Chinese Journal OF Rice Science ›› 2019, Vol. 33 ›› Issue (4): 313-322.DOI: 10.16819/j.1001-7216.2019.9043
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Peng XU, Hong WANG, Ranran TU, Qunen LIU, Weixun WU, Xiumin FU, Liyong CAO, Xihong SHEN*()
Received:
2019-04-10
Revised:
2019-05-24
Online:
2019-07-10
Published:
2019-07-10
Contact:
Xihong SHEN
徐鹏, 王宏, 涂燃冉, 刘群恩, 吴玮勋, 傅秀民, 曹立勇, 沈希宏*()
通讯作者:
沈希宏
基金资助:
CLC Number:
Peng XU, Hong WANG, Ranran TU, Qunen LIU, Weixun WU, Xiumin FU, Liyong CAO, Xihong SHEN. Orientation Improvement of Blast Resistance in Rice via CRISPR/Cas9 System[J]. Chinese Journal OF Rice Science, 2019, 33(4): 313-322.
徐鹏, 王宏, 涂燃冉, 刘群恩, 吴玮勋, 傅秀民, 曹立勇, 沈希宏. 利用CRISPR/Cas9系统定向改良水稻稻瘟病抗性[J]. 中国水稻科学, 2019, 33(4): 313-322.
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URL: http://www.ricesci.cn/EN/10.16819/j.1001-7216.2019.9043
Fig. 1. Schematic diagram of the targeted sites in Pita, Pi21 and ERF922. The underlined letters are the initiation codons. The bold letters are the protospacer adjacent motif (PAM) sequences.
引物名称 Primer name | 引物序列 Sequence(5'-3') | 引物名称 Primer name | 引物序列 Sequence(5'-3') | |
---|---|---|---|---|
Pita-g-F | GGCACCATGGCGCCGGCGGTCATTGC | Pi21-F | CTGAATTCCACGGGAATTGC | |
Pita-g-R | AAACGCAATGACCGCCGGCGCCATGG | Pi21-R | CTCCTCTGCGTTCATATTC | |
Pi21-g-F | GGCAGGTATATTGGTCATCTTGGTGG | ERF922-F | GCGACAACATCACGCATCGCC | |
Pi21-g-R | AAACCCACCAAGATGACCAATATACC | ERF922-R | TCAGGCCAAGTGCCTACTGCAA | |
ERF922-g-F | GGCAGTGGACGTGTCTCTGTCGCTGG | Hyg-F | GCTGTTATGCGGCCATTGTC | |
ERF922-g-R | AAACCCAGCGACAGAGACACGTCCAC | Hyg-R | GACGTCTGTCGAGAAGTTTC | |
Pita-F | CAGCCTCTCTGCATGATTTC | Cas9-F | ACCAGACACGAGACGACTAA | |
Pita-R | TCTAGCAGGTGTCGGAGATC | Cas9-R | ATCGGTGCGGGCCTCTTC |
Table 1 Primers used in this study.
