CN101798341B - Heat shock factor protein relevant to resistance of plants as well as coding gene and application thereof - Google Patents
Heat shock factor protein relevant to resistance of plants as well as coding gene and application thereof Download PDFInfo
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- CN101798341B CN101798341B CN 200910078056 CN200910078056A CN101798341B CN 101798341 B CN101798341 B CN 101798341B CN 200910078056 CN200910078056 CN 200910078056 CN 200910078056 A CN200910078056 A CN 200910078056A CN 101798341 B CN101798341 B CN 101798341B
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Abstract
The invention discloses a protein relevant to the resistance of plants as well as a coding gene and application thereof. The protein is a protein in (1) or (2) as follows: (1) a protein comprising an amino acid sequence shown as a sequence 2 in a sequence table; and (2) a protein which is formed by substituting and/or deleting and/or adding an amino acid residue sequence of the sequence 2 in the sequence table by one or several amino acid residues, is relevant to the resistance of the plants and is derived from the (1). A transgenic experiment introducing the coding gene of the protein relevant to the resistance of the plants into tobaccos and arabidopsis thaliana proves that the resistance of the plants introduced with the BhHsf1 gene is obviously enhanced, which shows that the BhHsf1 is a protein relevant to the resistance of the plants. The method for culturing transgenic plants the resistance of which is enhanced has an important theoretical and actual meaning and can be used for the culture and the identification of a resistant plant variety required by the farming and animal husbandry and the governing of an ecological environment.
Description
Technical field
The present invention relates to a kind of albumen relevant with plant stress-resistance and encoding gene and application, particularly relate to a kind of heat shock factor protein relevant with plant stress-resistance and encoding gene and application.
Background technology
Along with the rising of Global Temperature and the growth of extreme weather events, the adverse circumstance such as hot, non-irrigated is on the increasingly significant that affects of the yield and quality of crop.Therefore, aobvious important by the resistance day of molecule manipulation technology raising crop.
After organism is forced, can produce a series of emergency reaction, wherein heat shock protein (heat shockprotein, HSP) can in vivo run-up, and helps that associated protein is folding, distributes and degraded in the cell.Further research finds that the content of HSPs and organism thermotolerance are proportionate, and can improve the stress ability of organism and (the Harf ﹠amp of the survival rate in adverse circumstance; Hayer-Hartl, Protein folding:molecular chaperones inthe cytosol:from nascent chain to folded protein.Science.295; 1852～1858.2002; Young, Barral, Hard, More than folding:localized functions of cytosolic cham pemnes.Trends.Biochem.Sci.28:541～547,2003).
The expression of heat shock protein gene is the transcription factor that is subjected to a class special--thermal excited transcryption factor (heat stresstranscription factor, Hsf) regulate and control, they when the HSP genetic transcription can with its promoter region heat shock element (heat shock element, HSE) this conservative motif specific binding, thereby the open and close of regulation and control heat shock protein gene, finish its corresponding biological function (Pelham.A regulatory upstream promoter element inthe Drosophila hsp70 heat-shock gene.Cel1.30:517-528,1982; Pefisic, Xiao, Lis.Stable binding of Drosophila heat shock factor to head-tohead and tail-to-tail repeats of aconserved 5 bp recognition unit.Cel1.59:797～806,1989).Existing studies show that heat shock protein to improve the plant stress-resistance ability of coercing have obvious effect (Prieto-Dapena,
Almoguera, Jordano, The ectopic overexpression of a seed-specific transcription factor, HaHSFA9, conferstolerance to severe dehydration in vegetative organs.Plant J.54:1004-1014,2008; Chao Xu, Gong Zhenhui, the bright brightness of Lu, Ma Chao, Li Dawei, Zhao Jun, Meng Changjun, the clone of Arabidopis thaliana thermal excited transcryption factor AtHsfA6a and cellular localization analysis, Journal of Northwest Sci Tech University of Agriculture and Forestry (natural science edition) 4 phases in 2008: 94-98).
Summary of the invention
The purpose of this invention is to provide a kind of albumen relevant with stress resistance of plant and encoding gene thereof.
The albumen relevant with stress resistance of plant provided by the present invention, name is called BhHsf1, what derive from Gesneriaceae revolves capsule lettuce tongue (Boea hygrometrica), is following 1) or 2) protein:
1) protein that is formed by the aminoacid sequence shown in the sequence in the sequence table 2;
2) with the amino acid residue sequence of sequence in the sequence table 2 through the replacement of one or several amino-acid residue and/or disappearance and/or interpolation and relevant with stress resistance of plant by 1) protein of deriving.
In order to make 1) in BhHsf1 be convenient to purifying, label as shown in table 1 on N-terminal that can the protein that the aminoacid sequence shown in the sequence 2 forms in by sequence table or C-terminal connect.
The sequence of table 1. label
| Label | Residue | Sequence |
| Poly-Arg | 5-6 (being generally 5) | RRRRR |
| Poly-His | 2-10 (being generally 6) | HHHHHH |
| FLAG | 8 | DYKDDDDK |
| Strep-tag II | 8 | WSHPQFEK |
| c-myc | 10 | EQKLISEEDL |
Above-mentioned 2) but in the BhHsf1 synthetic, also can synthesize first its encoding gene, carry out again biological expression and obtain.Above-mentioned 2) encoding gene of the BhHsf1 in can be by lacking sequence in the sequence table 1 codon of one or several amino-acid residue in the dna sequence dna shown in the 5 ' terminal 15-1116 bit base, and/or carry out the missense mutation of one or several base pair, and/or obtain at the encoding sequence that its 5 ' end and/or 3 ' end connects the label shown in the table 1.
