CN102586268A - Cloning and application of arabidopsis oidium disease resistance suppressor gene EDTS3 - Google Patents
Cloning and application of arabidopsis oidium disease resistance suppressor gene EDTS3 Download PDFInfo
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Abstract
The invention relates to separated cloning, functional verification and application of arabidopsis oidium disease resistance suppressor gene EDTS3, and further relates to a mutant gene of the EDTS3 gene, protein encoded by the EDTS3 gene and application of the EDTS3 gene.
Description
Technical field
The application relates to the plant gene engineering technology field, is specifically related in the model plant Arabidopis thaliana clone, functional verification and application to the positive regulatory factor of powdery mildew resistance.
Background technology
Powdery Mildew is the Plant diseases that is caused by the Erysiphaceae in the fungi.When plant is infected morbidity, can produce the macroscopic white meal that constitutes by mycelium, conidiophore and conidium in a large number and also gain the name therefrom.Powdery Mildew distributes in the whole world extensively, and the harm dicotyledons is particularly general.It also infects multiple grass simultaneously, like wheat, barley, oat and multiple herbage.For example wheat powdery mildew is to be caused by specialization parasitical fungi standing grain Bu Shi powdery mildew, is one of main disease that influences Wheat Production.Wheat powdery mildew once was very popular continuously several times in China, had caused great financial loss.
Because there are various pathogenic bacterias in occurring in nature, plant has formed multi-level defense system gradually in secular coevolution, and it mainly comprises the resistance of non-host-resistance, basic resistance and disease-resistant gene mediation.
Non-host-resistance is mainly through plant self structure and secondary metabolite defence; Like PLANT CYTOSKELETON and de novo synthesis phytoalexin-antibiotic complex; So have wide spectrum, resistance is strong and persistent characteristics, is the major cause that plant can be resisted most of potential pathogen infections.
The basis resistance have another name called based on pathogen-associated molecular pattern (Pathogen-associated molecularpatterns, the resistance that PAMPs) excites (PAMP-Triggered Immunity, PTI).Pathogen-associated molecular pattern is that host self does not have, but in pathogenic bacteria, extensively exists, and the characterization of molecules of and high conservative most important again to its life history, like flagellin, the elongation factor of bacterium, LPS in the fungi and Regitex FA etc.The physical efficiency that receives on host cell surface is discerned these molecules; And the cascade signal path of the MAPK in activation downstream, signal is transmitted into nuclear the most at last, activates the protein induced course of disease (Pathogen-Responsive of WKRY; PR) expression of gene; (Reactive OxygenSpecies, generation ROS) and VISOSE (callose) be in the accumulation (1) of p of E, thereby resist the invasion of pathogenic bacteria for active oxygen.The characteristics of basis resistance are that anti-spectrum is wide, but resistance level is lower usually.
The resistance of disease-resistant gene mediation have another name called the resistance that effector excites (Effector TriggeredImmunity, ETI).After host plant is evolved out PTI, the infection mechanism that the part pathogenic bacteria develops and makes new advances.For example, bacterium utilizes III type excretory system that effector is directly imported vegetable cell inside, suppresses the generation of PTI.In order to resist infecting of these pathogenic bacterias; Plant evolution has gone out disease-resistant gene (Resistance gene; The R gene) comes the recognition effect factor (at this moment just can be called Avr albumen) and induced hypersensitivity reaction (Hypersensitive Response; Limiting growth of pathogenic bacteria and spreading (Chisholm et al., 2006) HR).The characteristics of the resistance of R gene mediated are high specificities, and anti-spectrum is narrow, and resistance is higher.
The disease resistance response of plant often receives the adjusting and the control of intravital three kinds of signaling molecules, and these signaling molecules are respectively Whitfield's ointment SA, jasmonic JA and ethene ET.Wherein SA is the center regulatory factor of various plant disease-resistant reactions, is to set up (System Acquire Resistance, basis SAR).And the disease-resistant signal path and the SA path that are mediated by JA/ET are mutual antagonisms.Comparatively typical example is exactly that false pseudomonas bacillus a kind of plant growth substance psendomonas syringae (coronatine) of producing similar JA/MeJA suppresses the resistance of SA-mediation, thereby reduces the resistance (2) of plant.
