中国水稻科学 ›› 2021, Vol. 35 ›› Issue (5): 439-448.DOI: 10.16819/j.1001-7216.2021.210305
冯志明1,2, 王广达1, 赵剑华1, 居冉1, 李梦臣1, 高鹏1, 胡珂鸣1,2, 陈宗祥1,2, 左示敏1,2,*()
收稿日期:
2021-03-06
修回日期:
2021-03-25
出版日期:
2021-09-10
发布日期:
2021-09-10
通讯作者:
左示敏
基金资助:
Zhiming FENG1,2, Guangda WANG1, Jianhua ZHAO1, Ran JU1, Mengchen LI1, Peng GAO1, Keming HU1,2, Zongxiang CHEN1,2, Shimin ZUO1,2,*()
Received:
2021-03-06
Revised:
2021-03-25
Online:
2021-09-10
Published:
2021-09-10
Contact:
Shimin ZUO
摘要:
【目的】明确水稻富含半胱氨酸类受体激酶(Cysteine-rich receptor-like kinases,CRK)家族基因对纹枯病菌侵染和植物激素的响应特征,是解析CRK在水稻纹枯病抗性中的功能的重要前期工作。【方法】利用生物信息学方法构建了水稻45个CRK的系统发育树,并利用qPCR分析了它们对纹枯病菌和对植物激素乙烯(Ethylene,ET)、茉莉酸(Jasmonic acid,JA)、水杨酸(Salicylic acid,SA)和细胞分裂素(Cytokinin,CK)等的响应特征,以及在水稻不同组织中的表达模式。【结果】水稻CRK家族可分为4个组,在染色体上成簇或紧密分布的CRK大部分来自同一组或同一分支。41个CRK响应纹枯病菌侵染,其中17个响应强烈;结合组织表达模式,发现这17个CRK基因中,CRK15、CRK23、CRK24、CRK26、CRK27、CRK28、CRK29、CRK30、CRK31和CRK33等10个基因在叶鞘和叶片中表达较强,暗示这些基因可能参与了对纹枯病的抗性;大部分同一分支的两个同源性较高的CRK对纹枯病菌的响应特征类似, 表明这些CRK基因在调控纹枯病抗性上可能存在功能冗余。40个CRK对3种或4种植物激素均有响应,它们对不同激素的响应存在差异性,说明CRK可能广泛参与这些激素介导的防御信号途径;对JA和SA响应相反的有17个,对JA和SA、ET和JA、ET和SA响应类似的分别有21、21、23个。这不仅反映了ET、JA、SA信号途径间的协同和拮抗作用,也说明这些基因可能参与ET、JA和SA间的交互作用。【结论】鉴定到一些可能参与调控水稻纹枯病抗性的CRK基因,且它们可能在植物激素介导的防御途径中起作用。这为我们进一步探索CRK在调控水稻纹枯病抗性上的功能提供了科学线索。
冯志明, 王广达, 赵剑华, 居冉, 李梦臣, 高鹏, 胡珂鸣, 陈宗祥, 左示敏. 水稻富含半胱氨酸类受体激酶家族基因对纹枯病菌和植物激素的响应特征分析[J]. 中国水稻科学, 2021, 35(5): 439-448.
Zhiming FENG, Guangda WANG, Jianhua ZHAO, Ran JU, Mengchen LI, Peng GAO, Keming HU, Zongxiang CHEN, Shimin ZUO. Response Characteristics of Rice Cysteine-rich Receptor-like Kinases Family Genes to Rhizoctonia solani and Plant Hormones[J]. Chinese Journal OF Rice Science, 2021, 35(5): 439-448.