Above-mentioned and cDNA gene plant adversity resistance related protein also belongs to protection scope of the present invention.
Specifically can be following 1 with the cDNA gene of plant adversity resistance related protein)-4) in arbitrary described gene:
1) its encoding sequence be in the sequence table sequence 1 from 5 ' terminal 15-1166 position deoxyribonucleotide;
2) its nucleotide sequence is the sequence 1 in the sequence table;
3) under stringent condition with sequence table in the dna molecular from 5 ' terminal 15-1166 position deoxyribonucleotide hybridization and the described albumen of coding claim 1 of sequence 1;
4) with 1) or 2) gene have the dna molecular of the homology 90% or more and the described albumen of claim 1 of encoding.
Above-mentioned stringent condition can be that (or the solution of 0.1 * SSC), 0.1%SDS is hybridized under 65 ℃ and washed film with 0.1 * SSPE in DNA or RNA hybrid experiment.
The primer of above-mentioned BhHsf1 full length gene or its arbitrary fragment of increasing is to also belonging to protection scope of the present invention.
Contain above-mentioned and recombinant vectors, transgenic cell line and recombinant bacterium the plant adversity resistance related protein encoding gene and also belong to protection scope of the present invention.
Available existing plant expression vector construction contains the recombinant expression vector of BhHsf1 gene.Described plant expression vector comprises the double base agrobacterium vector and can be used for the carrier etc. of plant micropellet bombardment, such as pCAMBIA3301, pCAMBIA1300, pBI121, pBin19, pCAMBIA2301, pCAMBIA1301-UbiN or other plant expression vector of deriving.
When using the gene constructed recombinant expression vector of BhHsf1, can before its transcription initiation Nucleotide, add any enhancement type, composing type, organizing specific type or inducible promoter, such as cauliflower mosaic virus (CAMV) 35S promoter, ubiquitin (Ubiquitin) gene promoter (pUbi) etc., they can use separately or be 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, these enhanser zones can be ATG initiator codon or neighboring region initiator codon etc., but must be identical with the reading frame of encoding sequence, to guarantee the correct translation of whole sequence.The source of described translation control signal and initiator codon is widely, can be natural, also can synthesize.Translation initiation region can be from transcription initiation zone or structure gene.
For the ease of transgenic plant cells or plant being identified and screening, can process used plant expression vector, as be added in the plant to express and to produce the enzyme of colour-change or the gene of luminophor (gus gene, GFP gene, luciferase genes etc.), have the antibiotic marker thing (gentamicin marker, kantlex marker etc.) of resistance or anti-chemical reagent marker gene (such as anti-weedkiller gene) etc.From the security consideration of transgenic plant, can not add any selected marker, directly with adverse circumstance screening transformed plant.
Described recombinant expression vector specifically can be and insert the above-mentioned recombinant expression vector that obtains with encoding gene plant adversity resistance related protein between the multiple clone site of plant expression vector pBin19, such as pBin19-35S-BhHsf1.
Another object of the present invention provides a kind of method of cultivating the transgenic plant of resistance raising.
The method of the transgenic plant that cultivation resistance provided by the present invention improves is that above-mentioned encoding gene BhHsf1 with plant adversity resistance related protein is imported in the plant, obtains the transgenic plant that resistance improves.
Described encoding gene BhHsf1 with plant adversity resistance related protein imports in the plant by described recombinant expression vector.
Above-mentioned resistance specifically can be heat impedance (thermotolerance) and drought resistance (drought tolerance).
Carry that of the present invention and plant expression vector plant adversity resistance related protein encoding gene BhHsf1 can be led by Ti-plasmids, Ri plasmid, plant viral vector, directly delivered DNA, microinjection, electricity, the conventional biological method such as agriculture bacillus mediated is transformed in vegetable cell or the tissue.The plant host that is converted both can be the farm crop such as paddy rice, wheat, soybean, tobacco, corn, rape, Chinese sorghum, cotton, also can be the fruits and vegetables flower plants such as the herbages such as clover, trifolium, wheatgrass and strawberry, tomato.
The present invention screens a BhHsf1 gene that is subjected to drought-induced expression from revolve capsule lettuce tongue (Boea hygrometrica), the transgenic plant heat stress that this gene is imported tobacco and Arabidopis thaliana experimental results show that, the T2 that changes BhHsf1 over to is all 100% for the survival rate of transgene tobacco strain, and the survival rate of genetically modified wild-type tobacco is not 0%; The T2 that changes BhHsf1 over to is respectively 100% (strain 7) for the survival rate of transgenic arabidopsis homozygote strain, 100% (strain 15) and 78% (strain 21), and the survival rate of not genetically modified wild-type Arabidopis thaliana is 0%.The transgenic plant drought stress that this gene is imported Arabidopis thaliana experimental results show that, the T2 that changes BhHsf1 over to is respectively 50% (strain 15) and 38% (strain 21) for the survival rate of transgenic arabidopsis homozygote strain, and the survival rate of not genetically modified wild-type Arabidopis thaliana is 0%.Illustrate that BhHsf1 is the albumen relevant with stress resistance of plant.Have important theory and practical significance with plant anti-adversity associated protein BhHsf1 and encoding gene thereof for new variety such as the crop of cultivating the resistance raising, woods grass, can be used for cultivation and the evaluation of the required resistance plant kind of husbandry and ecological environment treatment.