At present, the cloned resistance gene method mainly contains: subtractive hybridization, yeast two-hybrid system, map based cloning technology, transposon tagging and homologous gene clone etc.For the model plant Arabidopis thaliana, the map based cloning isolated genes becomes one of topmost method, and the simplification day by day of the especially discovery of the completion of genome sequencing plan and various molecule markers, and DNA extraction method is significantly shortened the clone gene spended time.
Present disease-resistant research shows that Arabidopis thaliana is similar with the unifacial leaf grass with other dicotyledonss on disease-resistant molecule mechanism.Compare with the R gene, disease-resistant related gene can provide more wide spectrum and long lasting resistance for plant.Utilizing the purpose of model plant clone plant disease-resistant related gene, is the homologous gene that has similar functions in the important plant in order to seek, and through plant gene engineering technology, improves the resistance of wide spectrum of plant to pathogenic bacteria.
The disease-resistant gene of from Arabidopis thaliana, cloning has the potential using value in production.For example, RPW8 is a disease-resistant gene of on Arabidopis thaliana, cloning, and NPR1 is the important regulating and controlling factor of the disease resistance response of in Arabidopis thaliana, cloning.Expressing the RPW8 gene can improve the resistance of tobacco to the white powder germ excessively in tobacco; Expressing arabidopsis gene NPR1 can improve the resistance of paddy rice to bacterium excessively in paddy rice.
EDR1, EDR2 and EDR3 are the genes involveds of regulation and control powder mildew resistance in the Arabidopis thaliana, the sudden change of these genes cause plant that the disease resistance of powdery mildew is strengthened and the necrocytosis (Program Cell Death) of inducing generation (3-5).Wherein, the EDR1 protein kinase of encoding has very high homology with CTR1 in the ethylene reaction; The EDR3 macromolecular GTPase that encodes.The edr2 two mutants, with edr1, edr3 has similar phenotype, also has the old and feeble reaction of enhanced powder mildew resistance and ethene, and its disease resistance also needs the Whitfield's ointment signal path, does not need jasmonic or Ethylene Signal path.The EDR2 novel albumen of encoding; Have three structural domains; Comprise a Pleckstrin Homology (PH) structural domain, STeroidogenic Acute Regulatoryprotein-related lipid-Transfer (START) structural domain and Unknown Function but the conservative carboxyl terminal structural domain DUF1336 of sequence.The function in preceding two zones possibly be and contaminated with lipid that the carboxyl terminal structural domain then possibly participated in albumen and proteic interaction.EDR2 has two lipid molecule lands, and the disease resistance of hint lipid metabolism and Powdery Mildew and the programmed death of vegetable cell are closely related.
With edr2 is point of penetration; The research that utilizes the model plant Arabidopis thaliana is the complete sum system comparatively; The comparatively advanced and sophisticated advantage of technology; Excavate disease-resistant related gene, accelerate the understanding of enantiopathy gene function and the understanding of signal path and even network structure and expand, with probably theoretical foundation and technical support being provided for the disease-resistant control of farm crop and molecular designing breeding.
Summary of the invention
The application is included in the model plant Arabidopis thaliana clone, functional verification and the application to the positive regulatory factor EDTS3 of powdery mildew resistance.The sudden change of EDTS3 gene can suppress the powdery mildew resistance of arabidopsis mutant body edr2, and suppress simultaneously powdery mildew induce produce programmed cell death (Programmedcell death, PCD).
(1) a kind of Arabidopis thaliana mildew-resistance suppressor gene EDTS3 sequence, CDS full length sequence 264bp, intronless.Its nucleotide sequence is shown in SEQ ID No.1.
(2) a kind of Arabidopis thaliana mildew-resistance genes involved EDTS3 encoded protein sequence, 87 amino acid of encoding, big or small 9.784kD.Its aminoacid sequence is shown in SEQ ID No.2.
(3) after EDTS3 transgenation forfeiture self function, show the powdery mildew resistance and the inhibition that suppress the edr2 two mutants and receive powdery mildew to induce the generation necrocytosis.Its phenotype is seen Fig. 1.