引物名称 Primer name | 引物序列 Sequence(5'-3') | 引物名称 Primer name | 引物序列 Sequence(5'-3') | |
---|---|---|---|---|
Pita-g-F | GGCACCATGGCGCCGGCGGTCATTGC | Pi21-F | CTGAATTCCACGGGAATTGC | |
Pita-g-R | AAACGCAATGACCGCCGGCGCCATGG | Pi21-R | CTCCTCTGCGTTCATATTC | |
Pi21-g-F | GGCAGGTATATTGGTCATCTTGGTGG | ERF922-F | GCGACAACATCACGCATCGCC | |
Pi21-g-R | AAACCCACCAAGATGACCAATATACC | ERF922-R | TCAGGCCAAGTGCCTACTGCAA | |
ERF922-g-F | GGCAGTGGACGTGTCTCTGTCGCTGG | Hyg-F | GCTGTTATGCGGCCATTGTC | |
ERF922-g-R | AAACCCAGCGACAGAGACACGTCCAC | Hyg-R | GACGTCTGTCGAGAAGTTTC | |
Pita-F | CAGCCTCTCTGCATGATTTC | Cas9-F | ACCAGACACGAGACGACTAA | |
Pita-R | TCTAGCAGGTGTCGGAGATC | Cas9-R | ATCGGTGCGGGCCTCTTC |
基因 Gene | 前引物 Forward primer | 后引物 Reverse primer |
---|---|---|
OsActin | GTGGACGTACTACTGGTATTGT | GAATCAGTCAGATCCCTACCAG |
OsPR1a | GGCCAATCTCCCTACTGATTAA | GCATAAACACGTAGCATAGCAT |
OsPR10 | CAGTGGTCAGTAGAGTGATCAG | GGGTTAAGCTTCATGGTGTAGA |
OsAOS2 | CAATACGTGTACTGGTCGAATG | CTTATTGCATATGCGTAGGACG |
OsPAL1 | GACCCTGTATTTTCTTCGTTCG | AGTAGCAATACTTTCACCCCAA |
OsRaP2.6 | AGAGAAAACCAGAAAAACGCTG | TGCTTTGGGGATGGAATATGTA |
β-1,3-glucanase | GGATCGGAGGAGTGTAGATAGA | GCAGAAAAGAAGGCAAAGTATACG |
qPita | CAGCAACTAACGAGGCATAATC | AGGTAGTAGTCATCTAGCAGGT |
qPi21 | GGCAAGATCATCAAGGAGATCC | CTTGGGCTTCTCGCAGTGA |
qERF922 | TGGACGTGTCTCTGTCGCT | GACGTCGAAGAGGACCATGTC |
Table 2 Primers used for the qRT-PCR of rice defense-related genes.
基因 Gene | 前引物 Forward primer | 后引物 Reverse primer |
---|---|---|
OsActin | GTGGACGTACTACTGGTATTGT | GAATCAGTCAGATCCCTACCAG |
OsPR1a | GGCCAATCTCCCTACTGATTAA | GCATAAACACGTAGCATAGCAT |
OsPR10 | CAGTGGTCAGTAGAGTGATCAG | GGGTTAAGCTTCATGGTGTAGA |
OsAOS2 | CAATACGTGTACTGGTCGAATG | CTTATTGCATATGCGTAGGACG |
OsPAL1 | GACCCTGTATTTTCTTCGTTCG | AGTAGCAATACTTTCACCCCAA |
OsRaP2.6 | AGAGAAAACCAGAAAAACGCTG | TGCTTTGGGGATGGAATATGTA |
β-1,3-glucanase | GGATCGGAGGAGTGTAGATAGA | GCAGAAAAGAAGGCAAAGTATACG |
qPita | CAGCAACTAACGAGGCATAATC | AGGTAGTAGTCATCTAGCAGGT |
qPi21 | GGCAAGATCATCAAGGAGATCC | CTTGGGCTTCTCGCAGTGA |
qERF922 | TGGACGTGTCTCTGTCGCT | GACGTCGAAGAGGACCATGTC |
基因 Gene | 株数 No. of plants | 突变基因型比率Ratio of mutation genotypes / % | 突变类型比率Ratio of mutation types / % | |||||
---|---|---|---|---|---|---|---|---|
纯合突变率 Homozygous | 杂合突变率 Heterozygous | 双等位突变率 Bi-allele | 碱基插入 Insert | 碱基缺失 Deletion | 碱基替换 Substitution | 替换和缺失 Substitution and deletion | ||
Pita | 17 | 41.2(7/17) | 0.0(0/17) | 47.1(8/17) | 73.5(25/34) | 14.7(5/34) | 0.0(0/34) | 0.0(0/34) |
Pi21 | 17 | 35.3(6/17) | 5.9(1/17) | 58.8(10/17) | 8.8(3/34) | 85.3(29/34) | 2.9(1/34) | 0.0(0/34) |
ERF922 | 17 | 5.9(1/17) | 17.7(3/17) | 64.7(11/17) | 17.6(6/34) | 50.0(17/34) | 8.8(3/34) | 2.9(1/34) |
Table 3 Ratios of mutant genotype and mutation types in T0 plants.
