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作物学报

作物学报 ›› 2024, Vol. 50 ›› Issue (1): 237-250.doi: 10.3724/SP.J.1006.2024.34076

• 研究简报 • 上一篇    下一篇

甘蓝型油菜盐胁迫响应基因表达谱分析及共表达网络的构建

杨闯(), 王玲, 全成滔, 余良倩, 戴成, 郭亮, 傅廷栋, 马朝芝*()   

  1. 华中农业大学作物遗传改良全国重点实验室 / 国家油菜工程技术研究中心 / 洪山实验室, 湖北武汉 430070
  • 收稿日期:2023-04-27 接受日期:2023-09-13 出版日期:2024-01-12 网络出版日期:2023-09-20
  • 通讯作者: *马朝芝, E-mail: yuanbeauty@mail.hzau.edu.cn
  • 作者简介:E-mail: 908311956@qq.com
  • 基金资助:
    国家重点研发计划项目(2021YFD160014101-02);湖北省重点研发计划项目(2020BBB061)

Relative expression profiles of genes response to salt stress and constructions of gene co-expression networks in Brassica napus L.

YANG Chuang(), WANG Ling, QUAN Cheng-Tao, YU Liang-Qian, DAI Cheng, GUO Liang, FU Ting-Dong, MA Chao-Zhi*()   

  1. National Key Lab oratory of Crop Genetic Improvement, Huazhong Agricultural University / National Engineering Research Center of Rapeseed / Hongshan Laboratory, Wuhan 430070, Hubei, China
  • Received:2023-04-27 Accepted:2023-09-13 Published:2024-01-12 Published online:2023-09-20
  • Contact: *E-mail: yuanbeauty@mail.hzau.edu.cn
  • Supported by:
    National Key Research and Development Program of China(2021YFD160014101-02);National Key Research and Development Program of Hubei Province(2020BBB061)

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摘要：

甘蓝型油菜是重要的油料作物, 而盐胁迫是影响油菜生长发育的主要环境条件之一, 可能会造成油菜减产、品质下降甚至死亡。本研究利用半冬性油菜ZS11作为试验材料, 对盐胁迫处理0、0.25、0.5、1、3、6、12和24 h的叶片和根系组织进行转录组测序, 通过测得的90份RNA-seq数据, 获得了油菜响应盐胁迫的高分辨率时间动态转录表达谱。相关性分析发现, 样本在盐胁迫处理1 h前后具有明显的早期响应与后期响应的聚类差异。利用DESeq2进行差异基因分析, 鉴定出根系以及叶片组织响应差异基因分别为20,462个和29,334个, 表明油菜叶片组织的响应程度整体上比根系更剧烈。进一步利用WGCNA分别构建根系以及叶片组织响应盐胁迫的基因共表达网络, 从中筛选出与盐胁迫早期响应阶段显著相关的tan和yellow模块, 以及与盐胁迫后期响应阶段显著相关的green和red模块, 对其进行GO富集分析, 并从中分别筛选出早期以及后期响应盐胁迫的核心转录因子41个和26个。功能注释显示4个模块中均存在已知的拟南芥同源基因参与不同阶段的盐胁迫响应, 还发现BnWRKY46和BnWRKY57等核心基因在505份盐胁迫处理的油菜群体变异数据中具有丰富的SNPs变异和单倍体类型, 表明这些核心转录因子可能是油菜响应盐胁迫的关键候选基因。本研究可为甘蓝型油菜耐盐性改良提供可靠的数据参考和候选基因资源。

关键词: 甘蓝型油菜, RNA-seq, 盐胁迫, WGCNA, 基因共表达网络

Abstract:

