CN110283238A - Paddy disease-resistant albumen RWR1 and its application - Google Patents

Abstract

The invention discloses paddy disease-resistant albumen RWR1 and its applications.The present invention provides following 1) -3) in application of any substance in regulation disease resistance of plant: 1) albumen RWR1；2) DNA molecular of albumen RWR1 is encoded；3) recombinant vector, expression cassette, transgenic cell line or the recombinant bacterium of the DNA molecular containing coding albumen RWR1.The experiment proves that the transgenic paddy rice of RWR1 gene overexpression when being encroached on by rice blast fungus, compares wild rice, the growth of rice blast fungus mycelia is significantly inhibited, and significantly improves it to the resistance of rice blast fungus.Therefore, RWR1 gene is related to resistance of the rice to rice blast fungus；The cultivation and identification of disease-resistant plant variety needed for RWR1 albumen and its encoding gene can be used for agricultural and ecological environment treatment, play a significant role in Resistant breeding field.

Description

Rice disease resistance protein RWR1 and its application

technical field

The invention belongs to the field of plant genetic engineering and relates to a rice disease resistance protein RWR1 and its application.

Background technique

Rice is the most important food crop in my country, and rice diseases caused by pathogenic microorganisms seriously affect the yield and quality of rice. With the continuous deepening of research on the interaction mechanism between rice and pathogenic bacteria, the use of disease-resistant genes to breed rice disease-resistant varieties has become the most effective, safest and most economical way for rice breeding experts to prevent and control rice diseases.

Prediction of genes related to rice disease resistance: Using comparative bioinformatics methods, four whole-genome sequencing rice species, AA genotype wild rice Oryza_glaberrima, FF genotype wild rice Oryza_brachyanth, and two Asian cultivated rice (japonica Nipponbare and indica 9311) were selected for comparison Genome analysis research, using PanOCT software to conduct collinearity analysis on the genomes of four varieties of rice, determine the collinearity region, identify disease resistance-related genes, and obtain the collinearity region of the target gene and the repeated gene cluster Cluster; combined with OrthMCL homologous protein analysis to obtain Homologous proteins; divide the predicted adjacent target genes in the collinear region into Cluster gene families (the distance between two adjacent target genes is less than 10 genes); combined with phylogenetic tree analysis to determine which target genes in the same gene family are homologous tandem repeats, Which are heterologous tandem repeats; finally, the historical events of homologous recombination of the target gene in the gene family were analyzed by RDP3 software.

Screening of rice broad-spectrum disease-resistant genes: Based on the results of comparative biological gene information, according to the evolution of rice disease-resistant genes and the characteristics of disease-resistant genes that have been reported so far, multiple genes that may have disease-resistant functions were selected for functional verification (including Contains 8 homologous genes of published and functional disease resistance genes).

Gene chip analysis technology was used to screen and clone rice disease resistance genes on a large scale. After transforming rice through transgenic technology, the resistance of transgenic rice plants to rice blast and other pathogens was identified, and the role of target genes in broad-spectrum disease resistance was clarified. Long-lasting and broad-spectrum rice disease resistance provides new gene and material resources.

Contents of the invention

One object of the present invention is to provide the use of any one of the following 1)-3).

The present invention provides the application of any one of the substances in the following 1)-3) in regulating and controlling plant disease resistance:

1) protein RWR1;

2) DNA molecule encoding protein RWR1;

3) Recombinant vectors, expression cassettes, transgenic cell lines or recombinant bacteria containing DNA molecules encoding protein RWR1;

The protein RWR1 is as follows (1) or (2):

(1) A protein consisting of the amino acid sequence shown in Sequence 2 in the sequence listing;

(2) A protein derived from (1) that undergoes substitution and/or deletion and/or addition of one or several amino acid residues to the amino acid sequence shown in Sequence 2 in the sequence listing and has the same function.

In the above application, the DNA molecule is any one of the following 1)-4):

1) The coding region is the DNA molecule shown in sequence 1 in the sequence listing;

2) The coding region is the DNA molecule shown in sequence 3 in the sequence listing;

3) a DNA molecule that hybridizes to the DNA sequence defined in 1) under stringent conditions and encodes a protein with the same function;

4) at least 70%, at least 75%, at least 80%, at least 85%, at least 90%, at least 95%, at least 96%, at least 97%, at least DNA molecules having 98% or at least 99% homology and encoding proteins with the same function.

In the above application, the disease resistance is rice blast resistance.

In the above application, the plant is a dicotyledonous plant or a monocotyledonous plant;

Or the plant is a monocotyledonous plant, and the monocotyledonous plant is specifically rice.

The application of any one of the substances in the above 1)-3) in cultivating disease-resistant plants is also within the protection scope of the present invention.

In the above application, the disease resistance is rice blast resistance;

Or, the plant is a dicotyledonous plant or a monocotyledonous plant;

Or the plant is a monocotyledonous plant, and the monocotyledonous plant is specifically rice.

The second object of the present invention is to provide a method for breeding disease-resistant transgenic plants.

A method for cultivating disease-resistant transgenic plants provided by the present invention comprises the steps of: increasing the expression level and/or activity of the DNA molecule encoding protein RWR1 in the target plant to obtain a transgenic plant, the disease resistance of the transgenic plant is higher than the plant of interest;

The protein RWR1 is as follows (1) or (2):

(1) A protein consisting of the amino acid sequence shown in Sequence 2 in the sequence listing;

(2) A protein derived from (1) that undergoes substitution and/or deletion and/or addition of one or several amino acid residues to the amino acid sequence shown in Sequence 2 in the sequence listing and has the same function.

