小麦IQM基因家族鉴定及非生物胁迫下表达分析

doi:10.13304/j.nykjdb.2022.0612

摘要/Abstract

摘要：

IQM基因是钙调素结合蛋白家族中的重要分支，在植物生长发育和应激反应中发挥重要作用。本研究利用生物信息学方法在小麦全基因组中鉴定出23个IQM基因家族成员，对其染色体位置、理化性质、系统进化关系、基因结构、蛋白保守结构域、启动子顺式作用元件和基因表达特性等进行了系统分析。结果表明，TaIQM基因家族成员随机分布在小麦18条染色体上，亚细胞定位结果显示所有基因均位于细胞核中，系统进化分析将其分为3个亚类，同一亚类间基因结构、蛋白保守结构域相似度较高，推测其功能相似；顺式作用元件分析表明，TaIQM基因家族成员启动子中含有多种与逆境胁迫及生长发育相关顺式作用元件；转录组数据分析显示，TaIQM基因家族成员在不同时期的根、茎、叶、穗和籽粒中表达量存在显著性差异；qRT-PCR分析显示，在小麦苗期地上部和地下部，TaIQM基因响应干旱、低温、高温、NaCl、ABA等多种胁迫，显示上调或下调表达。初步推断这些基因可能通过Ca²⁺信号通路参与非生物胁迫调控，研究结果为全面解析TaIQM基因结构与生物学功能、非生物胁迫响应分子机制提供依据。

关键词: 小麦, IQM家族, 基因鉴定, 非生物胁迫

Abstract:

IQM genes are important branch of the calmodulin-binding protein family that plays an important role in plant growth，development and stress response. In this study， 23 IQM gene family members were identified in the whole wheat genome using bioinformatics methods， and their chromosomal location， physicochemical properties， phylogenetic relationships， gene structure， protein conserved structural domains， promoter cis-acting elements and gene expression characteristics were systematically analysed. The results showed that the TaIQM gene family members were randomly distributed on the 18 chromosomes of wheat， and their subcellular localization showed that all genes were located in the nucleus. Phylogenetic analysis classified them into three subclasses with high similarity in gene structure and protein conserved structural domains among， and presumably similar functions； cis-acting element analysis showed that the promoters of TaIQM gene family members contained a variety of cis-acting elements related to stress， growth and development. Transcriptome analysis showed that the TaIQM gene family members were significantly differentially expressed in roots， stems， leaves， spikes and seeds at different times. qRT-PCR analysis showed that TaIQM genes were up- or down-regulated in response to various stresses such as drought， low and high temperature， NaCl and ABA in the above- and below-ground parts of wheat seedlings. It was tentatively concluded that these genes might be involved in abiotic stress regulatory through the Ca²⁺ signalling pathway. The results of this study provided clues for a comprehensive analysis of the structure and biological functions of TaIQM genes， the molecular mechanisms of abiotic stress response.

Key words: wheat, IQM gene family, gene indentification, abiotic stress

中图分类号:

梁婷, 左静红, 陆青, 杨东, 唐益苗, 郭春曼, 汪德州, 王伟伟. 小麦IQM基因家族鉴定及非生物胁迫下表达分析[J]. 中国农业科技导报, 2023, 25(2): 27-37.

Ting LIANG, Jinghong ZUO, Qing LU, Dong YANG, Yimiao TANG, Chunman GUO, Dezhou WANG, Weiwei WANG. Identification and Expression Analysis Under Abiotic Stress of IQM Gene Family in Wheat （Triticum aestivum L.）[J]. Journal of Agricultural Science and Technology, 2023, 25(2): 27-37.

图/表 7

表1 qRT-PCR引物 (续表Continued)