Brassica napus L. is an important oil crop. Salt stress is one of the major environmental conditions affecting the growth and development of B. napus, which may lead to yield reduction, quality deterioration, and even the death of B. napus. In this study, the B. napus semi-winter cultivar ZS11 was used as the experimental material to perform transcriptome sequencing on the leaf and root tissues with salt stress treatment (0, 0.25, 0.5, 1, 3, 6, 12, and 24 h). The measured 90 RNA-seq data provided the high-resolution time dynamic transcriptional expression spectrum of rapeseed tissues responding to salt stress was obtained. Correlation analysis showed that the samples exhibited significant clustering differences in early response and late response before and after 1 h salt stress treatment. Using DESeq2 for differential gene analysis, we identified 20,462 and 29,334 differential genes for the response of root and leaf tissues, respectively, indicating that the overall response of leaf tissue in rapeseed was more severe than the root. Furthermore, WGCNA was used to construct gene co-expression networks for the salt stress response in root and leaf tissues, respectively, and tan and yellow modules were significantly related to the early response to salt stress, and green and red modules were significantly related to the late response to salt stress. GO enrichment analysis was then performed, and 41 and 26 core transcription factors responding to salt stress at the early and late stages, respectively, were selected from these networks. Functional annotation showed that the known Arabidopsis homologous genes involved in salt stress responses at different stages existed in all four modules, and core genes, such as BnWRKY46 and BnWRKY57, had abundant SNPs variation and haplotypes in 505 salt stress-treated rape population variation data, suggesting that these core transcription factors might be key candidate genes for rape salt stress response. This study provides a reliable data reference and candidate gene resources for improving salt tolerance in B. napus.

Key words: Brassica napus L., RNA-seq, salt stress, WGCNA, gene co-expression networks

图1

盐胁迫样品处理条件"

图2

盐胁迫响应marker基因的表达模式"

图3

盐胁迫的相关性分析 A: 主坐标分析; B: 层次聚类分析。主坐标分析时使用对照组, 为了直观看到聚类情况, 层次聚类时未加对照组。"

图4

盐胁迫处理24 h内的差异基因数目及变化趋势 A: 差异基因数目随处理时间的变化趋势; B: 各时间点上下调基因数目统计。"

图5

最佳软阈值的选择 A, B: 不同软阈值的网络拓扑分析。图A代表叶片组织, 图B代表根系组织, 左侧面板显示了无尺度拟合指数(Y轴)作为软阈值(X轴)的函数, 右侧面板显示了平均连通度(Y轴)作为软阈值(X轴)的函数。"

附表1

叶片组织响应盐胁迫的基因共表达网络模块信息"

模块
Module		 基因数目(转录因子数目)
Gene_number (TF_number)		 模块
Module		 基因数目(转录因子数目)
Gene_number (TF_number)
cyan 198 (18) blue 4945 (209)
salmon 242 (36) pink 775 (22)
green 1654 (138) purple 466 (59)
red 1148 (73) tan 260 (75)
turquoise 8651 (559) black 801 (27)
greenyellow 405 (33) magenta 556 (45)
brown 3510 (291) yellow 3101 (184)

附表2

根系组织响应盐胁迫的基因共表达网络模块信息"

模块
Module		 基因数目(转录因子数目)
Gene_number (TF_number)		 模块
Module		 基因数目(转录因子数目)
Gene_numbe (TF_number)
greenyellow 234 (27) yellow 2322 (276)
pink 824 (85) purple 365 (12)
brown 2515 (134) red 1238 (99)
green 2077 (195) black 1066 (89)
magenta 651 (41) turquoise 4058 (393)
blue 3590 (426)

图6

基因模块与样本时间点的相关性热图 A: 叶片组织的相关性热图; B: 根系组织的相关性热图。颜色越深表明相关性越大。"

图7

早期及后期响应模块的基因表达谱 A: 叶片组织早期响应阶段的tan模块; B: 根系组织早期响应阶段的yellow模块; C: 叶片组织后期响应阶段的green模块; D: 根系组织后期响应阶段的red模块。热图显示了模块中所有共表达基因的表达谱。条形图显示了模块特征基因的表达模式。"

图8

早期及后期响应模块的GO富集分析 A: 叶片组织早期响应阶段的tan模块; B: 根系组织早期响应阶段的yellow模块; C: 叶片组织后期响应阶段的green模块; D: 根系组织后期响应阶段的red模块。"