In the above method, said increasing the expression level and/or activity of the DNA molecule encoding the protein RWR1 in the target plant is introducing the DNA molecule encoding the protein RWR1 into the target plant.

In the above method, the disease resistance is rice blast resistance;

In the above method, the plant is a dicotyledonous plant or a monocotyledonous plant;

Or the plant is a monocotyledonous plant, and the monocotyledonous plant is specifically rice.

The disease resistance of the above-mentioned transgenic plants is higher than that of the target plants in at least one of the following:

1) The transgenic plant has fewer necrotic spots than the target plant;

2) The disease-susceptible grade of the transgenic plant is smaller than that of the target plant;

3) The relative expression level of MoPot2 DNA of the transgenic plant is lower than that of the target plant;

The transgenic plant is a homozygous plant (+/+) of the transgenic rice.

The experiment of the present invention proves that when the transgenic rice overexpressing the RWR1 gene is attacked by blast fungus, compared with wild type rice, the growth of blast fungus hyphae is significantly inhibited, so that the resistance to blast fungus is significantly improved . Therefore, the RWR1 gene is related to the resistance of rice to blast fungus; the RWR1 protein and its encoding gene can be used for the cultivation and identification of disease-resistant plant varieties required for agricultural and ecological environment management, and play an important role in the field of plant disease-resistant breeding .

Description of drawings

Figure 1 shows the disease resistance phenotype of p1300: _RWR1 transgenic rice of the T2 generation after being inoculated with Magnaporthe oryzae RB22.

Fig. ₂ is the statistical result of the disease progression of p1300:RWR1 transgenic rice of T2 generation after being inoculated with Magnaporthe oryzae RB22.

Fig. 3 is the relative value of fungal DNA in rice leaves of T2 generation p1300: _RWR1 transgenic rice after being inoculated with Magnaporthe oryzae.

Detailed ways

The experimental methods used in the following examples are conventional methods unless otherwise specified.

The materials and reagents used in the following examples can be obtained from commercial sources unless otherwise specified.

The quantitative experiments in the following examples were repeated three times, and the results were averaged.

In the following examples, rice adopts conventional variety TP309 (hereinafter also referred to as wild type rice) (recorded in Wang J, Qu B, Dou S, Li L, Yin D, Pang Z, Zhou Z, Tian M, Liu G, Xie Q, Tang D, Chen X, Zhu L., The E3ligase OsPUB15 interacts with the receptor-like kinase PID2 and regulates plant cell death and innate immunity. BMC Plant Biol. 2015, 13; 15., public available from Chinese Academy of Sciences Genetics and Developmental Biology acquired by the Institute).

Embodiment 1, the acquisition of RWR1 gene

Extraction of rice (Oryza sativa) Oyzabrachyantha (recorded in Chen J, Huang Q, Gao D, WangJ, Lang Y, Liu T, Li B, Bai Z, Luis Goicoechea J, Liang C, Chen C, Zhang W, Sun S, LiaoY , Zhang X, Yang L, Song C, Wang M, Shi J, Liu G, Liu J, Zhou H, Zhou W, Yu Q, An N, Chen Y, Cai Q, Wang B, Liu B, Min J, Huang Y ,Wu H,Li Z,Zhang Y,Yin Y,Song W,Jiang J,Jackson SA,Wing RA,Wang J,Chen M.Whole-genome sequencing of Oryzabrachyanthreveals mechanisms underlying Oryza genome evolution.Nat Commun.2013; 1595, the public can obtain from the Institute of Genetics and Developmental Biology, Chinese Academy of Sciences) leaf DNA, design primers for PCR amplification.

RWR1-F: 5'ACGGCCAGTGCCAAGCTTGAGAGAGAGGGAGAGATGAA 3'

RWR1-R: 5'CGCGCCTCGAGATCCACCAACGGCGAGTAATAGCAT 3'

A 5273bp PCR product was amplified.

The above PCR product was sent for sequencing, and its nucleotide sequence was sequence 3 in the sequence listing, including a promoter sequence of about 1026 bp and a coding region sequence of 4247 bp, and the gene shown in positions 1-5273 of sequence 3 in the sequence listing was named RWR1 . Sequence 1 in the sequence listing is the CDS sequence of the gene, with a total of 3645 bases, encoding 1214 amino acids, named RWR1 protein (SEQ ID NO: 2).

Example 2, Functional Verification of RWR1 Gene

1. Obtaining the expression vector

The original plasmid pCAMBIA1300-flag was digested with Hind III and XbaI (recorded in Wang Z, Li N, Jiang S, Gonzalez N, Huang X, Wang Y, Inzé D, Li Y. SCFSAP controls organ size bytargeting PPD proteins for degradation in Arabidopsisthaliana. Nat .Commun.2016; 7:11192, the public can obtain from the Institute of Genetics and Developmental Biology, Chinese Academy of Sciences), recovered a 9.6kb large fragment, and obtained the pCAMBIA1300-221-flag vector backbone.

The 5273bp PCR product obtained in Example 1 was subjected to In-fusion reaction (TAKARA Company) with the vector backbone pCAMBIA1300-flag to obtain the binary expression vector pCAMBIA1300::RWR1.