基因号 Gene ID | 基因名称 Gene name | 蛋白大小 Protein length/aa | 蛋白分子量 MW/kDa | 等电点 pI | 引物序列 Primer sequence | 产物大小 Product size/bp | |
---|---|---|---|---|---|---|---|
LOC_Os01g23970 | OsCRK1 | 423 | 45.60 | 7.57 | F: R: | TTTGGCTCCTACGGTTCTGACC GCTAGTCCAGTGCTCCCATGTAAC | 66 |
LOC_Os01g38910 | OsCRK2 | 196 | 21.37 | 5.77 | F: R: | GACAATGGCGTTGCCTCTTCTTC AACCGAATGGCATCATCCGCATC | 71 |
LOC_Os02g06160 | OsCRK3 | 649 | 71.16 | 8.71 | F: R: | TGCGAGGTCATCGAATTCTGTCG ACAACATCTGGAGTCTGCTTGC | 101 |
LOC_Os02g12130 | OsCRK4 | 179 | 20.06 | 8.60 | F: R: | GCATGGAAGACAAGTCGGAAGG TCCAGTGGGCATTACTCCCTTG | 137 |
LOC_Os02g48080 | OsCRK5 | 426 | 48.08 | 7.47 | F: R: | ACAAACCCAGATGACATTCAGAGC ACCTTCGGGAAGAACTCCCTTG | 144 |
LOC_Os03g31260 | OsCRK6 | 314 | 35.25 | 9.51 | F: R: | TGCAAGCGTTAGTGGAACACTGG CACCTCGTCCAGACTCACTCATTC | 140 |
LOC_Os04g01860 | OsCRK7 | 217 | 23.41 | 4.38 | F: R: | AAGGGCCGTTTGACTACGAACC TGAATCCCTGTGCAGAATTTGGG | 70 |
LOC_Os04g03370 | OsCRK8 | 328 | 36.04 | 7.00 | F: R: | AGCAGTGGTTCACTCAGCTTTG CATGGTCCAGTGCTCCCATACAAG | 69 |
LOC_Os04g03530 | OsCRK9 | 430 | 49.02 | 10.07 | F: R: | GCTACACGTCATCTTTGGCTGTG AGCAAGGACGATGTCTCAGTGTG | 79 |
LOC_Os04g06090 | OsCRK10 | 183 | 21.23 | 10.58 | F: R: | TCGGTATCGTCCTCCTCGAGATTG TTGGTGTTCCGGTGCCATTACC | 62 |
LOC_Os04g06210 | OsCRK11 | 196 | 22.13 | 4.57 | F: R: | TTCAGGGACGCCACGATCTACTAC GCCGGTCACGTTGTTCATGTTC | 120 |
LOC_Os04g09780 | OsCRK12 | 278 | 28.83 | 5.26 | F: R: | TACGAGTACCTCCCTAACAGAAGC ATTGAGCCCTCTTTCTGGAATCG | 64 |
LOC_Os04g25060 | OsCRK13 | 262 | 27.29 | 6.94 | F: R: | CTGCACACACATCGATCGCTAATC TTGCAATGGCAGTGATCGTTAAGC | 60 |
LOC_Os04g25650 | OsCRK14 | 281 | 29.18 | 8.02 | F: R: | CTTACATGGCCTACGAGGTACAAG CGAGCTTTGCGTTCATCTCTTCG | 145 |
LOC_Os04g30030 | OsCRK15 | 463 | 52.89 | 6.88 | F: R: | CCCACCGAATTGTTGGAACCTATG TCCTTCGTCCGCTCACAATCTC | 126 |
LOC_Os04g54190 | OsCRK16 | 379 | 42.66 | 8.20 | F: R: | ACTCACCTGGCCAGAGAGATAC GCTCCCACTCTTGATGAAGGTAAC | 77 |
LOC_Os04g56360 | OsCRK17 | 494 | 55.00 | 7.21 | F: R: | GCTCCTACTACAGCGAGCAATC TGGTCCAGTGCTCCCATATAATGC | 60 |
LOC_Os04g56430 | OsCRK18 | 258 | 27.31 | 4.73 | F: R: | GCTTGGGTGATGGCAAGAACAAC AGCTGCCCTGAACCTTTGATGG | 76 |
LOC_Os05g34950 | OsCRK19 | 644 | 68.40 | 6.89 | F: R: | TCCTGGCCTTCATCGTGATTCTC TGAAATACTCGGGTACCTCTGACG | 135 |
