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作物学报 ›› 2023, Vol. 49 ›› Issue (1): 24-35.doi: 10.3724/SP.J.1006.2023.14233

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

大豆GmPIN2家族基因调控根系发育功能初探

梁政1,3(), 柯美玉1,3, 陈志威2, 陈栩3,*(), 高震3,*()   

  1. 1福建农林大学生命科学学院, 福建福州 350002
    2福建农林大学资源与环境学院, 福建福州 350002
    3福建农林大学海峡联合研究院园艺植物生物学及代谢组学研究中心, 福建福州 350002
  • 收稿日期:2021-12-08 接受日期:2022-05-05 出版日期:2023-01-12 网络出版日期:2022-05-13
  • 通讯作者: 陈栩,高震
  • 作者简介:E-mail: 1879949813@qq.com
  • 基金资助:
    福建农林大学校杰青项目(XJQ201921)

Function of GmPIN2 family gene in regulating root development in soybean

LIANG Zheng1,3(), KE Mei-Yu1,3, CHEN Zhi-Wei2, CHEN Xu3,*(), GAO Zhen3,*()   

  1. 1College of Life Science, Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China
    2College of Resources and Environment, Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China
    3Haixia Institute of Science and Technology, Horticultural Plant Biology and Metabolomics Center, Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China
  • Received:2021-12-08 Accepted:2022-05-05 Published:2023-01-12 Published online:2022-05-13
  • Contact: CHEN Xu,GAO Zhen
  • Supported by:
    Distinguished Young Scholar Program of Fujian Agriculture and Forestry University(XJQ201921)

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摘要：

生长素作为重要的植物激素之一。生长素的转运参与了植物各组织器官发育调控。在拟南芥中, 生长素转运调控主要由PIN家族蛋白所介导, 其中拟南芥AtPIN2主要通过介导生长素向基运输调控根的向重力性。大豆中PIN2家族蛋白及其功能研究尚未有报道。本研究通过构建系统进化树和蛋白质结构域分析发现, 大豆GmPIN2a和GmPIN2b为AtPIN2的同源基因。组织表达分析研究发现, GmPIN2a和GmPIN2b在根、根瘤原基和根瘤等组织中高表达。GmPIN2a和GmPIN2b在根部主要表达在根尖表皮和外部皮层细胞, 在根瘤中GmPIN2a、GmPIN2b均定位于根瘤基部维管束区, 此外, GmPIN2a定位在根瘤顶部表皮及外皮层。利用基因编辑技术CRISPR/Cas9 (Clustered Regularly Interspersed Short Palindromic Repeats/CRISPR associated 9)同时敲除GmPIN2a和GmPIN2b后发现, Gmpin2ab突变体根具有明显的重力缺失表型。与野生型相比, Gmpin2ab和35S::GmPIN2b的根面积和侧根长度显著降低, Gmpin2ab侧根夹角显著上升, 而35S::GmPIN2b侧根夹角不变。综上所述, GmPIN2a和GmPIN2b通过介导生长素向基运输对大豆根型调控具有重要作用。本研究为深入探究大豆PIN蛋白介导生长素极性运输在根形态建成的作用机制奠定了一定研究基础。

关键词: 大豆, 生长素, PIN蛋白, 根

Abstract:

Auxin is one of the important plant hormones. Auxin transport is involved in the regulation of plant tissue and organ development. In Arabidopsis thaliana, auxin transport is mainly regulated by PIN family proteins, among which AtPIN2 controls auxin basipetal transport and regulates root gravitation in Arabidopsis. However, GmPIN2 family proteins and their functions are still largely unknown in soybean. In this study, we identified two AtPIN2 homologous genes of GmPIN2a and GmPIN2b through phylogenetic and protein domain analysis. The relative expression level showed that GmPIN2a and GmPIN2b were highly expressed in root and nodule. Both genes were highly expressed in root epidermis and cortex, and the vascular bundles of the root nodule. Only GmPIN2a expressed in the epidermis and outer cortex at the tip of the root nodule. Furthermore, Gmpin2ab mutants were generated by CRISPR/Cas9 (Clustered Regularly Interspersed Short Palindromic Repeats/CRISPR Associated 9) gene editing technology. Gmpin2ab mutants had a significant loss of root gravitropism. Moreover, the loss of Gmpin2 resulted in a significant decrease of root area and lateral root length, while the lateral root angle of Gmpin2ab significantly increased. Overexpression of GmPIN2b also resulted in a decrease in root area and lateral root length, but the lateral root angle unchanged. In conclusion, GmPIN2a and GmPIN2b play important roles in regulation of soybean root development via mediating auxin basipetal transport. This study laid a foundation for further analysis of the function and mechanism of GmPIN proteins in soybean root morphogenesis.

