CN105154453A - Pathogenicity related botrytis cinerea gene BcSep4 and application thereof - Google Patents
Pathogenicity related botrytis cinerea gene BcSep4 and application thereof Download PDFInfo
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- CN105154453A CN105154453A CN201510672004.9A CN201510672004A CN105154453A CN 105154453 A CN105154453 A CN 105154453A CN 201510672004 A CN201510672004 A CN 201510672004A CN 105154453 A CN105154453 A CN 105154453A
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Abstract
The invention provides a pathogenicity related botrytis cinerea gene BcSep4 and an application thereof, belonging to the technical field of microbial gene engineering. The gene BcSep4 which is provided by the invention, comes from botrytis cinerea and is used for controlling infection structure development and pathogenicity has a DNA (deoxyribonucleic acid) sequence which is shown in SEQ ID No:1 in a sequence table and is formed by 2793 nucleotides. A protein coded by the gene BcSep4 provided by the invention has an amino acid sequence which is shown in SEQ ID No:2 in the sequence table and is formed by 343 amino acids. The gene BcSep4 can be applied in the field of plant botrytis cinerea resistant gene engineering. Defects of an infection structure and the pathogenicity occur by deleting, mutating or modifying the protein BcSep4 for controlling infection structure development and the pathogenicity of botrytis cinerea. The protein BcSep4 can be applied to designing and screening of antifungal agents as a target.
Description
Technical field
The invention belongs to technical field of microbial genetic engineering, be specifically related in plant protection art, to control fungal infection structure and formed and the gene of virulence and the application of coded protein thereof.
Background technology
Ash arrhizus bacteria (Botrytiscinerea), usually also known as doing Botrytis cinerea, belongs to Ascomycota (Ascomycota) fungi, is the pathogenic bacteria of gray mold, can infects 200 various plants, comprise nearly all vegetables and fruit tree crop.Host all can fall ill from seedling stage to the phase of bearing fruit, and each position of plant all can be infected by ash arrhizus bacteria, and the classical symptom of leaf portion morbidity shows as " V " shape scab, and flower portion main manifestations is rotten and tune withers, and fruit main manifestations is for rotting and coming off.The generation of disease and spread and there is close relationship with the humidity of environment, temperature, at 20 DEG C-23 DEG C, occurs serious when relative humidity more than 90%.Therefore, gray mold belongs to low temperature and high relative humidity type disease, very easily occurs in rainy season or Protected production, and the financial loss caused because of this disease is every year up to 100-1000 hundred million dollars.Because host range is extensive, in production, harm is serious, and add associated molecule investigative technique maturation, botrytis cinerea has become one of most important model plant pathogenic fungi, is subject to extensive research.
Ash arrhizus bacteria is typical necrotrophic pathogenic fungi, multiple virulence factor can be generated participate in causing a disease, mainly comprise cell wall degradation enzyme, at, toxin, plant hormone, the enzyme of opposing defense enzymes, tiny RNA and small-molecule substance etc., these factors mutually cooperate and enable ash arrhizus bacteria kill host cell and decompose dead host tissue as nutrition.Under natural condition, botrytis cinerea mainly with conidium as infecting the First aggression of host and infecting source again.Ash arrhizus bacteria is often attached to mycelium, conidium or sclerotium to survive the winter on plant invalid body or in soil and to get over the summer, becomes the primary infection inoculum of next Growing season.When condition is suitable for, sclerotial germination aerial mycelium and conidiophore, and produce a large amount of conidiums.Ripe conidium can be propagated by wind, rainwater, the general water of filling and farming operation etc.Under low temperature and high relative humidity condition, conidia germination forms germ tube, and germ tube end expands slightly to develop into appressorium or formed further and infects the Infection structures such as pad, mainly invades from the floral organ of decaying, wound and necrotic tissue.