LOC_Os06g30130 | OsCRK20 | 434 | 47.86 | 7.04 | F: R: | CGAAGACCTCTTAACCCTTGTGTG ATCACGTCGCTCCACGAGAAAC | 102 |
LOC_Os07g26180 | OsCRK21 | 163 | 18.63 | 7.57 | F: R: | TCGCCGGAACATATGGATACATGG GCTGAACACGTCCGATTTGATTG | 77 |
LOC_Os07g35290 | OsCRK22 | 348 | 37.45 | 8.90 | F: R: | AAGTTCAGGCACTGACGAACAG TGGAGCACTGAATCCAATTGAGC | 64 |
LOC_Os07g35310 | OsCRK23 | 659 | 71.05 | 6.81 | F: R: | TGGGAGCTTTATCTTGTGTTGGTG TCTCACACAGTGAGTTTAGAAGCG | 84 |
LOC_Os07g35580 | OsCRK24 | 696 | 74.78 | 7.90 | F: R: | CGTATACAGCTTCGGTGTGTTGC TTGCCCAATGTTCCCATACGAG | 116 |
LOC_Os07g35650 | OsCRK25 | 682 | 74.58 | 6.98 | F: R: | ACTCCTCTGAATGTTGTCGATGGC TGCCTGCAGTGATGGTTGAGAC | 63 |
LOC_Os07g35660 | OsCRK26 | 711 | 77.02 | 6.70 | F: R: | TGCCTACTCGCTGACGTGCTACA CTCTTGATTCCGTGGGGAGCAAAGC | 134 |
LOC_Os07g35680 | OsCRK27 | 698 | 75.47 | 6.26 | F: R: | AGCTATGCACAAGTTGCTGACG TAGGGCGCATGGAAGCTTTAGG | 115 |
LOC_Os07g35690 | OsCRK28 | 695 | 76.47 | 6.59 | F: R: | AGATGCGACCTATGATGGAGCAAG AGTTTAGGCAACGGCCCATCAC | 64 |
LOC_Os07g35700 | OsCRK29 | 645 | 73.68 | 7.23 | F: R: | AGAACAGACGTTTCTGCCCTACG AACCATACCGAACCCTCCTCTG | 130 |
LOC_Os07g43560 | OsCRK30 | 677 | 73.68 | 6.94 | F: R: | ATAATCATGGACGTGGCGAAGGG TGATGTCCCTGTGGTAGATCGC | 76 |
因号 Gene ID | 基因名称 Gene name | 蛋白大小 Protein length/aa | 蛋白分子量 MW/kDa | 等电点 pI | 引物序列 Primer sequence | 产物大小 Product size/bp | |
LOC_Os08g25430 | OsCRK31 | 591 | 65.53 | 5.15 | F: R: | AAGGTGCAAGAACAAGAAGGTTGC AAGCAGTCCCTCAGCTATCCCATC | 80 |
LOC_Os09g16590 | OsCRK32 | 333 | 37.19 | 6.85 | F: R: | AGGAAGGCACTTATAGGCCAGAGC ACATCCCTGTCCTTCTCAAATGGG | 101 |
LOC_Os09g16950 | OsCRK33 | 516 | 56.46 | 8.20 | F: R: | TGCGTGGCGCATTCAAGTGTAG TCTCTCGCTCTGCCAACCTTTG | 104 |
LOC_Os11g11780 | OsCRK34 | 305 | 34.70 | 6.99 | F: R: | CAGTGGCCAAAGAGGTTTGAACG TGGCCACTATCAGCAGTTTGCC | 92 |
LOC_Os11g26140 | OsCRK35 | 408 | 45.39 | 7.95 | F: R: | GGTGAACACCATGTAGCAGGTC ACCTCTGCACTTTGGACAGCAG | 64 |
LOC_Os11g29510 | OsCRK36 | 307 | 34.85 | 8.98 | F: R: | CAAACCAATTGCTGTCCCGATGC GGAGCTTGAGCTTAGATGGTGTGG | 95 |
LOC_Os11g39450 | OsCRK37 | 456 | 51.83 | 7.52 | F: R: | CAGTCACATTGCTGGAACATACGG TCCTTCCCGTGACAATCTCTAGG | 122 |
LOC_Os11g44690 | OsCRK38 | 479 | 53.95 | 6.82 | F: R: | GATATGCAGCTCTTCGCCAACTTC CAGTGAGTTTAGAAGCGCACCAAC | 134 |
