Identification of HD-Zip Gene Family in Camellia oleifera and Analysis of Its Expression under Abiotic Stress

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

Abstract

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

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.

Figures/Tables 12

References 47

1	Sharif R, Raza A, Chen P, et al. HD-ZIP gene family: potential roles in improving plant growth and regulating stress-responsive mechanisms in plants [J]. Genes, 2021, 12(8): 1256.
2	Ariel FD, Manavella PA, Dezar CA, et al. The true story of the HD-zip family [J]. Trends Plant Sci, 2007, 12(9): 419-426.
3	Palena CM, Tron AE, Bertoncini CW, et al. Positively charged residues at the N-terminal arm of the homeodomain are required for efficient DNA binding by homeodomain-leucine zipper proteins [J]. J Mol Biol, 2001, 308(1): 39-47.
4	Elhiti M, Stasolla C. Structure and function of homodomain-leucine zipper (HD-Zip) proteins [J]. Plant Signal Behav, 2009, 4(2): 86-88.
5	Söderman E, Mattsson J, Engström P. The Arabidopsis homeobox gene ATHB-7 is induced by water deficit and by abscisic acid [J]. Plant J, 1996, 10(2): 375-381.
6	Zhang SX, Haider I, Kohlen W, et al. Function of the HD-Zip I gene Oshox22 in ABA-mediated drought and salt tolerances in rice [J]. Plant Mol Biol, 2012, 80(6): 571-585.
7	王宏, 李刚波, 张大勇, 等. 植物HD-Zip转录因子的生物学功能 [J]. 遗传, 2013, 35(10): 1179-1188.
	Wang H, Li GB, Zhang DY, et al. Biological functions of HD-zip transcription factors [J]. Hereditas, 2013, 35(10): 1179-1188.
8	Ciarbelli AR, Ciolfi A, Salvucci S, et al. The Arabidopsis homeodomain-leucine zipper Ⅱ gene family: diversity and redundancy [J]. Plant Mol Biol, 2008, 68(4-5): 465-478.
9	Turchi L, Baima S, Morelli G, et al. Interplay of HD-Zip Ⅱ and Ⅲ transcription factors in auxin-regulated plant development [J]. J Exp Bot, 2015, 66(16): 5043-5053.
10	Gao J, Chen JY, Feng LY, et al. HD-zip Ⅲ gene family: identification and expression profiles during leaf vein development in soybean [J]. Plants, 2022, 11(13): 1728.
11	Nakamura M, Katsumata H, Abe M, et al. Characterization of the class Ⅳ homeodomain-leucine zipper gene family in Arabidopsis [J]. Plant Physiol, 2006, 141(4): 1363-1375.
12	Xie QM, Gao YN, Li J, et al. The HD-Zip IV transcription factor SlHDZIV8 controls multicellular trichome morphology by regulating the expression of Hairless-2 [J]. J Exp Bot, 2020, 71(22): 7132-7145.
13	谭晓风. 油茶分子育种研究进展 [J]. 中南林业科技大学学报, 2023, 43(1): 1-24.
	Tan XF. Advances in the molecular breeding of Camellia oleifera [J]. J Cent South Univ For Technol, 2023, 43(1): 1-24.
14	董斌, 李荣喜, 洪文泓, 等. 油茶干旱胁迫响应机制的研究进展 [J]. 生物技术通报, 2020, 36(1): 144-149.
	Dong B, Li RX, Hong WH, et al. Research progress on drought stress response mechanism in Camellia oleifera [J]. Biotechnol Bull, 2020, 36(1): 144-149.
15	Zhu HG, Wang FQ, Xu ZP, et al. The complex hexaploid oil-Camellia genome traces back its phylogenomic history and multi-omics analysis of Camellia oil biosynthesis [J]. Plant Biotechnol J, 2024, 22(10): 2890-2906.
16	Wang YX, Liu ZW, Wu ZJ, et al. Transcriptome-wide identification and expression analysis of the NAC gene family in tea plant [Camellia sinensis (L.) O. kuntze] [J]. PLoS One, 2016, 11(11): e0166727.