Key words: Glycine max (soybean), auxin, PIN protein, root

表1

引物序列"

引物用途
Purpose of primers		 引物名称
Primer name		 引物序列
Primer sequence (5°-3°)
启动子序列扩增引物
Promoter amplification primers		 pGmPIN2a-F AGATGGTTAGAGAGGCATGAGCCACAATGGAAC
pGmPIN2a-R AGCTCTTATACTCGACATGGTTCATGCGGG
pGmPIN2b-F AGATGGTTAGAGAGGTGATGTTCTTGGCCACTG
pGmPIN2b-R AGCTCTTATACTCGAGATCATGGTTAATGCAGGG
定量PCR引物序列
RT-PCR primers		 GmPIN2a-F CCACACGAAACTGTTGCCTC
GmPIN2a-R AAGCTCCCTCTGATCTCCGT
GmPIN2b-F GGGAATTTGAGGCATGCAGT
GmPIN2b-R CAGGCTCCCTCTCTCCACTC
GmActin-11-F GAGCTATGAATTGCCTGATGG
GmActin-11-R CGTTTCATGAATTCCAGTAGC
sgRNA序列 sgRNA sequence sgRNA-GmPIN2ab TGGAACACCTTCACAAAACATGG
验证编辑载体的引物序列
Primer sequence for verifying the gene editing vectors		 pCas9-F GCCAAGAGCGAGCAGGAAAT
pCas9-R GGTGGATGATATTCTCGGCCT

图1

大豆GmPIN和拟南芥AtPIN蛋白家族系统进化树 红色框表示大豆GmPIN2和拟南芥AtPIN2。"

图2

拟南芥、大豆、蒺藜苜蓿、百脉根、水稻和玉米PIN2蛋白家族生物信息学分析 A: 拟南芥、大豆、蒺藜苜蓿、百脉根、水稻和玉米PIN2蛋白家族基因结构。B: 拟南芥、大豆、蒺藜苜蓿、百脉根、水稻和玉米PIN2蛋白家族基因启动子区域分析。C: 拟南芥、大豆、蒺藜苜蓿、百脉根、水稻和玉米PIN2蛋白家族蛋白序列比对。AT5G57090: AtPIN2; Glyma.13G101900: GmPIN2a; Glyma.17G057300: GmPIN2b; LOC_Os06g44970: OsPIN2; Medtr4g127100: MtPIN2a; Medtr4g127090: MtPIN2b; Lj4g3v2139970: LjPIN2; Zm00001d046893_P001: ZmPIN2。"

表2

拟南芥、大豆、蒺藜苜蓿、百脉根、水稻和玉米PIN2蛋白家族同源比对"

Seq-> AtPIN2 GmPIN2a GmPIN2b OsPIN2 MtPIN2a MtPIN2b LjPIN2 ZmPIN2
AtPIN2 100.0 76.5 77.2 61.7 75.0 63.7 76.5 61.0
GmPIN2a 76.5 100.0 96.8 63.9 85.1 68.3 90.7 61.9
GmPIN2b 77.2 96.8 100.0 63.5 85.5 68.8 90.8 61.9
OsPIN2 61.7 63.9 63.5 100.0 63.8 53.8 63.4 87.3
MtPIN2a 75.0 85.1 85.5 63.8 100.0 75.4 85.0 63.6
MtPIN2b 63.7 68.3 68.8 53.8 75.4 100.0 68.8 53.4
LjPIN2 76.5 90.7 90.8 63.4 85.0 68.8 100.0 61.7
ZmPIN2 61.0 61.9 61.9 87.3 63.6 53.4 61.7 100.0

图3

GmPIN2a和GmPIN2b基因组织表达模式分析和组织化学定位 A: GmPIN2a和GmPIN2b基因组织表达模式; B: GmPIN2a和GmPIN2b在根和根瘤中的组织化学定位, 标尺为100 μm。"

图4

Gmpin2ab-#1和Gmpin2ab-#2纯合突变体构建 A: GmPIN2a和GmPIN2b基因结构及sgRNA靶向位点示意图。黑色箭头代表sgRNA位置, 黑色框代表外显子, 红色框代表5′UTR和3′UTR, 黑色线条代表内含子。B: Gmpin2ab-L1和Gmpin2ab-L2中基因编辑载体的验证。白色数字从左往右分别表示: 野生型、35S::GmPIN2b、Gmpin2ab-L1和Gmpin2ab-L2。C: 野生型、35S::GmPIN2b、Gmpin2ab-L1和Gmpin2ab-L2中大豆GmPIN2家族表达量检测。P值由假设方差相等的双尾t检验确定, **表示Gmpin2ab-L1、Gmpin2ab-L2与野生型在P < 0.01水平差异显著; ***表示35S::GmPIN2b与野生型在P < 0.001水平差异显著; ****表示Gmpin2ab-L1、Gmpin2ab-L2与野生型在P < 0.0001水平差异显著。ns: 没有显著差异。D: Gmpin2ab-#1和Gmpin2ab-#2纯合突变体序列对比。绿色字母代表sgRNA靶向序列, 红色字母, 红色短线代表增添或缺失的密码子。E: 野生型, Gmpin2ab-L1和Gmpin2ab-L2纯合突变体蛋白序列。AA代表氨基酸(amino acid), 黑色箭头表示移码突变起始位点, 紫色框表示移码突变蛋白序列, 数字代表移码的氨基酸数量。"

图5

野生型、Gmpin2ab-L1、Gmpin2ab-L2和35S::GmPIN2b植株表型 A: Gmpin2ab-L1和Gmpin2ab-L2向重力性缺失表型。红色框表示突变体向地上部分生长的根。B: 生长7 d的野生型、Gmpin2ab-L1、Gmpin2ab-L2和35S::GmPIN2b的扫描图, 比例尺为1 cm。P值由假设方差相等的双尾t检验确定, ****表示P < 0.0001水平差异显著。ns: 没有显著差异, 样本数为7。"

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