CN105481959B - A method of plant is improved to NaCl tolerance by lowering PAB2 and PAB4 - Google Patents
A method of plant is improved to NaCl tolerance by lowering PAB2 and PAB4 Download PDFInfo
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Abstract
Plant is improved to the method for NaCl tolerance by lowering PAB2 and PAB4 the invention discloses a kind of.The present invention specifically provide protein 1 and protein 2 as target it is following it is any in application: a1) improve the salt tolerance of plant；A2) the plant variety that breeding salt tolerance improves；The protein 1 is PAB2 albumen；The protein 2 is PAB4 albumen.The research of the invention finds that PAB2 the and PAB4 Gene Double knockout mutations body obtained from arabidopsis biological study center enhances NaCl tolerance；Therefore, plant is adjusted to the tolerance of NaCl using PAB2 and PAB4.In addition, PAB2 and PAB4 low expression, salt tolerant genetically modified crops can also be obtained by genetic engineering means (RNAi technology).The present invention meets sustainable agriculture development demand, and for studying molecular mechanism, the improvement hereditary capacity of Plant Tolerance NaCl, cultivating Efficient salt-tolerant new varieties etc. has important practical value and market prospects.
Description
Technical field
The invention belongs to field of biotechnology, are related to a kind of by lowering PAB2 and PAB4 raising plant to NaCl tolerance
Method.
Background technique
As world population constantly increases, demand of the mankind to grain is increasing.Salt stress is influenced in world wide
One of principal element of crop yield.Salt stress mainly causes to damage as stress by sodium chloride to plant.Soil
The sodium ion of middle and high concentration will have a direct impact on protein synthesis, photosynthesis and energetic supersession of plant etc., lead to plant
Hypoevolutism, germination percentage reduce, the growth of organ and differentiation are suppressed, metabolism slows down, albumen synthesizes reduction etc., finally
Influence the yield of crop.Up to the present, the gross area in the saline and alkaline soil in the whole world is up to 8.4 hundred million hm for cut-off2, account for about world land area
6%, and press annual hundred million hm of 1-42Speed increase；The saline alkali land area in China is 0.27 hundred million hm2, account for about cultivated area
10%, just with the increase of annual more than 1000 mu of speed.
Under the ever-increasing severe situation of China's salt lick, it is effectively improved and utilizes salt lick resource and improve salt marsh
The crop yield changed on soil has major and immediate significance to China's agricultural development.So far, improve and utilize salt lick
Mode there are mainly two types of: first is that passing through the betterments of land engineering measure such as Rational Irrigation, the fresh water desalinization of soil by flooding or leaching；But these methods consume
It provides huge and has very strong side effect；Current freshwater resources become increasingly precious, the enforcement difficulty of these measures it is larger and
It is difficult to maintain.The method of another effective use salinization land is to cultivate Efficient salt-tolerant new varieties, this is also Future Development agriculture
The important directions of industry.
The salt tolerance of plant is to be controlled by polygenes quantitative character and complex character affected by environment；Therefore, tradition is educated
Kind method is extremely limited in the effect for improving salt tolerance of crop.With the development of molecular biology and scientists are to plant
Salt tolerant Study on Molecular Mechanism deepens continuously, and the salt tolerance for improving crop by genetic engineering means is in widespread attention.Gene
The advantage of engineering technology means be can accurately, stably and efficiently change gene expression, and then can effectively improve crop
Salt tolerance greatly accelerates breeding speed, becomes the effective way for cultivating Efficient salt-tolerant new varieties.And identify resistant gene of salt with
And further investigation plant salt tolerance molecular mechanism is to cultivate the top priority of Efficient salt-tolerant new varieties.
Poly (A) is almost present in all Eukaryotic ends mRNA3 ', and affects the almost all of metabolism of mRNA
Activity: transhipment, translation and degradation.Poly (A) binding protein [Poly (A)-binding protein, PABP/PAB] is RNA knot
A kind of highly conserved albumen subtribe, is widely present in the eucaryotes such as yeast, nematode, people and arabidopsis in hop protein superfamily
In different types of cell.PABP/PAB albumen is important one of translation initiation factor, it can be with the end mRNA poly (A) tail
It bar combines, controls the length of poly (A), participate in synthesis, degradation and stability and starting of translation of regulating mRNA etc..