Table 1 qRT-PCR primers

基因 Gene	正向引物 Forward primer （5’–3’）	反向引物 Reverse primer （5’–3’）
TaIQM1	GACGATTGGTGGTCAAAGATGG	ACGCCGTCTATCTCAACTTCTGTC
TaIQM2	TCGTTTCAGCACTCCAGTTTCC	TTGACATCGGTGAGGCTAACGT
TaIQM3	CAAGGGCAGTGCGTCAAGTA	TGGAGTCATCCGAAGTGTGC
TaIQM4	GGAGATGTTCACAAGGCTGGATTC	GGGTAGTCCCTGACGCAAATGAT
TaIQM5	ATGGAGTTCTGAAGGCTATCTGGC	TTCTTCGGGTTTGTCGGCATC
TaIQM6	GTGCCAAGGGAGAAGGTCATT	GGTAGTCCCTGACGCAAATGAT
TaIQM7	CGAGCCTACTGAAACGGAAGAAC	CCATTTGAAGGACGGGAGACTG
TaIQM8	TCAAGAACTGGGAGGCGGAG	AGACGAAGTGCGGCTTGGAC
TaIQM9	CGAGAGGGCAAGGTATGAGGTTA	ACCCTTCTTCTTCTGCCCGAT
TaIQM10	AACTCGGACAGGCCATTTGT	TAGAGCTCACCGCATCGAAC
TaIQM11	CCATAAGCACAACCACAGCCT	CAACAATGGACGACAAGCACAG
TaIQM12	CTCGCCAGCAGATGATGACA	AAGGACCCTTTTCTCGCAGG
TaIQM13	GCTCATCCACCAAGGAAGACTAC	GGTACCATGTTGGAAGGCAGT
TaIQM14	GGAACCCTGAAGGCTATTTGG	TGGTTCCTCCCGTTTCTCTGT
TaIQM15	TAATGTCCGTGTGGAGCAAGCA	CGCTTGCATTCTGAGCTCTATCG
TaIQM16	GTGGTCACTACCGCCCTACA	CCTCTGCTGGGCTCATCTTC
TaIQM17	TGGAGCAAGCGAGACCAACCTA	GGCCGAGCTGATAGGATTTTGAC
TaIQM18	GAGAGGACTATGAGGTCGTGATTG	CCATTTTCAGCAACCAGCCT
TaIQM19	CCCAACTGATTCTCCCTTCCAAC	CCACACGGAGATTATCGTTGACCT
TaIQM20	CACAACTGATTCTCCCTTCCAAC	ATTCCCAGTGCTCCATTTCAG
TaIQM21	AGTGGACATTACAAACCAAGTGCG	ATTGTGGAGGATTGGATGGCA
TaIQM22	TGGACATTACAAACCGAGTGCG	AGCAGCAGTAGGGTTCTGTTTGGT
TaIQM23	CGTCAGCCTCAACGCATCAA	GGTTATGCCCGTACCTATGTCTTG
18S	CGCGCGCTACGGCTTTGACCTA	CGGCAGATTCCCACGCGTTACG

表2 小麦IQM基因家族蛋白理化性质及亚细胞定位 (续表Continued)

Table 2 Characteristics and subcellular localization prediction of TaIQM genes

基因 Gene	基因号 Gene ID	染色体定位 Chromosome localization	氨基酸数 Number of amino acids	相对分子量 Molecular weight/Da	等电点 Point isoelectric	亚细胞定位 Subcellular localization
TaIQM1	TraesCS1A02G125400.1	Chr1A： 149352287-149356056	517	57 811.59	7.68	细胞核Nucleus
TaIQM2	TraesCS1B02G143800.1	Chr1B： 195555337-195560543	580	65 214.18	7.69	细胞核 Nucleus
TaIQM3	TraesCS1D02G128400.1	Chr1D： 141545585-141547817	424	47 667.67	9.49	细胞核Nucleus
TaIQM4	TraesCS2A02G154900.1	Chr2A： 102130189-102132879	564	63 624.94	6.47	细胞核Nucleus
TaIQM5	TraesCS2B02G180000.1	Chr2B： 154592307-154595019	572	64 351.52	6.36	细胞核Nucleus
TaIQM6	TraesCS2D02G160200.1	Chr2D： 103126893-103129599	571	64 331.63	6.26	细胞核Nucleus
TaIQM7	TraesCS3A02G206400.1	Chr3A： 363415357-363419189	556	62 188.59	9.13	细胞核Nucleus
TaIQM8	TraesCS3B02G238500.1	Chr3B： 373842628-373844908	541	60 758.12	9.07	细胞核Nucleus
TaIQM9	TraesCS3D02G209200.1	Chr3D： 276574646-276577885	530	59 496.51	8.97	细胞核Nucleus
TaIQM10	TraesCS4A02G021000.1	Chr4A： 14261744-14265762	610	67 879.37	9.60	细胞核Nucleus
TaIQM11	TraesCS4B02G282600.2	Chr4B： 565861371-565865127	669	74 779.20	9.41	细胞核Nucleus
TaIQM12	TraesCS4D02G281600.1	Chr4D： 452795390-452798654	613	68 128.53	9.67	细胞核Nucleus
TaIQM13	TraesCS5A02G129600.1	Chr5A： 290924878-290930090	461	51 276.55	6.88	细胞核Nucleus
TaIQM14	TraesCS5A02G375700.1	Chr5A： 573606336-573609441	534	59 693.35	8.60	细胞核Nucleus
TaIQM15	TraesCS5B02G128200.1	Chr5B： 234295411-234304552	480	53 236.89	6.14	细胞核Nucleus
TaIQM16	TraesCS5B02G377900.1	Chr5B： 555950716-555953748	538	60 186.87	7.57	细胞核Nucleus
TaIQM17	TraesCS5D02G137100.1	Chr5D： 218051693-218055671	480	53 123.76	6.30	细胞核Nucleus
TaIQM18	TraesCS5D02G385200.1	Chr5D： 454309041-454312175	533	59 832.36	7.17	细胞核Nucleus
TaIQM19	TraesCS6A02G108900.1	Chr6A： 77780102-77784144	485	53 610.33	8.58	细胞核Nucleus
TaIQM20	TraesCS6B02G133900.1	Chr6B： 130822739-130829829	477	52 647.25	6.44	细胞核Nucleus
TaIQM21	TraesCS6B02G137500.1	Chr6B： 135112248-135116198	483	53 349.14	8.82	细胞核Nucleus
TaIQM22	TraesCS6D02G093400.1	Chr6D： 58234159-58240593	480	53 062.87	8.04	细胞核Nucleus
TaIQM23	TraesCS6D02G097300.1	Chr6D： 61134482-61138500	484	53 425.06	8.06	细胞核Nucleus