Appressorium, infect the growth of the Infection structures such as pad, it is vital for infecting host for ash arrhizus bacteria, is affected if Infection structure is grown, and ash arrhizus bacteria will be difficult to intrusion host, and its hazard rating can be seriously undermined thereupon.The growth of ash arrhizus bacteria Infection structure is subject to the external source condition such as nutrition, interface and grows the combined regulating of signal, associated molecule mechanism it be unclear that, this field is furtherd investigate to the molecular mechanism not only contributing to disclosing the necrotrophic pathogenic fungi such as ash arrhizus bacteria and cause a disease, and for the medicament that research and development control comprises the plant pathogenic fungi of ash arrhizus bacteria, there is significant application value.
From conidia germination to the Infection structure of growing complete function, it is a meticulous regulation process, identify the important component of this regulation process, and verify the pathogenic function of genes involved, likely therefrom finding can as the protein of mycocide action target, and theory and technology basis established by the efficient medicament preventing and treating gray mold and other similar disease for exploitation.
Sep4 is a kind of gtp binding protein matter, extensively be present in the eukaryote (except plant) comprising fungi, this protein, by aggregating into hetero-oligomer protein complex or forming fibril further, participates in the vital process of the multiple key of regulating cell.By the analysis to ash arrhizus bacteria Sep4 encoding gene, evaluating the effect of this gene at ash arrhizus bacteria pathogenic course, being conducive to identifying potential control target, for screening novel fungicidal medicament.
Summary of the invention
Object of the present invention aims to provide a kind of protein controlling the growth of ash arrhizus bacteria Infection structure and pathogenic gene and coding thereof.
Control Infection structure provided by the present invention is grown and pathogenic gene derives from ash arrhizus bacteria, name is called BcSep4, it has SEQ ID No: the DNA sequence dna shown in 1, and this DNA sequence dna is made up of 2793 Nucleotide, comprises the promotor of gene, open reading frame and terminator.Be promoter sequence between 5 ' end the 1st to 931 Nucleotide; The open reading frame of BcSep4 is made up of 1147 Nucleotide, wherein comprise 3 exons, lay respectively at SEQIDNo:1 5 ' end the 932nd to 1067 Nucleotide between, between the 1134th to 1696 Nucleotide and between the 1746th to 2078 Nucleotide, the coding region length of composition adds up to 1032 Nucleotide; Be terminator sequence between 5 ' end the 2079th to 2793 Nucleotide.
The invention provides from ash arrhizus bacteria control Infection structure grow and pathogenic gene BcSep4 coded by protein, it has SEQ ID No: the aminoacid sequence shown in 2, and this sequence is made up of 343 amino acid.
Control Infection structure from ash arrhizus bacteria is grown and pathogenic gene BcSep4 can be applicable to Genes For Plant Tolerance gray mold genetically engineered field.
From ash arrhizus bacteria control Infection structure grow and pathogenic gene BcSep4 coded by protein, ash arrhizus bacteria control Infection structure growth and pathogenic PROTEIN B cSep4 are lacked, suddenly change or modified and make its Infection structure, virulence generation defect, can be used as target and apply in design and screening antifungal medicine.
Present invention demonstrates that disappearance or the sudden change of BcSep4 gene cause ash arrhizus bacteria Infection structure rate of formation and virulence significantly to reduce, illustrate that BcSep4 gene is that ash arrhizus bacteria causes the necessary gene of farm crop gray mold.Therefore, screening can stop the compound of this genetic expression and its protein expression, modification and location, effectively can control the generation of ash arrhizus bacteria conidia germination formation Infection structure and gray mold, thus contribute to development of new sterilant.Namely an important use of BcSep4 gene provided by the present invention is: the expression of the expression of this gene and its protein, modification and locate screening and the design that can be used for antifungal medicine (particularly botrytis resistant bacterium medicament) as important candidate targets site.
Accompanying drawing explanation
Fig. 1 is that each species Sep4 protein system grows tree analysis schematic diagram
Wherein: what underscore marked is ash arrhizus bacteria.
Fig. 2 be ash arrhizus bacteria BcSep4 gene knock out strategy (carrying out gene replacement by homologous recombination) schematic diagram
Wherein: WT is wild type strain B05.10, pSep4-ko is knockout carrier, BcSep4-KO is BcSep4 deletion mutant body.