LOC_Os11g44860 | OsCRK39 | 527 | 59.22 | 9.85 | F: R: | CGTCAACGCTAACAGCACACAC TTGATCGAGTTGCCATCCTCAAG | 78 |
LOC_Os12g14480 | OsCRK40 | 364 | 41.15 | 9.99 | F: R: | GTGGAGGCCCTAAGAAGAAAGC AGCTCTGGTAAGACTACCTGTGAC | 100 |
LOC_Os12g14610 | OsCRK41 | 485 | 53.55 | 8.20 | F: R: | TGCAAAGCTTCTCCAGGAAATCAC TTCGCAGCAATTTCTGTCCCATC | 117 |
LOC_Os12g41270 | OsCRK42 | 897 | 95.79 | 8.42 | F: R: | AGGACCTCGAAGGATTCAGAGC AGATGCCACTCCAAGCAAGATAGC | 68 |
LOC_Os12g41490 | OsCRK43 | 687 | 74.42 | 5.86 | F: R: | ATGTGCGGCTGGTGTTGGAATC ACCAATTACTCCCTCCGTGTTGG | 73 |
LOC_Os12g41510 | OsCRK44 | 839 | 91.65 | 6.31 | F: R: | GGATGGAGAGATCACAAGAAAGCC CACCATGAGAGAATGGACCACAAC | 95 |
LOC_Os12g41530 | OsCRK45 | 682 | 74.26 | 5.68 | F: R: | GTTGCTGATGGCACTGGAACTC TGCATAACCTCTGTGATGTGCAG | 63 |
表1 水稻OsCRK基因家族及其荧光定量PCR引物
Table 1 OsCRK gene family members and their qPCR primers.
基因号 Gene ID | 基因名称 Gene name | 蛋白大小 Protein length/aa | 蛋白分子量 MW/kDa | 等电点 pI | 引物序列 Primer sequence | 产物大小 Product size/bp | |
---|---|---|---|---|---|---|---|
LOC_Os01g23970 | OsCRK1 | 423 | 45.60 | 7.57 | F: R: | TTTGGCTCCTACGGTTCTGACC GCTAGTCCAGTGCTCCCATGTAAC | 66 |
LOC_Os01g38910 | OsCRK2 | 196 | 21.37 | 5.77 | F: R: | GACAATGGCGTTGCCTCTTCTTC AACCGAATGGCATCATCCGCATC | 71 |
LOC_Os02g06160 | OsCRK3 | 649 | 71.16 | 8.71 | F: R: | TGCGAGGTCATCGAATTCTGTCG ACAACATCTGGAGTCTGCTTGC | 101 |
LOC_Os02g12130 | OsCRK4 | 179 | 20.06 | 8.60 | F: R: | GCATGGAAGACAAGTCGGAAGG TCCAGTGGGCATTACTCCCTTG | 137 |
LOC_Os02g48080 | OsCRK5 | 426 | 48.08 | 7.47 | F: R: | ACAAACCCAGATGACATTCAGAGC ACCTTCGGGAAGAACTCCCTTG | 144 |
LOC_Os03g31260 | OsCRK6 | 314 | 35.25 | 9.51 | F: R: | TGCAAGCGTTAGTGGAACACTGG CACCTCGTCCAGACTCACTCATTC | 140 |
LOC_Os04g01860 | OsCRK7 | 217 | 23.41 | 4.38 | F: R: | AAGGGCCGTTTGACTACGAACC TGAATCCCTGTGCAGAATTTGGG | 70 |
LOC_Os04g03370 | OsCRK8 | 328 | 36.04 | 7.00 | F: R: | AGCAGTGGTTCACTCAGCTTTG CATGGTCCAGTGCTCCCATACAAG | 69 |
LOC_Os04g03530 | OsCRK9 | 430 | 49.02 | 10.07 | F: R: | GCTACACGTCATCTTTGGCTGTG AGCAAGGACGATGTCTCAGTGTG | 79 |
LOC_Os04g06090 | OsCRK10 | 183 | 21.23 | 10.58 | F: R: | TCGGTATCGTCCTCCTCGAGATTG TTGGTGTTCCGGTGCCATTACC | 62 |
LOC_Os04g06210 | OsCRK11 | 196 | 22.13 | 4.57 | F: R: | TTCAGGGACGCCACGATCTACTAC GCCGGTCACGTTGTTCATGTTC | 120 |
LOC_Os04g09780 | OsCRK12 | 278 | 28.83 | 5.26 | F: R: | TACGAGTACCTCCCTAACAGAAGC ATTGAGCCCTCTTTCTGGAATCG | 64 |