(4), can its expression amount be raise through genetic engineering technique, thereby improve the disease resistance of vegetables and crop according to the disease-resistant phenotype of the inhibition after the EDTS3 transgenation.
Particular content is following:
1. Arabidopis thaliana powdery mildew disease-resistant property suppressor gene EDTS3, 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 EDTS3 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 edts3 of above 1 gene EDTS3, the difference of the nucleotide sequence of its nucleotide sequence and SEQ ID No.1 is wherein to take place the A104G sudden change.
5. above 4 mutator gene edts3 encoded protein matter, the difference of the aminoacid sequence of its aminoacid sequence and SEQ ID No.2 are wherein to take place the R35K sudden change.
6. above 1 gene EDTS3 or above 2 or 3 protein are used for regulation and control (strengthen or suppress) plant mildew-resistance resistance or regulation and control (promote or suppress) powdery mildew and induce the application that produces programmed cell death; Wherein said plant comprises Arabidopis thaliana, barley, wheat, is preferably arabidopsis mutant body edr2.
7. above 6 application; Wherein enhancement of plant mildew-resistance resistance or the application that promotes powdery mildew to induce to produce programmed cell death through excessive in said plant or more than the overexpression 1 gene EDTS3 or above 2 or 3 protein realize; Suppress plant mildew-resistance resistance or suppress powdery mildew and induce the application that produces programmed cell death to realize through in said plant, suppressing above 1 gene EDTS3 or above 2 or 3 protein expression, the mistake function mutation can take place through the gene EDTS3 that makes claim 1 protein expression that wherein suppresses gene EDTS3 or claim 2 or 3 of claim 1 realize.
8. above 4 mutator gene edts3 is used to suppress plant mildew-resistance resistance and induces the application that produces programmed cell death with the inhibition powdery mildew, and wherein said plant comprises Arabidopis thaliana, barley, wheat, is preferably arabidopsis mutant body edr2.
Description of drawings
Fig. 1. wild-type and two mutants edr2, edts3/edr2 growth 4-6 inoculate after week the phenotype of white powder germ (G.cichoracearum) after 8 days (on), Trypan blue coloration result (descending).
Fig. 2. separate the sketch map of EDTS3 gene through the method for map based cloning.
Fig. 3. wild-type, two mutants edr2, edts3/edr2 and complementary carrier transform two mutants edts3/edr2 (T1) growth 4-6 and inoculate the phenotype of white powder germ (G.cichoracearum) after 8 days after week.
Embodiment
Embodiment one edts3 two mutants suppresses disease-resistant phenotype analytical
1.edts3/edr2 two mutants suppresses the powder mildew resistance and the necrocytosis of powdery mildew inductive of edr2 two mutants
(1) material and method
Edts3/edr2 (wherein edts3 is enhanced disease resistance two suppressor 3) is through with 0.03%EMS (available from sigma company) mutagenesis two mutants edr2 (4), from mutagenesis colony, filters out of obtaining and can suppress edr2 white powder germ resistance enhanced two mutants.An and then two mutants called after edts3/edr2 that will obtain.
At first; Two mutants edr2; The seed of edts3/edr2 and wild-type col-0 (available from ArabidopsisBiological Resource Center (ABRC)) is broadcast MS substratum (being purchased the Laboratories from PhytoTechnology); After 4 ℃ of vernalization 2-3 days, transfer to the phytotron under the 9h light/15hdark illumination condition.Approximately 7-10 days, move to seedling in the soil.After growth of seedling 4-6 week, inoculation white powder germ (G.cichoracearum) (6).This white powder germ preserves with a two mutants pad4 (7) to this white powder germ susceptible of Arabidopis thaliana.Have the plant of the pad4 of a large amount of white powder spores to scrape lightly growth when connecing bacterium and waiting to connect on the plant leaf of white powder, the plant that will connect bacterium then covers with preservative film earlier preserved moisture 1 day, after remove preservative film, the plant continued growth was carried out disease-resistant phenotypic evaluation after 7 days.The blade that will have typical phenotype is cut, and takes a picture and carries out trypan blue dyeing.