17	Horton P, Park KJ, Obayashi T, et al. WoLF PSORT: protein localization predictor [J]. Nucleic Acids Res, 2007, 35(Web Server issue): W585-W587.
18	Kumar S, Stecher G, Li M, et al. MEGA X: molecular evolutionary genetics analysis across computing platforms [J]. Mol Biol Evol, 2018, 35(6): 1547-1549.
19	Letunic I, Bork P. Interactive tree of life (iTOL) v5: an online tool for phylogenetic tree display and annotation [J]. Nucleic Acids Res, 2021, 49(W1): W293-W296.
20	Bailey TL, Boden M, Buske FA, et al. MEME SUITE: tools for motif discovery and searching [J]. Nucleic Acids Res, 2009, 37(Web Server issue): W202-W208.
21	Chen CJ, Wu Y, Li JW, et al. TBtools-Ⅱ: a “one for all, all for one” bioinformatics platform for biological big-data mining [J]. Mol Plant, 2023, 16(11): 1733-1742.
22	Lescot M, Déhais P, Thijs G, et al. PlantCARE, a database of plant Cis-acting regulatory elements and a portal to tools for in silico analysis of promoter sequences [J]. Nucleic Acids Res, 2002, 30(1): 325-327.
23	Gong WF, Song QL, Ji K, et al. Full-length transcriptome from Camellia oleifera seed provides insight into the transcript variants involved in oil biosynthesis [J]. J Agric Food Chem, 2020, 68(49): 14670-14683.
24	Maren NA, Duduit JR, Huang DB, et al. Stepwise optimization of real-time RT-PCR analysis [J]. Methods Mol Biol, 2023, 2653: 317-332.
25	Zhao Y, Ma Q, Jin XL, et al. A novel maize homeodomain-leucine zipper (HD-Zip) I gene, Zmhdz10, positively regulates drought and salt tolerance in both rice and Arabidopsis [J]. Plant Cell Physiol, 2014, 55(6): 1142-1156.
26	Ingram J, Bartels D. The molecular basis of dehydration tolerance in plants [J]. Annu Rev Plant Physiol Plant Mol Biol, 1996, 47: 377-403.
27	Yamaguchi-Shinozaki K, Shinozaki K. Transcriptional regulatory networks in cellular responses and tolerance to dehydration and cold stresses [J]. Annu Rev Plant Biol, 2006, 57: 781-803.
28	Kopecká R, Kameniarová M, Černý M, et al. Abiotic stress in crop production [J]. Int J Mol Sci, 2023, 24(7): 6603.
29	Ahmad S, Chen Y, Shah AZ, et al. The Homeodomain-Leucine zipper genes family regulates the jinggangmycin mediated immune response of Oryza sativa to Nilaparvata lugens, and Laodelphax striatellus [J]. Bioengineering, 2022, 9(8): 398.
30	Sharif R, Xie C, Wang J, et al. Genome wide identification, characterization and expression analysis of HD-ZIP gene family in Cucumis sativus L. under biotic and various abiotic stresses [J]. Int J Biol Macromol, 2020: S0141-8130(20)32981-0.
31	Liu K, Han XL, Liang ZL, et al. Genome-wide identification, classification, and expression analysis of the HD-zip transcription factor family in apple (Malus domestica borkh.) [J]. Int J Mol Sci, 2022, 23(5): 2632.
32	Yang QY, Xiang WB, Li ZH, et al. Genome-wide characterization and expression analysis of HD-ZIP gene family in Dendrobium officinale [J]. Front Genet, 2022, 13: 797014.
33	Li Z, Gao ZQ, Li RH, et al. Genome-wide identification and expression profiling of HD-ZIP gene family in Medicago truncatula [J]. Genomics, 2020, 112(5): 3624-3635.
