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作物学报 ›› 2021, Vol. 47 ›› Issue (2): 197-209.doi: 10.3724/SP.J.1006.2021.02034

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

OsPAL2;3对水稻化感抑制稗草能力的调控作用

李兰兰(), 母丹, 严雪, 杨陆可, 林文雄*(), 方长旬*()   

  1. 福建农林大学生命科学学院农业生态研究所 / 福建省农业生态过程与安全监控重点实验室, 福建福州 350002
  • 收稿日期:2020-05-14 接受日期:2020-08-19 出版日期:2021-02-12 网络出版日期:2020-11-19
  • 通讯作者: 林文雄,方长旬
  • 作者简介:E-mail: 1244311185@qq.com
  • 基金资助:
    国家自然科学基金项目(31871556);福建农林大学杰出青年科研人才计划项目(xjq201805);福建农林大学科技创新专项基金项目(CXZX2018042)

Effect of OsPAL2;3 in regulation of rice allopathic inhibition on barnyardgrass (Echinochloa crusgalli L.)

LI Lan-Lan(), MU Dan, YAN Xue, YANG Lu-Ke, LIN Wen-Xiong*(), FANG Chang-Xun*()   

  1. Institute of Agroecology, School of Life Sciences, Fujian Agriculture and Forestry University / Fujian Provincial Key Laboratory of Agroecological Processing and Safety Monitoring, Fuzhou 350002, Fujian, China
  • Received:2020-05-14 Accepted:2020-08-19 Published:2021-02-12 Published online:2020-11-19
  • Contact: LIN Wen-Xiong,FANG Chang-Xun
  • Supported by:
    National Natural Science Foundation of China(31871556);Outstanding Youth Scientific Fund of Fujian Agriculture and Forestry University(xjq201805);Science and Technology Innovation of Fujian Agriculture and Forestry University(CXZX2018042)

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摘要： 苯丙氨酸解氨酶(phenylalanine ammonia-lyase, PAL, EC 4.3.1.5)是调控酚酸类化感物质合成的关键酶。PAL在水稻中为多基因家族, 化感水稻 PI312777 和非化感水稻 Lemont 中, 相同 PAL 基因成员的启动子组成序列均存在差异, 并以 OsPAL2;3、OsPAL2;4 基因启动子序列的差异最大, 且 PI321777 的 OsPAL2;3 基因启动子的驱动活性高于Lemont。过表达 OsPAL2;3 基因, 使 PI312777 和 Lemont 的抑草率分别提高 11.11%和 5.56%。过表达 OsPAL2;3 同时增强 OsPAL2;3、OsC4H、OsCCA、OsCOL、OsOMT 等基因的表达, 增加水稻的原儿茶酸和香草酸含量。Co-IP 结合质谱鉴定结果显示, OsPAL2;3 蛋白与转酮醇酶、碳酸酐酶、果糖二磷酸醛缩酶、ATP 合酶 α 亚基、ATP 合酶 β 亚基等相互作用, 调控水稻的苯丙氨酸代谢途径。本研究表明, OsPAL2;3 的转录水平高低是 P1312777 和 Lemont 化感抑草能力差异的原因之一; OsPAL2;3 与多个蛋白互作共同调控酚酸类化合物合成, 该基因可用于提高水稻化感抑草能力的育种研究。

关键词: 水稻, 化感作用, PAL基因, 启动子, 酚酸, 蛋白互作

Abstract:

Phenylalanine ammonia-lyase (PAL, EC 4.3.1.5) is the key enzyme in regulation of the synthesis of phenolic acid allelochemicals. PAL gene in rice belongs to a multigene family. In allelopathic accession rice PI312777 and non-allelopathic rice accession Lemont, the promoter sequences of the same PAL gene member were different, and there was the largest difference in OsPAL2;3 and OsPAL2;4 gene promoter sequence. Gene promoter of OsPAL2;3 from PI3127777 showed higher activity than the corresponding promoter from Lemont. Overexpression of OsPAL2;3 in PI312777 and Lemont resulted in increasing allelopathic inhibition on barnyardgrass (Echinochloa crusgalli L.), and the inhibitory ratios was increased by 11.11% in PI312777 and 5.56% in Lemont. Gene expression level of OsPAL2;3, OsC4H, OsCCA, OsCOL, and OsOMT was up-regulated in the OsPAL2;3-overexpressed transgenic rice compared with that of wild-type rice, and the contents of protocatechuic acid and vanillic acid were also increased. The results from Co-IP combined with mass spectrometry showed that transketolase, carbonic anhydrase, fructose-bisphospate aldolase isozyme, ATP synthase subunit alpha and ATP synthase subunit beta were interacted with OsPAL2;3 protein, resulting in regulating the phenylalanine pathway in rice. Our study indicated that the transcriptional activity of OsPAL2;3 contributed to the alleloapthic activity between PI312777 and Lemont, OsPAL2;3 was interacted with a couple of proteins to jointly regulate the synthesis of phenolic acids, and OsPAL2;3 could be considered as a candidate gene to improve the allelopathy of rice in breeding.

