谷子果胶甲酯酶抑制子PMEI基因家族鉴定及其对非生物胁迫的响应分析

doi:10.13560/j.cnki.biotech.bull.1985.2025-0018

生物技术通报 ›› 2025, Vol. 41 ›› Issue (7): 150-163.doi: 10.13560/j.cnki.biotech.bull.1985.2025-0018

谷子果胶甲酯酶抑制子PMEI基因家族鉴定及其对非生物胁迫的响应分析

李新妮¹^,²(), 李俊怡¹^,², 马雪华², 何卫¹^,², 李佳丽¹^,², 于佳¹^,², 曹晓宁¹^,², 乔治军¹^,²(), 刘思辰¹^,²()

^1.山西农业大学农业基因资源研究中心农业农村部黄土高原作物基因资源与种质创制重点实验室，太原 030031
^2.山西农业大学农学院，太谷 030801

收稿日期:2025-01-08 出版日期:2025-07-26 发布日期:2025-07-22
通讯作者: ‍：‍刘思辰，女，博士，副研究员，研究方向：谷子种质资源评价利用；E-mail: lsch209@126.com
乔治军，男，博士，研究员，研究方向：旱作栽培与逆境生理；E-mail: qiaozhijun@sxagri.ac.cn
作者简介:李新妮，女，硕士，研究方向：作物遗传育种；E-mail: tangerine7423@163.com
基金资助:
?：?山西农业大学校科技提升工程(CXGC2023094);中央引导地方科技发展资金项目(YDZJSX20231A040);国家重点研发计划项目(2023YFD120270505);现代农业产业技术体系建设专项(CARS-06-14.5-A16)

Identification of the PMEI Gene Family of Pectin Methylesterase Inhibitor in Foxtail Millet and Analysis of Its Response to Abiotic Stress

LI Xin-ni¹^,²(), LI Jun-yi¹^,², MA Xue-hua², HE Wei¹^,², LI Jia-li¹^,², YU Jia¹^,², CAO Xiao-ning¹^,², QIAO Zhi-jun¹^,²(), LIU Si-chen¹^,²()

^1.Agricultural Gene Resources Research Center of Shanxi Agricultural University, Key Laboratory of Crop Gene Resources and Germplasm Enhancement on Loess Plateau of the Ministry of Agriculture and Rural Affairs, Taiyuan 030031
^2.College of Agriculture, Shanxi Agricultural University, Taigu 030801

Received:2025-01-08 Published:2025-07-26 Online:2025-07-22

摘要/Abstract

摘要：

目的 PMEI（pectin methylesterase inhibitor）是控制细胞壁结构与特性的重要组成部分，在植物逆境胁迫响应方面发挥重要作用。探究谷子PMEI基因在非生物胁迫下的响应机制，为谷子的抗逆机理研究提供理论基础。方法运用生物信息学方法对谷子PMEI基因家族进行鉴定，采用实时荧光定量PCR技术对其家族成员在低温、干旱、MeJA、ABA胁迫下的表达模式进行分析。结果谷子基因组中共鉴定出68个SiPMEI基因家族成员，不均匀地分布在9条染色体上，大多数成员定位于细胞壁或叶绿体上。SiPMEI家族成员主要有2种结构域，PMEI结构域和pectinesteras+PMEI结构域，含有同一结构域的成员间理化性质、亚细胞定位和基因结构均较相似。启动子顺式作用元件分析表明SiPMEI基因家族成员含有多种非生物胁迫和激素响应元件。本研究选择了8个含有2种以上胁迫响应元件且含数量较多的SiPMEI家族成员进行实时荧光定量表达分析。结果表明，SiPMEI在谷子的根、茎、叶和穗中差异表达；非生物胁迫（低温、干旱）和激素胁迫（MeJA、ABA）处理下，SiPMEI基因的表达量在0‒24 h内整体呈现上升趋势，含PMEI结构域且定位于细胞壁上的成员SiPMEI30、SiPMEI32、SiPMEI36、SiPMEI63在胁迫下响应的最高表达量集中于8‒24 h。亚细胞定位于叶绿体上的成员SiPMEI22、SiPMEI31、SiPMEI38、SiPMEI47最高表达量分布较为分散，且上调位点较多。结论 SiPMEI在低温、ABA和MeJA的胁迫下均具有正向响应，并在干旱和MeJA胁迫下具有类似的表达趋势。这些差异表达表明SiPMEI可能通过不同的分子机制响应非生物胁迫。

关键词: 谷子, PMEI基因家族, 干旱胁迫, 低温胁迫, 茉莉酸甲酯胁迫, 脱落酸胁迫, 亚细胞定位, 结构域

Abstract:

Objective PMEI (pectin methylesterase inhibitor) is a key component regulating the structure and properties of the cell wall, playing a significant role in plant stress response mechanisms. Therefore, studying the response mechanism of PMEI gene in foxtail millet under abiotic stress may provide a theoretical basis for the mechanism of resistance to stress in millet. Method Bioinformatics was used to identify PMEI gene family in foxtail millet, and quantitative real-time PCR was employed to analyze the expression patterns of family members under low temperature, drought, MeJA and ABA stress. Result A total of 68 SiPMEI gene family members were identified in the foxtail millet (Setaria italica) genome, unevenly distributed on nine chromosomes, and most of them were localized to the cell wall or chloroplasts. There were two main domains of SiPMEI family members, PMEI domain and pectinesteras + PMEI domain, as well as similar physicochemical properties, subcellular localization and gene structure of the family members containing the same domain. Promoter cis-acting element analysis suggested that the SiPMEI gene family members contained multiple abiotic stress and hormone responsive elements. In this study, eight SiPMEI family members containing more than two stress response elements were selected for quantitative real-time expression analysis. The results showed that SiPMEI was differentially expressed in the root, stem, leaf and ear of the foxtail millet. Under abiotic stress (low temperature, drought), hormone stress (MeJA, ABA), the expression of SiPMEI gene tended to increase within 0-24 h, and the highest expressions of member SiPMEI30, SiPMEI32, SiPMEI36 and SiPMEI63 containing PMEI domain and localized on the cell wall responded to stress at 8‒24 h. The highest expression distribution of member SiPMEI22, SiPMEI31, SiPMEI38, and SiPMEI47 with subcellular localization on chloroplasts was more dispersed and had more upregulation sites. Conclusion SiPMEI has a positive response under low temperature, ABA and MeJA stresses, and has a similar expression trend under drought and MeJA stresses. These differential expressions suggest that SiPMEI may respond to abiotic stress through different molecular mechanisms.

Key words: foxtail millet, PMEI gene family, drought stress, low-temperature stress, methyl jasmonate stress, abscisic acid stress, subcellular localization, domain

李新妮, 李俊怡, 马雪华, 何卫, 李佳丽, 于佳, 曹晓宁, 乔治军, 刘思辰. 谷子果胶甲酯酶抑制子PMEI基因家族鉴定及其对非生物胁迫的响应分析[J]. 生物技术通报, 2025, 41(7): 150-163.

LI Xin-ni, LI Jun-yi, MA Xue-hua, HE Wei, LI Jia-li, YU Jia, CAO Xiao-ning, QIAO Zhi-jun, LIU Si-chen. Identification of the PMEI Gene Family of Pectin Methylesterase Inhibitor in Foxtail Millet and Analysis of Its Response to Abiotic Stress[J]. Biotechnology Bulletin, 2025, 41(7): 150-163.

图/表 14

表1 用于RT-qPCR引物序列

Table 1 Primer sequences used for RT-qPCR

基因名称 Gene name	基因序列 Gene sequence （5′‒3′）
25S-F	AGGCAACAGAAACTCCATACG
25S-R	ATGGCATAGCATTCATCACG
SiPMEI22-F	TGTTCTCGGGACAGGTTTCG
SiPMEI22-R	GTACAGCGTGTCCTGGTACG
SiPMEI30-F	GACCTCGGCTACGACTACTG
SiPMEI30-R	GGTTGTGCTCTTGGAGGCTA
SiPMEI31-F	TCTTCAGAGAGAGGGGGATCG
SiPMEI31-R	TAGCGCGACAAGCGAACAAT
SiPMEI32-F	TTTTCCGCGAACGTGTCCA
SiPMEI32-R	ACACCACCCCATATCGCTTG
SiPMEI36-F	CTCGGTACAGTCTTGCACCA
SiPMEI36-R	ATCCAGTGCAATGTCTCCGT
SiPMEI38-F	CCGAGTACATGAACCGTGGA
SiPMEI38-R	GTGGAGTTGAGCCAGAGGTC
SiPMEI47-F	CAGCCTTCGTGTTCCCTGTT
SiPMEI47-R	TCTTATTGATGACCGCCCTCG
SiPMEI63-F	AGCTTCGTTCGCTCATGCAA
SiPMEI63-R	AGTAATGAAGGCTGCAACACAC