34	Bowers JE, Chapman BA, Rong JK, et al. Unravelling angiosperm genome evolution by phylogenetic analysis of chromosomal duplication events [J]. Nature, 2003, 422: 433-438.
35	Zhang KM, Lan YG, Shi YN, et al. Systematic analysis and functional characterization of the PLATZ transcription factors in moso bamboo (Phyllostachys edulis) [J]. J Plant Growth Regul, 2023, 42(1): 218-236.
36	周佩娜, 党静洁, 邵永芳, 等. 荆芥HD-Zip基因家族的全基因组鉴定及分析 [J]. 浙江农林大学学报, 2023, 40(1): 12-21.
	Zhou PN, Dang JJ, Shao YF, et al. Genome-wide identification and expression analysis of HD-Zip gene family in Schizonepeta tenuifolia [J]. J Zhejiang A F Univ, 2023, 40(1): 12-21.
37	祁存英, 李媛, 杨成兰, 等. 青稞HD-Zip基因家族鉴定及其在非生物胁迫下的表达特性 [J]. 西北植物学报, 2023, 43(1): 66-78.
	Qi CY, Li Y, Yang CL, et al. Identification of HD-zip gene family of Tibetan hulless barley and its expression characteristics under abiotic stress [J]. Acta Bot Boreali Occidentalia Sin, 2023, 43(1): 66-78.
38	Zhao Y, Zhou YQ, Jiang HY, et al. Systematic analysis of sequences and expression patterns of drought-responsive members of the HD-Zip gene family in maize [J]. PLoS One, 2011, 6(12): e28488.
39	Hu X, Liang JX, Wang WJ, et al. Comprehensive genome-wide analysis of the DREB gene family in Moso bamboo (Phyllostachys edulis): evidence for the role of PeDREB28 in plant abiotic stress response [J]. Plant J, 2023, 116(5): 1248-1270.
40	叶泗洪, 季美君, 陈奇, 等. 陆地棉HD-ZIP_N基因家族的全基因组分析 [J]. 农业生物技术学报, 2023, 31(12): 2454-2465.
	Ye SH, Ji MJ, Chen Q, et al. Genome-wide analysis of the HD-ZIP_N gene family in Gossypium hirsutum [J]. J Agric Biotechnol, 2023, 31(12): 2454-2465.
41	刘梦梦, 刘晓, 尤倩, 等. 半夏HD-Zip基因家族全基因组鉴定及表达分析 [J]. 农业生物技术学报, 2024, 32(11): 145-152.
	Liu MM, Liu X, You Q, et al. Genome wide identification and expression analysis of the HD-zip gene family in Pinellia ternata [J]. J Agric Biotechnol, 2024, 32(11): 145-152.
42	邵晨冰, 黄志楠, 白雪滢, 等. 辣椒HD-Zip基因家族鉴定、系统进化及表达分析 [J]. 中国农业科学, 2020, 53(5): 1004-1017.
	Shao CB, Huang ZN, Bai XY, et al. Identification, systematic evolution and expression analysis of HD-zip gene family in Capsicum annuum [J]. Sci Agric Sin, 2020, 53(5): 1004-1017.
43	Zhang XX, Chen ZX, Wang CN, et al. Genome-wide identification of HD-ZIP gene family and screening of genes related to prickle development in Zanthoxylum armatum [J]. Plant Genome, 2023, 16(1): e20295.
44	Wang Z, Wu XL, Zhang BB, et al. Genome-wide identification, bioinformatics and expression analysis of HD-Zip gene family in peach [J]. BMC Plant Biol, 2023, 23(1): 122.
45	Wang YJ, Wang HJ, Yu C, et al. Comprehensive bioinformation analysis of homeodomain-leucine zipper gene family and expression pattern of HD-Zip I under abiotic stress in Salix suchowensis [J]. BMC Genomics, 2024, 25(1): 182.
46	Xu YQ, Liu CL. Genome-wide identification and analysis of the HD-Zip transcription factor family in oats [J]. Front Mol Biosci, 2024, 11: 1475276.
47	Li SM, Chen N, Li FF, et al. Characterization of wheat homeodomain-leucine zipper family genes and functional analysis of TaHDZ5-6A in drought tolerance in transgenic Arabidopsis [J]. BMC Plant Biol, 2020, 20(1): 50.