More to the research of PABP gene in yeast and animal, all there is their primary structure and gene structure height to protect
Keeping property, and research of the higher plant in relation to PABP gene is less.In yeast cells, the deletion mutant of PABP gene is lethal
Type, illustrate that the gene pairs yeast cells is of crucial importance；Arabidopsis PABP5 gene is anther-specific expression, which exists
Important role may be also functioned in Microspore development, its missing may cause the abortion of plant.In arabidopsis
Gene number of the PAB2 gene in the website tair is At4g34110, and gene number of the PAB4 gene in the website tair is At2g23350
(https://www.arabidopsis.org/)；In addition, these three genes of PAB2, PAB4 and PAB8 are reported ginseng in arabidopsis
With the duplication of regulation virus.
With the expansion of plant salt tolerance Study on Molecular Mechanism goed deep into and apply, how to be changed using the resistant gene of salt having found
Become plant to the tolerance of NaCl and how to cultivate Efficient salt-tolerant new varieties as study frontier.
Summary of the invention
Plant is improved to the method for NaCl tolerance by lowering PAB2 and PAB4 the object of the present invention is to provide a kind of.
The present invention provides applications any in following A-C:
A. protein 1 and protein 2 as target it is following it is any in application:
A1 the salt tolerance of plant) is improved；
A2) the plant variety that breeding salt tolerance improves；
The protein 1 is PAB2 albumen；The protein 2 is PAB4 albumen.
In the application, described " protein 1 and protein 2 be used as target " concretely: with the protein 1 and institute
Protein 2 is stated as target, lowers the expression of the protein 1 and the protein 2.
B. be able to suppress protein 1 and protein 2 are expressed in plant substance it is following it is any in application:
A1 the salt tolerance of plant) is improved；
A2) the plant variety that breeding salt tolerance improves；
The protein 1 is PAB2 albumen；The protein 2 is PAB4 albumen.
C. the substance of the encoding gene of the encoding gene of protein 1 and protein 2 expression in plant is able to suppress as follows
Application in any:
A1 the salt tolerance of plant) is improved；
A2) the plant variety that breeding salt tolerance improves；
The protein 1 is PAB2 albumen；The protein 2 is PAB4 albumen.
Further, protein 1 (PAB2 albumen) amino acid sequence shown in sequence 3 in sequence table forms
Protein；The protein 2 (PAB4 albumen) is the protein that the amino acid sequence shown in sequence 6 in sequence table forms.
In the present invention, all of above a2) in the breeding salt tolerance improve plant variety method, can specifically wrap
Include using the protein 1 (PAB2 albumen) and the lower plant of the protein 2 (PAB4 albumen) expression quantity as parent into
The step of row hybridization.
The present invention also provides a kind of methods of genetically modified plants that cultivation salt tolerance improves.
The method provided by the present invention for cultivating the genetically modified plants that salt tolerance improves, specifically may include following steps:
Inhibition expression is carried out to the encoding gene of the encoding gene of protein 1 and protein 2 in recipient plant, obtains transgenosis plant
Object；Genetically modified plants salt tolerance compared with the recipient plant enhances；
The protein 1 is PAB2 albumen；The protein 2 is PAB4 albumen.
Further, protein 1 (PAB2 albumen) amino acid sequence shown in sequence 3 in sequence table forms
Protein；The protein 2 (PAB4 albumen) is the protein that the amino acid sequence shown in sequence 6 in sequence table forms.
It is wherein, described that " encoding gene of encoding gene and protein 2 in recipient plant to protein 1 presses down
Tabulation reaches " it can be any coding for being able to suppress the encoding gene of protein 1 and the protein 2 described in the recipient plant
The method of gene expression.It in the present invention, is from arabidopsis biological study center [Arabidopsis Biological
Resource Center (ABRC), http://www.arabidopsis.org/] obtain PAB2 and PAB4 Gene Double knock out
Mutant, the double-mutant enhance NaCl tolerance.It is of course also possible to be obtained by genetic engineering means (RNAi technology)
PAB2 and PAB4 low expression, the genetically modified plants that NaCl tolerance is enhanced.