LOC_Os04g25060 | OsCRK13 | 262 | 27.29 | 6.94 | F: R: | CTGCACACACATCGATCGCTAATC TTGCAATGGCAGTGATCGTTAAGC | 60 |
LOC_Os04g25650 | OsCRK14 | 281 | 29.18 | 8.02 | F: R: | CTTACATGGCCTACGAGGTACAAG CGAGCTTTGCGTTCATCTCTTCG | 145 |
LOC_Os04g30030 | OsCRK15 | 463 | 52.89 | 6.88 | F: R: | CCCACCGAATTGTTGGAACCTATG TCCTTCGTCCGCTCACAATCTC | 126 |
LOC_Os04g54190 | OsCRK16 | 379 | 42.66 | 8.20 | F: R: | ACTCACCTGGCCAGAGAGATAC GCTCCCACTCTTGATGAAGGTAAC | 77 |
LOC_Os04g56360 | OsCRK17 | 494 | 55.00 | 7.21 | F: R: | GCTCCTACTACAGCGAGCAATC TGGTCCAGTGCTCCCATATAATGC | 60 |
LOC_Os04g56430 | OsCRK18 | 258 | 27.31 | 4.73 | F: R: | GCTTGGGTGATGGCAAGAACAAC AGCTGCCCTGAACCTTTGATGG | 76 |
LOC_Os05g34950 | OsCRK19 | 644 | 68.40 | 6.89 | F: R: | TCCTGGCCTTCATCGTGATTCTC TGAAATACTCGGGTACCTCTGACG | 135 |
LOC_Os06g30130 | OsCRK20 | 434 | 47.86 | 7.04 | F: R: | CGAAGACCTCTTAACCCTTGTGTG ATCACGTCGCTCCACGAGAAAC | 102 |
LOC_Os07g26180 | OsCRK21 | 163 | 18.63 | 7.57 | F: R: | TCGCCGGAACATATGGATACATGG GCTGAACACGTCCGATTTGATTG | 77 |
LOC_Os07g35290 | OsCRK22 | 348 | 37.45 | 8.90 | F: R: | AAGTTCAGGCACTGACGAACAG TGGAGCACTGAATCCAATTGAGC | 64 |
LOC_Os07g35310 | OsCRK23 | 659 | 71.05 | 6.81 | F: R: | TGGGAGCTTTATCTTGTGTTGGTG TCTCACACAGTGAGTTTAGAAGCG | 84 |
LOC_Os07g35580 | OsCRK24 | 696 | 74.78 | 7.90 | F: R: | CGTATACAGCTTCGGTGTGTTGC TTGCCCAATGTTCCCATACGAG | 116 |
LOC_Os07g35650 | OsCRK25 | 682 | 74.58 | 6.98 | F: R: | ACTCCTCTGAATGTTGTCGATGGC TGCCTGCAGTGATGGTTGAGAC | 63 |
LOC_Os07g35660 | OsCRK26 | 711 | 77.02 | 6.70 | F: R: | TGCCTACTCGCTGACGTGCTACA CTCTTGATTCCGTGGGGAGCAAAGC | 134 |
LOC_Os07g35680 | OsCRK27 | 698 | 75.47 | 6.26 | F: R: | AGCTATGCACAAGTTGCTGACG TAGGGCGCATGGAAGCTTTAGG | 115 |
LOC_Os07g35690 | OsCRK28 | 695 | 76.47 | 6.59 | F: R: | AGATGCGACCTATGATGGAGCAAG AGTTTAGGCAACGGCCCATCAC | 64 |
LOC_Os07g35700 | OsCRK29 | 645 | 73.68 | 7.23 | F: R: | AGAACAGACGTTTCTGCCCTACG AACCATACCGAACCCTCCTCTG | 130 |
LOC_Os07g43560 | OsCRK30 | 677 | 73.68 | 6.94 | F: R: | ATAATCATGGACGTGGCGAAGGG TGATGTCCCTGTGGTAGATCGC | 76 |
因号 Gene ID | 基因名称 Gene name | 蛋白大小 Protein length/aa | 蛋白分子量 MW/kDa | 等电点 pI | 引物序列 Primer sequence | 产物大小 Product size/bp | |
LOC_Os08g25430 | OsCRK31 | 591 | 65.53 | 5.15 | F: R: | AAGGTGCAAGAACAAGAAGGTTGC AAGCAGTCCCTCAGCTATCCCATC | 80 |