The present invention will be further described below in conjunction with specific embodiment.
Description of drawings
Fig. 1 is the abduction delivering result that Northern detects BhHsf1
Fig. 2 is the physical map of BhHsf1 gene overexpression carrier
Fig. 3 changes the T2 of pBin19-35S-BhHsf1 over to for the RT-PCR detected result of positive transgene tobacco and Arabidopis thaliana
Fig. 4 is that the transgene tobacco of BhHsf1 gene overexpression and the heat resistance of Arabidopis thaliana are identified
Fig. 5 is that the drought-resistant ability of the Arabidopis thaliana of BhHsf1 gene overexpression is identified
Embodiment
Method is ordinary method if no special instructions described in the following embodiment, and it is synthetic that the primer and probe are given birth to the worker by Shanghai.
Extraction is processed (25 ℃, illumination places on the filter paper air-dry, lower same) total RNA that revolves capsule lettuce tongue (Boea hygrometrica) blade of 8 hours through drought stress, utilizes ZAP-
Library construction test kit (Stratagene, La Jolla, CA) construction cDNA library.Therefrom 4800 genes of random choose use BioGridrobot (BioRobotics Ltd, Cambridge, UK) automatization point sample at Hybond-N
+On the nylon membrane (AmershamBiosciences, Freiburg, Germany), make cDNA microarray (cDNA chip).With this cDNA chip respectively with process without arid normal growth revolve capsule lettuce tongue blade and process 8 hours the polyA-RNA that revolves capsule lettuce tongue blade through drought stress prepared
33The probe of P mark is hybridized, and analyzes them in the dynamic change of normal condition and drought-induced rear genetic expression.Found that wherein 1 dna fragmentation from 5 ' end 667-1453 position deoxyribonucleotide that the gene through drought-induced rise has sequence 1.
The design primer utilizes 5 '-RACE method to obtain the full length sequence of this gene, called after BhHsf1.5 '-RACE concrete grammar is as follows:
1) synthetic the first chain cDNA
Extraction is processed 8 hours the total RNA that revolves capsule lettuce tongue (Boea hygrometrica) blade and take it as template, is used gene-specific primer GSP-Race1:5 '-tatccgacggctgctt-3 ' to carry out reverse transcription through drought stress.Reaction system is: total RNA (2 μ g/ μ l) 1.0 μ l, GSP-Race1 (1 μ M) 2 μ l, dNTP (2mM) 5 μ l, DEPC-H
2O 6.0 μ l, 5 * M-MLV buffer, 4 μ l, RNase inhibitor (5u/ μ l is available from Takara company) 1 μ l, M-MLV (available from Promega company) 1 μ l.65 ℃ of 10min again behind 37 ℃ of reaction 1h, purifying add the C tail after reclaiming (three rich PCR products reclaim test kit).
2) tailing of cDNA
Reaction system is: cDNA 5 μ l, DEPC-H
2O 13.5 μ l, 5 * buffer, 5 μ l, dCTP (10mm) 0.5 μ l.Add 1 μ l TdT terminal enzyme (DNA) (available from takara company) behind 75 ℃ of reaction 5min, 37 ℃ are reacted 1h again, obtain adding the cDNA sequence of C tail.
3) PCR of tailing cDNA
Take the cDNA sequence that adds the C tail of above-mentioned acquisition as template, pcr amplification.Reaction system is: cDNA 1 μ l, 10 * PCR buffer, 1 μ l, dNTP (2mm) 1 μ l, contain the primer GSP-Race2:5 ' of BamHI restriction endonuclease recognition site-gggatcctttggagggatttgat-3 ' (10 μ m) 0.5 μ l, poly G anchor primer 5 '-ggccacgcgtcgactagtacg14-3 ' (10 μ m) the 0.5 μ l that contains SalI restriction endonuclease recognition site, Taq enzyme (available from takara company) 0.1 μ l.Reaction conditions is: first 95 ℃ of denaturation 4min, and then 94 ℃ of sex change are 30 seconds, 55 ℃ of annealing 30 seconds, 72 extended totally 35 circulations 2 minutes again; Last 72 ℃ were extended 10 minutes.
4) nest-type PRC amplification
Above-mentioned PCR product is diluted 100 times as template, pcr amplification.The PCR reaction system is: template 1 μ l, 10 * PCR buffer, 1 μ l, dNTP (2mm) 1 μ l, contain the primer GSP-Race3:5 ' of BamHI restriction endonuclease recognition site-ctggat cctccttttgatgtgtttc-3 ' 0.5 μ l, contain the anchor primer 5 ' of SalI restriction endonuclease recognition site-ggccacgcgtcgactagtac-3 ' (10 μ m) 0.5 μ l, Taq enzyme (available from takara company) 0.1 μ l.Reaction conditions is: first 95 ℃ of denaturation 4min, and then 94 ℃ of sex change are 30 seconds, 55 ℃ of annealing 30 seconds, 72 extended totally 35 circulations 2 minutes again; Last 72 ℃ were extended 10 minutes.