Fig. 3 is that the PCR of BcSep4 deletion mutant body verifies electrophorogram
Wherein: a, b, c, d are the primer, and Fig. 2 is seen in corresponding position; M1, M2 are the BcSep4 deletion mutant that two strains independently obtain.
Fig. 4 is that the PCR of genetic complementation bacterial strain verifies electrophorogram
Wherein: a, b, c, d are the primer, and Fig. 2 is seen in corresponding position; M1/Sep4 is the complemented strain proceeding to complete BcSep4 gene on mutant M1 basis.
Fig. 5 is that the deletion mutant of BcSep4 gene and the cultural characteristic of wild type strain B05.10 and complemented strain contrast photo
Wherein: used medium is PDA, WT is wild-type, M1, M2 are the BcSep4 deletion mutant that two strains independently obtain, and M1/Sep4 is the complemented strain proceeding to complete BcSep4 gene on mutant M1 basis.
Fig. 6 is that the deletion mutant of BcSep4 gene compares with the sporulation quantity of wild type strain and complemented strain.
Fig. 7 is that the deletion mutant of BcSep4 gene compares photo with the virulence of wild type strain and complemented strain
Wherein: selected host is tomato, the method for Isolated leaf inoculation bacterium cake is adopted.Inoculate and evaluate after 3 days.
Fig. 8 is that the deletion mutant of BcSep4 gene compares photo with the virulence of wild type strain and complemented strain
Wherein: selected host is rape, the method for Isolated leaf inoculation bacterium cake is adopted.Inoculate and evaluate after 3 days.
Fig. 9 is that the deletion mutant of BcSep4 gene compares Photomicrograph with the note fields ability of wild type strain and complemented strain
Wherein: by PDB spore suspension (5 × 10
4ml
-1) drop on slide glass, after 6h is cultivated in 20 DEG C of moisturizings, with Lu Kafu white dyes dyeing, carry out the microscopic examination of dark-field burst of ultraviolel.Arrow is depicted as appressorium, and scale is 20 μm.
Figure 10 is the contrast photo that the deletion mutant of BcSep4 gene and wild type strain and complemented strain infect the microscopic examination of onion epidermis
Wherein: with 10
4ml
-1spore suspension inoculation onion epidermis, hatch 24h for 20 DEG C, with lactophenol indigo plant dyeing after carry out microscopic examination, contaminate not Shang color for infecting mycelia, arrow be depicted as ash arrhizus bacteria intrusion site.Scale is 10 μm.
Figure 11 be deletion mutant and the wild type strain of BcSep4 gene and complemented strain infect the contrast Photomicrograph padding Forming ability and compare
Wherein: by PDB spore suspension (5 × 10
4ml
-1) drop on slide glass, after 36h is cultivated in 20 DEG C of moisturizings, microscopic examination, scale is 100 μm.
Embodiment
In order to describe the present invention better, being further described below by specific embodiment, the method in following embodiment, if no special instructions, being ordinary method.
The correlation analysis of embodiment 1BcSep4 gene
BcSep4 gene is positioned on ash arrhizus bacteria No. IV karyomit(e), and its open reading frame is made up of 1147 Nucleotide, comprises 3 exons, and coding region cDNA total length is 1032 Nucleotide, and the protein of coding is made up of 343 amino acid.Get BcSep4 protein sequence to compare (http://blast.ncbi.nlm.nih.gov/Blast.cgi), find that Sep4 is extensively present in fungi, animal, also have in green alga, but in higher plant, this degenerate gene has fallen.Phylogenetic tree analyze (http://phylogeny.lirmm.fr/phylo_cgi/simple_phylogeny.cgi) show, the homology between filamentous fungus Sep4 protein is higher, wherein BcSep4 and sclerotinite Sep4 sibship nearest; BcSep4 and yeast are slightly far away, are next animals, with green alga sibship farthest (see Fig. 1).
