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作物学报 ›› 2022, Vol. 48 ›› Issue (6): 1401-1415.doi: 10.3724/SP.J.1006.2022.12032

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

水稻OsLPL2/PIR基因抗旱耐盐机制研究

周文期1,2(), 强晓霞3, 王森4, 江静雯1, 卫万荣1,*()   

  1. 1西南野生动植物资源保护重点实验室 / 西华师范大学生命科学学院, 四川南充 637000
    2甘肃省农业科学院作物研究所, 甘肃兰州 730070
    3甘肃省兰州市第四中学, 甘肃兰州 730050
    4陕西省畜牧产业试验示范中心, 陕西咸阳 713702
  • 收稿日期:2021-05-01 接受日期:2021-10-19 出版日期:2022-06-12 网络出版日期:2021-11-20
  • 通讯作者: 卫万荣
  • 作者简介:E-mail: zhouwenqi850202@163.com
  • 基金资助:
    2020年甘肃省科协青年科技人才托举工程项目和甘肃省农业科学院农业科技创新专项——博士基金项目(2020GAAS34)

Mechanism of drought and salt tolerance of OsLPL2/PIR gene in rice

ZHOU Wen-Qi1,2(), QIANG Xiao-Xia3, WANG Sen4, JIANG Jing-Wen1, WEI Wan-Rong1,*()   

  1. 1Key Laboratory of Southwest China Wildlife Resources Conservation / College of Life Sciences, China West Normal University, Nanchong 637000, Sichuan, China
    2Crops Research Institute, Gansu Academy of Agricultural Sciences, Lanzhou 730070, Gansu, China
    3Lanzhou No. 4 High School, Lanzhou 730050, China
    4Shannxi Province Animal Husbandry Industry Experimental Demonstration Center, Xianyang 713702, Shaanxi, China
  • Received:2021-05-01 Accepted:2021-10-19 Published:2022-06-12 Published online:2021-11-20
  • Contact: WEI Wan-Rong
  • Supported by:
    Young Scientific and Technological Talents Support Project of Gansu Association for Science and Technology in 2020 and the Agricultural Science and Technology Innovation Program of Gansu Academy of Agricultural Sciences(2020GAAS34)

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摘要：

干旱威胁着全球农业生产, 限制了农业可持续发展的前景。植物叶表皮在生长发育、抵御逆境胁迫、与外界环境进行水分和气体交换中, 发挥了至关重要的作用。本研究中, 利用水稻(Oryza sativa) less pronounced lobe epidermal cell 2-1 (lpl2-1)和less pronounced lobe epidermal cell 2-2 (lpl2-2)突变体为研究材料, 与野生型中花11 (Zhonghua 11, ZH11)经干旱胁迫和不同浓度盐处理, 发现lpl2-1和lpl2-2对逆境胁迫响应更敏感, 复水后统计成活率极显著降低, 低于对照1/2。相比ZH11, lpl2-1和lpl2-2株高变矮, 根长变短, 相同叶序气孔密度、气孔开度均极显著增加, 且表皮扁平细胞边缘锯齿状凸出变平滑, 嵌套不紧密, 导致lpl2-1和lpl2-2比ZH11水分散失更多; 离体叶片失水实验也证明了lpl2-1和lpl2-2叶片在等时间内失水更快, 失水率更高; 且过表达OsLPL2转入lpl2-1中, OE-OsLPL2/lpl2-1转基因阳性植株恢复了lpl2-1平滑表皮及对干旱和盐胁迫的敏感表型。研究结果表明, OsLPL2基因不仅控制水稻表皮细胞形态建成, 而且通过调控气孔密度、气孔开度、根生长发育等在响应植物逆境胁迫过程发挥关键作用。本研究为揭示水稻OsLPL2参与干旱胁迫的应答分子调控机制提供了一定的理论基础。

关键词: 水稻, 抗旱, 耐盐性, SCAR/WAVE复合体, 植物表皮细胞

Abstract:

Drought threatens global agricultural production and limits the prospects for sustainable agricultural development. Plant leaf epidermis plays a vital role in the process of growth, development, and resistance to adversity stress, and water and gas exchange with the external environment. In this study, compared with the wild-type Zhonghua 11 (ZH11), we found that mutants less pronounced lobe epidermal cell 2-1 (lpl2-1) and less pronounced lobe epidermal cell 2-2 (lpl2-2) were more sensitive to drought and salt stress response, and the survival rate after rewatering was extremely significantly reduced, which was less than half of the control. Compared with ZH11, lpl2-1 and lpl2-2 had shorter plant height, shorter root length, significantly increased stomatal density and stomatal openings in the same phyllodes, and the serrated lobe of the epidermal cell margin becomes smoother, and the epidermal cell nesting was not tight, which might result in faster and more water loss of lpl2-1 and lpl2-2 than ZH11. The water loss experiment of separated leaves also proved that the water loss rate of lpl2-1 and lpl2-2 leaves was higher than that of the ZH11 in equal time. Overexpression of OsLPL2 was transferred into lpl2-1, and the OE-OsLPL2/lpl2-1 transgenic positive plants recovered the smooth epidermis of lpl2-1 and the sensitive phenotype to drought and salt stress. These results showed that OsLPL2 gene not only controlled the microfilament synthesis and morphogenesis of rice epidermal cells, but also played a key role in response to plant stress by regulating stomatal density, stomatal conductance, and root growth and development. This study provides a theoretical basis for revealing the molecular regulation mechanism of OsLPL2 in response to drought stress in rice.

Key words: rice, drought tolerance, salt resistance, SCAR/WAVE complex, plant leaf epidermis

表1

本研究使用的引物序列"

引物名称
Primer name		 引物序列
Primer sequence (5°-3°)		 酶切位点
Restriction enzyme cutting site
p1301-LPL2F
p1301-LPL2R		 GGTACCATGGCCATCCCCGTCGAGG
ACTAGTTCAAGTAGCTCTCTGTGGCAATG		 Kpn I /Spe I
RT-LPL2-F
RT-LPL2-R		 CCTGGAATCATCTCGTGTTATCC
GCTGGGCAGAGTCATTGTAAAG
TUB-F
TUB-R		 TTTCACTCTTGGTGTGAAGCAGAT
GACTTCCTTCACGATTTCATCGTAA

表2

OsLPL2在不同物种中的基因编号及功能注释"

物种
Specie		 同源基因
Orthologous gene		 功能预测
Putative function
水稻Oryza sativa LOC_Os03g05020 PIR, putative, expressed
拟南芥Arabidopsis thaliana AT5G18410 Transcription activators
玉米Zea mays GRMZM2G113174 PIROGI; similar to KLK/PIR/PIR121/PIRP (KLUNKER, PIROGI)
玉米Zea mays GRMZM2G170567 PIROGI; similar to KLK/PIR/PIR121/PIRP (KLUNKER, PIROGI)
二穗短柄草Brachypodium distachyum Bradi1g75470 Protein PIR
高粱Sorghum bicolor Sb01g047340 Protein PIR
白杨树Populus trichocarpa POPTR_0013s04800 PIR121; transcription activator
葡萄Vitis vinifera GSVIVG00026355001 GSVIVG00026355001
人类Homo sapiens HsPIR121/CYFIP2 KLK/PIR/PIR121/PIRP (KLUNKER, PIROGI)

图1

OsLPL2/PIR在不同物种中的同源性分析 AtPIR是拟南芥中PIR蛋白, OsLPL2是水稻中PIR的同源蛋白, HsPIR121是人类中PIR的同源蛋白, ZmBRK2是玉米中PIR的同源蛋白, 同源性序列分析表明, 氨基酸相似度高达72.9%。OsLPL2尤其与玉米中BRK2同源蛋白相似性高达93%。"

