CN102586267A - Cloning of plant resistance related gene EDOS1 and application of plant resistance related gene EDOS1 in plant disease resistance - Google Patents
Cloning of plant resistance related gene EDOS1 and application of plant resistance related gene EDOS1 in plant disease resistance Download PDFInfo
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Abstract
The invention relates to isolation, cloning, functional verification and application of an arabidopsis thaliana powdery mildew resistance related gene EDOS1. The invention also relates to a mutant gene of the EDOS1 gene, protein of an EDOS1 genetic code and an application of the gene EDOS1.
Description
Technical field
The present invention relates to the genetically engineered field, be specifically related to the clone of a kind of arabidopsis gene EDOS1, Function Identification and application.Comprise that this gene replys and regulate the application in the ethene inductive leaf senile the regulation and control plant to the resistance of pathogenic bacteria, relate to the application of this gene in the transportation of regulation and control mRNA caryoplasm simultaneously, and the application of this gene in creating the disease-resistant transgenic crop varieties.
Background technology
At occurring in nature, plant is facing to the invasion and attack of various pathogenic bacterias.These pathogenic bacterias roughly can be divided into two types, and one type is biotroph, need the vegetable cell of survival to accomplish growth cycle; Another kind of is the dead volume nutritional type, needs the kill plants cell to obtain nutrition, accomplishes life cycle.
In very long evolutionary process, plant develops and effective disease resistance response.Wherein plant hormone Whitfield's ointment and ethene, jasmonic plays an important role, and Whitfield's ointment mainly plays an important role in the resistance to the biotroph germ; Ethene and jasmonic are mainly being brought into play important effect in the resistance to dead volume nutritional type germ.
Powdery mildew is a kind of fungi that occurring in nature extensively exists, and can infect important farm crop such as wheat, barley, grape, worldwide causes the significant loss of agriculture prodn.
In our previous research, we have reported the phenotype of Arabidopis thaliana edr1 two mutants and function (Frye and Innes, 1998 of EDR1 gene; Frye et al., 2001).Raf-like protein kinase of EDR1 genes encoding, EDR1 albumen has kinase activity.The edr1 two mutants shows powdery mildew inductive necrocytosis phenotype and to the remarkable resistance of white powder germ.The experiment of two mutation analysises shows that the phenotype of edr1 two mutants can be by pad4, sid2, and two mutants such as npr1 suppress, and the phenotype that shows edr1 needs the effect of Whitfield's ointment signal pathway.
Simultaneously, compare with wild-type, the edr1 two mutants shows more significantly ethene inductive leaf senile phenotype.
But we still know little about it at molecular mechanism dead for the EDR1 mediated cell and the plant disease-resistant reaction at present.
Can find out that EDR1 is an outstanding candidate gene of creating disease resistance enhanced genetically modified crops and crop molecular designing.The molecular mechanism that research EDR1 regulates the plant disease-resistant reaction, other components of searching EDR1 signal path have important theory and are worth and wide application prospect.
Summary of the invention
The present invention relates to the clone and the application of Arabidopis thaliana EDOS1 gene, refer more particularly to the effect of EDOS1 gene in the mRNA nucleo-cytoplasmic transport, also relate to this gene simultaneously in of resistance and the function in the leaf senile and the application of regulation and control plant to pathogenic bacteria.
The present invention is through the method for screening mutant and map based cloning; The EDOS1 gene separation and Function Identification have been carried out; Established the EDOS1 gene plant to the effect in the necrocytosis of the resistance of Powdery Mildew and pathogenic bacterium inducing; Find effect and the molecular mechanism thereof of EDOS1 gene in ethene inductive leaf senile, simultaneously, established the mechanism of EDOS1 effect gene mRNA caryoplasm transportation.The present invention is that the molecular mechanism of understanding the transportation of plant mRNA caryoplasm provides important evidence; The disease resistance response and the hormone signal transduction that how to influence plant for the transportation of research mRNA caryoplasm provide new clue, for the crossover network of exploring plant hormone Whitfield's ointment and ethene provides new node.In agriculture prodn, the present invention can be applied to the disease resistance response of enhancement of plant and the initiative of the genetically modified crops of the aspects such as ripe old and feeble period of regulation and control crop.