Knocking out and genetic complementation of embodiment 2BcSep4 gene
1) structure of knockout carrier
Adopt primer Sep4-UP-F (5'-CTCGAGCTTCGTGGGCATTGGTCTTG-3') and Sep4-UP-R (5'-GAATTCACGAGGGTATTATCTTTGGATTG-3'), with the genomic dna of ash arrhizus bacteria bacterial strain B05.10 for template amplification BcSep4 upstream region of gene 776bp fragment; Sep4-DN-F (5'-GGATCCGATGGCGAGATGGTATTTGC-3') and Sep4-DN-R (5'-CTGCAGTGCTGAGGCTCCGAGAAGA-3') is adopted to increase ash arrhizus bacteria BcSep4 downstream of gene 636bp fragment, reaction system is: 10mmol/LdNTPMixture, 0.5 μ L; 10 × PCRbuffer, 2.5 μ L; The each 1 μ L of upstream and downstream primer (10 μm of ol/mL); Template DNA, 1 μ L; Ex-Taq, 0.2 μ L (5U); ddH
2o, 18.8 μ L; Amplification program is: 94 DEG C of denaturations 3 minutes, then (1) 94 DEG C, sex change 50 seconds; (2) 59 DEG C, anneal 50 seconds; (3) 72 DEG C, extend 60 seconds; (4) circulate 30 times; (5) 72 DEG C extend 10 minutes.Above-mentioned DNA cloning product is successively cloned into XhoI, EcoRI site and BamHI, PstI site of pXEH carrier, is built into knockout carrier pSep4-ko (as shown in Figure 2), and carries out sequence verification.
2) conversion of ash arrhizus bacteria
A. the cultivation of agrobacterium tumefaciens
Picking contains the mono-bacterium colony of Agrobacterium tumefaciens strain Agl-1 of binary vector pSep4-ko, be seeded to the MM liquid nutrient medium (dipotassium hydrogen phosphate 0.205% containing 50 μ g/ml kantlex, 10 μ g/ml Rifampins, potassium primary phosphate 0.145%, sodium-chlor 0.015%, magnesium sulfate heptahydrate 0.05%, calcium chloride hexahydrate 0.01%, iron vitriol 0.00025%, ammonium sulfate 0.05%, glucose 0.2%) in, 250rpm, 28 DEG C of shaking culture 48h; 4000rpm, centrifugal 5 minutes, abandons supernatant, IM liquid nutrient medium (dipotassium hydrogen phosphate 0.205%, potassium primary phosphate 0.145%, sodium-chlor 0.015%, magnesium sulfate heptahydrate 0.05%, calcium chloride hexahydrate 0.01%, iron vitriol 0.00025%, ammonium sulfate 0.05%, glucose 0.2%, 200 μMs of AS, MES0.854%, glycerine 0.5%) resuspended, centrifugal 5 minutes of 4000rpm, abandons supernatant; IM substratum is resuspended, 28 DEG C, and 250rpm shaking culture 6h carries out pre-induced.
B. the product spore of ash arrhizus bacteria is cultivated
Select B05.10 bacterial strain, the spore that takes a morsel coats (the well-done filtration of potato 20% of PDA substratum, glucose 2%, agar 1.5%), put 28 DEG C of cultivation 8h and make spore fast-germination, be then transferred to 20 DEG C and cultivate 3-5 days, after treating that phage surface is covered by grey spore, by IM liquid nutrient medium scraping, collect spore, microscopic examination, utilize Hematocyte Counter to regulate spore concentration to be 1 × 10
6/ mL.
C. agrobacterium tumefaciens and ash arrhizus bacteria conidium Dual culture and transformant screening
Agrobacterium bacterium liquid and the mixing of ash arrhizus bacteria spore liquid equal-volume of 6h will be induced in advance in IM liquid nutrient medium, and add AS, make final concentration reach 500 μMs, mixing, then by 250 ~ 350 μ L/ wares, uniform application on the IM substratum being covered with glassine paper, 22 DEG C of dark culturing 48h; After Dual culture, glassine paper is transferred on the PDA substratum containing 100 μ g/mL Totomycin, continues under the same terms to cultivate.After 4 ~ 7 days, the bacterium colony of picking expansion is to containing in same antibiotic screening culture medium.