After sequencing, the recombinant expression vector pCAMBIA1300::RWR1 is a vector obtained by replacing the 35S promoter between Hind III and Xba I of the pCAMBIA1300-flag plasmid with the DNA molecule shown in Sequence 3. In the DNA molecule shown in sequence 3, the 1-1026th position is the promoter sequence of the RWR1 gene, and the 1027-5273rd position is the sequence of the RWR1 gene coding region.

2. Obtained from RWR1 gene expression recombinant bacteria

The recombinant expression vector pCAMBIA1300::RWR1 of the above 1 was transformed into Agrobacterium EHA105, a single clone was picked, and cultured in LB liquid medium containing kanamycin and rifampicin antibiotics at 28°C with shaking overnight to obtain transformants.

The transformants were identified by bacterial liquid PCR (RWR1-6500F and Flag-R, the positive recombinant bacteria with about 600 bp were obtained), and the positive recombinant bacteria were named EHA105/p1300::RWR1, and stored at -70°C for future use.

RWR1-6500F: 5′GAGACCTGTCACATGCTCATA 3′

Flag-R: 5'CTAGTTAATTAAGACGCGTCCT 3'.

3. Obtaining rice with RWR1 gene expression

Using Agrobacterium tumefaciens-mediated transformation of rice, the specific method is as follows:

Ⅰ. Preparation of transformed receptors: Rice TP309 immature embryos 14 days after pollination were selected as receptors for Agrobacterium infection.

Ⅱ. Cultivation of Agrobacterium engineering strains: Get a small amount of bacterial liquid from the positive recombinant bacteria Agrobacterium EHA105/p1300::RWR1 stock solution (20% glycerol), and put it in YEB solid medium (containing kanamycin 50mg/l, rif Mycin 50 mg/l) was streaked and cultured at 28°C in the dark until a single colony with a diameter of 1 mm grew out. Take a single colony and inoculate it on the same medium, and cultivate it until the vigorous growth period. Pick a little bacterium and inoculate in 20ml YEB liquid medium (containing corresponding antibiotics), shake at 220rpm at 28°C overnight in the dark. On the next day, transfer to 20ml YEB liquid medium containing 100uM acetosyringone with 2% inoculum size, cultivate to mid-logarithmic growth phase, and dilute with more than 3 times of liquid basic medium until the naked eye is slightly turbid (OD600 approx. 0.1), for transformation purposes.

Ⅲ. Agrobacterium rhizogenes infection and co-cultivation: Take the bacterial liquid into a culture bottle, add the pre-cultured callus, shake slightly and let stand for 10 minutes. After drying the callus on sterile filter paper, put it on the medium supplemented with 100 uM acetosyringone, and cultivate it in the dark at 25°C for 3 days.

Ⅳ. Screening of resistant callus: the callus after the co-cultivation is transferred to the screening medium containing 25mg/l hygromycin (hygromycinB) and 500mg/l cephalosporin, and after screening 1-2 times, it can be seen that Resistant callus outgrows.

Ⅴ. Regeneration of resistant plants: Transfer the resistant callus to the medium containing 25mg/l hygromycin, and the light conditions are the same as before. When the differentiated resistant budlet grows to 2-3cm, it is transferred to the regenerated seedling subculture medium containing 25mg/l hygromycin, and when it grows to a complete plant, it is transferred from the solid medium to Open culture in nutrient solution. After new roots were formed, the seedlings were transferred to the greenhouse to obtain the T ₀ generation-transferred RWR1 rice.

Using the same method, the empty vector pCAMBIA1300-221-flag was transformed into rice TP309 to obtain the T ₀ generation of empty vector-transferred rice.

The above T ₀ generation plants were sown to obtain T ₁ generation plants.

4. Identification of transgenic RWR1 rice

Cut about 3 cm leaves from T1 _- transformed RWR1 rice that has grown for about two weeks, and quickly freeze them with liquid nitrogen. The total DNA was extracted, and the wild-type rice TP309 was used as a control for PCR amplification to detect the integration of the target gene into the genome.

The primers used for PCR detection of RWR1 expression (amplified to obtain the junction fragment located at the 3' end of RWR1 and the tag Flag) are:

RWR1-6500F: 5′GAGACCTGTCACATGCTCATA 3′

Flag-R: 5′CTAGTTAATTAAGACGCGTCCT 3′

The PCR amplification products were recovered and sequenced, indicating that RWR1 had been integrated into the rice genome, and these plants integrated into the rice genome were recorded as positive T1 _- transformed RWR1 rice.

Neither the empty vector rice of the T1 generation nor the wild _- type rice TP309 had RWR1 integration.

5. Obtaining of homozygous RWR1 gene expression rice

_The above identified as positive T1 generation _RWR1 rice single plant harvested to obtain T2 generation seeds, soak 30-40 seeds in aqueous solution containing 25mg/L hygromycin for germination, and judge positive by the growth state of plant roots The growth of young roots of the plants (transgenic negative plants was severely inhibited under the action of hygromycin, showing that the main root was shorter, and the lateral roots basically did not occur, and browning began to elongate to about 1 cm; while the transgenic positive materials had hygromycin resistance. The growth of the root is basically not inhibited, and the occurrence of the main root and the lateral root is all normal), and it is recorded as the T2 generation p1300: _RWR1 transgenic rice, and the transplanting experiment is carried out.

The rice of the T ₁ generation transformed into the empty vector was harvested and planted, and the T ₂ generation of transformed rice with the empty vector was obtained.