图1 SiPMEI序列特征分析A：SiPMEI基因家族成员进化树，B：保守基序分布；C：基因结构域分布；D：内含子与外显子结构

Fig. 1 Analysis of SiPMEI sequence characterizationA: SiPMEI evolutionary tree of gene family members; B: conserved motif distribution; C: gene domain distribution; D: intron and exon structure

图2 谷子（Si）、水稻（Os）、拟南芥（At）PMEI基因家族系统发育进化树

Fig. 2 Phylogenetic evolutionary tree of the PMEI gene family in Setaria italica (Si), Oryza sativa (Os), and Arabidopsis (At)

图3 SiPMEI基因家族在谷子染色体上的分布

Fig. 3 Distribution of the SiPMEI gene family on the foxtail millet chromosome

表2 SiPMEI基因进化中复制事件的选择压力参数

Table 2 Selection pressure parameters for replication events in the evolution of SiPMEI genes

重复基因1 Duplicated gene l	重复基因2 Duplicated gene 2	非同义替换率 Ka	同义替换率 Ks	比值 Ka/Ks
SiPMEI46	SiPMEI41	0.586 044 760	0.756 527 598	0.774 650 867
SiPMEI29	SiPMEI19	0.310 390 814	0.409 962 679	0.757 119 686
SiPMEI46	SiPMEI27	0.484 690 326	0.680 501 629	0.712 254 469
SiPMEI32	SiPMEI30	0.404 813 285	0.573 874 330	0.705 404 065
SiPMEI26	SiPMEI27	0.591 358 520	0.538 368 419	1.098 427 208
SiPMEI6	SiPMEI23	0.202 922 616	0.397 601 411	0.510 366 941
SiPMEI5	SiPMEI34	0.660 998 002	0.498 090 476	1.327 064 124
SiPMEI63	SiPMEI50	0.617 173 087	0.439 225 359	1.405 139 924

图4 谷子种内PMEI基因家族共线性分析1-9 代表谷子第1-9染色体；染色体内部以基因密度填充，颜色由蓝到红代表基因密度由小到大

Fig. 4 Collinearity analysis of intra-specific PMEI gene families in foxtail millet1-9 indicates chromosome 1-9 in foxtail millet respectively. The interior of the chromosome is filled with gene density, with colors ranging from blue to red indicating a gradual increase in gene density

图5 SiPMEI基因家族成员与水稻和拟南芥PMEI基因家族种间共线性分析

Fig. 5 Analysis of inter-specific collinearity between SiPMEI gene family members and PMEI gene families of rice and Arabidopsis

图6 SiPMEI基因家族启动子顺式元件分析

Fig. 6 Analysis of cis-element promoters of the SiPMEI gene family

表3 所选引物及胁迫响应元件的数目

Table 3 Number of selected primers and stress-responsive elements

基因名称 Gene name	低温响应元件 Low-temperature responsive element	脱落酸响应元件 ABA responsive element	茉莉酸甲酯响应元件 MeJA responsive element	干旱响应元件 Drought responsive element	亚细胞定位 Subcellular location	结构域 Structural domain
SiPMEI22	1	8	6	0	Chloroplast	Pectinesteras + PMEI
SiPMEI30	0	6	10	0	Cell wall	PMEI
SiPMEI31	0	4	12	0	Chloroplast	PMEI
SiPMEI32	2	0	0	4	Cell wall	PMEI
SiPMEI36	1	4	10	0	Cell wall	PMEI
SiPMEI38	0	1	4	2	Chloroplast	Pectinesteras + PMEI
SiPMEI47	1	10	16	1	Chloroplast	Pectinesteras + PMEI
SiPMEI63	0	7	0	2	Cell wall	PMEI

图7 SiPMEI基因家族在不同组织部位中的表达分析

Fig. 7 Expression analysis of the SiPMEI gene family in different tissue sites

图8 低温胁迫下不同基因的表达水平误差线为标准误，图中不同字母表示差异水平显著（P < 0.05），下同

Fig. 8 Expressions of different genes under low-temperature stressThe error line is the standard error, and the different letters in the figure indicate significant difference levels (P <0.05), the same below

图9 干旱胁迫下不同基因的表达水平

Fig. 9 Expressions of different genes under drought stress

图10 茉莉酸甲酯胁迫下不同基因的表达水平

Fig. 10 Expressions of different genes under methyl jasmonate stress

图11 脱落酸胁迫下不同基因的表达水平

Fig. 11 Expressions of different genes under abscisic acid stress

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谷子果胶甲酯酶抑制子PMEI基因家族鉴定及其对非生物胁迫的响应分析

Identification of the PMEI Gene Family of Pectin Methylesterase Inhibitor in Foxtail Millet and Analysis of Its Response to Abiotic Stress

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