In above-mentioned application or method, the encoding gene (i.e. PAB2 gene) of the protein 1 is following 1) to 5) in appoint
DNA molecular described in one:
1) coded sequence is the DNA molecular shown in the nucleotide of 5 ' end the 346th to 2235 of sequence 2 in sequence table；
2) DNA molecular shown in sequence 2 in sequence table；
3) DNA molecular shown in sequence 1 in sequence table；
4) under strict conditions with 1) -3) it is any defined by DNA molecular hybridize and encode as shown in sequence 3 in sequence table
Amino acid sequence composition protein DNA molecular；
5) with 1) -4) DNA molecular of any restriction has 90% or more homology and coding is as shown in sequence 3 in sequence table
Amino acid sequence composition protein DNA molecular.
In above-mentioned application or method, the encoding gene (i.e. PAB4 gene) of the protein 2 is following 6) to 10) in appoint
DNA molecular described in one:
6) coded sequence is the DNA molecular shown in the nucleotide of 5 ' end the 157th to 2145 of sequence 5 in sequence table；
7) DNA molecular shown in sequence 5 in sequence table；
8) DNA molecular shown in sequence 4 in sequence table；
9) under strict conditions with 6) -8) it is any defined by DNA molecular hybridize and encode as shown in sequence 6 in sequence table
Amino acid sequence composition protein DNA molecular；
10) with 6) -9) DNA molecular of any restriction has 90% or more homology and coding is by 6 institute of sequence in sequence table
The DNA molecular of the protein of the amino acid sequence composition shown.
Above-mentioned stringent condition can for 6 × SSC, the solution of 0.5%SDS hybridizes at 65 DEG C, then with 2 × SSC,
It is primary that 0.1%SDS and 1 × SSC, 0.1%SDS respectively wash film.
Wherein, sequence 1 is made of 3712 nucleotide, is PAB2 gene sequence in arabidopsis gene group, wherein
614-846,966-1152,1810-1898,1977-2051,2142-2408,2547-
2780,2952-3053,3204-3287 are intron sequences；Sequence 2 is made of 2441 nucleotide, for institute
The cDNA sequence of PAB2 gene is stated, wherein 346-2235 are coded sequence (ORF)；Sequence 1 and the equal polynucleotide of sequence 2
Protein shown in middle sequence 3, sequence 3 are made of 629 amino acid residues.
Sequence 4 is made of 3217 nucleotide, is PAB4 gene sequence in arabidopsis gene group, wherein 455-
666,786-869,1527-1623,1702-1794,1885-1965,2101-2201,
2370-2452,2636-2728 are intron sequences；Sequence 5 is made of 2373 nucleotide, is the PAB4 base
The cDNA sequence of cause, wherein 157-2145 are coded sequence (ORF)；Sequence 6 in sequence 4 and the equal polynucleotide of sequence 5
Shown in protein, sequence 6 is made of 662 amino acid residues.
In above-mentioned application or method, it can also be monocotyledon that the plant, which can be dicotyledon,.
In one embodiment of the invention, the plant is dicotyledon, is further crucifer, specifically
It is more specific for arabidopsis wild type for arabidopsis (Col-0 is environmental).
The research of the invention finds that PAB2 the and PAB4 Gene Double obtained from arabidopsis biological study center (ABRC) is knocked out and is dashed forward
Variant enhances NaCl tolerance；Therefore, plant is adjusted to the tolerance of NaCl using PAB2 and PAB4.In addition, can also lead to
It crosses genetic engineering means (RNAi technology) and obtains PAB2 and PAB4 low expression, salt tolerant genetically modified crops.The present invention meets sustainable
Agricultural development demand cultivates the side such as Efficient salt-tolerant new varieties for research Plant Tolerance NaCl molecular mechanism, improvement hereditary capacity
Face has important practical value and market prospects.
Detailed description of the invention
Fig. 1 is the identification that PAB2 and PAB4 gene T-DNA is inserted into double-mutant pab2-1pab4-1.WT represents quasi- south in figure
Mustard wild type, i.e. Col-0 are environmental；2/4 represents pab2-1pab4-1 double-mutant in figure.It can be seen from the figure that pab2-
1pab4-1 double-mutant plant is homozygote plant.
