CN103224555B - Plant development associated protein GhSOC1 and encoding gene thereof and application - Google Patents
Plant development associated protein GhSOC1 and encoding gene thereof and application Download PDFInfo
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Abstract
The invention discloses a kind of plant development associated protein GhSOC1 and encoding gene thereof and application.Protein provided by the invention, called after GhSOC1 albumen is following (a) or (b): the protein that (a) is made up of the aminoacid sequence shown in sequence in sequence table 1; (b) by the aminoacid sequence shown in sequence in sequence table 1 through the replacement of one or several amino-acid residue and/or disappearance and/or interpolation and with the protein derived by sequence 1 of flowering of plant time correlation.The present invention has cloned cotton GhSOC1 gene from upland cotton, and successfully builds plant over-express vector, adopts agriculture bacillus mediated inflorescence dip method to transform the plant that sets out, and transgenic plant ratio sets out flowering of plant in advance and can genetic stability.The present invention has substantial worth for the breeding of plant.
Description
Technical field
The present invention relates to a kind of plant development associated protein GhSOC1 and encoding gene thereof and application.
Background technology
Cotton is one of important cash crop of China, in national product, occupy critical role, grain and cotton strive in the development to a certain degree limiting cotton.The selection and popularization of short season cotton kind, can alleviate the contradiction that ground striven by grain and cotton, is the effective way realizing reaping a bumper harvest of grain and cotton.
Prematureness, as one of the important character of cotton, becomes the breeding objective that upland cotton is important.Research shows, blooms sooner or later closely related with the prematureness of cotton.Therefore, clone blooms genes involved, carries out expression analysis and transgenosis functional verification to it, can provide the genetic resources of high-quality for short season cotton breeding.
Summary of the invention
The object of this invention is to provide a kind of plant development associated protein GhSOC1 and encoding gene thereof and application.
Protein provided by the invention, available from cotton variety " CCRI 36 ", called after GhSOC1 albumen is following (a) or (b):
A protein that () is made up of the aminoacid sequence shown in sequence in sequence table 1;
(b) by the aminoacid sequence shown in sequence in sequence table 1 through the replacement of one or several amino-acid residue and/or disappearance and/or interpolation and with the protein derived by sequence 1 of flowering of plant time correlation.
In order to make the protein in (a) be convenient to purifying, the N-terminal of the protein that the aminoacid sequence shown in sequence 1 forms or C-terminal label as shown in table 1 can be connected in by sequence table.
The sequence of table 1 label
| Label | Residue | Sequence |
| Poly-Arg | 5-6(is generally 5) | RRRRR |
| Poly-His | 2-10(is generally 6) | HHHHHH |
| FLAG | 8 | DYKDDDDK |
| Strep-tag II | 8 | WSHPQFEK |
| c-myc | 10 | EQKLISEEDL |
Protein in above-mentioned (b) can synthetic, also can first synthesize its encoding gene, then carries out biological expression and obtain.The encoding gene of the protein in above-mentioned (b) is by the codon by lacking one or several amino-acid residue in the DNA sequence dna shown in sequence in sequence table 2, and/or carry out the missense mutation of one or several base pair, and/or the encoding sequence connecting the label shown in table 1 is held to obtain at its 5 ' end and/or 3 '.
The gene (called after GhSOC1 gene) of described GhSOC1 albumen of encoding also belongs to protection scope of the present invention.
Described GhSOC1 gene is the DNA molecular described in following (1) or (2) or (3) or (4) or (5):
(1) in sequence table sequence 2 from the DNA molecular shown in 5 ' end the 102 to 767 Nucleotide;
(2) in sequence table sequence 2 from the DNA molecular shown in 5 ' end the 102 to 785 Nucleotide;
(3) DNA molecular shown in sequence 2 in sequence table;
(4) DNA sequence dna limited with (1) or (2) or (3) is under strict conditions hybridized and is encoded and the DNA molecular of the albumen of flowering of plant time correlation;
(5) DNA sequence dna limited with (1) or (2) or (3) at least has more than 90% homology and encode and the DNA molecular of albumen of flowering of plant time correlation.