6. Analysis of RWR1 gene expression rice resistance to blast fungus

Before rice inoculation, T2 generation p1300: _RWR1 transgenic rice, T2 generation empty vector rice and wild _- type rice TP309 were all cultured under conventional conditions: 14 hours of light, 10 hours of darkness, and a temperature of 28°C.

When the plant grows to the stage of three leaves and one heart (about 20 days), the inoculation experiment of Magnaporthe oryzae RB22 is carried out, and the concentration of the spore liquid is 5× ^{10 5} /ml (RB22 is recorded in Kang H, Wang Y, Peng S, Zhang Y, Xiao Y, Wang D, Qu S, Li Z, Yan S, Wang Z, Liu W1, Ning Y, Korniliev P, Leung H, Mezey J, McCouch SR, Wang GL.. Dissection of the genetic architecture of rice resistance to the blastfungus Magnaportheoryzae. Mol Plant Pathol.2016,17(6):959-72., publicly available from Institute of Genetics and Developmental Biology, Chinese Academy of Sciences). After the leaves were completely sprayed wet, treated in the dark for 24 hours, and cultured in light for 5 days, the incidence was counted and recorded by taking pictures.

The results showed that, compared with wild-type rice TP309, the homozygous T2 generation p1300: _RWR1 transgenic rice had significantly improved resistance to blast fungus RB22 after spraying RB22 spore liquid, indicating that RWR1 protein and rice blast fungus related to resistance.

The specific performance is:

1) Statistical analysis of plant phenotypes after inoculation with RB22

The results are shown in Figure 1, after 6 days of spraying, the leaves of homozygous plants (+/+) and heterozygous plants (+/-) of T2 generation p1300: _RWR1 transgenic rice were basically not infected by Magnaporthe oryzae; The leaves of wild-type rice as a control (wild-type rice isolated in FIG. 1, denoted as -/-) had a large number of necrotic spots.

There was no significant difference in phenotype between the _T2 generation of empty vector rice and wild type rice.

2) Investigate and analyze the disease progression of rice material leaves

According to the technical specification for rice blast resistance field identification in Sichuan Province (DB51/T714-2007), the disease progression of rice materials was counted, and the results are shown in Figure _2. After 6 days of spraying, the pure The leaf susceptibility grades of synzygous (+/+) and heterozygous (+/-) plants were much lower than those of control wild-type rice and transgenic wild-type rice (-/-).

3) Detect the relative value of rice blast fungus DNA in rice leaves and rice leaf DNA

Six days after RB22 spraying, the relative DNA expression levels of the marker gene MoPot2 of Magnaporthe oryzae in the T2 generation p1300: _RWR1 transgenic rice and wild-type rice were detected.

The specific method is as follows:

A. DNA extraction from rice leaves

Total DNA was extracted by CATB method:

1) Take about 3 cm of the above-mentioned rice leaves of each strain, grind them into powder in liquid nitrogen, add 0.3 mL of CATB DNA extract, shake and mix well.

2) After the grinding solution was placed in an incubator at 65°C for 30 minutes, 0.3 mL of chloroform was added, the centrifuge tube was tightly capped, the centrifuge tube was shaken vigorously for 20 seconds, and centrifuged at 12,000 g for 5 minutes.

3) Take about 0.3 mL of the upper aqueous phase into a new centrifuge tube, add 0.3 mL of isopropanol, let stand at room temperature for 10 minutes, and centrifuge at 12,000 g for 10 minutes at 4°C.

4) Discard the supernatant, add 0.7mL of 75% ethanol, vortex and mix well, and centrifuge at 12000g for 1 minute at 4°C.

5) Carefully discard the supernatant, then dry at room temperature or in vacuum for 10 minutes, dissolve in 0.3 mL double-distilled water, and obtain DNA.

B. Fluorescent quantitative PCR analysis

1) Reaction system

SYBR Premix 10μL

2) Reaction program

95℃, 30s; 30cycles of(95℃, 10s; 60℃, 20s; 72℃, 20s)

3) Data analysis

The Ct value is the number of cycles experienced when the fluorescent signal in the PCR tube reaches the set threshold value.

ΔCt=Ct(Gene)-Ct(UBQ), the expression level of the gene is measured by the value of 2-ΔCt.

The genes expressed in quantitative PCR analysis and the primers used are as follows:

The internal reference gene is ubqutin, and the Q-PCR primers used to amplify the internal reference gene are:

UBQ-F: 5′-TTCTGGTCCTTCCACTTTCAG-3′

UBQ-R: 5′-ACGATTGATTTAACCAGTCCATGA-3′

The Q-PCR primers for detecting MoPot2 are:

MoPot2-F: 5′-ACGACCCGTCTTTTACTTATTTGG-3′

MoPot2-R: 5′-AAGTAGCGTTGGTTTTGTTGGAT-3′

The results are shown in Figure 3. Compared with the control plant wild-type rice TP309, the relative expression of _MoPot2 DNA in the T2 generation p1300: _RWR1 transgenic rice was significantly reduced; There was no significant difference in the relative expression of DNA.

The above experiments were repeated 3 times with consistent results.

The above results prove that the RWR1 gene is related to the resistance of rice to blast fungus, indicating that the RWR1 protein and its coding gene can be used in the cultivation and identification of disease-resistant plant varieties in agricultural production, and play an important role in the field of plant disease-resistant breeding.