Fig. 2 is to detect PAB2 gene expression in PAB2 gene T-DNA insertion mutation body pab2-1 with real-time fluorescence quantitative PCR
The analysis result of amount.It can be seen from the figure that pab2-1 mutant is PAB2 knock out mutants body.
Fig. 3 is to detect PAB4 gene expression in PAB4 gene T-DNA insertion mutation body pab4-1 with real-time fluorescence quantitative PCR
The analysis result of amount.It can be seen from the figure that pab4-1 mutant is PAB4 knock out mutants body.
Fig. 4 is to detect PAB2 and PAB4 gene expression amount in pab2-1pab4-1 double-mutant with real-time fluorescence quantitative PCR
Analysis result.WT represents arabidopsis wild type in figure, i.e. Col-0 is environmental；It is bis- to represent pab2-1pab4-1 by pab2/4 in figure
Mutant.It can be seen from the figure that pab2-1pab4-1 double-mutant is PAB2 and PAB4 Gene Double knockout mutations body.
Fig. 5 is impact analysis result of the NaCl to pab2-1pab4-1 double-mutant growth of seedling.Pab2/4 is represented in figure
Pab2-1pab4-1 double-mutant.
Specific embodiment
Experimental method used in following embodiments is conventional method unless otherwise specified.
The materials, reagents and the like used in the following examples is commercially available unless otherwise specified.
Arabidopsis wild type (Col-0 is environmental): arabidopsis wild type seeds (Arabidopsis thaliana,
Ecotype Columbia-0), it is arabidopsis biological study center [Arabidopsis Biological Resource
Center (ABRC), http://www.arabidopsis.org/] product.
PAB2 and PAB4 Gene Double knocks out T-DNA mutant pab2-1pab4-1, is arabidopsis biological study center
(ABRC) product, seed number are CS8177, and background is arabidopsis wild type (Col-0 is environmental)；Wherein, pab2-1 single mutation kind
Son number is SALK_026293, and pab4-1 single mutation seed number is SALK_113383.The bis- prominent body specifying informations of pab2-1pab4-1
(Dufresne, P.J., Ubalijoro, E., Fortin, M.G., and are also had a detailed description in the article delivered
Laliberte,J.F.(2008).Arabidopsis thaliana class II poly(A)-binding proteins
are required for efficient multiplication of turnip mosaic virus.Journal of
General Virology 89,2339-2348.)。
Embodiment 1, pab2-1 mutant, pab4-1 mutant and the identification of pab2-1pab4-1 double-mutant and expression quantity point
Analysis
One, the identification of pab2-1 mutant, pab4-1 mutant and pab2-1pab4-1 double-mutant
According to the primer combination in Fig. 1, the bis- mutant homozygous body plant of pab2-1pab4-1 are identified by round pcr；Separately
Outside, identical method identifies pab2-1 mutant and pab4-1 mutant homozygote plant.
The primer sequence is as follows:
Pab2-1LP:5 '-ATTCGAAAGTGTCAAACACGC-3 ' (Left primer, LP)
Pab2-1RP:5 '-TAATAAAAATGTTGCCAGCGC-3 ' (Right primer, RP)
Pab4-1LP:5 '-GAGAAGGCAATGCAGAAGTTG-3 ' (Left primer, LP)
Pab4-1RP:5 '-ACTACCTGCAGCTTAGCCCTC-3 ' (Right primer, RP)
LBb1:5 '-GCGTGGACCGCTTGCTGCAACT-3 ' (Left border primer, LB)
Two, gene expression amount analysis is corresponded in pab2-1 mutant, pab4-1 mutant and the bis- mutation of pab2-1pab4-1
Extract arabidopsis wild type (Col-0 is environmental), pab2-1 mutant, pab4-1 mutant and pab2-
1pab4-1 double-mutant strain total serum IgE detects PAB2 or/and PAB4 gene in each experimental material using real-time fluorescence quantitative PCR
Expression at transcriptional level.It is specific as follows:
With pab2-1 mutant, pab4-1 mutant, pab2-1pab4-1 double-mutant and arabidopsis wild type (Col-
0 is environmental) 12 days seedling are grown as experimental material in plate.The total serum IgE of each experimental material is extracted, reverse transcription is at single-stranded
Then cDNA analyzes expression feelings of the PAB2 or/and PAB4 gene in each experimental material by real time fluorescence quantifying PCR method
Condition.