(2) result and analysis
, cover with white powder on the blade of wild-type and double-mutant edts3/edr2, and have basically no visible powdery mildew spore on the blade of two mutants edr2 after 8 days at the inoculation powdery mildew, and the necrocytosis of performance powdery mildew inductive.Dyeing has also proved The above results: the blade of wild-type and double-mutant edts3/edr2 has been covered with mycelia and conidiophore, and dead mesophyll cell is dyed blueness (Fig. 1) in the two mutants.The above results shows that the edts3/edr2 two mutants has suppressed edr2 powdery mildew resistance.
The acquisition of embodiment two EDTS3 genes
1. utilize map based cloning to isolate the EDTS3 gene
We utilize map based cloning to separate the EDTS3 gene.Specific practice is: two mutants edts3/edr2 and environmental Lansberg (being purchased from ABRC) do hybridization, and the F1 of acquisition produces F2 generation for selfing.In F2 generation, select to contain the individual plant of the wild-type phenotype of edr2 sudden change, make its selfing produce F3 generation then., F3 selects the individuality of wild-type in for each strain being; Be evenly distributed on chromosomal telltale mark on 5 karyomit(e)s (http://signal.salk.edu/genome/SSLP_info/SSLPsordered.html) through PCR method and utilization, with the front end of gene Primary Location at the disconnected arm of the 1st karyomit(e).Subsequently, utilize about 3000 strain F3 again, (Fig. 2) with the zone (8) of gene Fine Mapping 40kb between F9H16 and F26F24 for individual plant.This zone comprises 12 genes.Find in the CDS of Gene A t1g21327, to contain a base mutation A104G through order-checking, and other genes have no variation, thereby confirm that this gene is the candidate gene of EDTS3 gene.
2.EDTS3 the structure of gene
EDTS3CDS total length 264bp does not contain intron; EDTS3 albumen is the abundant small protein of a Serine, wherein is made up of 23 amino acid a low complex degree structural domain.
The functional verification of embodiment three EDTS3 genes
For whether checking edts3 sudden change suppresses because the At1g21327 base mutation causes the resistance of edr2 two mutants powdery mildew; We have made up the genetic transformation carrier with the At1g21327 of self promotor; The method of utilizing flower to infect transforms double-mutant edts3/edr2 plant; In T1 generation, transgenic line has recovered the phenotype of edr2 two mutants.
We are template with the DNA of wild-type col-0, amplify the At1g21327 gene fragment of the 5.7kb that has KpnI and PstI with one couple of PCR primers F:5 '-AACTGCAGAAGTGGTGCTCACGTTGTTGTATTT-3 ' and R:5 '-AACTGCAGAAGTGGTGCTCACGTTGTTGTATTT-3 '.
The PCR reaction system
DNA： 2.0ul
10xbuffer： 5.0ul
dNTPs(2.0mM)： 5.0ul
MgSO4(25mM)： 2.0ul
Primers F (10uM): 1.6ul
Primer R (10uM): 1.6ul
KOD?Taq(1U)： 1.0ul
H2O：31.8ul
The PCR response procedures: in advance sex change, 95 ℃ 3 minutes; Thermal cycling is annealed 55 ℃ of 30 second in 95 ℃ of 30 second of sex change, extend 72 ℃ 5 minutes, 30 of cycle numbers.
Entire segment comprise 3.6Kb promoter region, 0.26kb coding region and 1.8Kb 3 '-UTR.The fragment of amplification and pCAMBIA1300 (9) be double digestion (KpnI and PstI simultaneously; Available from TaKaRa company; 37 ℃), connect (T4 ligase enzyme (available from NEB company); 16 ℃) and heat shock method transformed into escherichia coli E.coli DH10B (available from full formula King Company) after, identify through order-checking (the big genome company of China) to transform Agrobacterium GV3101 (9) with freeze-thaw method subsequently by correct plasmid.The method of contaminating with flower (10) transforms double-mutant edts3/edr2 plant, and 7 all backs obtain T1 for seed.To seed disinfection, cleaning, be seeded into then in the MS substratum that adds Totomycin (80 μ g/ml).After 7-10 days, select T1 for the transgenic positive plant.
