油茶HD-Zip基因家族鉴定及其在非生物胁迫下的表达分析

doi:10.13560/j.cnki.biotech.bull.1985.2024-1146

摘要/Abstract

摘要：

目的鉴定油茶HD-Zip基因家族成员，分析其结构、特性和功能，解析其在盐、干旱、低温和高温等非生物胁迫下的表达模式。方法基于六倍体普通油茶全基因组数据，利用生物信息学方法筛选鉴定CoHDZs基因家族成员，对其基因结构、保守基序、顺式作用元件、染色体定位和基因共线性等进行分析；基于转录组数据分析CoHDZs基因在油茶叶、芽、花瓣、雌蕊、种子、花萼、雄蕊和果实不同发育期中的表达模式；利用实时荧光定量PCR方法解析CoHDZs在不同逆境胁迫下的瞬时表达特征。结果 CoHDZs基因家族包含46个成员（CoHDZ1-CoHDZ46）分布在4个不同的亚家族（HD-Zip Ⅰ，HD-Zip Ⅱ，HD-Zip Ⅲ和HD-Zip Ⅳ）中，且同一亚家族中的CoHDZ基因具有相似的结构特征。顺式作用元件分析表明油茶HD-Zip基因家族成员广泛参与光响应、激素响应以及非生物胁迫响应。染色体定位和共线性分析可知，46个CoHDZs基因不均匀地分布在26条染色体和2条染色体骨架上，全基因组复制（WGD）事件或片段复制可能是CoHDZs基因进化的主要动力；转录组分析表明，CoHDZs基因存在明显的组织特异性表达；实时荧光定量PCR验证实验表明，油茶HD-Zip基因家族成员在盐、干旱、低温和高温胁迫下高度表达，不同CoHDZs基因在同一胁迫下的表达存在差异，且相同基因在不同胁迫下的表达情况也有不同，其中CoHDZ1、CoHDZ3、CoHDZ14、CoHDZ18、CoHDZ20、CoHDZ22、CoHDZ42、CoHDZ44和CoHDZ45这9个基因是油茶HD-Zip家族响应非生物胁迫的关键基因。结论 CoHDZs基因广泛参与油茶的生长发育、激素信号传导和非生物胁迫响应并发挥着重要的调控作用。

关键词: 油茶, HD-Zip基因家族, 组织特异性, 非生物胁迫

Abstract:

Objective To identify the HD-Zip gene family members in Camellia oleifera, analyze their structure, characteristics and functions, as well as examine their expression patterns under abiotic stress,including salt, drought, low temperature and high temperature. Method Based on the whole genome data of hexaploid common C. oleifera, members of CoHDZs gene family were screened and identified by bioinformatics method. The analysis included an examination of gene structure, conserved motifs, cis-acting elements, chromosomal locations, and gene colinearity. Additionally, expression patterns of CoHDZ genes were analyzed during various developmental stages of C. oleifera, including leaves, buds, petals, pistils, seeds, calyxes, stamens, and fruits. Furthermore, real-time fluorescence quantitative PCR was utilized to investigate the transient expression characteristics of CoHDZs under different stress conditions. Result The CoHDZ gene family comprises 46 members (CoHDZ1-CoHDZ46) that are categorized into four distinct subfamilies: HD-Zip Ⅰ, HD-Zip Ⅱ, HD-Zip Ⅲ, and HD-Zip Ⅳ. Members of the CoHDZ genes within the same subfamily have similar structural characteristics. Analysis of cis-acting elements indicated that members of the C. oleifera HD-Zip gene family are significantly involved in light response, hormone response, and abiotic stress response. Chromosomal localization and collinearity analysis revealed that the 46 CoHDZ genes are unevenly distributed across 26 chromosomes and 2 chromosome scaffolds. Whole genome duplication (WGD) events or segmental duplications are likely the primary drivers of the evolution of CoHDZ genes. Transcriptome analysis demonstrated that CoHDZ genes have pronounced tissue-specific expression. Real-time fluorescence quantitative PCR experiments confirmed that members of the C. oleifera HD-Zip gene family are highly expressed under conditions of salt, drought, low temperature, and high temperature stress. Furthermore, the expressions of different CoHDZ genes vary under the same stress conditions, and the expression of the same gene also differs across various stressors. Notably, CoHDZ1, CoHDZ3, CoHDZ14, CoHDZ18, CoHDZ20, CoHDZ22, CoHDZ42, CoHDZ44, and CoHDZ45 are identified as key genes in the C. oleifera HD-Zip family responding to abiotic stress. Conclusion CoHDZs genes are widely involved in the growth and development, hormone signaling and abiotic stress response of C. oleifera, and play an important regulatory role.