图2

lpl2-1幼苗期植株较矮, 根长及根细胞长度变短 A~D: 分别生长1周、2周、3周和4周的lpl2-1植株; E, F: 分别是中花11与lpl2-1生长1周的根细胞扫描图; G, H: 中花11与lpl2-1生长4周的根细胞扫描图; I: 测量1~4周的植株高度, N = 30; J: 测量1~4周的根长, N = 30, 1、2、3和4分别表示第1、2、3和4周。A~D: 标尺为1 cm; E~H: 标尺为100 μm。*表示与对照有显著性差异, P < 0.05; **表示与对照有极显著性差异(P < 0.01)。"

图3

lpl2-2突变体叶片表皮形态 A: ZH11幼叶表皮细胞, 细胞边缘锯齿状凸出(黑色斜箭头所示); B: lpl2-2幼叶表皮细胞变平滑(黑色斜箭头所示), 发育畸形的气孔, 类似于保卫细胞母细胞(箭头所示); C: 幼叶叶片气孔开度统计, N = 100; D: ZH11成熟叶气孔结构放大图, 成熟气孔由2个副卫细胞(SC)和2个保卫细胞(GC)构成, SP表示气孔开度; E: lpl2-2成熟叶气孔结构放大图, SL表示气孔长度, SW表示气孔宽度; F: 成熟叶叶片气孔开度统计, N = 100。G, H: ZH11与lpl2-2第4、5叶和剑叶、次剑叶气孔密度, N > 3000; A~E: 标尺为20 μm。**表示极显著性差异(P < 0.01)。"

图4

lpl2对干旱胁迫更敏感 A, E: 干旱处理前的ZH11和lpl2-1; B, F: 干旱后的ZH11和lpl2-1; C、G: 复水后的ZH11和lpl2-1; I: 干旱处理前的ZH11和lpl2-2; J: 干旱后的ZH11和lpl2-2; K: 复水后的ZH11和lpl2-2; D, H, L: 成活率。A~D, I~L: 苗期干旱, 标尺为10 cm; E-H, 抽穗期干旱, 标尺为20 cm。每次实验都定苗50株, 3次生物学重复实验。D、H、L统计N = 150, **表示极显著性差异(P < 0.01)。"

图5

lpl2对盐胁迫更敏感 A: ZH11与lpl2-1幼苗; B: 不同浓度NaCl处理, lpl2-1响应比对照更敏感; C: 浇水恢复后, lpl2-1成活植株明显少于CK; D: ZH11与lpl2-2幼苗; E: 不同浓度NaCl处理, lpl2-2响应比对照更敏感; F: 浇水恢复后, lpl2-2成活植株明显少于CK; A~F, 标尺为10 cm, 每次实验都定苗25株, 4次生物学重复。G, H: lpl2-1、lpl2-2和ZH11成活率统计, N = 100, **表示极显著性差异(P < 0.01)。"

图6

ZH11与lpl2-1和lpl2-2幼叶与成熟叶离体叶片失水率 A: 幼苗期ZH11与lpl2离体叶片失水率对比, 第4、第5叶; B: 抽穗期ZH11与lpl2离体叶片失水率对比, 剑叶和次剑叶。N = 50, **表示与对照有极显著性差异(P < 0.01)。lpl2-2和lpl2-1气孔密度及失水率无显著差异, 但是lpl2与CK差异极显著。"

图7

OE-OsLPL2/lpl2-1恢复了lpl2-1对干旱和盐胁迫耐受性 A: 干旱处理前幼苗期ZH11和Com #3; B: 干旱后的ZH11和Com #3; C: 复水后的ZH11和Com #3; D: 干旱复水后成活率统计, N = 150, 3次生物学重复, 每次定苗50株; E~H: 分别是ZH11与Com #3不同浓度NaCl处理前, 处理6 d、8 d及恢复5 d后植株表型; I: 浇水恢复后, Com #3成活率与ZH11无显著差异, 标尺为10 cm, N = 100, 4次生物学重复, 每次定苗25株, NS: not significant。"

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