Finally, it is noted that the above embodiments are only used to illustrate the technical solutions of the present invention and not to limit them. Although the present invention has been described with reference to the preferred embodiments of the present invention, those skilled in the art should understand that it can be described in the form Various changes may be made in matter and details thereof without departing from the spirit and scope of the invention as defined in the appended claims.

sequence listing

<110> Institute of Genetics and Developmental Biology, Chinese Academy of Sciences

<120> Rice disease resistance protein RWR1 and its application

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Met Gly Gly Leu Gly Lys Thr Thr Phe Ala Gln Leu Ile Tyr Asn Asp

195 200 205

Pro Glu Ile Leu Lys His Phe Gln Leu Arg Arg Trp Cys Cys Val Ser

210 215 220

Asp Glu Phe Asp Val Val Ser Ile Ala Asn Asn Ile Cys Val Ser Thr

225 230 235 240

Glu Arg Asn Arg Glu Arg Ala Leu Gln Asp Leu Gln Lys Glu Val Ser

245 250 255

Gly Lys Lys Phe Leu Ile Val Leu Asp Asp Val Trp Asn Arg Asp Ser

260 265 270

Asp Lys Trp Gly Lys Leu Met Thr Cys Leu Lys Gln Gly Ser Arg Gly

275 280 285

Ser Val Val Leu Thr Thr Thr Arg Asp Val Lys Val Ala Thr Ile Met

290 295 300

Ala Thr Ser Glu Val Glu Val Tyr Asn Leu Gly Lys Leu Gly Glu Val

305 310 315 320

Tyr Leu Lys Glu Ile Ile Gln Ser Lys Ala Ile Gly Leu Pro Gly Ser

325 330 335

Asp Glu His Leu Glu Val Leu Asn Lys Ile Val Gln Arg Cys Asp Gly

340 345 350

Ser Pro Leu Ala Ala Lys Ser Phe Gly Ser Val Leu Ser Ser Arg Ser

355 360 365

Thr Val Gln Glu Trp Lys Asp Ile Leu Ala Lys Ser Asn Ile Cys Asn

370 375 380

Glu Gly Glu Asp Thr Ile Phe Pro Ile Leu Arg Leu Ser Tyr Asp Asp

385 390 395 400

Leu Pro Phe Asp Met Lys Gln Cys Phe Ala Phe Cys Ala Ile Phe Pro

405 410 415

Lys Asp Tyr Val Ile Asp Val Glu Thr Leu Ile Lys Leu Trp Leu Ala

420 425 430

His Asp Phe Ile Pro Leu Gln Glu Asp Asp Asn Leu Glu Ser Ala Ala

435 440 445

Glu Asp Ile Phe Lys Glu Leu Val Trp Arg Ser Phe Phe Gln Asp Val

450 455 460

Lys Lys Ser Ser Ser Ile Trp Thr Thr Cys Lys Ile His Asp Leu Met His

465 470 475 480

Asp Ile Ala Gln Ser Val Met Gly Lys Glu Cys Val Ser Ile Ala Gly

485 490 495

Arg Ser Asn Phe Ile Ser Leu Leu Ser Glu His Pro Arg Tyr His Phe

500 505 510

His Ser Ser Tyr Lys Glu Thr Val Leu Leu Asp Asp Phe Met Arg Lys

515 520 525

Gln Ser Pro Thr Leu Arg Ser Leu Leu Phe Glu Arg Trp Phe Asn Tyr

530 535 540

Phe Ser Thr Ser His Leu Ser Lys Cys Ser Ser Leu Arg Ala Leu Lys

545 550 555 560

Leu Leu Arg Cys Ser Glu Phe Leu Pro Ile Gly His Leu Gln His Leu

565 570 575

Arg Tyr Leu Asn Ile Ser Ser Asn Ser Cys Ile Lys Lys Leu Pro Lys

580 585 590

Asp Ile Cys Ile Leu Tyr Asn Leu Gln Thr Leu Val Leu Ser Tyr Cys

595 600 605

Lys Asn Leu Val Glu Leu Pro Lys Asp Met Lys Tyr Met Lys Asn Leu

610 615 620

Arg His Leu Tyr Thr Asp Gly Cys Pro Lys Leu Lys Tyr Met Pro Pro

625 630 635 640