At 22 ℃, under the 9h light/15h dark condition, long to 4-6 week when seedling, we observe phenotype simultaneously wild-type, two mutants and transgenic T1 pickup kind powdery mildew after 8 days.The double-mutant edts3/edr2 phenotype that changed complete complementation over to of At1g21327 gene shows as edr2 phenotype (see figure 3) as a result.Show thus two mutants disease-resistant phenotype since the sudden change of At1g21327 gene cause.Therefore the EDTS3 gene is the At1g21327 gene.
The application of embodiment four Arabidopis thaliana powdery mildew resistance related gene EDTS3 in the crop disease-resistant improvement
After EDTS3 transgenation forfeiture self function, show the powdery mildew resistance and the inhibition that suppress the edr2 two mutants and receive powdery mildew to induce the generation necrocytosis.Can make its excessive or overexpression through genetic engineering technique according to this characteristic, thereby improve the disease resistance of vegetables and crop, cultivate crop varieties with disease-resistant proterties.
Reference
1.Chisholm?ST，Coaker?G，Day?B，&?Staskawicz?BJ(2006)Host-microbeinteractions：shaping?the?evolution?of?the?plant?immune?response.Cell124(4)：803-814.
2.Mudgett?MB(2005)New?insights?to?the?function?of?phytopathogenic?bacterialtype?III?effectors?in?plants.Annu?Rev?Plant?Biol?56：509-531.
3.Frye?CA，Tang?D，&?Innes?RW(2001)Negative?regulation?of?defense?responsesin?plants?by?a?conserved?MAPKK?kinase.Proc?Natl?Acad?Sci?USA98(1)：373-378.
4.Tang?D，Ade?J，Frye?CA，&?Innes?RW(2005)Regulation?of?plant?defenseresponses?in?Arabidopsis?by?EDR2，a?PH?and?START?domain-containingprotein.Plant?J?44(2)：245-257.
5.Tang?D，Ade?J，Frye?CA，&?Innes?RW(2006)A?mutation?in?the?GTP?hydrolysissite?of?Arabidopsis?dynamin-related?protein?1E?confers?enhanced?cell?death?inresponse?to?powdery?mildew?infection.(Translated?from?eng)Plant?J47(1)：75-84(in?eng).
6.Adam?L，et?al.(1999)Comparison?of?Erysiphe?cichoracearum?and?E.cruciferarum?and?a?survey?of?360?Arabidopsis?thaliana?accessions?for?resistanceto?these?two?powdery?mildew?pathogens.Mol?Plant?Microbe?Interact12(12)：1031-1043.
7.Jirage?D，et?al.(1999)Arabidopsis?thaliana?PAD4?encodes?a?lipase-like?gene?thatis?important?for?salicylic?acid?signaling.(Translated?from?eng)Proc?Natl?AcadSci?USA?96(23)：13583-13588(in?eng).
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Claims (8)
1. Arabidopis thaliana powdery mildew disease-resistant property suppressor gene EDTS3, 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 EDTS3 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 edts3 of the gene EDTS3 of claim 1, the difference of the nucleotide sequence of its nucleotide sequence and SEQID No.1 is wherein to take place the A104G sudden change.
5. the mutator gene edts3 encoded protein matter of claim 4, the difference of the aminoacid sequence of its aminoacid sequence and SEQID No.2 are wherein to take place the R35K sudden change.
6. the protein of the gene EDTS3 of claim 1 or claim 2 or 3 is used to regulate and control plant mildew-resistance resistance or regulates and control powdery mildew and induce the application that produces programmed cell death; Wherein said plant comprises Arabidopis thaliana, barley, wheat, is preferably arabidopsis mutant body edr2.
7. the application of claim 6; Wherein enhancement of plant mildew-resistance resistance or promotion powdery mildew induce the application that produces programmed cell death to realize through the gene EDTS3 of excessive in said plant or overexpression claim 1 or the protein of claim 2 or 3; Inhibition plant mildew-resistance resistance or inhibition powdery mildew induce the application that produces programmed cell death to realize through the gene EDTS3 of inhibition claim 1 or the protein expression of claim 2 or 3 in said plant, wherein suppress the gene EDTS3 of claim 1 or the protein expression of claim 2 or 3 and can realize through the gene EDTS3 generation mistake function mutation that makes claim 1.