3) checking of deletion mutant
Two pairs of primers are selected to be screened transformant by pcr amplification.What amplification met following result is defined as BcSep4 deletion mutant body: the primer a (5'-CTGCTCTAACTCTGCCTCTT-3') outside the homology arm of upstream on genome and the primer b (5'-ACAGACGTCGCGGTGAGTTCA-3') of hygromycin gene match to increase to and expect the recombinant fragment of size (1.3kb); And coding region primer c (5'-TCGAGGAGAACGGCGTAAAG-3') and d (5'-CAACAACAGCGAATGGGATG-3') are without amplified band (wild type strain can increase 0.5kb fragment).As a result, from transformant, 2 strain BcSep4 deletion mutant bodies are screened: M1, M2 (as shown in Figure 3), for follow-up function analysis.
4) genetic complementation of BcSep4 deletion mutant body
Adopt primer C-F (5'-CTGCTCTAACTCTGCCTCTT-3') and C-R (5'-TGCTGAGGCTCCGAGAAGA-3'), amplification ash arrhizus bacteria BcSep4 full length gene 2793bp (comprising promotor, open reading frame and terminator), first be cloned on pMD18-t carrier, and then subclone to pSULFgfp carrier (containing chlorimuronethyl resistant gene) SalI and XbaI site between, be built into genetic complementation carrier pSep4-ko-c.Carrier, through sequence verification, confirms do not have amino acid mutation.Adopt foregoing Agrobacterium-medialed transformation method, use 100 μ g/mL chlorimuronethyls to screen, this complementary fragment is proceeded in BcSep4 deletion mutant body M1 genome, obtain genetic complementation bacterial strain M1/Sep4.Once primer a and the b, c and the d that used when selecting mutant to verify carry out pcr amplification, result meets expection (as shown in Figure 4): identical with mutant M1, in complemented strain M1/Sep4, original BcSep4 gene is replaced by hygromycin gene HPH (primer a and b amplification are for positive), but additionally has a follow-up BcSep4 gene (coding region primer c and d amplification are similarly positive) proceeded to.
The effect of embodiment 3BcSep4 gene in ash arrhizus bacteria growth and development process
Adopt plating method, evaluate the variation situation of the Relevant phenotype such as to grow of BcSep4 mutant.Test strains is seeded in the center of PDA substratum with bacterium block mode, 20 DEG C of dark culturing.The colonial morphology of mutant and the speed of growth normally, compare with wild-type and do not have marked difference; But the ability that mutant forms sclerotium significantly strengthens (see Fig. 5).In addition, BcSep4 gene mutation body bacterial strain sporulation quantity significantly reduces (see Fig. 6).In genetic complementation bacterial strain, above-mentioned abnormal phenotype is all replied normally, proves that BcSep4 gene has the ability promoting illumination, suppress Sclerotia forming to a certain extent.
The effect of embodiment 4BcSep4 gene in ash arrhizus bacteria is pathogenic
Adopt Isolated leaf inoculation method, evaluate the virulence changing conditions of BcSep4 mutant.Gather the blade with certain leaf age from the host plant of hot-house culture, in horizontal positioned container, use punch tool to beat and get test strains bacterium cake, face down left-hand thread on blade, 20 DEG C of moisturizing dark culturing, the virulence of 3 days postevaluation test strains.Experimental result shows, BcSep4 mutant has completely lost pathogenecity, no matter is inoculating tomato (see Fig. 7), or inoculation rape (see Fig. 8), and BcSep4 mutant all can not cause disease.The virulence of genetic complementation bacterial strain returns to wild-type levels, and this shows, BcSep4 is a crucial Disease-causing gene, and it is necessary to be that ash arrhizus bacteria infects host.