Above-mentioned stringent condition can be in the solution of 6 × SSC, 0.5%SDS, hybridizes under 65oC, then uses 2 × SSC, 0.1%SDS and 1 × SSC, 0.1%SDS respectively to wash film once.
Recombinant vectors containing described GhSOC1 gene, expression cassette, transgenic cell line or recombinant bacterium all belong to protection scope of the present invention.
Available existing plant expression vector construction contains the recombinant expression vector of described gene.Described plant expression vector comprises double base agrobacterium vector and can be used for the carrier etc. of plant micropellet bombardment.Described plant expression vector also can comprise 3 ' end untranslated region of foreign gene, namely comprises the DNA fragmentation of polyadenylation signals and any other participation mRNA processing or genetic expression.The bootable polyadenylic acid of described polyadenylation signals joins 3 ' end of mRNA precursor.When using described gene constructed recombinant plant expression vector, can add any one enhancement type promotor or constitutive promoter before its transcription initiation Nucleotide, they can be used alone or are combined with other plant promoter; In addition, when using gene constructed plant expression vector of the present invention, also can use enhanser, comprise translational enhancer or transcriptional enhancer, but must be identical with the reading frame of encoding sequence, to ensure the correct translation of whole sequence.The source of described translation control signal and initiator codon is widely, can be natural, also can be synthesis.Translation initiation region can from transcription initiation region or structure gene.For the ease of identifying transgenic plant cells or plant and screening, can process plant expression vector used, the coding can expressed in plant as added can produce enzyme or the gene of luminophor, the antibiotic marker thing with resistance or the chemical resistance reagent marker gene etc. of colour-change.From the security consideration of transgenic plant, any selected marker can not be added.
Described recombinant vectors specifically can be to insert in the multiple clone site (as between Sal I and Sac I restriction enzyme site) of carrier pRI101-AN as described in the recombinant plasmid that obtains of GhSOC1 gene.
The present invention also protects a kind of method of cultivating transgenic plant, is described GhSOC1 channel genes object plant, obtains the transgenic plant of flowering time early than described object plant.The plant tissue of conversion by using Ti-plasmids, Ri plasmid, plant viral vector, directly delivered DNA, microinjection, conductance, conventional biology methods transformed plant cells or the tissue such as agriculture bacillus mediated, and is cultivated into plant by the expression vector carrying described GhSOC1 gene.Described GhSOC1 gene specifically imports in described object plant by described recombinant plasmid.Described object plant is monocotyledons or dicotyledons.Described dicotyledons specifically can be Arabidopis thaliana, as Columbia ecotype Arabidopis thaliana.
The present invention also protects described GhSOC1 albumen or the application of described GhSOC1 gene in regulating plant flowering time.Described plant is monocotyledons or dicotyledons.Described dicotyledons specifically can be Arabidopis thaliana, as Columbia ecotype Arabidopis thaliana.
The present invention has cloned cotton GhSOC1 gene from upland cotton, and successfully builds plant over-express vector, adopts agriculture bacillus mediated inflorescence dip method to transform the plant that sets out, and transgenic plant ratio sets out flowering of plant in advance and can genetic stability.The present invention has substantial worth for the breeding of plant.
Accompanying drawing explanation
Fig. 1 is the spatial and temporal expression pattern analysis result of GhSOC1 gene.
Fig. 2 is the structural representation of recombinant plasmid pRI101-GhSOC1.
Fig. 3 is that the PCR of part sample identifies electrophorogram.
Fig. 4 is the southern results of hybridization of part sample.
Fig. 5 is flowering time, the lotus throne number of sheets and stem leaf number under long-day conditions.
Fig. 6 is the phenotype photo of transfer-gen plant and Columbia ecotype Arabidopis thaliana.