The PCR product is carried out agarose gel electrophoresis detect, the result obtains the band about 500bp; Reclaim the band about this 500bp, and check order after carrier pGEM-T (available from Promega company) is connected.Splicing obtains BhHsf1 gene complete sequence according to sequencing result, its nucleotide sequence is shown in sequence in the sequence table 1, analyze and find that this sequence comprises open reading frame (ORF), 5 '-UTR (5 '-non-translational region) and the 3 '-UTR of complete BhHsf1 gene, its ORF is the deoxyribonucleotide from 5 ' terminal 15-1166 position, and the amino acid residue sequence of coding is shown in sequence in the sequence table 2.This amino acid residue sequence belongs to the Hsf protein family.
The abduction delivering analysis of embodiment 2, BhHsf1 gene
Northern blot carries out with reference to Amersham Phamacia Biotech product description.
Extract respectively and be not subjected to drought stress, drought stress to process 2,8,24,72 hours the total RNA that revolves capsule lettuce tongue (Boeahygrometrica) blade, carry out Northern blot take following dna fragmentation as probe: extract the total RNA that revolves capsule lettuce tongue (Boea hygrometrfca) blade that processed through drought stress 8 hours, with hsf1-f5 '-cgagatctatggtttctgtggtag-3 ' and
GSP-Race3:5 '-ctggatcctccttttgatgtgtttc-3 ' is for primer carries out RT-PCR, obtain having sequence 1 from the dna fragmentation of 5 ' terminal 15-1166 position deoxyribonucleotide as probe.
From left to right be followed successively by among the Northern blot (Fig. 1) and be not subjected to drought stress, drought stress 2h, 8h, 24h and 72h.Concrete detected result as shown in Figure 1.Wherein,
L
CDo not revolve capsule lettuce tongue blade for what be subjected to drought stress; Ld2 revolves capsule lettuce tongue blade behind the drought stress 2h;
Ld8 revolves capsule lettuce tongue blade behind the drought stress 8h; Ld24 revolves capsule lettuce tongue blade behind the drought stress 24h;
Ld72 revolves capsule lettuce tongue blade behind the drought stress 72h.
Upper figure is the Northern blot detected result of BhHsf1 mRNA; Figure below is total RNA formaldehyde gel electrophoretic image before the corresponding transferring film of above sample, and two bands are 18S rRNA and 28S rRNA.
Northern blot test experience is presented at and is not subjected to revolving of drought stress not detect BhHsf1 in the capsule lettuce tongue blade to express, BhHsf1 just has higher expression in drought stress 2h blade, and remain to drought stress 72h always, illustrate that BhHsf1 is the gene of abduction delivering of being dewatered really.
Embodiment 3, the resistance that turns the BhHsf1 genetic tobacco detect to be analyzed
1) turns the acquisition of BhHsf1 genetic tobacco and Arabidopis thaliana
A. preparation contains the recombinant expression vector of BhHsf1 gene
8 hours the total RNA that revolves capsule lettuce tongue (Boea hygrometrica) blade is processed in extraction through drought stress, carry out RT-PCR take hsf1-f 5 '-cgagatctatggtttctgtggtag-3 ' and hsf1-r 5 '-agtcgacagtaggcgaatcccat-3 ' as primer, obtain having the dna fragmentation from 5 ' terminal 15-1166 position deoxyribonucleotide of sequence 1, this dna fragmentation is connected on the pEasy-blunt carrier (the full formula in Beijing King Company), with the recombinant vectors called after pBhHsf1 that obtains.Behind recombinant vectors pBhHsf1 usefulness BamHI and SalI (available from the precious biotech firm in Dalian) double digestion, reclaim the fragment (BhHsf1) of 1152bp, with CaMV35S promotor and poly-A sequence (Odell, JT; Nagy, F; Chua, NH.Identification of DNA sequences required for activity of thecauliflower mosaic virus 35S promoter.Nature.313 (2005): 810-812) together be connected to carrier pBin19 (D.A.Frisch, L.W.Harris-Haller, N.T.Yokubaitis, T.L.Thomas, S.H.Hardin, T.C.Hall, Complete sequence of the binary vector Bin 19, Plant Mol.Biol.27 (1995) 405-409.), obtain recombinant plasmid pBin19-35S-BhHsf1 (physical map as shown in Figure 2).Transform the bacillus coli DH 5 alpha competent cell, go out required recon and carry out sequencing analysis with the LB Screening of Media that contains 10 μ g/ml sulphuric acid kanamycins, identify through order-checking, prove that the sequence of recombinant plasmid and structure are correct.
B. contain the preparation of reconstitution cell and the recombinant plant of BhHsf1 gene
Utilize electric shocking method that pBin19-35S-BhHsf1 is transformed in Agrobacterium LAB4404 (Beijing Baeyer enlightening company) cell, screen with the YEB substratum that contains 50 μ g/ml sulphuric acid kanamycins, 30 μ g/ml Streptomycin sulphates and 50 μ g/ml Rifampins, obtain to carry the restructuring agrobacterium strains LAB-pBin19-35S-BhHsf1 of 35S promoter and BhHsf1 gene.