In order to seek other gene on the EDR1 signal path, we have carried out edr1 and have suppressed son and enhanser mutant choice.We have obtained one of them two mutants edos1, and through map based cloning, we have obtained the EDOS1 gene.Through the functional analysis to EDOS1, we find that EDOS1 has important regulation to the caryoplasm transportation of plant disease-resistant reaction and plant mRNA.
On the other hand; As eukaryote; The RNA of plant is synthetic in nuclear; And RNA to translate into protein be in tenuigenin, to carry out, reaction plays important regulation with adverse circumstance to growth and development of plant in the caryoplasm transportation of RNA, several reports of minority have explained that influencing the two mutants that the mRNA caryoplasm transports can regulate the resistance of plant to pathogenic bacteria.Certainly, at present still very limited to the research of the concrete mechanism of the molecular mechanism of plant mRNA transportation and the reaction of mRNA transportation regulation and control plant disease-resistant.
Through research, can help us to understand the signal path of EDR1 to EDOS1.Simultaneously, through the research to EDOS1, we can further investigate the influence of mRNA caryoplasm transportation to the plant disease-resistant reaction.What is more important, EDOS1 itself also is the suitable target gene of crop transgene improvement and crop molecular designing.In sum, the clone and the functional analysis of EDOS1 gene had important theory and realistic meaning, and the EDOS1 gene there is wide application prospect.
1. the gene order of an arabidopsis gene EDOS1, full-length gene group dna sequence dna 4302bp, CDS total length 1800bp.Its CDS sequence is seen SEQ ID No.1.
2.EDOS1 protein sequence, 599 amino acid of encoding, big or small 68.3kD.Its aminoacid sequence is seen SEQ ID No.2.
3.EDOS1 after the sudden change, can suppress disease-resistant phenotype and the powdery mildew inductive necrocytosis phenotype of edr1.
4.EDOS1 after the sudden change, can strengthen the ethene inductive leaf senile phenotype of edr1.
5.EDOS1 after the sudden change, mRNA accumulates in nucleus.
6.EDOS1 gene extensively exists and high conservative in higher plant, can be applied to gene engineering technology field, disease resistance response and the mature period of regulation and control crop.
Particular content is following:
1. Arabidopis thaliana powder mildew resistance genes involved EDOS1, it is one of following nucleotide sequences:
1) nucleotide sequence of SEQ ID No.1;
2) nucleotide sequence with SEQ ID No.1 has 90% above homology and the proteinic nucleotide sequence of coding identical function.
2. protein, it is by gene EDOS1 coding of above 1.
3. above 2 protein, it is:
1) by the albumen shown in the aminoacid sequence of SEQ ID No.2; Or
2) with the aminoacid sequence of (1) through replacement, lack or add one or several amino acid and have and (1) identical functions by (1) deutero-albumen.
4. the mutator gene edos1 of above 1 EDOS1 gene, the difference of the nucleotide sequence of its nucleotide sequence and SEQ IDNo.1 is to have occurred to the position of 2886bp at 2879bp the disappearance of 7bp.
5. above 1 gene EDOS1 or above 2 or 3 protein are used to regulate and control the application that plant mildew-resistance resistance, powdery mildew are induced programmed cell death, ethene slaking or the transportation of mRNA caryoplasm of generation; Wherein said plant comprises Arabidopis thaliana, barley, grape, wheat, is preferably arabidopsis mutant body edr1.
6. above 5 application; Wherein suppress the plant powdery mildew resistance, suppress application that powdery mildew induces the programmed cell death of generation, promotes the ethene slaking or suppress the transportation of mRNA caryoplasm and realize, wherein suppress the protein expression of above 1 gene EDOS1 or above 2 or 3 and can realize through making above 1 gene EDOS1 that function mutation take place to lack through in said plant, suppressing above 1 gene EDOS1 or above 2 or 3 protein expression; And enhancement of plant mildew-resistance resistance, promote powdery mildew to induce the programmed cell death of generation, ethene suppressing slaking or the application that promotes the transportation of mRNA caryoplasm through excessive in said plant or more than the overexpression 1 gene EDOS1 or above 2 or 3 protein realize.
