CN113025621B - Application of CIPK14 gene in improving drought resistance of pigeon pea - Google Patents
Application of CIPK14 gene in improving drought resistance of pigeon pea Download PDFInfo
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Abstract
The invention provides application of CIPK14 genes in improving drought resistance of pigeon pea, and relates to the technical fields of genetic engineering and genetic breeding. The CIPK14 gene improves drought resistance of pigeon pea and growth and survival rate of pigeon pea under drought conditions by promoting accumulation of xyloside in pigeon pea.
Description
Technical Field
The invention relates to the technical fields of genetic engineering and genetic breeding, in particular to application of CIPK14 genes in improving drought resistance of pigeon pea.
Background
Drought has high occurrence frequency, long duration and wide influence range in the world, and is one of important reasons for restricting the development of economic forests. Under drought stress, a series of stress reactions can occur in the forest to adapt to the environment, and when the stress reactions are excessive, the forest is damaged to a certain extent: drought stress can alter the structural permeability of cell membranes, damage some tissues, affect metabolic processes, and the like. The method can improve the drought resistance of the economic forest, and the obtained new variety with good drought resistance has important significance for improving the yield of the economic forest, guaranteeing the quality of the economic forest, saving manpower and material resources and reducing loss. Compared with traditional breeding, the molecular breeding efficiency is high, the hybridization and screening processes of multiple generations are not needed, and the molecular breeding has strong directionality. Through a gene modification technology, a certain or a certain specific genes are edited or expressed and inherited stably, so that the stress resistance of the forest is improved, and the method is one of means for obtaining a new variety of the forest more efficiently. In addition, root transgene over-expression and gene editing technology can also be used for exploring different structures and functions of different genes acting on organisms, and provides theoretical basis for better using gene resources for cultivating new varieties.
The pigeon pea is perennial woody, grows in the edge environment with low investment and easy risk, and is an adaptive economic forest with dual purposes of medicine and food: the leaves contain secondary metabolites with important medicinal value, and the seeds are edible. Therefore, pigeon pea has become an ideal crop for sustainable development of agriculture in karst areas in south China. Although the pigeon pea has good stress resistance, long-term drought is still a main factor for limiting the yield of the pigeon pea, so that the gene for regulating and controlling drought resistance of the pigeon pea is screened by a genetic engineering means, the pigeon pea variety is improved, the drought resistance is rapidly improved, and the method has important significance for cultivation of the pigeon pea.
Disclosure of Invention
In view of the above, the invention aims to provide application of CIPK14 genes in improving drought resistance of pigeon pea, wherein the CIPK14 genes can improve drought resistance of pigeon pea.
In order to achieve the above object, the present invention provides the following technical solutions:
the invention provides application of CIPK14 genes in improving drought resistance of pigeon pea.
The invention also provides application of the CIPK14 gene in promoting growth of pigeon pea under drought conditions.
The invention also provides application of the CIPK14 gene in improving survival rate of pigeon pea under drought conditions.
The invention also provides application of the CIPK14 gene in promoting the accumulation of xyloside of pigeon pea.
Preferably, the method comprises the following steps:
1) Connecting cDNA of CIPK14 gene into a vector to obtain an expression vector;
2) Transforming the expression vector obtained in the step 1) into competent agrobacterium to obtain transformed bacteria;
3) And (3) infecting pigeon pea with the transformed bacteria obtained in the step (2).
Preferably, the vector of step 1) includes a pROK2 vector.
Preferably, the cDNA of the CIPK14 gene and the vector in the step 1) are subjected to double digestion by XbaI and KpnI and then are connected by using T4 ligase.
Preferably, the step 2) agrobacterium comprises agrobacterium K599.
Preferably, the expression vector is transformed into competent Agrobacterium by heat shock.
Preferably, the transformed bacteria in the step 3) infect the position 1cm above the pigeon pea basal part.
The invention provides application of a CIPK14 gene in improving the drought resistance of pigeon pea, wherein the CIPK14 gene improves the drought resistance of pigeon pea and the growth and survival rate of pigeon pea under drought conditions by promoting the accumulation of xyloside in pigeon pea.
Drawings
FIG. 1-A is an identification of all CIPK genes in Cajanus cajan; FIG. 1-B shows the variation of CIPK gene expression level in roots under drought stress, and shows that 28 CIPK genes are totally contained in pigeon pea, and the CIPK14 expression level is up-regulated most in the roots under drought stress;
FIG. 2-A is a graph showing the interaction of CIPK14 with CBL1 yeast; FIG. 2-B is a graph showing the fluorescent complementary interaction of CIPK14 and CBL 1;
FIG. 3 shows the measurement of the expression level of CIPK14 gene in the root of Cajanus cajan which is overexpressed by CIPK14 in example 2; wherein CK represents control pigeon pea, CIPK14-OE represents CIPK14 gene root overexpression pigeon pea, and A is transgenic root and original root phenotype; b is the expression quantity detection of CIPK14 genes in the control pigeon pea roots and the root transgenic pigeon pea roots;
FIG. 4 shows plant growth of CIPK14 root overexpressing pigeon pea after drought treatment in example 3; wherein CK represents control pigeon pea, and CIPK14-OE represents CIPK14 gene root overexpression pigeon pea;
FIG. 5 shows survival of CIPK14 root overexpressing pigeon pea after drought treatment in example 4; wherein CK represents control pigeon pea, and CIPK14-OE represents CIPK14 gene root overexpression pigeon pea;
FIG. 6 is a schematic diagram of the mechanism of enhancing drought resistance by promoting the accumulation of genistin, a flavonoid, by CIPK14-CBL1 protein complex of the invention (Agilent HPLC was used for detecting metabolites in transgenic and control roots).