8. the mutator gene edts3 of claim 4 is used to suppress plant mildew-resistance resistance and suppresses powdery mildew and induce the application that produces programmed cell death, and wherein said plant comprises Arabidopis thaliana, barley, wheat, is preferably arabidopsis mutant body edr2.
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| CN104211792A (en) * | 2013-05-31 | 2014-12-17 | 中国科学院遗传与发育生物学研究所 | Powdery mildew resistance associated protein, encoding gene thereof and application of both |
| CN104788552A (en) * | 2014-01-17 | 2015-07-22 | 中国科学院遗传与发育生物学研究所 | ScRGA-6RL2 protein and coding gene and application thereof |
| CN104788550A (en) * | 2014-01-17 | 2015-07-22 | 中国科学院遗传与发育生物学研究所 | ScRGA-6RL3 protein and coding gene and application thereof |
| CN104830872A (en) * | 2015-05-18 | 2015-08-12 | 山西省农业科学院棉花研究所 | Cotton GhEDR2 gene as well as encoding protein and application of cotton GhEDR2 gene |
| CN108690808A (en) * | 2018-05-29 | 2018-10-23 | 广东省农业科学院蔬菜研究所 | A kind of preservation of cucurbits powdery mildew bacterium and propagation method |
| CN108728477A (en) * | 2017-04-24 | 2018-11-02 | 华东理工大学 | A kind of efficient Transpositional mutation system and construction method |
| CN112979776A (en) * | 2021-03-29 | 2021-06-18 | 浙江省农业科学院 | Pear protein fragment PyEDR1-307 and coding sequence and application thereof |
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2011
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| CN104211792A (en) * | 2013-05-31 | 2014-12-17 | 中国科学院遗传与发育生物学研究所 | Powdery mildew resistance associated protein, encoding gene thereof and application of both |
| CN104211792B (en) * | 2013-05-31 | 2017-05-10 | 中国科学院遗传与发育生物学研究所 | Powdery mildew resistance associated protein, encoding gene thereof and application of both |
| CN104788552A (en) * | 2014-01-17 | 2015-07-22 | 中国科学院遗传与发育生物学研究所 | ScRGA-6RL2 protein and coding gene and application thereof |
| CN104788550A (en) * | 2014-01-17 | 2015-07-22 | 中国科学院遗传与发育生物学研究所 | ScRGA-6RL3 protein and coding gene and application thereof |
| CN104788552B (en) * | 2014-01-17 | 2017-11-03 | 中国科学院遗传与发育生物学研究所 | ScRGA 6RL2 and its encoding gene and application |
| CN104788550B (en) * | 2014-01-17 | 2017-11-28 | 中国科学院遗传与发育生物学研究所 | ScRGA 6RL3 and its encoding gene and application |
| CN104830872A (en) * | 2015-05-18 | 2015-08-12 | 山西省农业科学院棉花研究所 | Cotton GhEDR2 gene as well as encoding protein and application of cotton GhEDR2 gene |
| CN104830872B (en) * | 2015-05-18 | 2017-10-24 | 山西省农业科学院棉花研究所 | Cotton GhEDR2 genes and its encoding proteins and application |
| CN108728477A (en) * | 2017-04-24 | 2018-11-02 | 华东理工大学 | A kind of efficient Transpositional mutation system and construction method |
| CN108728477B (en) * | 2017-04-24 | 2022-02-22 | 华东理工大学 | Efficient transposition mutation system and construction method |
| CN108690808A (en) * | 2018-05-29 | 2018-10-23 | 广东省农业科学院蔬菜研究所 | A kind of preservation of cucurbits powdery mildew bacterium and propagation method |
| CN112979776A (en) * | 2021-03-29 | 2021-06-18 | 浙江省农业科学院 | Pear protein fragment PyEDR1-307 and coding sequence and application thereof |
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