Asp Leu Gly Gln Leu Thr Ser Leu Gln Ile Leu Thr Ser Phe Val Val

645 650 655

Gly Ala Arg Ser Gly Cys Ser Asn Leu Arg Glu Leu Arg Thr Leu Asn

660 665 670

Leu Cys Gly Arg Leu Gln Leu Cys Gly Leu Glu Asn Glu Lys Glu Glu

675 680 685

Asn Ala Lys Ala Ala Asn Leu Arg Asn Lys Glu Lys Leu Thr His Leu

690 695 700

Ser Leu Glu Trp Asn Ser Asn Cys His Leu Glu Gly Thr Asn Ser Pro

705 710 715 720

Tyr Lys Val Leu Asp Ala Leu Lys Pro His His Arg Leu Gln Met Leu

725 730 735

Lys Val Ile Ser Tyr Thr Gly Ser Ser Phe Pro Ala Trp Ile Thr Asp

740 745 750

Leu Gly Val Leu Gln Asn Leu Ile Glu Leu His Leu Glu Gly Cys Thr

755 760 765

Met Cys Gly Glu Phe Pro Gln Phe Ile Arg Phe Lys Phe Leu Gln Val

770 775 780

Leu Tyr Leu Ser Arg Leu Asp Asn Leu Gln Thr Leu Cys Arg Glu Glu

785 790 795 800

Gly Arg Gln Gly Thr Glu Gln Ala Phe His Gln Leu Glu Lys Val Val

805 810 815

Ile Asn Ile Cys Pro Lys Phe Gln Thr Leu Cys Ser Gly Val Ala Ser

820 825 830

Thr Ala Phe Pro Glu Leu Lys Glu Val Lys Leu Met Asp Leu Glu Ser

835 840 845

Phe Glu Thr Trp Val Ala Met Glu Gly Arg Gln Gly Tyr Met Pro Thr

850 855 860

Phe Pro Leu Leu Glu Glu Val Glu Ile Asn Lys Cys Pro Lys Leu Thr

865 870 875 880

Thr Leu Pro Glu Ala Pro Lys Leu Lys Ile Leu Asn Leu Asn Glu Asn

885 890 895

Lys Ala Gln Leu Ser Leu Ser Leu Leu Gln Ser Ser Tyr Ile Ser Ser

900 905 910

Leu Ser Lys Leu Arg Leu Glu Ile Asp Asp Lys Glu Thr Thr Leu Gln

915 920 925

Leu Leu Asp Gln Ile His Glu Ser Ser Leu Ser Glu Met Glu Leu Thr

930 935 940

His Cys Asn Ile Leu Phe Pro Leu Ser Pro Ser Gln Ser Lys Met Arg

945 950 955 960

Ile Trp Glu Trp Leu Gly Gln Leu Val Glu Leu Lys Ile Asp Ser Cys

965 970 975

Asp Ser Leu Ile Tyr Trp Pro Glu Glu Glu Phe Leu Cys Leu Val Ser

980 985 990

Leu Lys Lys Leu Thr Ile Lys Glu Cys Tyr Asn Leu Ile Gly Arg Pro

995 1000 1005

Thr Gln Val Thr Gly Asn Pro Thr Leu Leu Pro His Leu Thr Ser

1010 1015 1020

Leu Tyr Val Ser Lys Cys Val Arg Leu Arg Glu Leu Phe Val Leu

1025 1030 1035

Pro Pro Ser Ile Lys Tyr Ile Thr Ile Asn Asp Ser Ile Cys Leu

1040 1045 1050

Glu Ser Phe Ser Phe Pro Ser Tyr His Leu Pro Cys Leu Glu Arg

1055 1060 1065

Leu Ser Phe Trp Asn Cys Arg Ser Val Val Thr Leu Gln Asn Leu

1070 1075 1080

Pro Pro Cys Leu Met Leu Ser Ile Asp Ala Cys Trp Glu Leu Gln

1085 1090 1095

Ser Leu Ser Gly Gln Leu Asp Glu Leu Lys His Leu Gly Ile Val

1100 1105 1110

Arg Cys Asn Lys Leu Glu Ser Leu Asn Cys Leu Gly Glu Leu Pro

1115 1120 1125

Ser Leu Glu His Leu Asp Leu Lys Met Cys Lys Arg Leu Ala Ser

1130 1135 1140

Ala Pro Cys Gly Pro Arg Ser Tyr Ser Ser Ser Leu Leu Ser Ile Thr

1145 1150 1155

Ile Gln Asp Cys Pro Arg Met Asn Met Lys Lys Val Tyr Glu Trp

1160 1165 1170

Leu Arg Pro Arg Leu Asp Ser Leu Glu Glu Arg Asp Leu Ser His

1175 1180 1185

Ala His Thr Arg Val Ile Tyr Glu Glu Ser Lys Cys Pro Thr Leu

1190 1195 1200

Lys Ser Trp Lys Tyr Ala Ile Thr Arg Arg Trp

1205 1210

<210> 3

<211> 5273

<212>DNA

<213> Rice (Oryza sativa)