Key words: Camellia oleifera, HD-Zip gene family, tissue specificity, abiotic stress

黄丹, 彭兵阳, 张盼盼, 焦悦, 吕佳斌. 油茶HD-Zip基因家族鉴定及其在非生物胁迫下的表达分析[J]. 生物技术通报, 2025, 41(6): 191-207.

HUANG Dan, PENG Bing-yang, ZHANG Pan-pan, JIAO Yue, LYU Jia-bin. Identification of HD-Zip Gene Family in Camellia oleifera and Analysis of Its Expression under Abiotic Stress[J]. Biotechnology Bulletin, 2025, 41(6): 191-207.

图/表 12

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基因名称 Gene name	上游引物 Forward primer（5'-3'）	下游引物 Reverse primer（5'-3'）
CoHDZ1	CATCACCATCACCACCACCAGTTC	TGATACGCCTTCTGTGCATCCAAC
CoHDZ3	CCACCACCACTACCACCACCTC	TCGTCCACCACCGCACTTCC
CoHDZ4	CATCACCATCACCACCACCAGTTC	TGATACGCCTTCTGTGCATCCAAC
CoHDZ6	CCACCACCACTACCACCACCTC	TCGTCCACCACCGCACTTCC
CoHDZ7	CATCACCATCACCACCACCAGTTC	TGATACGCCTTCTGTGCATCCAAC
CoHDZ14	TCTCTGTTCCTCTCTGGCTCTTCC	AATTGGACTTGGTCAGGCGTAAGC
CoHDZ18	TCTCTGTTCCTCTCTGGCTCTTCC	AATTGGACTTGGTCAGGCGTAAGC
CoHDZ20	AAGAGGGAGAGGGAGAGGGAGATC	GGAGAAGGTGACGGAGGAGGAG
CoHDZ21	TCCGCCGATGCCTCCATAGTG	AGCCGTAGCAGCAACCTCTCC
CoHDZ22	CAGAACCACAGGTGCCTTGAGAAG	GCTTGTGTCGGTAGTAGCTGGAG
CoHDZ23	AGCAGCGAGAGGAAGAGGAGATTG	CGGCGATTTTCACCTTTTGCTCTG
CoHDZ26	ACGGTGGAGGAGGAGGAGGAG	GGAACCAAACAGCAACTTGACGAG
CoHDZ27	GGAGGAGGAGGAGGGAATGAGAAG	CGCTTGAGCTGCTTAGCCTTCC
CoHDZ30	CCCTCCAAGCTCAGAACCACAAAC	TTCACCGATCCGCCCTCTGTC
CoHDZ42	AGAACTTGTTGCCTCCTCCTCCTC	ACCCAAAGGCGTGATCTTCAACTG
CoHDZ44	GAAGAGGCGAGTGAGCGTTGC	GTTCCTGAGCGAGCTTCACTTTCC
CoHDZ45	CCACCACCACTACCACCACCTC	TCGTCCACCACCGCACTTCC
GAPDH	CTGCTGCCCACTTGAAGG	GACCCTCAACAATGCCAAAC

