| 狗头枣果皮GH9基因家族鉴定及不同发育期的表达与纤维素含量分析 |
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| 引用本文：王真珍，陈国梁，张亦弛，马瑞峰，张向前，李 爽.狗头枣果皮GH9基因家族鉴定及不同发育期的表达与纤维素含量分析[J].西北植物学报,2023,43(4):539~549 |
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| 基金项目:国家自然科学基金地区科学基金(31860535)；陕西省自然科学基础研究计划(2020JM-553)；陕西省红枣重点实验室科技计划(Sxhzdsys-zj21-03)；陕西省省级大学生创新创业训练计划(S202210719110)；延安大学校级大学生创新创业项目(D2021097)；延安大学博士科研启动项目(YDBK2018-45) |
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| 中文摘要:为了明确糖苷水解酶9 (glycoside hydrolase 9，GH9) 基因家族在枣果皮发育过程中的作用，该研究以狗头枣果实为材料，对其幼果期(StageⅠ)、青果期(StageⅡ)、膨大期(StageⅢ)、白熟期(StageⅣ)、半红期(StageⅤ)、全红期(StageⅥ)6个不同发育阶段的果皮纤维素含量、细胞组织结构、GH9基因家族的生物信息学以及基因表达进行分析。结果表明：(1)在枣果实发育过程中，果皮纤维素含量在StageⅠ~StageⅢ略有增加，于StageⅢ达到最高(217.33 mg/g)，随后快速显著下降并于StageⅥ降至最低(98.51 mg/g)，较StageⅢ下降了54.7%。(2)在枣果皮发育过程中果皮细胞在未着色时期紧密排列，在着色后明显皱缩且排列松散；番红-固绿染色后，从StageⅠ到Stage Ⅳ枣果皮角质层细胞和表皮细胞的细胞壁颜色由蓝绿色逐渐变为浅蓝进而转为粉色，胞内几乎没有颜色，说明狗头枣果皮细胞壁的纤维素含量从StageⅠ到StageⅣ逐渐减少。(3)枣果实转录组测序发现，与StageⅠ相比，StageⅡ~StageⅥ分别检测到1 090、2 039、3 300、4 594和7 855个差异表达基因；GO富集分析表明，在枣果实发育初期差异表达基因富集在与纤维素降解密切相关的水解酶活性分子功能中。(4)从枣基因组中鉴定到30个GH9基因家族成员，根据系统发育关系分为GH9A、GH9B和GH9C共3个亚族，其中GH9A亚族最大共有19个成员。(5)基因表达模式分析显示，所鉴定的30个GH9基因中有17个基因在果皮发育的StageⅠ~StageⅢ均有表达且表达量高，而在StageⅣ~StageⅥ表达量低；荧光定量PCR验证结果显示，8个差异表达基因的表达量均随枣果实发育进程逐渐快速降低，其变化趋势与枣果皮纤维素含量变化趋势一致，也与转录组测序结果的表达水平的变化趋势一致。研究推测，GH9基因家族在枣果皮发育过程中具有降解植物细胞壁的作用。 |
| 中文关键词:狗头枣 果皮 纤维素 组织结构 GH9基因家族 转录组测序 |
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| Identification of the GH9 Gene Family and Expression in Different Development Stages and Cellulose Content Analysis in Peel of Ziziphus jujuba ‘Goutouzao’ |
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| Abstract:In order to explore the role of glycoside hydrolase 9 gene family (glycoside hydrolase 9, GH9) in the development of jujube peel, this study used ‘Goutouzao’ as material, examined the cellulose content, histological structure, and bioinformatics of GH9 gene family and gene expression in peel of ‘Goutouzao’ at different developmental stages, including 6 developmental stages: young fruit stage (StageⅠ), green fruit stage (StageⅡ), enlarged stage (Stage Ⅲ), white ripening stage (StageⅣ), semi-red stage (StageⅤ), and full-red stage (StageⅥ). The results showed that: (1) during the development of jujube fruit, the cellulose content in peel of jujube increased slightly from StageⅠ to Stage Ⅲ, reached the highest level at Stage Ⅲ (217.33 mg/g), then decreased rapidly and significantly and reached the lowest level at StageⅥ (98.51 mg/g), which was 54.7% comparing with Stage Ⅲ. (2) The histological structure of peel of jujube were closely arranged in un-coloring stages, and obviously shrunk and arranged loosely after coloring stages during the fruit development process; After staining with saffron and solid green, the color of keratinocytes and epidermal in peel of jujube from StageⅠ to StageⅣ changed from blue-green to light blue and then to pink gradually, and there were almost no color in the cells, which indicated the cellulose content decreased gradually in the cell wall of the peel of jujube ‘Goutouzao’ from StageⅠ to StageⅣ. (3) The transcriptome sequencing analysis showed 1 090, 2 039, 3 300, 4 594 and 7 855 differentially expressed genes were identified from StageⅡ to StageⅥ compared with StageⅠ as control, respectively. The differentially expressed genes were enriched in the molecular function of hydrolase activity that related the cellulose degradation in the early developmental stages via GO enrichment. (4) Totally, 30 GH9 gene family members were identified from the jujube genome-wide. Phylogenetic analysis showed that the family was divided into three subfamilies GH9A, GH9B and GH9C, subfamily GH9A had the most of 19 family members. (5) The analysis of gene expression patterns showed that 17 of the 30 identified GH9 genes were expressed in StageⅠ to Stage Ⅲ of peel development with high expression level, while the expression level decreased in Stage Ⅳ and Stage Ⅵ. Real-time quantitative PCR assay showed that the expression levels of 8 differentially expressed genes decreased during the development of jujube fruit, and the trend was consistent with the cellulose content of jujube peel and transcriptome sequencing. The study shows that GH9 gene family played the role in degrading the cell wall of the peel during the development of jujube. |
| keywords:Ziziphus jujuba ‘Goutouzao’ fruit peel cellulose tissue structure GH9 gene family transcriptome sequencing |
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