大豆是重要的经济作物，富含油脂和蛋白质，对人类健康的改善、生物燃料的应用和土壤肥力的提高具有重要作用。然而，大豆在生长发育过程中经常会遭受各种逆境胁迫的伤害，其中干旱、盐害是影响大豆正常生长的两种主要逆境胁迫。干旱、盐害引起的不良反应包括渗透失衡、膜系统损伤、光合作用速率降低等，不仅抑制作物在各个阶段的生长发育和生理代谢，而且还影响作物的品质和产量。为了应对干旱、盐害，植物已经在形态、生理和分子水平上进化出多种生存策略，例如改变根、茎、叶组织的结构、改变代谢产物的组成以及调节胁迫相关基因的表达等。

转录组测序技术的发展，有助于从转录水平上了解植物对逆境胁迫的响应机理。本文研究了正常条件和干旱处理后大豆幼苗根、叶组织的转录组，分析了大豆响应干旱胁迫的分子机理；结合大豆转基因和基因编辑技术，对耐旱候选基因GmNAC8、GmNAC12及另一高度同源基因GmNAC100进行了功能研究。本研究的主要结果如下：

1. 干旱胁迫下大豆幼苗根、叶组织的转录组分析

对正常条件和干旱处理后天隆一号大豆幼苗根、叶组织进行转录组分析，在根中共筛选到10078个差异表达基因（DEGs），其中5179个基因显著上调表达，4899个基因显著下调表达。在叶中共筛选到8501个差异表达基因，其中4108个基因显著上调表达，4393个基因显著下调表达。KEGG富集分析显示，这些DEGs主要参与了植物激素信号转导途径、苯丙素合成代谢通路、植物病原菌互作通路。在根和叶的DEGs中分别发现了1113个和773个转录因子，包括AP2/ERF、AUX/IAA、bHLH、bZIP、C2H2、GATA、GRAS、MYB/MYB-Related、NAC、WRKY等转录因子家族。其中，在根、叶的DEGs中分别发现了61个、37个NAC转录因子，有13个NAC基因在根和叶中同时上调表达，有3个NAC基因在根和叶中同时下调表达。GmNAC8和GmNAC12是一对同源基因，在干旱胁迫下都显著上调表达，之前也多次被报道与大豆的耐旱性相关。因此，将GmNAC8和GmNAC12假定为调控大豆耐旱性的候选基因。

2. 耐旱候选基因GmNAC8的功能研究

从大豆中克隆了GmNAC8的编码序列，全长为1092 bp，编码363个氨基酸；亚细胞定位分析显示GmNAC8是一个核定位蛋白。通过qRT-PCR分析显示GmNAC8在干旱、高盐、高温胁迫下均显著上调表达，其中在干旱胁迫下上调表达最明显。GmNAC8在逆境相关植物激素ABA、ETH、SA处理后3 h表达上调到峰值。表达模式分析推测GmNAC8可能通过植物激素信号转导通路，参与大豆对逆境胁迫的调控。通过农杆菌介导的大豆遗传转化，获得4个GmNAC8过表达株系；通过CRISPR/Cas9介导的基因编辑，获得4种类型的GmNAC8的基因敲除株系，即GmNAC8-KO1（缺失8 bp）、GmNAC8-KO2（插入1 bp）、GmNAC8-KO3（缺失1 bp）、GmNAC8-KO4（缺失5 bp）。与野生型比较，GmNAC8-KO1、GmNAC8-KO2、GmNAC8-KO3、GmNAC8-KO4都发生了移码突变，导致翻译提前终止。干旱处理后，与野生型相比，GmNAC8过表达株系的存活率、SOD活性和PRO含量显著更高，表现出耐旱性；而GmNAC8基因敲除株系显著降低的存活率、SOD活性和PRO含量表现出干旱敏感性，表明GmNAC8正向调控大豆植株的耐旱性。通过酵母双杂技术筛选了119个GmNAC8的互作蛋白，通过GO注释和KEGG富集分析，发现这些互作蛋白参与铁离子响应，泛素依赖的代谢过程，ABA和水分缺失的响应等。基于GmDi19-3参与对干旱和ABA的响应，因此着重研究了GmNAC8与GmDi19-3之间的蛋白互作。通过双分子荧光互补实验和荧光素酶互补实验在烟草体内验证了GmNAC8与GmDi19-3之间存在相互作用。为了研究GmDi19-3的功能，通过农杆菌介导的大豆遗传转化获得8个GmDi19-3过表达株系，干旱处理后，GmDi19-3过表达植株表现出比野生型更强的耐旱性。综合以上结果，推测GmNAC8可能通过与GmDi19-3互作正向调控大豆的耐旱性。