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

[1]	LIU Yuan, ZHAO Ran, LU Zhen-fang, LI Rui-li. Research Progress in the Biological Metabolic Pathway and Functions of Plant Carotenoids [J]. Biotechnology Bulletin, 2025, 41(5): 23-31.
[2]	PENG Shao-zhi, WANG Deng-ke, ZHANG Xiang, DAI Xiong-ze, XU Hao, ZOU Xue-xiao. Cloning， Expression Characteristics and Functional Verification of the Pepper CaFD1 Gene [J]. Biotechnology Bulletin, 2025, 41(5): 153-164.
[3]	ZHANG Yi-xuan, MA Yu, WANG Tong-tong, SHENG Su-ao, SONG Jia-feng, LYU Zhao-yan, ZHU Xiao-biao, HOU Hua-lan. Genome-wide Identification and Expression Profiles of DIR Gene Family in Potato [J]. Biotechnology Bulletin, 2025, 41(3): 123-136.
[4]	WANG Chen, LIU Guo-mei, CHEN Chang, ZHANG Jin-long, YAO Lin, SUN Xuan, DU Chun-fang. Genome-wide Identification and Expression Analysis of CCDs Family in Brassia rapa L. [J]. Biotechnology Bulletin, 2025, 41(3): 161-170.
[5]	HAN Jiang-tao, ZHANG Shuai-bo, QIN Ya-rui, HAN Shuo-yang, ZHANG Ya-kang, WANG Ji-qing, DU Qing-jie, XIAO Huai-juan, LI Meng. Identification of β‍-amylase Gene Family in Melon and Their Response to Abiotic Stresses [J]. Biotechnology Bulletin, 2025, 41(3): 171-180.
[6]	KUANG Jian-hua, CHENG Zhi-peng, ZHAO Yong-jing, YANG Jie, CHEN Run-qiao, CHEN Long-qing, HU Hui-zhen. Expression Analysis of the GH3 Gene Family in Nelumbo nucifera underHormonal and Abiotic Stresses [J]. Biotechnology Bulletin, 2025, 41(2): 221-233.
[7]	GE Shi-jie, LIU Yi-de, ZHANG Hua-dong, NING Qiang, ZHU Zhan-wang, WANG Shu-ping, LIU Yi-ke. Identification and Expression Analysis of Protein Disulfide Isomerase Gene Family in Wheat [J]. Biotechnology Bulletin, 2025, 41(2): 85-96.
[8]	YIN Yuan, CHENG Shuang, LIU Ding-hao, DENG Xiao-xia, LI Kai-yue, WANG Jing-hong, LIN Ji-xiang. Research Progress in Exogenous Hydrogen Peroxide（H₂O₂）Affecting Plant Growth and Physiological Metabolism under Abiotic Stress [J]. Biotechnology Bulletin, 2025, 41(1): 1-13.
[9]	WU Zhi-jian, LIU Guang-yang, LIN Zhi-hao, SHENG Bin, CHEN Ge, XU Xiao-min, WANG Jun-wei, XU Dong-hui. Research Progress of Nano-regulation of Vegetable Seed Germination and Its Mechanism [J]. Biotechnology Bulletin, 2025, 41(1): 14-24.
[10]	LI Yu-xin, LI Miao, DU Xiao-fen, HAN Kang-ni, LIAN Shi-chao, WANG Jun. Identification and Expression Analysis of SiSAP Gene Family in Foxtail Millet（Setaria italica） [J]. Biotechnology Bulletin, 2025, 41(1): 143-156.
[11]	SHEN Peng, GAO Ya-Bin, DING Hong. Identification and Expression Analysis of SAT Gene Family in Potato（Solanum tuberosum L.） [J]. Biotechnology Bulletin, 2024, 40(9): 64-73.
[12]	MAN Quan-cai, MENG Zi-nuo, LI Wei, CAI Xin-ru, SU Run-dong, FU Chang-qing, GAO Shun-juan, CUI Jiang-hui. Identification and Expression Analysis of AQP Gene Family in Potato [J]. Biotechnology Bulletin, 2024, 40(9): 51-63.
[13]	LI Yong-hui, BAO Xing-xing, DUAN Yi-ke, ZHAO Yun-xia, YU Xiang-li, CHEN Yao, ZHANG Yan-zhao. Genome-wide Identification and Expression Features Analysis of the bZIP Family in Rhododendron henanense subsp. lingbaoense [J]. Biotechnology Bulletin, 2024, 40(8): 186-198.
[14]	CUI Yuan-yuan, WANG Zhao-yi, BAI Shuang-yu, REN Yu-zhao, DOU Fei-fei, LIU Cai-xia, LIU Feng-lou, WANG Zhang-jun, LI Qing-feng. Genome-wide Identification of Non-specific Phospholipase C Gene Family in Hordeum vulgare L. and Stress Expression Analysis at Seedling Stage [J]. Biotechnology Bulletin, 2024, 40(8): 74-82.
[15]	LIU Dan-dan, WANG Lei-gang, SUN Ming-hui, JIAO Xiao-yu, WU Qiong, WANG Wen-jie. Genome-wide Identification and Expression Pattern Profiling of the Trehalose-6-phosphate Synthase（TPS）Gene Family in Tea Plant（Camellia sinensis） [J]. Biotechnology Bulletin, 2024, 40(8): 152-163.