图9

早期及后期响应模块的基因共表达网络可视化 A: 叶片组织早期响应阶段的tan模块; B: 根系组织早期响应阶段的yellow模块; C: 叶片组织后期响应阶段的green模块; D: 根系组织后期响应阶段的red模块。转录因子用红色三角形表示, 非转录因子用蓝绿色圆形表示, 图形越大表明基因连通度越高。"

附表3

关键模块的核心转录因子功能注释"

关键模块
Hub modules		 核心转录因子
Core transcription factors		 基因别名
Gene aliases		 基因名字
Gene names		 基因注释
Gene annotation
tan BnaA03G0475900ZS WRKY53 AT4G23810 WRKY转录因子家族成员，是叶绿素合成/降解、衰老和气孔导度的拮抗调节因子。
A member of WRKY transcription factor family which is antagonistic regulators of chlorophyll synthesis/degradation, senescence, and stomatal conductance.
tan BnaC07G0058300ZS NAC036 AT2G17040 NAC转录因子家族成员，参与叶和花序茎的形态建成。
A member of the NAC transcription factor family involved in leaf and inflorescence stem morphogenesis.
tan BnaC03G0795800ZS ERF4 AT3G15210 作为JA应答防御基因和对坏死性真菌病原体尖孢镰刀菌抗性的负调节因子，可拮抗JA对根伸长的抑制作用。
Acts as a negative regulator of JA-responsive defense gene expression and resistance to the necrotrophic fungal pathogen Fusarium oxysporum and antagonizes JA inhibition of root elongation.
tan BnaA01G0133900ZS WRKY53 AT4G23810 WRKY转录因子家族成员，是叶绿素合成/降解、衰老和气孔导度的拮抗调节因子。
A member of WRKY transcription factor family which is antagonistic regulators of chlorophyll synthesis/degradation, senescence, and stomatal conductance.
tan BnaC01G0172400ZS WRKY53 AT4G23810 WRKY转录因子家族成员，是叶绿素合成/降解、衰老和气孔导度的拮抗调节因子。
A member of WRKY transcription factor family which is antagonistic regulators of chlorophyll synthesis/degradation, senescence, and stomatal conductance.
tan BnaC07G0454200ZS WRKY53 AT4G23810 WRKY转录因子家族成员，是叶绿素合成/降解、衰老和气孔导度的拮抗调节因子。
A member of WRKY transcription factor family。
tan BnaA09G0423300ZS ERF11 AT1G28370 ERF/AP2转录因子家族中ERF亚家族B-1的一员，该蛋白含有一个AP2结构域。
A member of the ERF subfamily B-1 of ERF/AP2 transcription factor family which contains one AP2 domain.
tan BnaC03G0651000ZS ERF11 AT1G28370 ERF/AP2转录因子家族中ERF亚家族B-1的一员，该蛋白含有一个AP2结构域。
A member of the ERF subfamily B-1 of ERF/AP2 transcription factor family which contains one AP2 domain.
tan BnaC03G0042100ZS NAC081 AT5G08790 具有NAC结构域的假定转录激活因子的大家族成员，可由创伤诱导表达。
A large family member of putative transcriptional activators with NAC domain, which can be Induced by wounding.
tan BnaA04G0247200ZS WRKY33 AT2G38470 WRKY转录因子家族成员，参与对各种非生物胁迫尤其是盐胁迫的反应。能控制根中凋亡屏障的形成，从而赋予耐盐性。
A member of WRKY transcription factor family. Involved in response to various abiotic stresses especially salt stress.Control apoplastic barrier formation in roots to confer salt tolerance.
tan BnaC03G0217300ZS WRKY33 AT2G38470 参与对各种非生物胁迫尤其是盐胁迫的反应。
Involved in response to various abiotic stresses especially salt stress.
tan BnaA06G0417500ZS ERF5 AT5G47230 ERF/AP2转录因子家族(ATERF-5)的ERF亚家族B-3的成员。
A member of the ERF subfamily B-3 of ERF/AP2 transcription factor family (ATERF-5).
tan BnaA03G0185200ZS WRKY33 AT2G38470 WRKY转录因子家族成员，参与对各种非生物胁迫尤其是盐胁迫的反应。