3. 耐旱候选基因GmNAC12的功能研究

从大豆中克隆了GmNAC12的编码序列，全长为1062 bp，编码353个氨基酸；亚细胞定位分析显示GmNAC12在细胞核表达。通过qRT-PCR分析显示GmNAC12在干旱、高盐、高温胁迫下均显著上调表达，在干旱胁迫下上调最明显。GmNAC12在ABA、ETH、SA处理后的3 h表达达到峰值，在MeJA处理后12 h表达达到峰值，推测GmNAC12可能通过植物激素信号转导通路，参与大豆对逆境胁迫的调控。通过农杆菌介导的大豆遗传转化，获得3个GmNAC12过表达株系；通过CRISPR/Cas9介导的基因编辑技术，获得5种类型的GmNAC12基因敲除株系，即GmNAC12-KO1（缺失1 bp）、GmNAC12-KO2（缺失2 bp）、GmNAC12-KO3（缺失3 bp）、GmNAC12-KO4（缺失4 bp）、GmNAC12-KO5（缺失5 bp）。与野生型比较，GmNAC12-KO1、GmNAC12-KO2、GmNAC12-KO4、GmNAC12-KO5都发生了移码突变，导致翻译提前终止。干旱处理后，与野生型相比，GmNAC12过表达株系的存活率显著升高，表现出耐旱性；而GmNAC12基因敲除株系的存活率显著降低，表现出对干旱的敏感，表明GmNAC12是大豆耐旱性的一个正向调控因子。通过酵母双杂技术筛选了185个GmNAC12的互作蛋白，通过互作蛋白的GO注释和KEGG富集分析，推测GmNAC12可能通过抗氧化防御通路来增强大豆的耐旱性。

4. 与GmNAC8和GmNAC12高度同源的GmNAC100的功能研究

从大豆中克隆了GmNAC100的编码序列，全长为1224 bp，编码407个氨基酸，亚细胞定位分析显示，GmNAC100定位于细胞核。通过qRT-PCR分析显示GmNAC100在干旱、高盐、高温胁迫下均显著上调表达，其中在高盐胁迫下上调表达最为迅速。GmNAC100的表达在ABA、ETH、SA处理后的3 h上调表达到达峰值，在MeJA处理后的12 h上调表达到达峰值，推测GmNAC100可能通过植物激素信号转导通路，参与大豆对逆境的调控。通过农杆菌介导的大豆遗传转化，获得9个GmNAC100过表达株系，通过CRISPR/Cas9介导的基因编辑技术，获得3种类型的GmNAC100基因敲除株系，即GmNAC100-KO1（缺失1 bp）、GmNAC100-KO2（缺失2 bp）、GmNAC100-KO3（缺失4 bp）。与野生型比较，GmNAC100-KO1发生移码突变，导致翻译提前终止；GmNAC100-KO2和GmNAC100-KO3发生移码突变，导致氨基酸序列改变。正常生长条件下，相对于野生型，GmNAC100的过表达株系有更发达的侧根，根鲜重更大，而GmNAC100基因敲除株系的侧根明显发育不良，表明GmNAC100调控大豆的侧根发育。150 mM NaCl处理后，与野生型相比，GmNAC100的过表达株系有更高的叶绿素含量和更低的相对电导率，表现出耐盐性，而GmNAC100基因敲除株系叶绿素含量更低、相对电导率更高，表明GmNAC100正向调控大豆的耐盐性。综合以上结果，推测GmNAC100可能通过调控大豆的侧根发育，从而影响大豆的耐盐性。

Soybean is an important economic crop, rich in oil and protein, and plays an important role in the improvement of human health and soil fertility, and the application of biofuels. However, soybeans often suffer from various stresses during growth and development. Among them, drought and salt stress are the two main stresses that affect the normal growth of soybeans. The adverse reactions caused by drought and salt stress include osmotic imbalance, membrane system damage, and reduced photosynthesis rate, etc., which not only inhibit the growth and physiological metabolism of crops at various stages, but also affect the quality and yield of crops. To adapt to drought and salt stress, plants have evolved various survival strategies at the morphological, physiological, and molecular levels, such as changing the structure of root, stem, and leaf tissues, changing the composition of metabolites, and regulating the expression of stress-related genes.