Wherein, the primer sequence of PAB2 gene is expanded are as follows:
PAB2RT-F:5 '-CAGGTTCAACTTCAGGGTCA-3 ' (352-371 of sequence 2)；
PAB2RT-R:5 '-CTGCGAGTCCGTCACATT-3 ' (481-498 reverse complementary sequences of sequence 2).
Expand the primer sequence of PAB4 gene are as follows:
PAB4RT-F:5 '-AGGTTATGCGGGACCCTAGT-3 ' (1229-1248 of sequence 5)；
PAB4RT-R:5 '-GTGGAGGTCCTTGACCGTAA-3 ' (1497-1516 reverse complemental sequences of sequence 5
Column).
Using Actin2/8 as reference gene, the primer sequence of internal reference Actin is expanded are as follows:
Actin-F:5 '-GGTAACATTGTGCTCAGTGGTGG-3 '；
Actin-R:5 '-AACGACCTTAATCTTCATGCTGC-3 '.
The reaction condition of above-mentioned primer is as follows:
(1) foundation of reaction system
Real-time fluorescence quantitative PCR reaction system
(2) three repetitions, gently get rid of mixing, are tested with Bio-Rad CFX96 fluorescence quantitative PCR instrument.
(3) setting of response procedures:
Real-time fluorescence quantitative PCR response procedures
(4) numerical analysis, with 2-ΔCtAs the relative difference for measuring gene transcription level, the expression of each gene is carried out
Analysis is compared.Ct value is that PCR reacts recurring number when fluorescence signal reaches given threshold, Δ Ct value be special primer Ct value with
The difference of Actin primer Ct value.
Real-time fluorescence quantitative PCR testing result is shown as shown in Figure 2, Figure 3 and Figure 4, and the expression of PAB2 and PAB4 gene is phase
To value, 100 are expressed as with PAB2 and PAB4 gene in arabidopsis wild type (Col-0).It can be seen from the figure that pab2-1 is prominent
PAB2 gene is not expressed on transcriptional level in variant, is PAB2 knock out mutants body (Fig. 2)；In pab4-1 mutant
PAB4 gene is not expressed on transcriptional level, is PAB4 knock out mutants body (Fig. 3)；In pab2-1pab4-1 double-mutant
PAB2 and PAB4 gene is not expressed on transcriptional level, is PAB2 and PAB4 Gene Double knockout mutations body (Fig. 4).
Embodiment 2, pab2-1pab4-1 double-mutant are analyzed NaCl tolerance and are tested
NaCl is able to suppress the growth of plant seedlings.With the raising of external source NaCl concentration, arabidopsis wild type (Col-0
It is environmental) by being inhibited to be enhanced, the growth of main root and blade can be all influenced to different extents, and experiment repeats 3
It is secondary.
Using arabidopsis wild type (Col-0 is environmental) and pab2-1pab4-1 double-mutant plant as experimental material.It will be each
The seed sowing of experimental material is on the MS plate without NaCl, and low-temperature stratification is put into illumination box after 3 days and cultivates at 4 DEG C
Then each experimental material seedling just sprouted is moved on to (0m Μ, 50m on the MS culture medium containing various concentration NaCl by 72h
Μ, 100m Μ, 150m Μ and 200m Μ), vertical growth observes Arabidopsis thaliana Seedlings growing state and takes pictures after 9 days.
As a result as shown in figure 5, on the culture medium of the NaCl containing 0mM, each genetic stocks seedling growth (leaf growth and master
Root long degree) it is almost the same；But on the MS culture medium containing various concentration NaCl, wild type seedlings growth receives obvious
Inhibition, and pab2-1pab4-1 double-mutant seedling relative to arabidopsis wild type (Col-0 environmental) to NaCl sensibility
Lower, growing way is significantly better than wild type, shows the phenotype of salt tolerant, i.e. pab2-1pab4-1 double-mutant increases NaCl tolerance
By force.