A member of the plant WRKY transcription factor family. Involved in response to various abiotic stresses - especially salt stress.
tan BnaA08G0213700ZS ERF11 AT1G28370 ERF/AP2转录因子家族中ERF亚家族B-1的一员。
A member of the ERF subfamily B-1 of ERF/AP2 transcription factor family.
tan BnaC04G0562300ZS WRKY33 AT2G38470 参与对各种非生物胁迫尤其是盐胁迫的反应。
Involved in response to various abiotic stresses especially salt stress.
yellow BnaC09G0432800ZS ZAT12 AT5G59820 锌指蛋白，参与高光和冷适应。
A zinc finger protein involved in high light and cold acclimation.
yellow BnaC08G0297500ZS NAC062 AT3G49530 作为冷信号和病原体抗性反应之间分子链接的转录因子。
Transcription factor that serves as a molecular link between cold signals and pathogen resistance responses.
yellow BnaC02G0323100ZS WRKY40 AT1G80840 病原体诱导转录因子，与WRKY18或WRKY60共表达会使植物对丁香假单胞菌和灰霉病菌更敏感。
Pathogen-induced transcription factor. Coexpression with WRKY18 or WRKY60 made plants more susceptible to both P. syringae and B. cinerea.
yellow BnaA03G0475900ZS WRKY53 AT4G23810 WRKY转录因子家族成员，是叶绿素合成/降解、衰老和气孔导度的拮抗调节因子。
A member of WRKY transcription factor family which is antagonistic regulators of chlorophyll synthesis/degradation, senescence, and stomatal conductance.
yellow BnaA08G0015900ZS ERF8 AT1G53170 ERF/AP2转录因子家族(ATERF-8)ERF亚家族B-1的一个成员，这种蛋白质含有一个AP2结构域。
A member of the ERF subfamily B-1 of ERF/AP2 transcription factor family (ATERF-8). The protein contains one AP2 domain.
yellow BnaC05G0100700ZS DDF1 AT1G12610 ERF/AP2转录因子家族(DD f1)DREB亚家族A-1的一个成员。这种蛋白质含有一个AP2结构域。该亚家族有6个成员，包括CBF1、CBF2和CBF3。该基因的过表达导致延迟开花和矮化，赤霉酸生物合成减少，以及对高水平盐的耐受性增加。
A member of the DREB subfamily A-1 of ERF/AP2 transcription factor family (DDF1). The protein contains one AP2 domain. There are six members in this subfamily, including CBF1, CBF2, and CBF3. Overexpression of this gene results in delayed flowering and dwarfism, reduction of gibberellic acid biosynthesis, and increased tolerance to high levels of salt.
yellow BnaC07G0454200ZS WRKY53 AT4G23810 WRKY转录因子家族成员，是叶绿素合成/降解、衰老和气孔导度的拮抗调节因子
A member of WRKY transcription factor family which is antagonistic regulators of chlorophyll synthesis/degradation, senescence, and stomatal conductance.
yellow BnaA01G0133900ZS WRKY53 AT4G23810 WRKY转录因子家族成员，是叶绿素合成/降解、衰老和气孔导度的拮抗调节因子
A member of WRKY transcription factor family which is antagonistic regulators of chlorophyll synthesis/degradation, senescence, and stomatal conductance.
yellow BnaC03G0795800ZS ERF4 AT3G15210 作为JA反应性防御应答基因和对坏死性真菌病原体尖孢镰刀菌抗性的负调节因子，能拮抗JA对根伸长的抑制作用。
Acts as a negative regulator of JA-responsive defense gene expression and resistance to the necrotrophic fungal pathogen Fusarium oxysporum and antagonizes JA inhibition of root elongation.
yellow BnaA06G0329700ZS NAC091 AT5G24590 NAC转录因子家族成员,参与调节拟南芥对TCV病毒的防御反应。
A member of NAC transcription factor family involved in regulating the defense response of Arabidopsis to turnip crinkle virus (TCV).