Embodiment
Following embodiment is convenient to understand the present invention better, but does not limit the present invention.Experimental technique in following embodiment, if no special instructions, is ordinary method.Test materials used in following embodiment, if no special instructions, is and purchases available from routine biochemistry reagent shop.Quantitative test in following examples, all arranges and repeats experiment for three times, results averaged.Cotton variety " CCRI 36 ": the Chinese Academy of Agriculture Science and Technologys Cotton Research Institute.Agrobacterium LBA4404: TAKARA, Code:9115.Carrier pRI101-AN:TAKARA, Code:3262.Columbia ecotype Arabidopis thaliana, English name is Arabidopsisthaliana (ecotypecolumbia): ABRCseedstock, OhioStateUniversityfacility.
The discovery of embodiment 1, GhSOC1 albumen and encoding gene thereof
One, RNA extracts
1,1ml extracting solution adds the mercaptoethanol of 2% final volume (20 μ l), 65 DEG C of preheatings.
2, being organized in liquid nitrogen of 100mg cotton variety " CCRI 36 " is claimed to grind, require to be organized in when leaving live body to put into Liquid nitrogen storage immediately, and mortar precooling, when liquid nitrogen is evaporated completely soon after grinding, to powder be organized to scrape in the extracting solution of 65 DEG C of preheatings fast, concuss at once, 65 DEG C of temperature bath 3-5min, shake 4-5 time therebetween.
3, isopyknic chloroform is added: primary isoamyl alcohol, concuss 10min, 4 DEG C, the centrifugal 5min of 13000rpm.
4, get supernatant liquor, add equal-volume chloroform: primary isoamyl alcohol, concuss 10min, 4 DEG C, the centrifugal 5min of 13000rpm.
5, get supernatant liquor, add the 10mol/LLicl of 1/4 volume, mix gently, 4 DEG C spend the night (3-6h), precipitated rna.
6, by solution 4 DEG C, 13000rpm, centrifugal 12min, outwells supernatant liquor, precipitation is dissolved in 400 μ lEPC water.
7, add isopyknic phenol, at once concuss, 4 DEG C, the centrifugal 5min of 13000rpm, suct clear liquid.
8, the chloroform of body is added etc.: primary isoamyl alcohol (24:1), at once concuss, 4 DEG C, the centrifugal 5min of 10000rpm.
9, get supernatant liquor, add the dehydrated alcohol of the sodium-acetate of the 3mol/L of 0.1 times of volume, 2.5 times of volumes ,-70 DEG C precipitate more than 30 minutes.
10,4 DEG C, the centrifugal 20min of 13000rpm, abandon supernatant liquor.
11,70% alcohol washes 1-2 time, air-dry, and be dissolved in 40 μ lDEPC water ,-80 DEG C of preservations, detect for subsequent use.
Two, cDNA preparation
Use the SMARTScribe of clontech company
tMreversetranscriptase, uses front centrifugal.
1, RNA-free pipe is put on ice, carry out mark, add 2 μ gTotalRNA, 1ul20 μM of oligo (dT)
20, add RNA-free water and be made into mix to 5 μ l, 72 DEG C, 3min is placed on ice.
2, cDNASynthesisMix:2 μ l5XFirst-StrandBuffer is prepared, 1 μ ldNTP, 1 μ l20mMDTT, 1 μ l100U/ μ lSMAATScribeRT; 10 μ lcDNAmix are joined in RNA/Primermixture, softly mixes, centrifugal;
3, hatch 60-90min for 42 DEG C, 70 DEG C of heating, 15 DEG C of termination reactions, 1 μ l detects, and-20 DEG C save backup.
Three, the cloning process of gene
The clone of GhSOC1 gene mainly utilizes information biology, RACE-PCR technology completes.
1, according to build cDNA library in conjunction with electronic cloning technology, sequence assembly is carried out to EST.
2,3 ＇ RACE are carried out to obtain the complete ORF of this gene.