LAB-pBin19-35S-BhHsf1 is adopted leaf dish method transformation of tobacco (Nicotiana tabacum cv.SR-1), floral organ infusion method arabidopsis thaliana transformation (Col-0), above-mentioned transgene tobacco and Arabidopis thaliana are moved to hot-house culture, the seed of selfing, collection transgene tobacco.Simultaneously, with wild-type tobacco and Arabidopis thaliana in contrast.
With the planting seed of above-mentioned transgene tobacco and Arabidopis thaliana on the MS substratum that contains 100 μ g/ml sulphuric acid kanamycins, will be on the above-mentioned MS substratum that contains 100 μ g/ml sulphuric acid kanamycins resistance separate than the T1 that is 3: 1 and move to hot-house culture for the surviving seedlings of transgenic plant, collection T1 is for the seed of transgenic plant.Get 100 T1 and be seeded in for transgenic plant seed in the MS substratum that contains 100 μ g/ml sulphuric acid kanamycins, the result obtains three T2 altogether for turning pBin19-35S-BhHsf1 tobacco strain: 2-9,4-10,8-2.Three T2 are for turning pBin19-35S-BhHsf1 Arabidopis thaliana strain 7,15,21.
C. the BhHsf1 gene expression analysis of transgenic plant
Extract the T2 of above-mentioned each strain of a month seedling age for total RNA of transgenic plant, carry out RT-PCR take hsf1-f5 '-cgagatctatggtttctgtggtag-3 ' and hsf1-r 5 '-agtcgacagtaggcgaatcccat-3 ' as primer, simultaneously with not genetically modified wild-type tobacco and Arabidopis thaliana in contrast.The pcr amplification program is: 95 ℃ of denaturation 4min, and 94 ℃ of sex change 45 seconds, 56 ℃ of annealing 30 seconds, 72 ℃ were extended totally 27 circulations 1 minute; At last again 72 ℃ extended 10 minutes; Simultaneously with 18s rRNA gene as confidential reference items, the pcr amplification program is: 95 ℃ of denaturation 4min, 94 ℃ of sex change 45 seconds, 56 ℃ of annealing 30 seconds, 72 ℃ were extended totally 18 circulations 1 minute; At last again 72 ℃ extended 10 minutes.The PCR product is carried out agarose gel electrophoresis detect, every group of sample carries out 2 times to be repeated.The result shows, does not amplify band in the not genetically modified wild-type tobacco (WT), turns among pBin19-35S-BhHsf1 tobacco strain 2-9,4-10 and the 8-2, all has BhHsf1 to express.Do not amplify band in the not genetically modified wild-type Arabidopis thaliana (WT), turning all has BhHsf1 to express (Fig. 3) in pBin19-35S-BhHsf1 Arabidopis thaliana strain 7,15 and 21.
Among Fig. 3, increased the stripe size of the BhHsf1 gene fragment that obtains as 1164bp take primer hsf1-f and hsf1-r, 18S rRNA gene is 195bp as the increase stripe size of the dna fragmentation that obtains of confidential reference items.
Left side WT refers to not turn the Arabidopis thaliana wild-type of BhHsf1 gene, show among the figure with PCR method gene specific amplification BhHsf1 gene (on) and 18s rRNA shown Nucleotide histogram picture in sepharose.
Right side WT refers to not turn the tobacco wild-type of BhHsf1 gene, show among the figure with PCR carry out gene specific amplification BhHsf1 gene (on) and 18s rRNA shown Nucleotide histogram picture in sepharose.
2) detection turns the resistance of transgene tobacco and the transgenic arabidopsis of BhHsf1 gene
With embodiment 3 steps 1) T2 of 3 strains that change plasmid pBin19-35S-BhHsf1 over to of obtaining among the A is sowed on the MS substratum that contains 100 μ g/ml sulphuric acid kanamycins for transgenic arabidopsis, not genetically modified wild-type Arabidopis thaliana is sowed on the MS substratum, cultivation is under the condition of 22 ℃, 50% humidity, illumination 16h and dark 8h, after 6 days all seedlings all are transferred on the MS substratum, carrying out heat shock after 3 days processes: 37 ℃ of heat shocks 1 hour, normal culture condition (22 ℃) 3 hours, then 49 ℃ of heat shocks are 1 hour.Then be put in normal culture condition (22 ℃).Take a picture after 3 days and add up survival rate.Three repetitions are established in experiment altogether, repeat equal 50 strains of each strain at every turn.The result is shown in Fig. 4 A and B.Without the heat shock treatment group, the equal normal growth of wild-type and transgenic arabidopsis (Fig. 4 A); And the heat shock treatment group, the survival rate of not genetically modified wild-type Arabidopis thaliana is 0%; The T2 that changes plasmid pBin19-35S-BhHsf1 over to is respectively 100% (7), 100% (15) and 78% (21) (Fig. 4 B) for the survival rate of transgenic arabidopsis strain.Show that the heat impedance that turns BhHsf1 gene Arabidopis thaliana obviously improves.