基因名称 Gene name	上游引物 Forward primer（5'-3'）	下游引物 Reverse primer（5'-3'）
CoHDZ1	CATCACCATCACCACCACCAGTTC	TGATACGCCTTCTGTGCATCCAAC
CoHDZ3	CCACCACCACTACCACCACCTC	TCGTCCACCACCGCACTTCC
CoHDZ4	CATCACCATCACCACCACCAGTTC	TGATACGCCTTCTGTGCATCCAAC
CoHDZ6	CCACCACCACTACCACCACCTC	TCGTCCACCACCGCACTTCC
CoHDZ7	CATCACCATCACCACCACCAGTTC	TGATACGCCTTCTGTGCATCCAAC
CoHDZ14	TCTCTGTTCCTCTCTGGCTCTTCC	AATTGGACTTGGTCAGGCGTAAGC
CoHDZ18	TCTCTGTTCCTCTCTGGCTCTTCC	AATTGGACTTGGTCAGGCGTAAGC
CoHDZ20	AAGAGGGAGAGGGAGAGGGAGATC	GGAGAAGGTGACGGAGGAGGAG
CoHDZ21	TCCGCCGATGCCTCCATAGTG	AGCCGTAGCAGCAACCTCTCC
CoHDZ22	CAGAACCACAGGTGCCTTGAGAAG	GCTTGTGTCGGTAGTAGCTGGAG
CoHDZ23	AGCAGCGAGAGGAAGAGGAGATTG	CGGCGATTTTCACCTTTTGCTCTG
CoHDZ26	ACGGTGGAGGAGGAGGAGGAG	GGAACCAAACAGCAACTTGACGAG
CoHDZ27	GGAGGAGGAGGAGGGAATGAGAAG	CGCTTGAGCTGCTTAGCCTTCC
CoHDZ30	CCCTCCAAGCTCAGAACCACAAAC	TTCACCGATCCGCCCTCTGTC
CoHDZ42	AGAACTTGTTGCCTCCTCCTCCTC	ACCCAAAGGCGTGATCTTCAACTG
CoHDZ44	GAAGAGGCGAGTGAGCGTTGC	GTTCCTGAGCGAGCTTCACTTTCC
CoHDZ45	CCACCACCACTACCACCACCTC	TCGTCCACCACCGCACTTCC
GAPDH	CTGCTGCCCACTTGAAGG	GACCCTCAACAATGCCAAAC

基因名称 Gene name	基因ID Gene ID	亚家族 Group	染色体号 Chr	蛋白质长度 Protein length （aa）	分子量 MW （Da）	等电点 PI	亚细胞定位 Subcellular localization
CoHDZ1	YCChr1a_018730.1	Ⅰ	Chr1a	320	35142.84	5.7	细胞核
CoHDZ2	YCChr1a_024300.1	Ⅱ	Chr1a	225	25151.16	5.24	细胞核
CoHDZ3	YCChr1a_028340.1	Ⅰ	Chr1a	282	31781.2	4.65	细胞核
CoHDZ4	YCChr1b_015690.1	Ⅰ	Chr1b	233	25128.64	5.34	细胞核
CoHDZ5	YCChr1b_022120.1	Ⅱ	Chr1b	191	21187.5	5.66	细胞核
CoHDZ6	YCChr1b_027440.1	Ⅰ	Chr1b	269	30716.12	4.81	细胞核
CoHDZ7	YCChr1c_029510.1	Ⅰ	Chr1c	321	35324.02	5.75	细胞核
CoHDZ8	YCChr1c_033970.1	Ⅱ	Chr1c	229	25567.69	5.24	细胞核
CoHDZ9	YCChr2a_005360.1	Ⅱ	Chr2a	190	21142.64	6.8	细胞核
CoHDZ10	YCChr2b_004640.1	Ⅱ	Chr2b	186	20523.57	6.05	细胞核
CoHDZ11	YCChr2b_005710.1	Ⅱ	Chr2b	307	33107.72	7.76	细胞核
CoHDZ12	YCChr2c_005960.1	Ⅱ	Chr2c	179	19654.72	5.67	细胞核
CoHDZ13	YCChr2c_008630.1	Ⅱ	Chr2c	377	41108.66	7.56	细胞核