3, three special primers are designed according to this gene one section of est sequence obtained: GSP1, GSP2 and GSP3; With the compound sample of each tissue of cotton for template, carry out the synthesis of a chain according to the SMARTRACEcDNAAmplificationKit specification sheets of Clontech company.
4, TOUCHDOWN-PCR is carried out according to the requirement of test kit, then NEST-PCR is carried out after its product being carried out 50 times of dilutions, the specific target stripe obtained is reclaimed and connects pGEMT-easy carrier, then send Invitrogen company to check order, obtain the complete from cDNA sequence of this gene finally by splicing.
By the protein called after GhSOC1 albumen shown in the sequence 1 of sequence table, be made up of 221 amino-acid residues.Be GhSOC1 gene by the unnamed gene of coding GhSOC1 albumen, its cDNA is as shown in the sequence 2 of sequence table, and the sequence 2 that open reading frame is sequence table is from shown in 5 ' end the 102 to 767 Nucleotide.
The spatial and temporal expression pattern analysis of embodiment 2, GhSOC1 gene
One, the tissue expression pattern of GhSOC1 gene
1, extract the total serum IgE of the different tissues of cotton variety " CCRI 36 ", and reverse transcription is cDNA respectively.
5 times of volume dilution liquid of the cDNA 2, obtained with step 1, for template, carry out quantitative fluorescent PCR.
Primer for the expression amount detecting GhSOC1 gene is as follows:
F:5’-AGCATGCAGTGGCAGCATCTGA-3’；
R:5’-TGGCTCTGACGCGGGTTACG-3’。
Primer pair for the expression amount detecting reference gene (actin) is as follows:
F:5’-ATCCTCCGTCTTGACCTTG-3’；
R:5’-TGTCCGTCAGGCAACTCAT-3’。
Quantitative fluorescent PCR reaction system: 10 μ lSYBRPremixExTag, 0.8 μ lF, 0.8 μ lR, 2 μ lcDNA templates, 6 μ lddH
2o, 0.4 μ lROXReferenceDye, cumulative volume 20ul.
Quantitative fluorescent PCR response procedures: 95 DEG C of denaturation 1min; 95 DEG C of sex change 5s, 60 DEG C of annealing 30s, 72 DEG C of extension 30s, 40 circulations; 72 DEG C extend 10min; At 72 DEG C, fluorescent signal is collected at 30s place.
Relative quantification Δ Δ Ct method is adopted to analyze.
The expression pattern (relative expression quantity) in each tissue of GhSOC1 gene is shown in Figure 1A.Result shows: GhSOC1 gene is predominant expression in terminal bud and blade, root, stem, spends middle expression amount low.
Two, GhSOC1 gene is at the expression pattern of terminal bud different development stage
Method is with step one.
GhSOC1 gene is shown in Figure 1B at the expression pattern of terminal bud different development stage.In Figure 1B, I, II, III, IV, V, VI represents rough leaf flattens period terminal bud to the 6th true leaf respectively.
The application of embodiment 3, GhSOC1 gene
One, the structure of recombinant plasmid
1, extract cotton variety " CCRI 36 " blade total serum IgE and reverse transcription is cDNA.
2, with the cDNA of step 1 for template, with F1 and R1 composition primer pair carry out pcr amplification, obtain pcr amplification product.
F1：5’-ACATATGCCC
GTCGACATGGTGAGGGGAAAGACTC-3；
R1：5’-GATCGGGGAAATTC
GAGCTCGATAAGTTGGCAATGCCATC-3’。
3, use the pcr amplification product of restriction enzyme Sal I and Sac I double digestion step 2, reclaim digestion products.
4, with restriction enzyme Sal I and Sac I double digestion carrier pRI101-AN, the carrier framework of about 10330bp is reclaimed.