表2 小麦IQM基因家族蛋白理化性质及亚细胞定位 (续表Continued)

Table 2 Characteristics and subcellular localization prediction of TaIQM genes

基因 Gene	基因号 Gene ID	染色体定位 Chromosome localization	氨基酸数 Number of amino acids	相对分子量 Molecular weight/Da	等电点 Point isoelectric	亚细胞定位 Subcellular localization
TaIQM1	TraesCS1A02G125400.1	Chr1A： 149352287-149356056	517	57 811.59	7.68	细胞核Nucleus
TaIQM2	TraesCS1B02G143800.1	Chr1B： 195555337-195560543	580	65 214.18	7.69	细胞核 Nucleus
TaIQM3	TraesCS1D02G128400.1	Chr1D： 141545585-141547817	424	47 667.67	9.49	细胞核Nucleus
TaIQM4	TraesCS2A02G154900.1	Chr2A： 102130189-102132879	564	63 624.94	6.47	细胞核Nucleus
TaIQM5	TraesCS2B02G180000.1	Chr2B： 154592307-154595019	572	64 351.52	6.36	细胞核Nucleus
TaIQM6	TraesCS2D02G160200.1	Chr2D： 103126893-103129599	571	64 331.63	6.26	细胞核Nucleus
TaIQM7	TraesCS3A02G206400.1	Chr3A： 363415357-363419189	556	62 188.59	9.13	细胞核Nucleus
TaIQM8	TraesCS3B02G238500.1	Chr3B： 373842628-373844908	541	60 758.12	9.07	细胞核Nucleus
TaIQM9	TraesCS3D02G209200.1	Chr3D： 276574646-276577885	530	59 496.51	8.97	细胞核Nucleus
TaIQM10	TraesCS4A02G021000.1	Chr4A： 14261744-14265762	610	67 879.37	9.60	细胞核Nucleus
TaIQM11	TraesCS4B02G282600.2	Chr4B： 565861371-565865127	669	74 779.20	9.41	细胞核Nucleus
TaIQM12	TraesCS4D02G281600.1	Chr4D： 452795390-452798654	613	68 128.53	9.67	细胞核Nucleus
TaIQM13	TraesCS5A02G129600.1	Chr5A： 290924878-290930090	461	51 276.55	6.88	细胞核Nucleus
TaIQM14	TraesCS5A02G375700.1	Chr5A： 573606336-573609441	534	59 693.35	8.60	细胞核Nucleus
TaIQM15	TraesCS5B02G128200.1	Chr5B： 234295411-234304552	480	53 236.89	6.14	细胞核Nucleus
TaIQM16	TraesCS5B02G377900.1	Chr5B： 555950716-555953748	538	60 186.87	7.57	细胞核Nucleus
TaIQM17	TraesCS5D02G137100.1	Chr5D： 218051693-218055671	480	53 123.76	6.30	细胞核Nucleus
TaIQM18	TraesCS5D02G385200.1	Chr5D： 454309041-454312175	533	59 832.36	7.17	细胞核Nucleus
TaIQM19	TraesCS6A02G108900.1	Chr6A： 77780102-77784144	485	53 610.33	8.58	细胞核Nucleus
TaIQM20	TraesCS6B02G133900.1	Chr6B： 130822739-130829829	477	52 647.25	6.44	细胞核Nucleus
TaIQM21	TraesCS6B02G137500.1	Chr6B： 135112248-135116198	483	53 349.14	8.82	细胞核Nucleus
TaIQM22	TraesCS6D02G093400.1	Chr6D： 58234159-58240593	480	53 062.87	8.04	细胞核Nucleus
TaIQM23	TraesCS6D02G097300.1	Chr6D： 61134482-61138500	484	53 425.06	8.06	细胞核Nucleus

图1 IQM蛋白进化树分析注：At—拟南芥；Os—水稻；Ta—小麦。

Fig. 1 Phylogeny analysis of IQMproteinsNote： At—Arabidopsis thaliana； Os—Oryza sativa； Ta—Triticum aestivum L..