yellow BnaC04G0080600ZS WRKY33 AT2G38470 参与应对各种非生物胁迫，尤其是盐胁迫。
Involved in response to various abiotic stresses especially salt stress.
yellow BnaA06G0356400ZS WRKY48 AT5G49520 一种胁迫和病原体诱导的转录激活因子，会抑制植物的基础防御。
A stress- and pathogen-induced transcriptional activator that represses plant basal defense.
yellow BnaC05G0398500ZS AZF2 AT3G19580 锌指蛋白家族成员，其mRNA水平在响应ABA、高盐和轻度脱水时会上调表达。
A member of zinc finger protein family whose mRNA levels are upregulated in response to ABA, high salt, and mild desiccation.
yellow BnaA09G0110600ZS NAC036 AT2G17040 NAC转录因子家族成员，参与叶和花序茎的形态发生。
A member of the NAC transcription factor family that involved in leaf and inflorescence stem morphogenesis.
yellow BnaC09G0112900ZS NAC036 AT2G17040 NAC转录因子家族成员，参与叶和花序茎的形态发生。
A member of the NAC transcription factor family involved in leaf and inflorescence stem morphogenesis.
yellow BnaC07G0433500ZS LSD1 AT4G20380 锌指蛋白家族成员,负调节基础防御途径,监控超氧化物依赖性信号并负调节植物细胞死亡途径。
A member of zinc finger family protein that negatively regulates a basal defense pathway, monitors a superoxide-dependent signal and negatively regulates a plant cell death pathway.
yellow BnaC03G0563400ZS NAC062 AT3G49530 NAC转录因子家族成员,作为冷信号和病原体抗性反应之间分子链接。
A member of the NAC transcription factor family that serves as a molecular link between cold signals and pathogen resistance responses.
yellow BnaC05G0161100ZS ERF017 AT1G19210 ERF/AP2转录因子家族的DREB亚家族A-5的一个成员,这种蛋白质含有一个AP2结构域。
A member of the DREB subfamily A-5 of ERF/AP2 transcription factor family. The protein contains one AP2 domain.
yellow BnaC04G0512500ZS POSF21 AT2G31370 碱性亮氨酸拉链(bZIP)转录因子家族蛋白。
Basic-leucine zipper (bZIP) transcription factor family protein.
yellow BnaA07G0269000ZS MYBD AT1G70000 MYB样结构域转录因子，在响应光和细胞分裂素的花青素积累中发挥正调控作用。
A MYB-like Domain transcription factor that plays a positive role in anthocyanin accumulation in response to light and cytokinin.
yellow BnaA03G0321600ZS DREB2B AT3G11020 ERF/AP2转录因子家族(DREB2B)的DREB亚家族A-2的成员。这种蛋白质含有一个AP2结构域。
A member of the DREB subfamily A-2 of ERF/AP2 transcription factor family (DREB2B). The protein contains one AP2 domain.
yellow BnaC03G0264000ZS WRKY46 AT2G46400 WRKY转录因子，通过调节ABA信号和生长素体内平衡对渗透/盐胁迫依赖性LR抑制进行前馈抑制。
A WRKY transcription factor that contributes to the feedforward inhibition of osmotic/salt stress-dependent LR inhibition via regulation of ABA signaling and auxin homeostasis.
yellow BnaA02G0116700ZS MYB36 AT5G57620 MYB转录因子家族成员，在根发育过程中促进内皮层的分化。
A member of MYB transcriptional factor family that acts to promote differentiation of the endodermis during root development.
yellow BnaA04G0034100ZS BHLH107 AT3G56770 碱性螺旋-环-螺旋(bHLH) DNA结合超家族蛋白质。
basic helix-loop-helix (bHLH) DNA-binding superfamily protein.
yellow BnaA06G0158600ZS NAC062 AT3G49530 NAC转录因子家族成员,作为冷信号和病原体抗性反应之间分子链接。