5, the digestion products of step 3 is connected with the carrier framework of step 4, obtains recombinant plasmid pRI101-GhSOC1.According to sequencing result, structrual description carries out to recombinant plasmid pRI101-GhSOC1 as follows: between the Sal I and Sac I restriction enzyme site of carrier pRI101-AN, insert the sequence 2 of sequence table from the double chain DNA molecule shown in 5 ' end 102-785 position Nucleotide.The structural representation of recombinant plasmid pRI101-GhSOC1 is shown in Fig. 2.
Two, the acquisition of GhSOC1 gene Arabidopis thaliana
1, recombinant plasmid pRI101-GhSOC1 is imported Agrobacterium LBA4404 competent cell, obtain recombinational agrobacterium bacterium.
2, dry after the seed disinfection of Columbia ecotype Arabidopis thaliana on aseptic filter paper, then chosen on MS solid medium, with sealed membrane sealing, then wrap masking foil and place 4 DEG C of vernalization 48h, then remove masking foil and be transferred to light temperature incubator (22 DEG C; 16h illumination/8h is dark) cultivate, to transplant after 7 days to soil and to be transferred to culturing room (22 DEG C; 16h illumination/8h is dark).
3, the recombinational agrobacterium that step 1 obtains is seeded to the LB liquid nutrient medium of 5ml containing 50 μ g/ml kantlex, 28 DEG C, 180rpm shaken overnight; Then 200mlLB liquid nutrient medium is transferred to the volume ratio of 1:50,28 DEG C, 180rpm is cultured to OD
600nm=1.2-1.6; Then the centrifugal 10min of 5000rpm, collects thalline, thalline is resuspended in permeabilization buffer (being the aqueous solution of the silwet-77 of 0.02% containing 5g/100mL sucrose, volumn concentration), obtains OD
600nmthe bacteria suspension of=0.6-0.8.
Contaminate 50s, then dark culturing 24h in the bacteria suspension that the flower of the Arabidopis thaliana 4, step 2 obtained obtains in step 3, then continue to cultivate (22 DEG C in culturing room; 16h illumination/8h is dark), after fruit pod maturation, collect seed, be T0 for seed.
5, by T0 for the 1/2MS solid medium be seeded in after seed disinfection vernalization containing 50 μ g/ml kantlex, the plant grown is T0 for plant.
6, the blade of T0 for plant is got, extract total serum IgE reverse transcription is cDNA, take cDNA as template, carry out PCR qualification (target sequence is about 1200bp) with the primer pair that F2 (5 '-TTCCCACTGAATCAAAGGCCAT-3 ') and R2 (5 '-CAATCTGTTGAGTTGGGTTGCAC-3 ') forms.The PCR of part sample identifies that electrophorogram is shown in Fig. 3 (CK1 is take water as the negative control of template, the positive control that CK2 is is template with recombinant plasmid pRI101-GhSOC1).
7, PCR being accredited as positive T0 for plant selfing gathers in the crops T1 for seed, by T1 for the 1/2MS solid medium be seeded in after seed disinfection vernalization containing 50 μ g/ml kantlex, the plant grown is T1 for plant, adopt identical method obtain successively T2 for plant, T3 for plant and T4 for plant.For certain T0 for plant, if the T1 that its stochastic sampling detects all can grow containing on the substratum of kantlex for plant for plant and T4 for plant, T3 for plant, T2, this T0 is being 1 for plant and filial generation thereof and stable is turning GhSOC1 gene strain.Obtain 5 altogether and stable turn GhSOC1 gene strain, called after S-8 strain, S-6 strain, S-4 strain, S-2 strain and S-1 strain successively.
8, the genomic dna of T4 for plant of S-8 strain, S-6 strain, S-4 strain, S-2 strain and S-1 strain is extracted respectively, adopt the DNA molecular shown in sequence 2 of sequence table to carry out southern hybridization as probe, partial results is shown in Fig. 4 (positive control that P is is template with recombinant plasmid pRI101-GhSOC1).Result shows, in each strain, goal gene is single copy.
Three, the acquisition of empty carrier plant is turned
The plasmid that will set out replaces recombinant plasmid to carry out step 2, obtains turning empty carrier plant.