The development of transcriptome sequencing technology helps to understand the mechanism of plants response to stress at the transcription level. In this study, the transcriptome of the root and leaf tissues of soybean seedlings under normal conditions and drought treatment was analyzed, and the molecular mechanism of soybean response to drought stress was analyzed; combined with soybean transgene and gene editing techniques, the candidate genes for drought tolerance GmNAC8 and GmNAC12, and another highly homologous gene GmNAC100, were conducted for functional study. The main results of this study are as follows:

1. Transcriptome analysis of root and leaf tissues of soybean seedlings under drought stress

Transcriptome analysis was performed using root and leaf tissues of soybean seedlings under normal conditions and drought treatment. A total of 10078 differentially expressed genes (DEGs) were screened in the roots, of which 5179 genes were significantly up-regulated and 4899 genes were significantly down-regulated. A total of 8501 DEGs were screened in leaves, of which 4108 genes were significantly up-regulated and 4393 genes were significantly down-regulated. KEGG enrichment analysis showed that these DEGs were mainly involved in plant hormone signal transduction pathway, phenylpropanoid biosynthesis pathway, and plant-pathogen interaction pathway. 1113 and 773 transcription factors were found respectively in DEGs of roots and leaves, including AP2/ERF, AUX/IAA, bHLH, bZIP, C2H2, GATA, GRAS, MYB/MYB-Related, NAC, WRKY and other transcription factor families. 61 and 37 NAC transcription factors were found in the DEGs of roots and leaves, respectively. 13 NAC genes were up-regulated in roots and leaves, and 3 NAC genes were down-regulated in roots and leaves. GmNAC8 and GmNAC12 are a pair of homologous genes, both of which showed significant up-regulation expression under drought stress. It has also been reported many times that they related to soybean drought tolerance. Therefore, GmNAC8 and GmNAC12 were assumed to be candidate genes for regulating drought tolerance in soybean.

2. Functional study of drought tolerance candidate gene GmNAC8

The CDs of GmNAC8 was cloned from soybean, with a total length of 1092 bp, and encodes 363 amino acids. Subcellular localization analysis showed that GmNAC8 is a nuclear localization protein. GmNAC8 was significantly up-regulated under drought, high salt and high temperatures, of which GmNAC8 was up-regulated most under drought stress. GmNAC8 was up-regulated to the peak value at 3 h after stress-related plant hormones ABA, ETH and SA treatment. Analysis of expression patterns speculated that GmNAC8 may be involved in the response to stress through plant hormone signal transduction pathways. Through Agrobacterium-mediated genetic transformation of soybean, four overexpression lines of GmNAC8 were obtained. Through CRISPR/Cas9-mediated gene editing, four types of GmNAC8 gene knockout lines were obtained, namely GmNAC8-KO1 (deletion 8 bp), GmNAC8-KO2 (insertion 1 bp), GmNAC8-KO3 (deletion 1 bp), GmNAC8-KO4 (deletion 5 bp). Compared with the wild-type, GmNAC8-KO1, GmNAC8-KO2, GmNAC8-KO3, and GmNAC8-KO4 all experienced frameshift mutations, leading to early termination of translation. After drought treatment, compared with wild-type, GmNAC8 overexpression lines had significantly higher survival rate, SOD activity and PRO content, showing drought tolerance; while GmNAC8 gene knockout lines showed significantly lower survival rate, SOD activity and PRO content, indicating that GmNAC8 positively regulates the drought tolerance of soybean. Through yeast two-hybrid analysis, 119 proteins interacting with GmNAC8 were screened. Through GO annotation and KEGG enrichment, these interaction proteins were found to be involved in iron response, ubiquitin-dependent metabolic processes, response to ABA and water deficit etc. Based on GmDi19-3 response to water deficit and ABA, the study focused on the interaction between GmNAC8 and GmDi19-3. The interaction between GmNAC8 and GmDi19-3 was verified in tobacco by bimolecular fluorescence complementation and luciferase complementation experiments. In order to study the function of GmDi19-3, eight overexpression lines of GmDi19-3 were obtained through Agrobacterium-mediated genetic transformation of soybean. After drought treatment, GmDi19-3 overexpression plants showed stronger drought tolerance than wild-type. Based on the above results, it is speculated that GmNAC8 positively regulate soybean drought tolerance probably by interacting with GmDi19-3.