CoHDZ14	YCChr4b_013830.1	Ⅰ	Chr4b	328	36236.27	6.17	细胞核
CoHDZ15	YCChr4b_019640.1	Ⅳ	Chr4b	601	66874.14	6.33	细胞核
CoHDZ16	YCChr4b_019700.1	Ⅲ	Chr4b	778	85841.14	5.93	细胞核
CoHDZ17	YCChr4b_025610.1	Ⅱ	Chr4b	162	18288.36	5.54	细胞核
CoHDZ18	YCChr4c_015970.1	Ⅰ	Chr4c	225	29217.36	4.81	细胞核
CoHDZ19	YCChr4c_027810.1	Ⅱ	Chr4c	166	18786.98	6.19	细胞核
CoHDZ20	YCChr4c_030590.1	Ⅰ	Chr4c	279	30935.94	6.81	细胞核
CoHDZ21	YCChr5a_011690.1	Ⅰ	Chr5a	208	23163.6	4.8	叶绿体、细胞核
CoHDZ22	YCChr7a_015220.1	Ⅰ	Chr7a	106	12381.09	8.47	细胞核
CoHDZ23	YCChr8c_012310.1	Ⅰ	Chr8c	177	20756.37	5.48	细胞核
CoHDZ24	YCChr9a_004470.1	Ⅳ	Chr9a	587	65387.15	6.14	细胞核
CoHDZ25	YCChr9a_017880.1	Ⅳ	Chr9a	688	75790.99	5.47	细胞核
CoHDZ26	YCChr10a_007180.1	Ⅰ	Chr10a	126	14257.15	9.55	细胞核
CoHDZ27	YCChr10b_009490.1	Ⅰ	Chr10b	123	14239.31	9.96	细胞核
CoHDZ28	YCChr11a_017100.1	Ⅲ	Chr11a	812	88970.68	6.31	细胞核
CoHDZ29	YCChr11a_020800.1	Ⅱ	Chr11a	308	34896.48	7.62	细胞核
CoHDZ30	YCChr11b_008940.1	Ⅰ	Chr11b	189	21602.32	6.23	细胞核
CoHDZ31	YCChr11b_015650.1	Ⅱ	Chr11b	527	60449.42	6.59	细胞核
CoHDZ32	YCChr11c_017250.1	Ⅱ	Chr11c	281	31909.08	8.16	细胞核
CoHDZ33	YCChr12a_013940.1	Ⅲ	Chr12a	132	15639.05	9.81	细胞核
CoHDZ34	YCChr12b_016460.1	Ⅲ	Chr12b	882	97008.44	6.11	细胞核
CoHDZ35	YCChr12c_014450.1	Ⅲ	Chr12c	718	78967.97	5.7	细胞核
CoHDZ36	YCChr13a_010080.1	Ⅱ	Chr13a	251	28136.51	8.35	细胞核
CoHDZ37	YCChr13a_014620.1	Ⅳ	Chr13a	507	56653.47	5.4	细胞核
CoHDZ38	YCChr13b_013930.1	Ⅳ	Chr13b	653	74082.16	5.96	细胞核
CoHDZ39	YCChr13c_011380.1	Ⅱ	Chr13c	305	34301.66	8.41	细胞核
CoHDZ40	YCChr13c_013630.1	Ⅳ	Chr13c	653	74062.17	5.96	细胞核
CoHDZ41	YCChr13c_016960.1	Ⅳ	Chr13c	481	53576	5.32	细胞核
CoHDZ42	YCChr14c_017780.1	Ⅰ	Chr14c	279	31582.16	5.1	细胞核
CoHDZ43	YCChr15a_016280.1	Ⅲ	Chr15a	707	77241.76	6.1	细胞核
CoHDZ44	YCChr15c_001540.1	Ⅰ	Chr15c	108	12482.2	6.4	细胞核
CoHDZ45	YCScaffold_003590.1	Ⅰ	Scaffold8	282	31780.11	4.65	细胞核
CoHDZ46	YCScaffold_018850.1	Ⅲ	Scaffold184	777	85065.99	5.78	细胞核