Embodiment 5BcSep4 gene is at ash arrhizus bacteria Infection structure---the effect in appressorium growth course
Adopt slide culture, observe the ability that BcSep4 mutant develops into appressorium.By PDB (the well-done filtration of potato 20%, glucose 2%) spore suspension (5 × 10
4mL
-1) drop on slide glass, after 6h is cultivated in 20 DEG C of moisturizings, with Lu Kafu white dyes dyeing (cell walls specificity can be caught color by this dyestuff, is convenient to observe).Microscopic examination finds, the germ tube that the spore germination of wild type strain goes out, and bend, end expands usually more, forms Infection structure---appressorium.The spore of BcSep4 mutant then loses the ability developing into appressorium substantially, rapid linear extension after spore germination, and end is difficult to expand; BcSep4 gene is imported mutant again, above-mentioned abnormal phenotype can be returned to wild-type levels (see Fig. 9).
Adopt onion epidermis inoculation method, observe the infection ability after the spore germination of BcSep4 mutant.With 10
4mL
-1spore suspension inoculation onion epidermis, hatch 24h for 20 DEG C, with lactophenol indigo plant dyeing after carry out microscopic examination, do not contaminate color for infecting mycelia.Microscopic examination finds, the appressorium that wild type strain can rely on germ tube end to be formed invades onion epidermis, and the ability that BcSep4 mutant loss invades, after spore germination, only present simple interest, cannot appressorium be formed and invade epidermis; Genetic complementation bacterial strain has normal infection ability (see Figure 10).
Above result of study shows, BcSep4 gene take part in ash arrhizus bacteria Infection structure---the formation of appressorium, and play a key effect.
Embodiment 6BcSep4 gene is at ash arrhizus bacteria Infection structure---infect the effect in pad growth course
Adopt slide culture, observe BcSep4 mutant and develop into the ability infecting pad.By PDB spore suspension (5 × 10
4mL
- 1) drop on slide glass, 36h is cultivated in 20 DEG C of moisturizings.Microscopic examination finds, the mycelium of wild type strain can form a large amount of multi-branched mat-like structure---infect pad, mutant has then completely lost the ability developing into this complicated Infection structure; BcSep4 gene is imported mutant again, can be recovered it and infect pad developmental potency (see Figure 11).This result of study shows, BcSep4 gene not only take part in the formation of ash arrhizus bacteria appressorium, and infecting the effect that in pad growth course, same performance is very important, this gene is that ash arrhizus bacteria Infection structure is grown necessary, if this gene or its coded protein loss of activity, ash arrhizus bacteria infects losing the ability that host causes disease.
Claims (4)
1. the control Infection structure from ash arrhizus bacteria (Botrytiscinerea) is grown and a pathogenic gene BcSep4, it is characterized in that having SEQ ID No: the DNA sequence dna shown in 1.
2. the growth of the control Infection structure from ash arrhizus bacteria according to claim 1 and the protein coded by pathogenic gene BcSep4, is characterized in that having SEQ ID No: the aminoacid sequence shown in 2.
3. the control Infection structure from ash arrhizus bacteria according to claim 1 is grown and the application of pathogenic gene BcSep4 in Genes For Plant Tolerance gray mold genetically engineered field.
4. the growth of the control Infection structure from ash arrhizus bacteria according to claim 2 and the protein coded by pathogenic gene BcSep4, ash arrhizus bacteria control Infection structure growth and pathogenic PROTEIN B cSep4 are lacked, suddenly change or modified and make its Infection structure, virulence generation defect, as the application of target in design and screening antifungal medicine.