With embodiment 3 steps 1) T2 of 3 strains that change plasmid pBin19-35S-BhHsf1 over to of obtaining among the A is sowed on the MS substratum for transgene tobacco and not genetically modified wild-type tobacco, cultivation is under the condition of 22 ℃, 50% humidity, illumination 16h and dark 8h, carrying out heat shock after 2 weeks processes: 49 ℃ of heat shocks are put in normal culture condition after 2 hours, take a picture after 7 days and add up survival rate.Three repetitions are established in experiment altogether, repeat equal 100 strains of each strain at every turn.The results are shown in Figure the front tobacco photo of 4, Fig. 4 C heat shock, Fig. 4 D is the tobacco photo of growing again 7 days after heat shock is processed, and shows that wild-type tobacco survival rate after heat shock of normal growth is 0%; The T2 that changes plasmid pBin19-35S-BhHsf1 over to is all 100% for transgenosis homozygote tobacco strain survival rate after heat shock.Illustrate that the heat impedance that turns the BhHsf1 genetic tobacco obviously improves.
With embodiment 3 steps 1) T2 of 3 strains that change plasmid pBin19-35S-BhHsf1 over to of obtaining among the A is sowed on the MS substratum that contains 100 μ g/ml sulphuric acid kanamycins for transgenic arabidopsis, not genetically modified wild-type Arabidopis thaliana is sowed on the MS substratum, cultivation (lower with) under 22 ℃, the condition of 50% humidity, photoperiod 16/8h, all be transferred to all seedlings in the soil after 5 days, culture condition is the same, stop to water after 3 weeks, carrying out drought stress processes: arid was taken a picture after 2 weeks, recovery is watered, and takes a picture after 3 days and adds up survival rate.Three repetitions are established in experiment altogether, repeat equal 48 strains of each strain at every turn.The result as shown in Figure 5.When processing without arid, the equal normal growth of wild-type and transgenic arabidopsis (Fig. 5 A); And after the arid processing, it is withered that wild-type and transgenic arabidopsis are all wilted, but transgenic plant are significantly better than wild-type (Fig. 5 B); After the rehydration, the survival rate of not genetically modified wild-type Arabidopis thaliana is 0%; The T2 that changes plasmid pBin19-35S-BhHsf1 over to is respectively 50% (15) and 38% (21) (figure for the survival rate of transgenic arabidopsis strain
C).Show that the drought resistance that turns BhHsf1 gene Arabidopis thaliana obviously improves.
The experimental result of this embodiment shows that the resistance that changes the plant of BhHsf1 gene over to obviously is better than not turning the plant of BhHsf1 gene, illustrates that BhHsf1 is one of albumen relevant with stress resistance of plant.
Sequence table
＜110〉Institute of Botany, Chinese Academy of Sciences
＜120〉heat shock factor protein relevant with stress resistance of plant and encoding gene thereof and application
<160>2
<210>1
<211>1453
<212>DNA
＜213〉revolve capsule lettuce tongue genus and revolve capsule lettuce tongue (Boea hygrometrica)
<221>CDS
<222>(15)..(1166)
<400>1
aaacatactt gtac atg gtt tct gtg gta gct gca ggg agt aaa caa gaa 50
Met Val Ser Val Val Ala Ala Gly Ser Lys Gln Glu
1 5 10
atg gtg gcg agt gaa gct ctg att ggt gtg aag gag gag ccc ttg gtg 98
Met Val Ala Ser Glu Ala Leu Ile Gly Val Lys Glu Glu Pro Leu Val
15 20 25
ttt ctt gat gag gct gag tat ctt gga ggt ggg ttc agt ggc tgc agg 146
Phe Leu Asp Glu Ala Glu Tyr Leu Gly Gly Gly Phe Ser Gly Cys Arg
30 35 40
agt gga ggt gag gag gag gag tgg ggg gat gcg gcg gag gag cat ctt 194
Ser Gly Gly Glu Glu Glu Glu Trp Gly Asp Ala Ala Glu Glu His Leu
45 50 55 60
ccg aaa ccg ttg gag ggg ctg agg gat atc ggg cct ccc ccg ttt ctg 242
Pro Lys Pro Leu Glu Gly Leu Arg Asp Ile Gly Pro Pro Pro Phe Leu
65 70 75
aag aag acg ttt gag atg gtg gat gat cct cgt acg gac tcg ata ctg 290
Lys Lys Thr Phe Glu Met Val Asp Asp Pro Arg Thr Asp Ser Ile Leu
80 85 90
tcg tgg agt ggc gcc ggc aat agc ttc gtt gtg tgg gat ccc cac acc 338
Ser Trp Ser Gly Ala Gly Asn Ser Phe Val Val Trp Asp Pro His Thr
95 100 105
ttt gcc acc gat ctg ctc ccc aag cat ttc aag cac aac aac ttc tcc 386