Key words: rice, allelopathy, PAL gene, promoter, phenolic acid, protein interaction

表1

本研究中所用到的引物"

功能
Function		 基因名称
Gene name		 正向引物
Forward primer (5'-3')		 反向引物
Reverse primer (5'-3')
启动子克隆
Promoter cloning		 OsPAL2;1 GGTAGTGCTAGATATGAAGGGCTGC CACGAGGAGAAGAGAGGATTCGAT
OsPAL2;2 GTTTTCCGAACGGGTAACCGATGTG CTCACAGATGCAGTAGTAGCACACG
OsPAL2;3 GGTACTGGTACCGCCAGTACGCATA AGTTAGCTAGACGGCCGGAGAGAAC
OsPAL2;4 GGGTCATATGTGACACGTCATCACA CAGAAGATGGATCGGTTAACCTAGC
OsPAL4;1 ATGAGCTGTGAGGCCCAGCAAGCAG AACAACGACGAGGAGGAGGAGGAGG
OsPAL4;2 CGTCCCGTAATACAGGGGATTTTGG GAAGCAGAGGGGAGATCGATGTGTA
OsPAL11 TAGAAGATCAGTTCGCTCCCAGGCT TACAGGTCGCCGAGTACGTACGTAT
荧光定量PCR
qRT-PCR		 OsPAL2;3 CCGTGCTCTTTGAGGCTAAC GCTTGTGAGTCAGGTGGTCG
OsC4H ACCGCAGCGTCTCCTTC ACCACCCGAGCATCCAG
OsCOL TGGTGGAGTGCGTGCTG AGGCGTTGGCGTAGATG
OsOMT TGTCCTGTGAAATGGGTG CCTCGGAACAAGAACTG
OsCAD TCGGCGTCGCTAATTTCATCC TCGATGGAAGAACGGGCAGAG
β-actin CTGCGGGTATCCATGAGACT GCAATGCCAGGGAACATAGT
基因过表达载体
Gene overexpression vector		 OsPAL2;3 cgactctagaggatccATGGCGTGCGAGA ACGGTCAG ccatactagtggatccGCAGATGGGCAGGG GCGCGC
转基因水稻PCR鉴定
PCR identification of transgenic rice line		 OsPAL2;3-OX CTGAAGCTCATGTCCTCCACGTTC GATGTTGCCGTCCTCCTTGAAGTC
双分子荧光互补载体
Bimolecular fluorescent complimentary vector		 OsPAL2;3 atggcgcgccactagtATGGCGTGCGAGA ACGGTCAG cacctcctccactagtGCAGATGGGCAGGGG CGCGC
ATP synthase subunit alpha atggcgcgccactagtATGGCAACCCTTCG AGTCGACGA cacctcctccactagtAAGGGAAAACCGTTC GAGTTGTTCC
ATP synthase subunit beta atggcgcgccactagtATGGCGACTCGCCG GGCCCTCT cacctcctccactagtTGAAGCCGACTCCTTG GCGATC
Transketolase atggcgcgccactatATGGCCGCGCACTCC GTCG cacctcctccactagtCAGGCTCTTTGCTGTT GCGAT
Carbonic anhydrase atggcgcgccactagtATGTCGACCGCCGC CGCCGC cacctcctccactagtGGGCTCCCATAAGTCC AAGTTG
Frutose-bisphosphate
aldolase		 atggcgcgccactagtATGGCGTCTGCTAC TCTCCTCAA cacctcctccactagtGTAGACGTAGTTCTTG ACGAACA

图1

PI312777与Lemont的PAL基因成员启动子片段的凝胶电泳检测及部分差异序列 (a) PAL基因启动子片段的凝胶电泳检测; (b) PI312777与Lemont的OsPAL2;3及OsPAL2;4基因启动子的差异序列。"

图2

PI312777与Lemont的OsPAL2;3、OsPAL2;4基因启动子驱动的GFP荧光强度检测"

图3

OsPAL2;3过表达水稻中的eYFP融合蛋白的表达以及eYFP的荧光检测 (a) Western-blotting检测eYFP表达丰度; (b) 激光共聚焦显微镜检测eYFP荧光强度。"

图4

OsPAL2;3过表达水稻与野生型株系的OsPAL、OsOMT、OsC4H、OsCOL、OsCAD基因表达差异"

表2

OsPAL2;3过表达水稻与野生型植株中的酚酸类化合物的含量"