A member of the NAC transcription factor family that serves as a molecular link between cold signals and pathogen resistance responses.
yellow BnaA06G0081500ZS DDF1 AT1G12610 ERF/AP2转录因子家族(DD f1)DREB亚家族A-1的一个成员,该基因的过表达导致延迟开花和矮化，对高水平盐的耐受性增加。
A member of the DREB subfamily A-1 of ERF/AP2 transcription factor family (DDF1). Overexpression of this gene results in delayed flowering and dwarfism, and increased tolerance to high levels of salt.
green BnaA05G0016800ZS GBF3 AT2G46270 一种bZIP G盒结合蛋白，其表达受ABA、寒冷和缺水诱导。
A bZIP G-box binding protein, induced by ABA, cold and water deprivation.
green BnaC03G0260300ZS SOC1 AT2G45660 控制开花，是一氧化碳促进开花所必需的。
Controls flowering and is required for CO to promote flowering.
green BnaC05G0560000ZS ATHB-1 AT3G01470 一个参与叶和下胚轴发育的同源结构域亮氨酸拉链I类(HD-Zip I)转录激活因子。
A homeodomain leucine zipper class I (HD-Zip I) transcriptional activator involved in leaf and hypocotyl development.
green BnaA07G0347200ZS ATY13 AT1G74650 R2R3因子基因家族成员;与生殖发育有关的蜡调节因子。
A member of the R2R3 factor gene family; wax regulator associated with reproductive development.
green BnaC01G0510700ZS NAC047 AT3G04070 含NAC结构域的蛋白47。
NAC domain containing protein 47.
green BnaA09G0409000ZS NFYA7 AT1G30500 核因子Y，亚单位A7。
nuclear factor Y, subunit A7.
green BnaA01G0191600ZS ARR1 AT3G16857 一种拟南芥反应调节蛋白(ARR )，在细胞分裂素信号途径中与其他B型ARR协同作用。还参与细胞分裂素依赖的下胚轴伸长抑制和细胞分裂素依赖的组织培养中的绿化和抽芽。ARR1、ARR10和ARR12是干旱反应的冗余调节因子，其中ARR1最为关键。
An Arabidopsis response regulator (ARR) protein that acts in concert with other type-B ARRs in the cytokinin signaling pathway. Also involved in cytokinin-dependent inhibition of hypocotyl elongation and cytokinin-dependent greening and shooting in tissue culture. ARR1, ARR10, and ARR12 are redundant regulators of drought response, with ARR1 being the most critical.
green BnaC04G0017100ZS GBF3 AT2G46270 一种bZIP G盒结合蛋白，受ABA、寒冷和缺水诱导。
A bZIP G-box binding protein induced by ABA, cold and water deprivation.
green BnaC06G0084400ZS NFYA5 AT1G54160 CBF-B/NF-YA转录因子家族的成员。响应ABA和干旱，功能丧失突变对干旱高度敏感。
A member of the CBF-B/NF-YA transcription factor family. Expression is upregulated in response to ABA and drought. Loss of function mutations are hypersensitive to drought.
green BnaA01G0036700ZS ATB2 AT4G34590 碱性结构域亮氨酸拉链(bZIP)转录因子bZIP11。蔗糖抑制翻译，导致uORF2中的核糖体停滞。直接调节参与氨基酸代谢的酶编码基因ASN1和ProDH2的基因表达。
A basic domain leucine zipper (bZIP) transcription factor bZIP11. Translation is repressed by sucrose that results in ribosome stalling in the uORF2. Directly regulates gene expression of ASN1 and ProDH2, which are enzyme-coding genes involved in amino acid metabolism.
green BnaC05G0008500ZS —— AT1G01250 ERF/AP2转录因子家族的DREB亚家族A-4的成员。这种蛋白质含有一个AP2结构域。
A member of the DREB subfamily A-4 of ERF/AP2 transcription factor family. The protein contains one AP2 domain.
green BnaA09G0663800ZS NFYC9 AT1G08970 核因子Y C (NF-YC)同系物NF-YC9，调节GA和ABA介导的种子萌发。