In addition, the present inventor has detected pab2-1 mutant and pab4-1 mutant pair using identical method
The tolerance of NaCl, as a result, it has been found that, no matter in the culture medium without NaCl or in the MS culture medium containing various concentration NaCl
On, pab2-1 mutant and pab4-1 mutant and arabidopsis wild type (Col-0 is environmental) seedling growth are almost the same.
Claims (8)
1. protein 1 and protein 2 as target it is following it is any in application:
A1 the salt tolerance of plant) is improved；
A2) the plant variety that breeding salt tolerance improves；
The protein 1 is the protein that the amino acid sequence shown in sequence 3 in sequence table forms；The protein 2 is served as reasons
The protein that amino acid sequence shown in sequence 6 forms in sequence table；
The plant is arabidopsis.
2. be able to suppress protein 1 and protein 2 are expressed in plant substance it is following it is any in application:
A1 the salt tolerance of plant) is improved；
A2) the plant variety that breeding salt tolerance improves；
The protein 1 is the protein that the amino acid sequence shown in sequence 3 in sequence table forms；The protein 2 is served as reasons
The protein that amino acid sequence shown in sequence 6 forms in sequence table；
The plant is arabidopsis.
3. being able to suppress the substance of the encoding gene of the encoding gene of protein 1 and protein 2 expression in plant following any
In application:
A1 the salt tolerance of plant) is improved；
A2) the plant variety that breeding salt tolerance improves；
The protein 1 is the protein that the amino acid sequence shown in sequence 3 in sequence table forms；The protein 2 is served as reasons
The protein that amino acid sequence shown in sequence 6 forms in sequence table；
The plant is arabidopsis.
4. application according to claim 1 to 3, it is characterised in that: the encoding gene of the protein 1 is as follows
1) to the DNA molecular any in 3):
1) coded sequence is the DNA molecular shown in the nucleotide of 5 ' end the 346th to 2235 of sequence 2 in sequence table；
2) DNA molecular shown in sequence 2 in sequence table；
3) DNA molecular shown in sequence 1 in sequence table.
5. application according to claim 1 to 3, it is characterised in that: the encoding gene of the protein 2 is as follows
4) to the DNA molecular any in 6):
4) coded sequence is the DNA molecular shown in the nucleotide of 5 ' end the 157th to 2145 of sequence 5 in sequence table；
5) DNA molecular shown in sequence 5 in sequence table；
6) DNA molecular shown in sequence 4 in sequence table.
6. a kind of method for cultivating the genetically modified plants that salt tolerance improves, includes the following steps: in recipient plant to protein 1
Encoding gene and the encoding gene of protein 2 carry out inhibition expression, obtain genetically modified plants；The genetically modified plants and institute
State recipient plant enhances compared to salt tolerance；
The protein 1 is the protein that the amino acid sequence shown in sequence 3 in sequence table forms；The protein 2 is served as reasons
The protein that amino acid sequence shown in sequence 6 forms in sequence table；
The plant is arabidopsis.
7. according to the method described in claim 6, it is characterized by: the encoding gene of the protein 1 is following 1) in 3)
Any DNA molecular:
1) coded sequence is the DNA molecular shown in the nucleotide of 5 ' end the 346th to 2235 of sequence 2 in sequence table；
2) DNA molecular shown in sequence 2 in sequence table；
3) DNA molecular shown in sequence 1 in sequence table.
8. according to the method described in claim 6, it is characterized by: the encoding gene of the protein 2 is following 4) in 6)
Any DNA molecular:
4) coded sequence is the DNA molecular shown in the nucleotide of 5 ' end the 157th to 2145 of sequence 5 in sequence table；
5) DNA molecular shown in sequence 5 in sequence table；
6) DNA molecular shown in sequence 4 in sequence table.
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| WO2014112997A1 (en) * | 2013-01-16 | 2014-07-24 | 1,4 Group, Inc. | Treatment of crops with a lower alkyl naphthalene to alter cell cycle and water regulation |
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