图2 TaIQM家族成员结构特征和保守基序分析A：基因结构；B：保守基序；C：IQM保守基序序列

Fig. 2 Structure and conservative motif analysis of the TaIQMA： Gene structure；B： Conserved motif；C： Conserved motif sequences of IQM

图3 TaIQM启动子顺式作用元件预测注：ABRE—脱落酸顺式作用元件；ARE—无氧诱导所必需的顺式作用调节元件；Circadian—参与昼夜节律控制的顺式作用调节元件；G-box—参与光反应的顺式作用调控元件；TGACG-motif—与 MeJA 反应有关的顺式作用调节因子；GT1-motif—光响应元件；TC-rich repeats—参与防御和应激反应的顺式作用元件；G-box—参与光反应的顺式作用调控元件；CGTCA-motif—缺氧特异性诱导中的增强子样元件；CAT-box—与分生组织表达有关的顺式作用调控元件；P-box—赤霉素反应元件；GARE-motif—赤霉素反应元件；TGA-element—生长素反应元件；ACE—参与光反应的顺式作用元件；GCN4_motif—参与胚乳表达顺式作用元件；TCA-element—参与水杨酸反应的顺式作用元件；TATC-box—赤霉素反应顺式作用元件；MBS—干旱诱导MYB结合位点；AuxRR-core—参与生长素反应的顺式作用元件；LTR—参与低温响应的顺式作用元件。

Fig. 3 Cis-acting elements analysis of TaIQM genesNote： ABRE—Cis-acting element involved in the abscisic acid responsiveness； ARE—Cis-acting regulatory element essential for the anaerobic induction； Circadian—Cis-acting regulatory element involved in circadian control； G-box—Cis-acting regulatory element involved in light responsiveness； TGACG-motif—Cis-acting regulatory element involved in the MeJA-responsiveness； GT1-motif—Light responsive element； TC-rich repeats—Cis-acting element involved in defense and stress responsiveness； G-box—Cis-acting regulatory element involved in light responsiveness； CGTCA-motif—Cis-acting regulatory element involved in the MeJA-responsiveness； CAT-box—Cis-acting regulatory element related to meristem expression； P-box—Gibberellin-responsive element； GARE-motif—Gibberellin-responsive element； TGA-element—Auxin-responsive element； ACE—Cis-acting element involved in light responsiveness； GCN4_motif—Cis-regulatory element involved in endosperm expression； TCA-element—Cis-acting element involved in salicylic acid responsiveness； TATC-box—Cis-acting element involved in gibberellin-responsiveness； MBS—MYB binding site involved in drought-inducibility； AuxRR-core—Cis-acting regulatory element involved in auxin responsiveness； LTR—Cis-acting element involved in low-temperature responsiveness.

图4 TaIQM基因在不同组织中的表达量热图注：红色表示表达上调，蓝色表示表达下调。Z10—1叶期；Z13—3叶期；Z23—分蘖早期；Z30—起身期；Z32—拔节早期；Z39—拔节晚期；Z65—开花中期；Z71—开花后2 d；Z75—开花后10 d；Z85—开花后30 d。

Fig. 4 Heat map of relative expression level of TaIQM genes in different tissuesNote： Red color indicates up-regulation expression， blue color indicates down-regulation expression. Z10—1 leaf period； Z13—3 leaves stage； Z23—Early tillering； Z30—Standing stage； Z32—Early jointing stage； Z39—Late jointing stage； Z65—Middle flowering； Z71—2 d after flowering； Z75—10 d after flowering； Z85—30 d after flowering.

图5 小麦IQM基因家族在非生物胁迫下的表达分析A：ABA、NaCl、热、冷处理表达量分析； B：干旱处理表达量分析。红色表示表达上调，蓝色表示表达下调

Fig. 5 Expression analysis of IQM genes in wheat under abiotic stressesA： Relative expressionunder ABA， NaCl， heat and clod； B： Relative expression under drought.Red color indicates up-regulation expression， blue color indicates down-regulation expression

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