243932-123247784	NLS	819	273	26.72	6.43
SOV6g040700.1	Chr. 6: 140433598-140434404	NLS, Cyt	807	269	28.36	9.07
SOV6g047370.1	Chr. 6: 148547185-148548063	Chl, NLS	879	293	30.18	6.04
SOV2g030150.1	Chr. 2: 101786656-101787666	NLS, Chl	1011	337	34.80	7.22
SOV6g041850.1	Chr. 6: 141988749-141989711	NLS, Chl	963	321	32.97	5.97
SOV2g038950.1	Chr. 2: 112654862-112655896	Cyt, NLS	1035	345	36.80	6.85
SOV4g018880.1	Chr. 4: 52004210-52011645	NLS	1041	347	36.10	8.46
SOV2g031340.1	Chr. 2: 103380711-103385136	NLS	990	330	34.05	9.58
SOV6g040610.1	Chr. 6: 140337052-140344304	NLS, Cyt	1044	348	35.29	9.89
SOV2g029980.1	Chr. 2: 101504484-101511484	NLS, Chl	1032	344	35.92	9.44
SOV6g041780.1	Chr. 6: 141900514-141905585	NLS	1026	342	35.70	8.33
SOV3g001110.1	Chr. 3: 1145951-1151558	Chl, NLS	1089	363	37.96	5.73
SOV5g021850.1	Chr. 5: 39407625-39411618	NLS, Chl	1089	363	37.31	7.87
SOV5g038610.1	Chr. 5: 122545368-122554893	NLS	1092	364	37.09	9.87
SOV6g047350.1	Chr. 6: 148521377-148529101	NLS, Chl	1413	471	50.13	8.79
SOV4g002370.1	Chr. 4: 2764521-2765501	NLS, Cyt	981	327	36.06	6.04

图1 菠菜、拟南芥和水稻AHL蛋白家族进化树分析紫色表示I亚族, 蓝色表示II亚族。

Figure 1 Phylogenetic tree of AHL family members of spinach, Arabidopsis and rice Purple indicates subfamily I, and blue indicates subfamily II.

图2 菠菜SoAHL家族成员保守结构域分析

Figure 2 Conserved domain analysis of SoAHL family members in spinach

图3 SoAHL家族成员保守基序(A)及基因结构分析(B)

Figure 3 Analysis of conserved motifs (A) and gene structure (B) in the SoAHL family members

图4 菠菜SoAHL基因的区域共线性关系具有同源性的染色体区域用红线连接SoAHL基因, 外环为基因密度热图。

Figure 4 Regional collinearity of the SoAHL gene in spinach The homologous chromosome regions were connected with the SoAHL genes by red lines, and the outer ring represents the gene density heatmap.

图5 AHL基因家族物种间共线性分析红色线表示物种间AHL基因家族成员的共线性基因, 浅灰线表示各物种间包含的所有共线性基因。At: 拟南芥; Os: 水稻; So: 菠菜

Figure 5 Collinearity analysis of the AHL gene family in different species The red lines between the bars indicate collinear genes among members of the AHL gene family across species; the fine gray line shows all the collinear genes contained between species. At: Arabidopsis thaliana; Os: Oryza sativa; So: Spinacia oleracea

图6 SoAHL基因家族成员顺式作用元件分析

Figure 6 Analysis of cis-acting elements of the SoAHL gene family members

图7 菠菜3种组织叶酸含量(A)与SoAHL基因表达谱(B) L: 叶片; P: 叶柄; R: 根。不同小写字母表示差异显著(P<0.05)。

Figure 7 Folate content (A) and expression pattern of SoAHL gene (B) in three tissues of spinach L: Leaf; P: Petiole; R: Root. Different lowercase letters indicate significant differences(P<0.05).

表3 水杨酸(SA)处理对菠菜生物量的影响

Table 3 Effects of salicylic acid (SA) treatment on the biomass of spinach

Salicylic acid	Days of treatments (d)	Shoot fresh biomass (g)	Root fresh biomass (g)	Shoot dry biomass (g)	Root dry biomass (g)
Mock	0	0.737±0.171 d	0.053±0.004 b	0.083±0.013 b	0.004±0.001 a
	1	1.042±0.049 c	0.076±0.016 b	0.083±0.013 b	0.030±0.036 a
	3	1.097±0.125 c	0.066±0.008 b	0.095±0.014 b	0.008±0.002 a
	5	1.304±0.017 b	0.079±0.016 b	0.132±0.028 a	0.012±0.008 a
	7	1.511±0.054 a	0.104±0.016 a	0.111±0.020 ab	0.010±0.003 a
SA50	0	0.800±0.190 c	0.061±0.003 a	0.087±0.015 a	0.007±0.003 a
	1	1.123±0.116 b	0.077±0.015 a	0.123±0.032 a	0.012±0.005 a
	3	1.083±0.115 b	0.088±0.056 a	0.339±0.441 a	0.010±0.003 a
	5	1.303±0.080 ab	0.069±0.010 a	0.096±0.018 a	0.010±0.006 a
	7	1.501±0.085 a	0.077±0.015 a	0.094±0.038 a	0.049±0.070 a

图8 水杨酸(SA)处理下菠菜表型(A)、叶酸含量(B)与SoAHL表达谱(C) CKD5: 对照组处理5天; SAD5: SA处理5天; CKD7: 对照组处理7天; SAD7: SA处理7天。星号表示同一时间下处理间差异显著(P<0.05, Tukey检验); 热图方框中数字为基因相对表达量。Bars=5 cm

Figure 8 Phenotype (A), total folate content (B) of spinach and expression analysis of SoAHL (C) under salicylic acid (SA) treatment CKD5: Control group for 5 days; SAD5: SA treatment for 5 days; CKD7: Control group for 7 days; SAD7: SA treatment for 7 days. The stars indicate significant differences between treatments at the same time (P<0.05, Tukey test); the numbers in the heat map boxes respresent gene relative expression. Bars=5 cm

图9 SoAHLa瞬时过表达后基因的相对表达量(A)及菠菜叶片叶酸含量变化(B) CK: 注射空载体pEG100-SW的菠菜叶片; OE: 注射含SOV4g018880.1重组过表达载体的菠菜叶片。* 表示同一时间下处理间差异显著(P<0.05, Tukey检验)。

Figure 9 Changes in the relative expression level of the SoAHLa (A) and the folate content of leaf (B) after transient overexpression of SoAHLa CK: Spinach leaves injected with empty pEG100-SW; OE: Spinach leaves injected with vector containing SOV4g018880.1. * indicate significant differences between treatments at the same time (P<0.05, Tukey test).

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菠菜AT-hook基因家族鉴定与表达谱分析

Identification of the Spinach AT-hook Gene Family and Analysis of Expression Profiles

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