基因名称 Gene name	基因ID Gene ID	亚家族 Group	染色体号 Chr	蛋白质长度 Protein length （aa）	分子量 MW （Da）	等电点 PI	亚细胞定位 Subcellular localization
CoHDZ1	YCChr1a_018730.1	Ⅰ	Chr1a	320	35142.84	5.7	细胞核
CoHDZ2	YCChr1a_024300.1	Ⅱ	Chr1a	225	25151.16	5.24	细胞核
CoHDZ3	YCChr1a_028340.1	Ⅰ	Chr1a	282	31781.2	4.65	细胞核
CoHDZ4	YCChr1b_015690.1	Ⅰ	Chr1b	233	25128.64	5.34	细胞核
CoHDZ5	YCChr1b_022120.1	Ⅱ	Chr1b	191	21187.5	5.66	细胞核
CoHDZ6	YCChr1b_027440.1	Ⅰ	Chr1b	269	30716.12	4.81	细胞核
CoHDZ7	YCChr1c_029510.1	Ⅰ	Chr1c	321	35324.02	5.75	细胞核
CoHDZ8	YCChr1c_033970.1	Ⅱ	Chr1c	229	25567.69	5.24	细胞核
CoHDZ9	YCChr2a_005360.1	Ⅱ	Chr2a	190	21142.64	6.8	细胞核
CoHDZ10	YCChr2b_004640.1	Ⅱ	Chr2b	186	20523.57	6.05	细胞核
CoHDZ11	YCChr2b_005710.1	Ⅱ	Chr2b	307	33107.72	7.76	细胞核
CoHDZ12	YCChr2c_005960.1	Ⅱ	Chr2c	179	19654.72	5.67	细胞核
CoHDZ13	YCChr2c_008630.1	Ⅱ	Chr2c	377	41108.66	7.56	细胞核
CoHDZ14	YCChr4b_013830.1	Ⅰ	Chr4b	328	36236.27	6.17	细胞核
CoHDZ15	YCChr4b_019640.1	Ⅳ	Chr4b	601	66874.14	6.33	细胞核
CoHDZ16	YCChr4b_019700.1	Ⅲ	Chr4b	778	85841.14	5.93	细胞核
CoHDZ17	YCChr4b_025610.1	Ⅱ	Chr4b	162	18288.36	5.54	细胞核
CoHDZ18	YCChr4c_015970.1	Ⅰ	Chr4c	225	29217.36	4.81	细胞核
CoHDZ19	YCChr4c_027810.1	Ⅱ	Chr4c	166	18786.98	6.19	细胞核
CoHDZ20	YCChr4c_030590.1	Ⅰ	Chr4c	279	30935.94	6.81	细胞核
CoHDZ21	YCChr5a_011690.1	Ⅰ	Chr5a	208	23163.6	4.8	叶绿体、细胞核
CoHDZ22	YCChr7a_015220.1	Ⅰ	Chr7a	106	12381.09	8.47	细胞核
CoHDZ23	YCChr8c_012310.1	Ⅰ	Chr8c	177	20756.37	5.48	细胞核
CoHDZ24	YCChr9a_004470.1	Ⅳ	Chr9a	587	65387.15	6.14	细胞核
CoHDZ25	YCChr9a_017880.1	Ⅳ	Chr9a	688	75790.99	5.47	细胞核
CoHDZ26	YCChr10a_007180.1	Ⅰ	Chr10a	126	14257.15	9.55	细胞核
CoHDZ27	YCChr10b_009490.1	Ⅰ	Chr10b	123	14239.31	9.96	细胞核
CoHDZ28	YCChr11a_017100.1	Ⅲ	Chr11a	812	88970.68	6.31	细胞核
CoHDZ29	YCChr11a_020800.1	Ⅱ	Chr11a	308	34896.48	7.62	细胞核
CoHDZ30	YCChr11b_008940.1	Ⅰ	Chr11b	189	21602.32	6.23	细胞核
CoHDZ31	YCChr11b_015650.1	Ⅱ	Chr11b	527	60449.42	6.59	细胞核
CoHDZ32	YCChr11c_017250.1	Ⅱ	Chr11c	281	31909.08	8.16	细胞核
CoHDZ33	YCChr12a_013940.1	Ⅲ	Chr12a	132	15639.05	9.81	细胞核
CoHDZ34	YCChr12b_016460.1	Ⅲ	Chr12b	882	97008.44	6.11	细胞核
CoHDZ35	YCChr12c_014450.1	Ⅲ	Chr12c	718	78967.97	5.7	细胞核
CoHDZ36	YCChr13a_010080.1	Ⅱ	Chr13a	251	28136.51	8.35	细胞核
CoHDZ37	YCChr13a_014620.1	Ⅳ	Chr13a	507	56653.47	5.4	细胞核
CoHDZ38	YCChr13b_013930.1	Ⅳ	Chr13b	653	74082.16	5.96	细胞核
CoHDZ39	YCChr13c_011380.1	Ⅱ	Chr13c	305	34301.66	8.41	细胞核
CoHDZ40	YCChr13c_013630.1	Ⅳ	Chr13c	653	74062.17	5.96	细胞核
CoHDZ41	YCChr13c_016960.1	Ⅳ	Chr13c	481	53576	5.32	细胞核
CoHDZ42	YCChr14c_017780.1	Ⅰ	Chr14c	279	31582.16	5.1	细胞核
CoHDZ43	YCChr15a_016280.1	Ⅲ	Chr15a	707	77241.76	6.1	细胞核
CoHDZ44	YCChr15c_001540.1	Ⅰ	Chr15c	108	12482.2	6.4	细胞核
CoHDZ45	YCScaffold_003590.1	Ⅰ	Scaffold8	282	31780.11	4.65	细胞核
CoHDZ46	YCScaffold_018850.1	Ⅲ	Scaffold184	777	85065.99	5.78	细胞核

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