Detailed Description
The invention provides application of CIPK14 genes in improving drought resistance of pigeon pea.
The invention also provides application of the CIPK14 gene in promoting growth of pigeon pea under drought conditions.
The invention also provides application of the CIPK14 gene in improving survival rate of pigeon pea under drought conditions.
The invention also provides application of the CIPK14 gene in promoting the accumulation of xyloside of pigeon pea.
In the present invention, the CIPK14 gene is numbered xm_020370625.1 in the pigeon pea genome database.
In the present invention, the application preferably includes the steps of:
1) Connecting cDNA of CIPK14 gene into a vector to obtain an expression vector;
2) Transforming the expression vector obtained in the step 1) into competent agrobacterium to obtain transformed bacteria;
3) And (3) infecting pigeon pea with the transformed bacteria obtained in the step (2).
The invention connects cDNA of CIPK14 gene into carrier to obtain expression carrier.
In the present invention, the CIPK14 gene is preferably obtained by: extracting total RNA of pigeon pea, carrying out reverse transcription to obtain cDNA, and amplifying by using the cDNA as a template and using a primer pair to obtain CIPK14 genes. The conditions for the amplification are not particularly limited, and those skilled in the art can perform the conventional procedures. In the present invention, the primer sequences upstream and downstream of the primer pair are as follows:
upstream primer (SEQ ID No. 1): ATGAATGGGCAAAAAATTAAGCGCC;
downstream primer (SEQ ID No. 2): CAGGAAACCAATACCAAAATCATAT.
In the present invention, the vector preferably includes a pROK2 vector. In the present invention, the cDNA of the CIPK14 gene and the vector are preferably ligated by using T4 ligase after double digestion with XbaI and KpnI. The conditions for double cleavage and ligation are not particularly limited, and those skilled in the art can perform the conventional operation.
The invention transforms the obtained expression vector into competent agrobacterium to obtain transformed bacteria.
In the present invention, the agrobacterium preferably includes agrobacterium K599. In the present invention, the expression vector is preferably transformed into competent Agrobacterium by heat shock.
The obtained transformed bacteria infest pigeon pea. In the present invention, the transformant is preferably infected 1cm above the pigeon pea basal part.
The invention is described in detail below in connection with examples for further illustration of the invention, but they are not to be construed as limiting the scope of the invention.
Materials, reagents, and the like used in the following examples, unless otherwise specified, were commercially available, and the main reagents include: restriction enzymes, DNA polymerase, T4 ligase, reverse transcription kits of Thermo biosystems; an RNA extraction kit from Promega corporation; quantitative PCR reagent of Takara company; plasmid extraction kits and DNA recovery kits were purchased from nuuzan company; the MS culture medium, agar powder, agarose, ampicillin, kanamycin, gentamicin sulfate, rifampicin and other antibiotics are purchased from sigma; the various other chemical reagents used in the examples are imported or homemade analytically pure reagents; primer synthesis and sequencing was accomplished by Zhongmeitai and company.
Example 1
The pigeon pea is planted in the pot, 150g of soil is planted in each pot, and the seedling age is 20 days. The control group was watered with 30ml of water daily, the experimental group was watered with 30ml of water only on the first day, and then water was stopped. The roots of the experimental group and the control group were taken at days 7 and 14, respectively, and after marking, they were quick frozen with liquid nitrogen and stored in a-80℃refrigerator. Then extracting RNA of the root by using an RNA extraction kit of Promega company; detecting the quality of the RNA by using Thermo Nanodrop and agarose gel electrophoresis; the RNA was then reverse transcribed using a Takara kit and quantitatively detected by fluorescence using a Takara SYBR kit, the instrument being Bio-Rad CFX connector. The results are shown in FIG. 1-A and FIG. 1-B, and FIG. 1-A is the identification of all CIPK genes in pigeon pea; FIG. 1-B shows the variation of CIPK gene expression level in roots under drought stress, and shows that 28 CIPK genes are in total in pigeon pea, and the CIPK14 expression level is up-regulated most in the roots under drought stress. Wherein, fig. 1-a: 26 CIPK protein sequences in Arabidopsis were used as the search for the pigeon pea genome using the BLASTP method of the NCBI website, website https:// www.ncbi.nlm.nih.gov, followed by domain analysis using NCBI-CD search, yielding a total of 28 CcCIPK proteins. Then, by taking soybean and Arabidopsis CIPK proteins as references, carrying out phylogenetic analysis by using MEGA6, wherein a phylogenetic tree is an NJ method, and 1000 times of bootstrap and p-distance methods are adopted. Fig. 1-B: planting pigeon pea with seedling age of 20 days in 150g of soil of each pot, watering 30ml of a control group every day, stopping watering after watering the drought group for the first day, and respectively taking roots of the control group and the drought group on the 7 th and 14 th days of drought, and extracting RNA of the roots by using an RNA extraction kit of Promega company; detecting the quality of the RNA by using Thermo Nanodrop and agarose gel electrophoresis; the RNA was then reverse transcribed using a Takara kit and quantitatively detected by fluorescence using a Takara SYBR kit, the instrument being Bio-Rad CFX connector.