7. above 4 mutator gene edos1 is used to suppress plant mildew-resistance resistance, suppresses the application that powdery mildew is induced the programmed cell death of generation, promoted ethene slaking and the transportation of inhibition mRNA caryoplasm; Wherein said plant comprises Arabidopis thaliana, barley, grape, wheat, is preferably arabidopsis mutant body edr1.
Description of drawings
Fig. 1. the wild-type Col-0 in 5 weeks of growth, edr1, the phenotype of edos1/edr1 inoculation white powder germ after 8 days (on) and the result (descending) of Trypan Blue Staining.
Fig. 2. the wild-type in 5 weeks of growth, phenotype after the 100ppm ethylene gas was handled 3 days of edr1, edos1/edr1 (on) and the result (descending) that measures of chlorophyll content.
Fig. 3. clone the sketch map of EDOS1 gene, position and the gene structure and the complementary result of edos1 sudden change through the method for map based cloning.
Fig. 4. the comparison of the phenotype of two mutants and Col-0 performance.
Fig. 5. the compare of analysis of EDOS1 protein sequence in the plants such as Arabidopis thaliana, paddy rice, corn, clover.
Fig. 6 .Col-0, among edos1 and the edos1/edr1, mRNA accumulates in nucleus.
Embodiment
Embodiment one edos1 can suppress the disease-resistant phenotype of edr1 two mutants
(1) material and method
Edr1 two mutants (Frye and Innes, 1998; Frye et al., 2001) seed soaked 16 hours with 0.3%EMS (Ethyl methanesulfonate, available from SIGMA-ALDRICH, article No. M-0880), used ddH
20 washes 12 times.Be sowed at seed in the soil, 22 ℃, illumination in 16 hours was grown for 7 weeks.Receive seed by 20 plant as a Pool, obtain 240 Pool altogether.200 of each Pool sowings, 22 ℃, grew for 5 weeks in the illumination greenhouse in 9 hours, seeks the individuality that the edr1 phenotype do not occur, and called after edos series mutation body is also incited somebody to action a wherein strain called after edos1/edr1.
Wild-type Col-0 (available from Arabidopsis Biological Resource Center), edr1, the edos1/edr1 two mutants is in soil; 22 ℃; Grew after 5 weeks in 9 hours illumination greenhouses, inoculation white powder germ (Golovinomyces cichoracearum) (Frye and Innes, 1998).This white powder germ preserves with the two mutants pad4 to this white powder germ susceptible (Jirage et al., 1999) of Arabidopis thaliana.Have the plant of the pad4 of a large amount of white powder spores to scrape lightly treating growth when connecing bacterium and connect on the plant leaf of white powder, connect bacterium after, the plant that has connect bacterium was covered 24 hours with preservative film earlier, after remove preservative film, disease-resistant phenotypic evaluation is carried out in continued growth 7 days.Typical blade is taken a picture, and representational blade is carried out Trypan Blue Staining, the necrocytosis spot of the mycelial growth of mark powdery mildew and blade.
(2) result and analysis
The inoculation powdery mildew grew a large amount of powdery mildew (Fig. 1 is upper left) after 8 days on the wild-type plant; Have only very faint powdery mildew growth on the edr1 two mutants plant, and the spot (among Fig. 1) of visible necrocytosis is arranged; The phenotype of edos1/edr1 two mutants is similar to wild-type, and the growth of tangible powdery mildew is promptly arranged, and does not have the spot (Fig. 1 is upper right) of tangible necrocytosis.
The result of Trypan Blue Staining has more clearly illustrated wild-type, edr1, and edos1/edr1 is to the reaction of powdery mildew.Can see a large amount of mycelia (under Fig. 1 left side) on the blade of wild-type plant; Almost do not have the visible mycelia on the edr1 plant, clearly navy blue necrotic spot (under among Fig. 1) is but arranged; Phenotype and the wild-type of edos1/edr1 similar (Fig. 1 bottom right).