<400> 3

gagagagagg gagagatgaa tgcataaaaa gagaaaagtt ttagtgggac tcacattaag 60

aatggtgcac tatggagctt gtatcctatg tgtggctttg tcctacgtga catctctttt 120

tttatgagag agtggcctgc atagtgtatg tccgtggtac ttttttagtt atggaggcct 180

ctctatgtca ccatcccttga gcaggtacca ctgttttcta tgtaaaattt ggtacctctt 240

gttaccttag gtactagaag gtaccaaatt ttaagtttta ctctcatccc tcctttttta 300

tcttaaggta ccggtatctc gcggtaccaa atcatttatg atcgttggat caaacagtgc 360

acatcctatt tagctagatc caatggtgag aaacgatttg gtatctcgag gtactggtac 420

ctcgaggtat aaaaggaagg ataagagtaa aactcaccaa attttacata taaaacagtg 480

gtacctcttg taccttctta aggatggaaa aaaaactctt ttaacaacat atattgtcta 540

aaaaatacca gcaggatgct tgcatcggat cgttggtaac gaaaaattac ggggcattta 600

actttttgtc actcttaaaa ttggttaata ataaatttat cactcattat atatgatata 660

tacgctctga tatgttaggt ccaatgataa atatgttaat tatgccggt tatgtacgcc 720

gtgtgtccag agcatctcat taaagtagga atcactctat cttctctccc acatcgatcg 780

accacctctc tctcttccat ctccatccac agcatcccct cggagattag ggcctccgtt 840

ctctgatcga tctagatcga tcccccacaa catctagatc gggtctcgac gaaggttagt 900

gccgctagct agctgcctgc gatatcatat catttcagtt ctgcaatgtg gagtaattaa 960

ccgtgttccc tctcccatcc atgatgcaga gaagagaacg aagcatcatc ttcagaggga 1020

gcaacgatgg ctgagttttt ggttcggccg ctgctgtcca cggtgcagaa cgcttccagc 1080

tatcttgcag gccagtacag ggtgatggaa ggcatggagg agcagcgcaa agctctggag 1140

cgcatgcttc cactcatcct caccgtcatc cacgacgcac agaacagaac caaacaatcc 1200

caagtaggcg cttggctgca agagctcaag aaggtgtcct acgaggcgac cgacgtgttc 1260

gacgagttca gatacgaggc gctccggcgc gaagccagga ggaaagggca cggcgctgta 1320

agcctcttct cctctcgtaa cccaatcgtg tttcgctaca ggatgggcaa gaagctgcgg 1380

aagatcgtgc agagaatcaa ggaacttgtc gaggagatga attcctttgg gctcgtacac 1440

cggcaggaaa caccgaggca gtcgaggcaa actgattcag tgatgcttga ttttgagaag 1500

gatattgtta gcagatccag agatgaggag aagaggaagg ttgtcaagat attggtggat 1560

gaagctagcg acaggggagct cacagtcctt cctgttgttg gaatgggtgg tcttggcaag 1620

actacatttg cacagctcat ctacaatgac cctgaaatcc tgaagcattt tcagcttcgc 1680

aggtggtgtt gtgtgtctga tgaatttgat gtcgttagca tcgcaaacaa catatgtgtg 1740

agcacagaga gaaatcgtga aagggcactg caagatctgc agaaggaagt aagtggaaag 1800

aagtttctga tagtgttgga tgatgtgtgg aatagggatt ctgacaagtg gggaaagtta 1860

atgacctgcc ttaagcaggg ctccaggggc agtgtggtac taacaacaac tcgggatgtc 1920

aaagtcgcta caattatggc taccagtgaa gttgaagtgt ataatcttgg taagctagga 1980

gaagtgtatt tgaaggaaat aatccaaagt aaagcaattg gtttgccagg aagtgatgag 2040

catttggaag ttcttaataa aattgttcag agatgtgatg gctctccttt agctgcaaaa 2100

tcctttggct ctgtgttgtc tagcaggagt actgtacaag aatggaagga tatattagcc 2160

aaaagtaaca tttgcaatga gggggaggac acaatttttc ctatacttcg tctcagctat 2220

gacgacttac catttgacat gaagcaatgc tttgctttct gtgctatatt cccaaaagat 2280

tatgtgattg atgtggagac tttgattaag ctatggttgg cacatgactt cataccatta 2340

caagaggatg acaatctaga atcggcagcc gaagatatct tcaaggagct agtttggagg 2400

tcattttttc aagatgtaaa gaaatcttct atatggacca catgcaagat acatgatctt 2460

atgcacgaca ttgctcaatc tgttatggga aaagaatgtg tcagcatagc tggaaggtcc 2520

aattttataa gtctgttatc agaacatcct aggtatcact ttcactcatc atacaaagag 2580

actgttctct tagatgactt tatgagaaaa caatctccaa ctctccggag tttattgttt 2640

gaacgatggt ttaattactt cagcacatca catttatcca agtgcagttc tctgcgagca 2700

ctgaagctcc tacgatgcag cgaattctta ccaatcgggc accttcagca cctaagatat 2760

ctcaatatct catcaaacag ttgtatcaaa aagcttccta aagatatatg catactctac 2820

aatctacaga ctttggtcct ctcttattgt aaaaatcttg tcgaacttcc aaaggatatg 2880

aagtatatga aaaatctgcg acacctttat acggatggat gtccaaaatt gaagtacatg 2940

cctccggacc ttggacagtt aacttccctg cagatattaa catcttttgt ggtgggagct 3000

aggtctggtt gcagtaacct tagagaattg cgtaccttaa acctttgtgg caggctacag 3060

ttatgtggcc tagaaaatga aaaggaagaa aatgcgaaag cagccaatct tcgaaacaaa 3120

gagaaactta cacatttgtc tcttgagtgg aatagcaact gccatcttga aggaacaaat 3180

tccccttata aggttcttga tgctcttaaa cctcatcaca ggctgcagat gcttaaggta 3240

atttcctata caggcagttc ttttccagca tggataacag accttggtgt cctgcaaaac 3300

ttgatagagc tccattaga gggctgtaca atgtgtggag aatttcctca gttcattcgt 3360

ttcaagtttc ttcaggttct ttatctgagt agacttgata acttgcaaac cctatgtcgc 3420

gaggaaggaa gacaaggaac agaacaagca tttcatcagc ttgagaaggt tgtcatcaac 3480

atctgtccaa agtttcaaac attgtgctct ggtgtggcat ccactgcatt tccagaacta 3540

aaggaagtca agttaatgga tttggagagc tttgagacat gggtggcaat ggaagggagg 3600

caaggttaca tgccaacatt tcctctgctt gaggaggttg aaatcaacaa gtgcccaaaa 3660

ttgacaactc tacctgaagc accaaagctc aagattttaa atctaaatga aaacaaagcg 3720

cagctgtcct tgtcattgct tcaatccagc tatatatcct cattgtccaa gctaagattg 3780

gaaatagatg acaaagaaac aaccctgcag ctgcttgatc agatccacga atcatctctc 3840

tcagaaatgg agttaacaca ttgcaacatt ctcttcccct tgagcccatc acagtcaaaa 3900

atgaggatct gggaatggct tggacaactt gttgagctga aaatcgactc ctgcgattcg 3960

ctcatctact ggccagaaga agagttccta tgcttggtat ccctgaagaa attgaccatc 4020