LOC_Os09g16590 | OsCRK32 | 333 | 37.19 | 6.85 | F: R: | AGGAAGGCACTTATAGGCCAGAGC ACATCCCTGTCCTTCTCAAATGGG | 101 |
LOC_Os09g16950 | OsCRK33 | 516 | 56.46 | 8.20 | F: R: | TGCGTGGCGCATTCAAGTGTAG TCTCTCGCTCTGCCAACCTTTG | 104 |
LOC_Os11g11780 | OsCRK34 | 305 | 34.70 | 6.99 | F: R: | CAGTGGCCAAAGAGGTTTGAACG TGGCCACTATCAGCAGTTTGCC | 92 |
LOC_Os11g26140 | OsCRK35 | 408 | 45.39 | 7.95 | F: R: | GGTGAACACCATGTAGCAGGTC ACCTCTGCACTTTGGACAGCAG | 64 |
LOC_Os11g29510 | OsCRK36 | 307 | 34.85 | 8.98 | F: R: | CAAACCAATTGCTGTCCCGATGC GGAGCTTGAGCTTAGATGGTGTGG | 95 |
LOC_Os11g39450 | OsCRK37 | 456 | 51.83 | 7.52 | F: R: | CAGTCACATTGCTGGAACATACGG TCCTTCCCGTGACAATCTCTAGG | 122 |
LOC_Os11g44690 | OsCRK38 | 479 | 53.95 | 6.82 | F: R: | GATATGCAGCTCTTCGCCAACTTC CAGTGAGTTTAGAAGCGCACCAAC | 134 |
LOC_Os11g44860 | OsCRK39 | 527 | 59.22 | 9.85 | F: R: | CGTCAACGCTAACAGCACACAC TTGATCGAGTTGCCATCCTCAAG | 78 |
LOC_Os12g14480 | OsCRK40 | 364 | 41.15 | 9.99 | F: R: | GTGGAGGCCCTAAGAAGAAAGC AGCTCTGGTAAGACTACCTGTGAC | 100 |
LOC_Os12g14610 | OsCRK41 | 485 | 53.55 | 8.20 | F: R: | TGCAAAGCTTCTCCAGGAAATCAC TTCGCAGCAATTTCTGTCCCATC | 117 |
LOC_Os12g41270 | OsCRK42 | 897 | 95.79 | 8.42 | F: R: | AGGACCTCGAAGGATTCAGAGC AGATGCCACTCCAAGCAAGATAGC | 68 |
LOC_Os12g41490 | OsCRK43 | 687 | 74.42 | 5.86 | F: R: | ATGTGCGGCTGGTGTTGGAATC ACCAATTACTCCCTCCGTGTTGG | 73 |
LOC_Os12g41510 | OsCRK44 | 839 | 91.65 | 6.31 | F: R: | GGATGGAGAGATCACAAGAAAGCC CACCATGAGAGAATGGACCACAAC | 95 |
LOC_Os12g41530 | OsCRK45 | 682 | 74.26 | 5.68 | F: R: | GTTGCTGATGGCACTGGAACTC TGCATAACCTCTGTGATGTGCAG | 63 |
图2 水稻OsCRK对纹枯病菌和激素ET、JA、SA和CK的响应基因表达变化倍数(FC)为纹枯病菌接种后或各激素处理后相对于未处理对照的每个时间点的表达比值。热图是基于Log2 FC生成。红色和蓝色分别表示基因上调和下调。
Fig. 2. Responses of rice OsCRK to R. solani, ET, JA, SA and CK.Fold change(FC) is expressed as the mean expression value of R. solani-infected or plant hormones-treated relative to mock-treated at each time point. The heat map was generated using Log2 FC. Red and blue colors indicate up- and down-regulated genes, respectively.
图3 OsCRK基因在水稻不同组织中的表达模式R,S,L,LS和P分别表示根,茎,叶,叶鞘和穗组织。
Fig. 3. The expression pattern of OsCRK in different rice tissues. R, S, L, LS and P indicate root, stem, leaf, leaf sheath and panicle, respectively.
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