Embodiment two edos1 can strengthen the ethene inductive leaf senile phenotype of edr1
1. induce down at ethene, the edos1/edr1 two mutants shows the phenotype of blade yellow more seriously
(1) materials and methods
Arabidopis thaliana plant Col-0, edr1 and edos1/edr1 are in soil, and 22 ℃, grew after 5 weeks in the greenhouse of illumination in 9 hours, transfer to one pellucidly in the closed glass container, the ethylene gas that reinjects, and concentration is 100ppm.After 72 hours, observe phenotype.Simultaneously, take by weighing 0.2 gram blade respectively,, measure chlorophyll content wherein again with straight alcohol extraction chlorophyll wherein.Simultaneously, detect untreated plant as contrast.
(2) result and analysis
The plant of not handling with ethene all do not have yellow phenotype significantly (Fig. 2 a, b, c).After ethene was handled, yellow (Fig. 2 d) had appearred in the blade of wild-type Col-0; The edr1 two mutants shows yellow phenotype more seriously (Fig. 2 e); The edos1/edr1 two mutants demonstrates the old and feeble phenotype (Fig. 2 f) of extremely serious yellow.
Can find out also that from the experiment of measuring chlorophyll content when not handling, the chlorophyll content of all plant is similar; And after ethylene gas is handled, compare with wild-type Col-0, the chlorophyll content of edr1 obviously reduces; And compare with edr1, the chlorophyll content of edos1/edr1 sharply reduces (Fig. 2 figure below).
The map based cloning and the complementation of embodiment three EDOS1 genes
(1) material and method
Be clone EDOS1 gene, we are edos1/edr1 two mutants and the environmental plant of Lansberg (available from Arabidopsis Biological Resource Center) hybridization, and the F1 of acquisition produces F2 generation for selfing.From F2 for selecting 80 individual plants similar the plant with the edos1/edr1 mutation type surface; Carry out genotype identification (Konieczny and Ausubel, 1993) with coarse positioning mark (http://signal.salk.edu/genome/SSLP_info/SSLPsordered.html).The result of coarse positioning shows that EDOS1 is positioned at chromosomal front end No. 5.Subsequently, we have designed the mark of Fine Mapping, and the F2 plant about 4000 is carried out genotype identification, navigate to the BAC of No. five karyomit(e) front ends to EDOS1, and (Fig. 3 a) for the zone about a 30kb among the F17I14.All genes in this zone are checked order, and through order-checking, we have found the disappearance (Fig. 3 b) of one section 7bp in Gene A T5G09860.
For having this mutational site, the phenotype of verifying edos1 causes; Our from the genomic dna of wild-type Col-0, increase Gene A T5G09860 fragment of a 5.7kb; Comprise the promoter region of 800bp, the genomic dna of 4.3kb, the flanking sequence of 200bp.Amplimer is LeftPrimer:5 '-cagagcaaatcgaattacgtaacca-3 '; Right Primer:5 '-ggagatcggtggtgtttatgga-3 '.56 ℃ of annealing temperatures are extended 6min, 35 circulations.After cutting with Kpn I and Hind III enzyme, be connected on the pCAMBIA1300 carrier (GenBank sequence number AF234296, (Heo et al., 1999)).Use the heat shock method to be transformed into Agrobacterium GV3101 (Hajdukiewicz et al., 1994 to plasmid; Sambrook et al., 1989) in, identify that correct back transforms the edos1/edr1 plant, obtains the T0 seed in generation.
(0.8% agar pH=5.8) screens the T0 seed in generation, obtains the T1 transfer-gen plant for 4.33g MS salt/L, 1% sucrose with the MS substratum that contains the 120mg/L Totomycin.Handle with ethylene gas, carry out phenotypic evaluation, identify the phenotype that can transfer-gen plant whether complementary edos1/edr1.
(2) result and analysis
After checking order through the cDNA to this gene, we find that variation has taken place in the montage of deletion segment, and the insertion (Fig. 3 c) of one section 11bp has appearred in the cDNA of two mutants.CDNA to after the sudden change carries out the prediction of coding protein sequence, and after the fragment of inserting 11bp, two successive terminator codons have appearred in encoding sequence in the mutational site, has caused in the two mutants albumen (Fig. 3 d) of a brachymemma of EDOS1 coding of sudden change.The EDOS1 gene is a gene (Fig. 3 e) with a plurality of exons.Grow after 4 weeks; T1 is different with edos1/edr1 for the phenotype of complementary plant (AT5G09860 in edos1/edr1); And with edr1 closely similar (Fig. 3 f); The phenotype that changing over to of Gene A T5G09860 can complementary edos1/edr1 is described, and the phenotype of edos1/edr1 is just because of the 14 7bp disappearance that exon is terminal among the AT5G09860, promptly lacks to the 7bp of 2886bp position and cause at 2879bp.Therefore the EDOS1 gene is Gene A T5G09860.