A NUCLEAR FACTOR-Y C (NF-YC) homologue NF-YC9 that modulate GA- and ABA-mediated seed germination.
green BnaC09G0556700ZS FLC AT5G10140 由开花基因座C转录因子编码的MADS-box蛋白，作为花转变的阻遏物并有助于昼夜节律钟的温度补偿。冷处理过程中表达下调。春化、FRI和自主途径都会影响FLC染色质的状态。在体内与SOC1和FT染色质相互作用。
MADS-box protein encoded by FLOWERING LOCUS C - transcription factor that functions as a repressor of floral transition and contributes to temperature compensation of the circadian clock. Expression is downregulated during cold treatment. The small RNAs are most likely derived from an antisense transcript of FLC. Interacts with SOC1 in vivo.
green BnaC07G0522500ZS GATA3 AT4G34680 锌指转录因子GATA因子家族成员。
A member of the GATA factor family of zinc finger transcription factors.
red BnaA02G0180600ZS WRKY57 AT1G69310 WRKY转录因子成员。WRKY57的激活赋予了耐旱性。
A member of the WRKY Transcription Factor. Activation of WRKY57 confers drought tolerance.
red BnaC02G0139100ZS LRL3 AT5G58010 调节根毛发育的碱性螺旋-环-螺旋(bHLH)蛋白。
A basic helix-loop-helix (bHLH) protein that regulates root hair development.
red BnaA03G0557900ZS ATHB40 AT4G36740 同源结构域亮氨酸拉链I类(HD-Zip I)蛋白。
A homeodomain leucine zipper class I (HD-Zip I) protein.
red BnaA09G0422600ZS NAC010 AT1G28470 含NAC结构域的蛋白10。
NAC domain containing protein 10.
red BnaC07G0322500ZS MYB121 AT3G30210 R2R3因子基因家族成员。
A member of the R2R3 factor gene family.
red BnaA05G0156300ZS NFYA5 AT1G54160 CCAAT结合转录因子(CBF-B/NF-YA)家族的成员。响应ABA和干旱。
A member of the CCAAT-binding transcription factor (CBF-B/NF-YA) family. Expression is upregulated in response to ABA and drought.
red BnaA07G0311300ZS ZFHD1 AT1G69600 锌指同源结构域转录因子家族成员ZFHD1。干旱、高盐和脱落酸诱导ZFHD1的表达。
A member of the zinc finger homeodomain transcriptional factor family. Expression of ZFHD1 is induced by drought, high salinity and abscisic acid.
red BnaA08G0184300ZS ATHB40 AT4G36740 同源结构域亮氨酸拉链I类(HD-Zip I)蛋白。
A homeodomain leucine zipper class I (HD-Zip I) protein.
red BnaC06G0308100ZS WRKY57 AT1G69310 WRKY转录因子成员。WRKY57的激活赋予了耐旱性。
A member of the WRKY Transcription Factor. Activation of WRKY57 confers drought tolerance.
red BnaC08G0452900ZS SRS3 AT2G21400 SHI基因家族成员，能促进拟南芥中雌蕊、雄蕊和叶的发育。
A member of SHI gene family that promote gynoecium, stamen and leaf development in Arabidopsis.
red BnaA07G0271900ZS WRKY57 AT1G69310 WRKY转录因子成员。WRKY57的激活赋予了耐旱性。
A member of the WRKY Transcription Factor. Activation of WRKY57 confers drought tolerance.
red BnaC09G0556700ZS FLC AT5G10140 冷处理过程中表达下调。
Expression is downregulated during cold treatment.

图10

核心基因BnWRKY46和BnWRKY57在505份群体数据中的SNPs及单倍型分析 A: BnWRKY46的连锁不平衡图。B: BnWRKY46在505份群体数据中的单倍型箱线图, 纵坐标是油菜幼苗期盐胁迫处理组与对照组的根长比值, 不同字母表示使用双向方差分析的P ≤ 0.05的显著性差异。C: BnWRKY57的连锁不平衡图。D: BnWRKY57在505份群体数据中的单倍型箱线图, 纵坐标是油菜幼苗期盐胁迫处理组与对照组的根长比值, 不同字母表示使用双向方差分析的P ≤ 0.05的显著性差异。"

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