基因 Gene | 株数 No. of plants | 突变基因型比率Ratio of mutation genotypes / % | 突变类型比率Ratio of mutation types / % | |||||
---|---|---|---|---|---|---|---|---|
纯合突变率 Homozygous | 杂合突变率 Heterozygous | 双等位突变率 Bi-allele | 碱基插入 Insert | 碱基缺失 Deletion | 碱基替换 Substitution | 替换和缺失 Substitution and deletion | ||
Pita | 17 | 41.2(7/17) | 0.0(0/17) | 47.1(8/17) | 73.5(25/34) | 14.7(5/34) | 0.0(0/34) | 0.0(0/34) |
Pi21 | 17 | 35.3(6/17) | 5.9(1/17) | 58.8(10/17) | 8.8(3/34) | 85.3(29/34) | 2.9(1/34) | 0.0(0/34) |
ERF922 | 17 | 5.9(1/17) | 17.7(3/17) | 64.7(11/17) | 17.6(6/34) | 50.0(17/34) | 8.8(3/34) | 2.9(1/34) |
Fig. 3. Screening of transgenic-free mutant plants by PCR in T1. M, DNA marker; 1, ddH2O; 2, Wild type; 3, Positive control; Lines 4 to 24, T1 mutation plants; P1, Primer Hyg-F and Hyg-R; P2, Primer Cas9-F and Cas9-R.
Fig. 4. Three mutation types of homozygotes in T2. The PAM sequence is highlighted in blue and insertions are represented by red letters. The deletions are shown by red hyphens. The initiation codon is highlighted by underline in black. D2101 and D2102 are monogenic mutation of Pi21, mutations with 1 bp and 4 bp deletions, respectively. D0301 is co-mutation of Pita, Pi21 and ERF922, mutations with 1 bp insertion in Pita and Pi21, respectively, and the mutation with 23 bp deletion in ERF922.
Fig. 5. Expression analyses of Pita, Pi21 and ERF922 in homozygous mutant plants. * and ** represent significant difference between the wild type and the homozygous lines at the 0.05 and 0.01 levels, respectively(t-test).
Fig. 6. Pita, Pi21 and ERF922 gene amino acid sequences in homozygous mutant lines and the wild type. A indicate the alignment of Pi21 gene amino acid sequences in homozygous mutant lines D2101 and D2102; B, C and D show the alignment of Pita, Pi21 and ERF922 gene amino acid sequences in homozygous mutant line D0301.
Fig. 7. Identification of homozygous mutant lines and the control varieties after inoculation. A, Homozygous mutant lines and wild type inoculated at the seedling stage. B, Statistical lesion area of wild type and homozygous mutant lines(Mean±SE, n=3). Different lowercase letters above the bars indicate significant difference between the mutant and the wild type at the 0.01 level(t-test). LTH, Lijiangxintuanheigu; WT, Wild type; D2101 and D2102, Homozygous lines of Pi21; D0301, Homozygous lines of Pita, Pi21 and ERF922.
Fig. 8. Relative expression levels of defense-related genes in the wild type (L1014, WT) and the homozygous mutant lines inoculated with M. coryza (Mean±SE, n=3). * and ** represent significant difference between the wild type and the homozygous lines at the 0.05 and 0.01 levels by t-test with the same time point.
Fig. 9. Relative expression level of defense-related genes in the wild type(L1014) and the homozygous mutant line D0301, inoculated with M. coryza (Mean±SE, n=3). *and** represent significant difference between the wild type and the homozygous lines at the 0.05 and 0.01 levels by t-test at the same stage, respectively.
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