The phenotype of embodiment four edos1 single mutation
The phenotype of edos1 single mutation plant
(1) material and method
Be to obtain the edos1 single mutant, we hybridize with edos1/edr1 and Col-0, F2 for plant in searching do not have the plant of edr1 background.What detection edr1 background was used is Derived CleavedAmplified Polymorphic Sequences (dCAPS) method (Neff et al., 2002).Primer is LP:5 '-AGGCTGAAAGGACAGATTCTTCATG-3 '; RP:5 '-TGTTGAGGAATTGTTCTCCAACTGA-3 '.It is HaeIII (available from Dalian TAKARA company) that enzyme is cut used enzyme.
Wild-type Col-0 and edos1 two mutants are seeded in the soil.22 ℃, illumination in 9 hours was grown after 5 weeks, the false monospore bacillus (Pseudomonas syringae) of injection inoculation cloves (Chisholm et al., 2006; Weigel and Glazebrook, 2002), concentration is OD
600=5 * 10
-4After 3 hours and 72 hours, take off blade, use ddH with punch tool
2O cleans, and grinds the blade sample, presses gradient dilution, is coated with flat board respectively, counting after 3 days.
Wild-type Col-0, edr1, edos1/edr1 and edos1 two mutants are seeded in the soil.22 ℃, illumination in 9 hours was grown after 5 weeks, was put in the sealing ground transparent vessel, injected the 100ppm ethylene gas.After 72 hours, get the 0.2g blade, use the straight alcohol extracting chlorophyll, measure chlorophyll content again.Simultaneously, detect untreated plant as contrast.
(2) result and analysis
Can find out that from the last figure of Fig. 4 in the blade sample of the false monospore bacillus of injection inoculation cloves after 3 hours, pathogen growth does not have notable difference in the blade of edos1 and Col-0.After 72 hours, in the edos1 two mutants, pathogen growth is a little more than Col-0.
Can find out that from figure below of Fig. 4 after ethylene gas is handled 72 hours, compare with edr1 with Col-0, the chlorophyll content of edos1 single mutant and edos1/edr1 double-mutant all significantly reduces.And in contrasting, Col-0, edr1, edos1/edr1, the chlorophyll content of edos1 is very approaching. with being untreated
Above presentation of results, Col-0 compares with wild-type, and the edos1 single mutant has reduction by a small margin to the resistance of pathogenic bacteria; Simultaneously, the edos1 single mutant significantly improves ethene inductive leaf senile susceptibility.
The aminoacid sequence of embodiment five EDOS1 is conservative at the higher plant camber
EDOS1 encoded protein sequence high conservative.With paddy rice (Oryza sativa, NP_001048715), willow (Populus trichocarpa; XP_002299188), and castor-oil plant (Ricinuscommunis, XP_002529986); Grape (Vitis vinifera, XP_002263874), corn (Zeamays; NP_001159168) the homogenic aminoacid sequence comparison in can find that the similarity of its sequence reaches more than 80%.This result has hinted that EDOS1 has very important function; Also be illustrated in these plants that economic worth is arranged; EDOS1 very maybe be also closely related with aging with plant disease-resistant, also explained to be applied to the huge possibility in the plant genetic engineering to the EDOS1 gene.As shown in Figure 5.