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| CN105483146A (en) * | 2016-01-22 | 2016-04-13 | 吉林大学 | Botrytis cinerea gene BcAls1 relative to pathogenicity and application of botrytis cinerea gene BcAls1 |
| CN105483138A (en) * | 2016-01-22 | 2016-04-13 | 吉林大学 | Botrytis cinerea gene BcAtm1 relative to pathogenicity and application of botrytis cinerea gene BcAtm1 |
| CN105483106A (en) * | 2016-01-22 | 2016-04-13 | 吉林大学 | Botrytis cinerea gene BcFch1 relative to pathogenicity and application of botrytis cinerea gene BcFch1 |
| CN105567708A (en) * | 2016-01-22 | 2016-05-11 | 吉林大学 | Botrytis cinerea gene BcNop53 related to pathogenicity and application thereof |
| CN106399328A (en) * | 2016-11-04 | 2017-02-15 | 吉林大学 | Pathogenicity correlated Botrytis cinerea gene BcSEP6 and application thereof |
| CN106497943A (en) * | 2016-11-04 | 2017-03-15 | 吉林大学 | A kind of ash arrhizus bacteria gene BcSEP5 related to pathogenicity and its application |
| CN106754977A (en) * | 2016-12-12 | 2017-05-31 | 中国农业科学院植物保护研究所 | Rice blast fungus agricultural chemicals target gene MoR1 and its encoding proteins and application |
| CN111118039A (en) * | 2020-02-07 | 2020-05-08 | 浙江大学 | Pathogenicity-related botrytis cinerea genes Bcmet3 and Bcmet16 and application |
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| CN105483138A (en) * | 2016-01-22 | 2016-04-13 | 吉林大学 | Botrytis cinerea gene BcAtm1 relative to pathogenicity and application of botrytis cinerea gene BcAtm1 |
| CN105483106A (en) * | 2016-01-22 | 2016-04-13 | 吉林大学 | Botrytis cinerea gene BcFch1 relative to pathogenicity and application of botrytis cinerea gene BcFch1 |
| CN105567708A (en) * | 2016-01-22 | 2016-05-11 | 吉林大学 | Botrytis cinerea gene BcNop53 related to pathogenicity and application thereof |
| CN105483146A (en) * | 2016-01-22 | 2016-04-13 | 吉林大学 | Botrytis cinerea gene BcAls1 relative to pathogenicity and application of botrytis cinerea gene BcAls1 |
| CN105567708B (en) * | 2016-01-22 | 2019-05-03 | 吉林大学 | One kind ash arrhizus bacteria gene BcNop53 relevant to pathogenicity and its application |
| CN105483138B (en) * | 2016-01-22 | 2019-03-26 | 吉林大学 | One kind ash arrhizus bacteria gene BcAtm1 relevant to pathogenicity and its application |
| CN105483106B (en) * | 2016-01-22 | 2019-03-26 | 吉林大学 | One kind ash arrhizus bacteria gene BcFch1 relevant to pathogenicity and its application |
| CN106399328A (en) * | 2016-11-04 | 2017-02-15 | 吉林大学 | Pathogenicity correlated Botrytis cinerea gene BcSEP6 and application thereof |
| CN106497943A (en) * | 2016-11-04 | 2017-03-15 | 吉林大学 | A kind of ash arrhizus bacteria gene BcSEP5 related to pathogenicity and its application |
| CN106497943B (en) * | 2016-11-04 | 2019-05-07 | 吉林大学 | One kind ash arrhizus bacteria gene BcSEP5 relevant to pathogenicity and its application |
| CN106399328B (en) * | 2016-11-04 | 2019-06-28 | 吉林大学 | One kind ash arrhizus bacteria gene BcSEP6 relevant to pathogenicity and its application |
| CN106754977A (en) * | 2016-12-12 | 2017-05-31 | 中国农业科学院植物保护研究所 | Rice blast fungus agricultural chemicals target gene MoR1 and its encoding proteins and application |
| CN106754977B (en) * | 2016-12-12 | 2020-02-14 | 中国农业科学院植物保护研究所 | Magnaporthe grisea pesticide target gene MoR1 and encoding protein and application thereof |
| CN111118039A (en) * | 2020-02-07 | 2020-05-08 | 浙江大学 | Pathogenicity-related botrytis cinerea genes Bcmet3 and Bcmet16 and application |
| CN111118039B (en) * | 2020-02-07 | 2021-07-06 | 浙江大学 | Pathogenicity-related botrytis cinerea genes Bcmet3 and Bcmet16 and application |
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Application publication date: 20151216 |