水稻株系
Rice lines		 酚酸含量Contents of phenolic acid (μg g-1)
原儿茶酸Protocatechuic acid 对羟基苯甲酸
p-hydroxybenzoic acid		 香草酸
Vanillic acid		 丁香酸
Syringic acid		 4-香豆酸
4-coumaric acid		 阿魏酸
Ferulic acid		 水杨酸
Salicylic acid		 肉桂酸
Cinnamic acid
PI32777野生型
Wild type of PI312777		 161.78±32.74 2.18±0.14 6.98±1.68 3.19±0.71 25.26±1.65 19.65±0.57 15.34±2.43 8.19±0.62
过表达OsPAL2;3转基因PI312777
OsPAL2;3-OX
transgenic PI312777		 165.11±2.41 1.74±0.41 10.05±0.97 4.77±2.40 31.42±2.30 15.56±0.60 10.64±0.92 8.07±0.20
Lemont野生型
Wild type of Lemont		 18.19±1.38 2.04±0.47 5.47±0.40 6.64±0.35 11.31±0.15 21.07±1.89 15.03±0.13 1.60±0.13
过表达OsPAL2;3转基因Lemont
OsPAL2;3-OX
transgenic Lemont		 26.29±1.81 1.83±0.14 6.45±0.65 3.61±0.37 9.01±4.28 17.64±1.74 17.06±5.61 0.85±0.56

图5

OsPAL2;3过表达水稻及其野生型株系根系分泌液中种植的稗草的生长情况"

表3

OsPAL2;3转基因水稻及其野生型植株根系分泌液培养的稗草形态指标"

处理方式
Treatment		 根长
Root length (cm)		 根长抑制率
IR of root length (%)		 株高
Plant height (cm)		 株高抑制率
IR of plant height (%)		 干重
Dry weight
(g)		 干重抑制率
IR of dry weight (g)
PI312777根系分泌物培养的稗草
Barnyardgrass cultured in the solution with PI31277 root exudates		 9.20±1.43 b 18.73 27.84±3.77 b 14.69 0.12±0.02 b 33.33
过表达OsPAL2;3转基因PI312777根系分泌物培养的稗草
Barnyardgrass cultured in the solution with OsPAL2;3-OX PI31277 root exudates		 6.87±2.00 c 39.31 27.57±2.87 b 15.61 0.10±0.02 b 44.44
对照组稗草
Control group of barnyardgrass		 11.32±2.11 a — 32.67±3.18 a — 0.18±0.01 a —
Lemont根系分泌物培养的稗草
Barnyardgrass cultured in the solution with Lemont root exudates		 9.94±1.20 b 12.19 28.89±3.19 b 11.57 0.14±0.02 b 22.22
过表达OsPAL2;3转基因Lemont根系分泌物培养的稗草
Barnyardgrass cultured in the solution with OsPAL2;3-OX Lemont root exudates		 10.36±1.28 b 8.48 27.32±3.33 b 16.38 0.13±0.01 b 27.78

图6

OsPAL2;3蛋白在过表达水稻体内的互作蛋白"

表4

OsPAL2;3互作蛋白的鉴定结果"

蛋白登录号
Protein accession number		 蛋白名称
Protein name		 全部肽段
Peptide count		 匹配的肽段
Unique peptide count		 覆盖率
Coverage
(%)		 分子量
Molecular weight (kD)		 等电点
Isoelectric point
LOC_Os02g41670.1 Phenylalanine ammonia-lyase 31 26 36.19 76.58 5.84
LOC_Os10g21268.1 Ribulose bisphosphate carboxylase large chain precursor 29 16 30.99 53.71 6.58
LOC_Os04g16740.1 ATP synthase subunit alpha 15 15 27.22 55.66 5.95
LOC_Os01g45274.3 Carbonic anhydrase, chloroplast precursor 25 14 42.37 28.04 6.99
LOC_Os02g41650.3 Phenylalanine ammonia-lyase 15 13 17.69 77.75 6.04
LOC_Os10g21266.1 ATP synthase subunit beta 14 13 29.92 53.98 5.38
LOC_Os11g07020.1 Fructose-bisphospate aldolase isozyme 11 11 27.06 42.00 6.39
LOC_Os12g10580.1 Ribulose bisphosphate carboxylase large chain precursor 18 10 18.94 56.06 9.04
LOC_Os04g38600.2 Glyceraldehyde-3-phosphate
dehydrogenase		 12 10 24.94 43.01 7.61
LOC_Os04g43800.1 Phenylalanine ammonia-lyase 11 10 13.59 76.93 5.93
LOC_Os06g04270.1 Transketolase, chloroplast precursor 11 10 14.67 80.02 6.12
LOC_Os05g41640.2 Phosphoglycerate kinase protein 10 10 21.28 50.13 6.29
LOC_Os12g37260.1 Lipoxygenase 2.1, chloroplast precursor 10 10 9.00 10.46 5.87

图7

BiFC检测OsPAL2;3及其互作蛋白的有效互作"

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