Phe Ala Thr Asp Leu Leu Pro Lys His Phe Lys His Asn Asn Phe Ser
110 115 120
agc ttc gtc cgc caa cta aac acc tat aga ttc agg aag att gat tcg 434
Ser Phe Val Arg Gln Leu Asn Thr Tyr Arg Phe Arg Lys Ile Asp Ser
125 130 135 140
gac aga tgg gag ttt gcc aac gaa ggg ttc cgg agg aac aag aag cat 482
Asp Arg Trp Glu Phe Ala Asn Glu Gly Phe Arg Arg Asn Lys Lys His
145 150 155
ttg ctg aaa cac atc aaa agg agg aag cag agc ccc caa atg atg cgg 530
Leu Leu Lys His Ile Lys Arg Arg Lys Gln Ser Pro Gln Met Met Arg
160 165 170
cca cac gag gca gca gca gca gca cag ccg tgg cag tat ccc acc aat 578
Pro His Glu Ala Ala Ala Ala Ala Gln Pro Trp Gln Tyr Pro Thr Asn
175 180 185
cat ggt gtg gat tcc gag att tat aag ctc gga gcc gac cag agc ctt 626
His Gly Val Asp Ser Glu Ile Tyr Lys Leu Gly Ala Asp Gln Ser Leu
190 195 200
ctg agg caa gaa atc gtg aag cta agg cag caa caa gag tgc tcg cag 674
Leu Arg Gln Glu Ile Val Lys Leu Arg Gln Gln Gln Glu Cys Ser Gln
205 210 215 220
cgc tac atc gcc gcc atg gaa gag cgt ctc cac gcc tcc gag atg cag 722
Arg Tyr Ile Ala Ala Met Glu Glu Arg Leu His Ala Ser Glu Met Gln
225 230 235
caa aaa cac atg atc gtt ttc atg atc aaa tcc ctc aag gat ccc atg 770
Gln Lys His Met Ile Val Phe Met Ile Lys Ser Leu Lys Asp Pro Met
240 245 250
ttt tta ctg gac tgt gtg gac agg atc aac agg aaa agg gca ctg agc 818
Phe Leu Leu Asp Cys Val Asp Arg Ile Asn Arg Lys Arg Ala Leu Ser
255 260 265
agc gaa gaa gtc gcc ttc aaa aga agg cgg ttg tct gaa aac atg gaa 866
Ser Glu Glu Val Ala Phe Lys Arg Arg Arg Leu Ser Glu Asn Met Glu
270 275 280
tcc aat att ggt att gat cag gac agg agg ttc caa gcc cag gag gag 914
Ser Asn Ile Gly Ile Asp Gln Asp Arg Arg Phe Gln Ala Gln Glu Glu
285 290 295 300
ttg agt acc ata cca tcc gaa ata cag aca cta ttc tct ccc gat tca 962
Leu Ser Thr Ile Pro Ser Glu Ile Gln Thr Leu Phe Ser Pro Asp Ser
305 310 315
tcc ggg agc cct gta cag gac cac aag gcc gaa aca gaa ttg cac agc 1010
Ser Gly Ser Pro Val Gln Asp His Lys Ala Glu Thr Glu Leu His Ser
320 325 330
tct gat gtc tgc tcc gac aac ttc ata ttg tgg gag aaa ctc atg gaa 1058
Ser Asp Val Cys Ser Asp Asn Phe Ile Leu Trp Glu Lys Leu Met Glu
335 340 345
gat gac atg atc tac gat gaa gaa caa ggg ccg gaa aag cag ccg tcg 1106
Asp Asp Met Ile Tyr Asp Glu Glu Gln Gly Pro Glu Lys Gln Pro Ser
350 355 360
gat acc cgt agg ata gga gaa ttc gat ccc aaa acc gtc gca atg gga 1154
Asp Thr Arg Arg Ile Gly Glu Phe Asp Pro Lys Thr Val Ala Met Gly
365 370 375 380
ttc gcc tac tag agaccggccg gaaggaggtc gtttccgaca tataaaatcc 1206
Phe Ala Tyr
agatcgaggt acaaatggaa agtgttacat atatatgatc atattttata taatttcctt 1266
gtcattgttg tatgctaaat tattagttta ttagatagga attaggattt tagttgaaag 1326
tggcattttt gcagagaacg ggagtctgcc gataatgtca tcatgtataa tatccatatt 1386
tttctcactt tgtaagtttg gtctatagac ttcatcatat atcatgtgaa tgtatcttga 1446
actctga 1453
<210>2
<211>383
<212>PRT
＜213〉revolve capsule lettuce tongue genus and revolve capsule lettuce tongue (Boea hygrometrica)
<400>2
Met Val Ser Val Val Ala Ala Gly Ser Lys Gln Glu Met Val Ala Ser
1 5 10 15
Glu Ala Leu Ile Gly Val Lys Glu Glu Pro Leu Val Phe Leu Asp Glu
20 25 30
Ala Glu Tyr Leu Gly Gly Gly Phe Ser Gly Cys Arg Ser Gly Gly Glu
35 40 45
Glu Glu Glu Trp Gly Asp Ala Ala Glu Glu His Leu Pro Lys Pro Leu
50 55 60
Glu Gly Leu Arg Asp Ile Gly Pro Pro Pro Phe Leu Lys Lys Thr Phe
65 70 75 80
Glu Met Val Asp Asp Pro Arg Thr Asp Ser Ile Leu Ser Trp Ser Gly
85 90 95
Ala Gly Asn Ser Phe Val Val Trp Asp Pro His Thr Phe Ala Thr Asp
100 105 110
Leu Leu Pro Lys His Phe Lys His Asn Asn Phe Ser Ser Phe Val Arg
115 120 125
Gln Leu Asn Thr Tyr Arg Phe Arg Lys Ile Asp Ser Asp Arg Trp Glu
130 135 140
Phe Ala Asn Glu Gly Phe Arg Arg Asn Lys Lys His Leu Leu Lys His
145 150 155 160
Ile Lys Arg Arg Lys Gln Ser Pro Gln Met Met Arg Pro His Glu Ala
165 170 175
Ala Ala Ala Ala Gln Pro Trp Gln Tyr Pro Thr Asn His Gly Val Asp
180 185 190
Ser Glu Ile Tyr Lys Leu Gly Ala Asp Gln Ser Leu Leu Arg Gln Glu
195 200 205
Ile Val Lys Leu Arg Gln Gln Gln Glu Cys Ser Gln Arg TyrIle Ala
210 215 220
Ala Met Glu Glu Arg Leu His Ala Ser Glu Met Gln Gln Lys His Met
225 230 235 240
Ile Val Phe Met Ile Lys Ser Leu Lys Asp Pro Met Phe Leu Leu Asp
245 250 255
Cys Val Asp Arg Ile Asn Arg Lys Arg Ala Leu Ser Ser Glu Glu Val