aaggagtgct ataacctaat tggccgtcct accccaggtga caggaaatcc aactctcctg 4080

ccacatctca catcgcttta tgtttctaag tgtgtcaggt tgagagagct ctttgttctt 4140

ccaccatcta tcaaatatat tacaattaat gactccattt gtcttgagag cttctcattc 4200

ccctcctatc atctgccatg cctagaacgt ctaagtttct ggaattgtcg ttcagtggta 4260

acacttcaga acctaccacc atgccttatg ctgtccattg atgcatgttg ggagcttcaa 4320

tcgctgtcag ggcagctgga tgaactcaag catttgggca ttgtacgctg caataaactg 4380

gagtcactga attgcttggg agaattgcca tcactggaac atcttgacct taagatgtgc 4440

aaacgtctag catcggcgcc atgtggccca aggagttact catctctttt gagtattaca 4500

atccaagact gcccaagaat gaatatgaag aaggtatatg agtggctccg gccacggctg 4560

gatagccttg aggaaagaga cctgtcacat gctcatacaa gagtaattta tgaaggtaca 4620

ctccattgct agcattttcc ttctttttcc gtttttcgtt tcttttacat agatgaatag 4680

attttatgtt cctgtgccag tgtatatctg cactcaaagt tccaatcatc aggccctgtg 4740

ttttataact gaataagata gaaattgtgtttcttggtta aagagattct agtctaacct 4800

tgatcgaaca aaggttcgtt ccaccaaaac cataggagct ccatcactgt ttattgacaa 4860

gtttgtgtct acctgtgagg tgtgctatgt gactgcattt gtagcttcct ataatattca 4920

acataattta gtcggtgtcg ataccatgca tgaaatgagc atttttttta tcctcgagaa 4980

actattataa ggaagcactg ttttttatat aaaatttggt actttctagt atctaatata 5040

ctaacagtgg tacctcttag taccttctca aggatcgtaa tatcgctcgc atggaatagt 5100

tcctttccct ctgtactcca ttttattttt tctctttttc tgcagtttat ttcgattgtg 5160

agatattatc taaaaaaaat tcggttgtga gactgacatt tatttttcat gctacagagt 5220

ctaaatgccc gacactaaaa tcatggaaat atgctattac tcgccgttgg tga 5273

Claims (10)

1. The application of any one of the following 1)-3) substances in the regulation and control of plant disease resistance: 1) protein RWR1; 2) DNA molecule encoding protein RWR1; 3) Recombinant vectors, expression cassettes, transgenic cell lines or recombinant bacteria containing DNA molecules encoding protein RWR1; The protein RWR1 is as follows (1) or (2): (1) A protein consisting of the amino acid sequence shown in Sequence 2 in the sequence listing; (2) A protein derived from (1) that undergoes substitution and/or deletion and/or addition of one or several amino acid residues to the amino acid sequence shown in Sequence 2 in the sequence listing and has the same function. 2. The application according to claim 1, characterized in that: The DNA molecule is any DNA molecule in the following 1)-4): 1) The coding region is the DNA molecule shown in sequence 1 in the sequence listing; 2) The coding region is the DNA molecule shown in sequence 3 in the sequence listing; 3) a DNA molecule that hybridizes to the DNA sequence defined in 1) under stringent conditions and encodes a protein with the same function; 4) at least 70%, at least 75%, at least 80%, at least 85%, at least 90%, at least 95%, at least 96%, at least 97%, at least DNA molecules having 98% or at least 99% homology and encoding proteins with the same function. 3. The application according to claim 1 or 2, characterized in that the disease resistance is rice blast resistance. 4. The application according to any one of claims 1-3, characterized in that: The plant is a dicot or a monocot; Or the plant is a monocotyledonous plant, and the monocotyledonous plant is specifically rice. 5. Use of any one of the substances in the following 1)-3) in any one of claims 1-4 in breeding disease-resistant plants. 6. The application according to claim 5, characterized in that: the disease resistance is rice blast resistance; Or, the plant is a dicotyledonous plant or a monocotyledonous plant; Or the plant is a monocotyledonous plant, and the monocotyledonous plant is specifically rice. 7. A method for cultivating disease-resistant transgenic plants, comprising the steps of: increasing the expression level and/or activity of the DNA molecule encoding protein RWR1 in the target plant to obtain a transgenic plant, the disease resistance of the transgenic plant is higher than that of the target plants; The protein RWR1 is as follows (1) or (2): (1) A protein consisting of the amino acid sequence shown in Sequence 2 in the sequence listing; (2) A protein derived from (1) that undergoes substitution and/or deletion and/or addition of one or several amino acid residues to the amino acid sequence shown in Sequence 2 in the sequence listing and has the same function. 8. The method of claim 7, wherein: Said increasing the expression level and/or activity of the DNA molecule encoding the protein RWR1 in the target plant is introducing the DNA molecule encoding the protein RWR1 into the target plant. 9. The method according to claim 7 or 8, characterized in that the disease resistance is rice blast resistance. 10. The method according to any one of claims 7-9, wherein the plant is a dicotyledonous plant or a monocotyledonous plant; Or the plant is a monocotyledonous plant, and the monocotyledonous plant is specifically rice.