Embodiment six is in the edos1/edr1 two mutants, and mRNA accumulates in nucleus
(1) materials and methods
Arabidopis thaliana plant Col-0, edr1 and edos1/edr1 after 2 weeks of growth on the MS substratum, choose length less than the blade of 5mm with 50% stationary liquid (120Mm NaCl, 7mM Na
2HPO4,3mMNaH
2PO
4, 2.7mM KCl, 0.1%Tween 20,80mM EGTA, 5%formaldehyde, and 10%DMSO) fixing.Use 50% ethanol after fixing the completion, 50% N.F,USP MANNITOL replaces wash-out 3 times.Join blade in the hybridization buffer (available from Sigma-Aldrich, article No. H-7033), add 5 '-fluorescein-labeled oligo dT, hybridization is spent the night.After using the ultrapure water wash-out again, observe with laser confocal microscope.Exciting light wavelength is 488nm.
(2) result and analysis
Can find out, all have only in the cell of wild-type Col-0 and edr1 fluorescent signal in the very faint nucleus (Fig. 6 a, b).And in the edos1/edr1 two mutants, the interior signal of nucleus is significantly arranged, show that mRNA accumulates (Fig. 6 c) in nucleus.This result explains that also EDOS1 plays important regulation in the transportation of mRNA caryoplasm.
Reference
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Claims (7)
1. Arabidopis thaliana powder mildew resistance genes involved EDOS1, it is one of following nucleotide sequences:
1) nucleotide sequence of SEQ ID No.1;
2) nucleotide sequence with SEQ ID No.1 has 90% above homology and the proteinic nucleotide sequence of coding identical function.
2. protein, it is by gene EDOS1 coding of claim 1.
3. the protein of claim 2, it is:
1) by the albumen shown in the aminoacid sequence of SEQ ID No.2; Or
2) with the aminoacid sequence of (1) through replacement, lack or add one or several amino acid and have and (1) identical functions by (1) deutero-albumen.
4. the mutator gene edos1 of the EDOS1 gene of claim 1, the difference of the nucleotide sequence of its nucleotide sequence and SEQID No.1 is to have occurred to the position of 2886bp at 2879bp the disappearance of 7bp.
5. the protein of the gene EDOS1 of claim 1 or claim 2 or 3 is used to regulate and control the application that plant mildew-resistance resistance, powdery mildew are induced the programmed cell death of generation, ethene slaking or the transportation of mRNA caryoplasm; Wherein said plant comprises Arabidopis thaliana, barley, grape, wheat, is preferably arabidopsis mutant body edr1.
6. the application of claim 5; Wherein suppress plant mildew-resistance resistance, suppress application that powdery mildew induces the programmed cell death of generation, promotes the ethene slaking or suppress the transportation of mRNA caryoplasm through in said plant, suppressing the gene EDOS1 of claim 1 or the protein expression of claim 2 or 3 is realized, the disappearance function mutation can take place through the gene EDOS1 that makes claim 1 protein expression that wherein suppresses gene EDOS1 or claim 2 or 3 of claim 1 realize; And enhancement of plant mildew-resistance resistance, the application that promotes powdery mildew to induce the programmed cell death of generation, ethene suppressing slaking or promotion mRNA caryoplasm to transport realize through the gene EDOS1 of excessive in said plant or overexpression claim 1 or the protein of claim 2 or 3.
7. the mutator gene edos1 of claim 4 is used to suppress the plant powdery mildew resistance, suppresses the application that powdery mildew is induced the programmed cell death of generation, promoted ethene slaking and the transportation of inhibition mRNA caryoplasm; Wherein said plant comprises Arabidopis thaliana, barley, grape, wheat, is preferably arabidopsis mutant body edr1.
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| CN107236746A (en) * | 2017-07-28 | 2017-10-10 | 福建农林大学 | Wheat powdery mildew resistant gene PmR2 and its clone and application |
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Cited By (4)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN102942621A (en) * | 2012-10-30 | 2013-02-27 | 中国农业大学 | Plant powdery mildew resistance related protein TaCAF1 and its coding gene and application |
| CN102942621B (en) * | 2012-10-30 | 2014-03-05 | 中国农业大学 | Plant powdery mildew resistance related protein TaCAF1 and its coding gene and application |
| CN107236746A (en) * | 2017-07-28 | 2017-10-10 | 福建农林大学 | Wheat powdery mildew resistant gene PmR2 and its clone and application |
| CN107236746B (en) * | 2017-07-28 | 2020-03-24 | 福建农林大学 | Wheat powdery mildew resistance gene PmR2 and clone and application thereof |
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