260 265 270
Ala Phe Lys Arg Arg Arg Leu Ser Glu Asn Met Glu Ser Asn Ile Gly
275 280 285
Ile Asp Gln Asp Arg Arg Phe Gln Ala Gln Glu Glu Leu Ser Thr Ile
290 295 300
Pro Ser Glu Ile Gln Thr Leu Phe Ser Pro Asp Ser Ser Gly Ser Pro
305 310 315 320
Val Gln Asp His Lys Ala Glu Thr Glu Leu His Ser Ser Asp Val Cys
325 330 335
Ser Asp Asn Phe Ile Leu Trp Glu Lys Leu Met Glu Asp Asp Met Ile
340 345 350
Tyr Asp Glu Glu Gln Gly Pro Glu Lys Gln Pro Ser Asp Thr Arg Arg
355 360 365
Ile Gly Glu Phe Asp Pro Lys Thr Val Ala Met Gly Phe Ala Tyr
370 375 380
Claims (12)
1. albumen, the protein that is formed by the aminoacid sequence shown in the sequence in the sequence table 2.
2. the encoding gene of the described albumen of claim 1.
3. encoding gene according to claim 2, it is characterized in that: the cDNA gene of described albumen is following 1)-2) in arbitrary described gene:
1) its encoding sequence be in the sequence table sequence 1 from 5 ' terminal 15-1166 position deoxyribonucleotide;
2) its nucleotide sequence is the sequence 1 in the sequence table.
4. the recombinant expression vector that contains claim 2 or 3 described genes.
5. recombinant expression vector according to claim 4 is characterized in that: the recombinant expression vector that described recombinant expression vector obtains for insert claim 2 or 3 described genes between the multiple clone site of pBin19.
6. the transgenic cell line that contains claim 2 or 3 described genes.
7. the recombinant bacterium that contains claim 2 or 3 described genes.
8. a method of cultivating transgenic plant is that claim 2 or 3 described encoding genes are changed in the plant, obtains the resistant transgenic plant.
9. method according to claim 8 is characterized in that: claim 2 or 3 described encoding genes are to import in the plants by claim 4 or 5 described recombinant expression vectors.
10. according to claim 8 or 9 described methods, it is characterized in that: described plant is paddy rice, wheat, soybean, tobacco, Arabidopis thaliana, corn, rape, Chinese sorghum, cotton, clover, trifolium, wheatgrass, strawberry or tomato.
11. according to claim 8 or 9 described methods, it is characterized in that: described resistant transgenic plant is heat resistanceheat resistant and/or drought resisting transgenic plant.
12. method according to claim 10 is characterized in that: described resistant transgenic plant is heat resistanceheat resistant and/or drought resisting transgenic plant.
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| CN102363790B (en) * | 2011-10-11 | 2013-10-16 | 昆明理工大学 | Plant expression vector of arabidopsis heat shock factor gene AtHsfA1d, and application thereof |
| CN103897047B (en) * | 2012-12-27 | 2016-01-20 | 中国科学院植物研究所 | Protein B hHSP70-1 and encoding gene thereof and application |
| CN104119430B (en) * | 2013-04-26 | 2016-08-10 | 中国科学院植物研究所 | A kind of albumen relevant to drought tolerance in plants and encoding gene thereof and application |
| CN103665128B (en) * | 2013-12-18 | 2015-06-17 | 中国农业大学 | Protein related with heat resistance of plants as well as encoding gene and application of protein |
| CN105301092B (en) * | 2015-12-04 | 2017-12-22 | 云南省烟草农业科学研究院 | A kind of application of albumen in the detection of tobacco drought tolerance |
| NO344051B1 (en) * | 2017-05-04 | 2019-08-26 | Patogen As | Novel virus in Fish and Method for detection |
| CN108034664B (en) * | 2018-01-26 | 2020-10-23 | 四川天艺生态园林集团股份有限公司 | Ornithogalum caudatum Qt DCAF8 gene and application thereof |
| CN112410347A (en) * | 2019-08-21 | 2021-02-26 | 中国农业大学 | Corn ZmHsf21 gene and application thereof |
| CN112251462B (en) * | 2020-10-26 | 2022-05-17 | 南京农业大学 | Application of soybeans GmHSFA2 and GmHSP20a in enhancing heat resistance of plants in flowering period |
| CN113604480B (en) * | 2021-09-17 | 2023-07-04 | 四川农业大学 | Corn transcription factor ZmHsf28 and application thereof |
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| Almoguera,C..A seed-specific heat-shock transcription factor involved in.《GenBank》.2002, * |
| 伊淑莹.植物热激因子网络.《生命的化学》.2008, * |
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