Four, phenotypic evaluation
Respectively the T4 of S-6 strain, S-4 strain and S-2 strain is proceeded as follows for seed for the seed of seed, Columbia ecotype Arabidopis thaliana, the T4 that turns empty vector control plant: be seeded in 1/2MS solid medium, long-day conditions (22 DEG C; 16h illumination/8h is dark) under cultivate, statistics flowering time (namely the flowers are in blossom puts gone through number of days from sowing to first), this plant are bloomed for the first time and were added up the lotus throne number of sheets and stem leaf number the same day.Carry out repeating experiment for three times, repeat each strain in experiment at every turn and get 20 seeds.
The results are shown in Table 1 and Fig. 5 (* * represents P<0.05, and * * * represents P<0.001).
Statistic data under table 1 long-day conditions
| Phenotype | Flowering time | The lotus throne number of sheets | Stem leaf number |
| Columbia ecotype Arabidopis thaliana | 29.29±1.312 | 10.94±0.827 | 2.76±0.437 |
| Turn empty vector control plant | 29.36±1.216 | 10.89±0.628 | 2.73±0.492 |
| S-6 strain | 24.94±1.983 | 7.24±0.903 | 3.82±0.636 |
| S-4 strain | 26.56±1.822 | 7.61±0.850 | 3.06±0.539 |
| S-2 strain | 26.00±1.745 | 7.74±0.548 | 4.05±0.900 |
Result shows, under long-day conditions, transfer-gen plant is bloomed 3.75 days in advance, and lotus throne leaf reduces 3.52, and stem leaf increases by 0.85, all reaches pole conspicuous level.
Certain repeats in experiment, the phenotype after the plant of S-4 strain blooms 3 days for the first time, and sees Fig. 6 with the phenotype that S-4 strain carries out Colombia's Arabidopsis thaliana ecotype of parallel laboratory test.In Fig. 6, in A, the left side is Columbia ecotype Arabidopis thaliana, and the right is the plant (35S::GhSOC1) of S-4 strain, and B is the situation that bears pods of Columbia ecotype Arabidopis thaliana, and C is the situation that bears pods of the plant of S-4 strain.
Claims (1)
1.GhSOC1 albumen, or the gene of coding GhSOC1 albumen, the application in regulating plant flowering time; Described plant is Arabidopis thaliana;
The protein that described GhSOC1 albumen is made up of the aminoacid sequence shown in sequence in sequence table 1;
The gene of described coding GhSOC1 albumen is the DNA molecular described in following (1) or (2) or (3):
(1) in sequence table sequence 2 from the DNA molecular shown in 5 ' end the 102 to 767 Nucleotide;
(2) in sequence table sequence 2 from the DNA molecular shown in 5 ' end the 102 to 785 Nucleotide;
(3) DNA molecular shown in sequence 2 in sequence table.
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| CN104861051B (en) * | 2014-02-25 | 2018-01-09 | 中国科学院遗传与发育生物学研究所 | Plant development associated protein AtUBP15 and its encoding gene and application |
| CN108251432B (en) * | 2018-01-02 | 2021-01-05 | 昆明理工大学 | Notoginseng disease course related protein genePnPRlikeAnd applications |
| CN108753796A (en) * | 2018-07-02 | 2018-11-06 | 中国农业科学院棉花研究所 | A kind of the DB genes and its coding albumen and application of the control limited fruit branch character of cotton |
| CN108949781A (en) * | 2018-08-15 | 2018-12-07 | 中国农业科学院棉花研究所 | A kind of limited fruit branch DB gene of cotton, coding albumen and its application |
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| Title |
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| 开花整合子SOC1花期调控的分子机制;鲜登宇等;《中国蔬菜》;20130315(第6期);1-8 * |
| 棉花花发育相关基因GhSOC1，GhFBO基因的克隆表达分析;卫江辉;《中国优秀硕士学位论文全文数据库农业科技辑》;20120515(第05期);12-32 * |
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