3. Functional study of drought tolerance candidate gene GmNAC12

The CDs of GmNAC12 was cloned from soybean, with a total length of 1062 bp, encodes 353 amino acids. Subcellular localization analysis revealed that GmNAC12 is a nuclear localization protein. GmNAC12 was significantly up-regulated under drought, high salt and high temperature stress. Among them, GmNAC12 was up-regulated most under drought stress. GmNAC12 was up-regulated to the peak value at 3 h after ABA, ETH, SA treatment, and at 12 h after MeJA treatment. It is speculated that GmNAC12 may participate in the regulation of stress through the plant hormone signal transduction pathway. Through Agrobacterium-mediated genetic transformation of soybean, three overexpression lines of GmNAC12 were obtained. Through CRISPR/Cas9-mediated gene editing, five types of GmNAC12 gene knockout lines were obtained, namely GmNAC12-KO1 (deletion 1 bp), GmNAC12-KO2 (deletion 2 bp), GmNAC12-KO3 (deletion 3 bp), GmNAC12-KO4 (Deletion 4 bp), GmNAC12-KO5 (deletion 5 bp). Compared with the wild-type, frameshift mutations occurred in GmNAC12-KO1, GmNAC12-KO2, GmNAC12-KO4, and GmNAC12-KO5, leading to early termination of translation. After drought treatment, compared with wild-type, the survival rate of GmNAC12 overexpression lines was significantly higher, showing drought tolerance; and the survival rate of GmNAC12 gene knockout lines was significantly lower, showing sensitivity to drought. This indicates that GmNAC12 is a positive regulator of soybean drought tolerance. 185 proteins interacting with GmNAC12 were screened by yeast two-hybrid technology. Through GO annotation and KEGG enrichment analysis of interaction proteins, it is speculated that GmNAC12 may enhance soybean drought tolerance through antioxidant defense pathways.

4. Functional study of GmNAC100 highly homologous to GmNAC8 and GmNAC12

The CDs of GmNAC100 was cloned from soybean, with a total length of 1224 bp, encodes 407 amino acids. Subcellular localization analysis revealed that GmNAC100 is a nuclear localization protein. GmNAC100 was up-regulated under drought, high salt and high temperature stress. Among them, the expression of GmNAC100 was up-regulated most rapidly under high salt stress. The expression of GmNAC100 was up-regulated to peak value at 3 h after ABA, ETH, SA treatment and at 12 h after MeJA treatment. Expression pattern analysis speculated that GmNAC100 may also participate in the regulation of stress through plant hormone signal transduction pathways. Through Agrobacterium-mediated genetic transformation of soybean, 9 overexpression lines of GmNAC100 were obtained. Through CRISPR/Cas9-mediated gene editing, three types of GmNAC100 gene knockout lines were obtained, namely GmNAC100-KO1 (deletion 1 bp), GmNAC100-KO2 (deletion 2 bp), GmNAC100-KO3 (deletion 4 bp). Compared with the wild-type, frameshift mutations occurred in GmNAC100-KO1, leading to early termination of translation; frameshift mutations occurred in GmNAC100-KO2 and GmNAC100-KO3, resulting in amino acid sequence changes. Under normal growth conditions, GmNAC100 overexpression lines had stronger lateral roots and larger root fresh weight than wild-type, and the lateral roots of GmNAC100 gene knockout lines were obviously underdeveloped, indicating that GmNAC100 regulates the development of soybean lateral roots. After 150 mM NaCl treatment, compared to wild type, GmNAC100 overexpression lines had higher chlorophyll content and lower relative electrical conductivity, showing salt tolerance; while GmNAC100 gene knockout lines had lower chlorophyll content, higher relative electrical conductivity, showing salt sensitivity. It indicated that GmNAC100 positively regulates soybean salt tolerance. In summary, it is speculated that GmNAC100 may regulate soybean salt tolerance by affecting the development of lateral roots.

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