Fig. 2-a: CIPK14 and CBL1 are respectively constructed into pGBKT7 and pGADT7 vectors by a double enzyme digestion method, then are transformed into Y2HGold yeast cells together, are coated on SD/-Leu-Trp and SD/-Leu-Trp-His-Ade flat plate culture mediums, and observe the growth condition. Fig. 2-B: CIPK14 and CBL1 were constructed into YFPc and YFPn vectors, respectively, by double digestion, and then transformed into GV3101 Agrobacterium competent cells, respectively, and the plasmid-transformed Agrobacterium was cultured overnight at 28℃in YEP+50mg/L Kana+25mg/L Rif medium, 150 rpm. After overnight incubation, the cells were collected by centrifugation at 5000rpm for 10min and suspended with BIFC suspension. The two agrobacteria are mixed according to the proportion of 1:1, the tobacco leaves are injected together, and fluorescence is observed under a Leica SP8 laser confocal microscope after three days of culture.
Construction and detection of CIPK14 gene overexpression vector
Extracting total RNA from pigeon pea, carrying out reverse transcription to obtain cDNA, using the cDNA as a template, using F and R as primers, amplifying CIPK14 genes, wherein the primers are provided with enzyme cutting sites, and connecting the primers to an over-expression vector after enzyme cutting. The construction method of the CIPK14 gene overexpression vector comprises the following steps:
(1) Total RNA was extracted from Cajanus cajan using the RNA extraction kit from Promega corporation, and the specific procedure was as described in the kit.
(2) Reverse transcription of RNA to cDNA was performed using a reverse transcription kit from Thermo company, and the specific procedure was referred to the kit instructions.
(3) Using cDNA as a template, F and R as primers, amplifying cDNA of CIPK14 genes, and recovering amplified products by electrophoresis cutting gel, wherein the recovery method refers to a kit of Noruzan company;
the primer sequences used for CIPK14 gene amplification are as follows:
SEQ ID No.1: upstream primer F: ATGAATGGGCAAAAAATTAAGCGCC;
SEQ ID No.2: the downstream primer R: CAGGAAACCAATACCAAAATCATAT.
(4) The cDNA of the recovered CIPK14 gene and pROK2 vector are digested with XbaI and KpnI, the digested product is recovered by running electrophoresis, the recovered product is connected with T4 ligase, and the CIPK14 gene is connected to the vector to drive the over-expression of the CIPK14 gene by a 35S promoter.
(5) Taking 5 μl of the product of the cleavage-ligation system of step (4), and transforming E.coli competence. Screening was performed on LB plates containing 50. Mu.g/mL kanamycin. Colony PCR identifies single clone, and positive clone is selected for sequencing. The obtained recombinant expression vector with correct sequencing was named pROK2-CIPK14. Colony PCR and sequencing universal primers were as follows:
SEQ ID NO.3: upstream primer F: GGATTGATGTGATATCTCCACTGACGTAA;
SEQ ID NO.4: the downstream primer R: CGATTAAGTTGGGTAACGCC.
Example 2
Construction and detection of CIPK14 gene root overexpression pigeon pea
Will be solidThe pROK2-CIPK14 overexpressing plasmid constructed in example 1 was transformed into competent Agrobacterium K599 strain by heat shock and colony PCR identified positive clones. Inoculating single colony of correctly identified agrobacterium in 2-3 mL liquid culture medium containing 100 μg/mL kanamycin and 50 μg/mL rifampicin, shake culturing overnight at 28 ℃, transferring a large amount of liquid culture medium containing antibiotics for the next day, shake culturing, collecting thallus after transferring several times, and re-suspending to OD 600 Between 0.8 and 1.0. The resuspended bacterial liquid was aspirated by a syringe, and injection was performed at a position 1cm above the pigeon pea basal portion, and transgenic roots were grown around the injection port after about 30 days. And (3) extracting RNA of the transgenic root, reversely transcribing cDNA, and quantitatively detecting the over-expression condition of the root transgenic plant by PCR, wherein the expression quantity of CIPK14 genes in the root transgenic plant is far higher than that of a control plant, CK is 1, and CIPK14-OE is 4.3.
Example 3
CIPK14 gene root overexpression pigeon pea drought treatment phenotype detection
150g of soil was added to each pot, water was added to the tray, 1 plant of root transgenic pigeon pea of example 2 was placed in each pot, after the water was absorbed, the water remaining in the tray was poured out, drought treatment was started, and control (control was transformed pROK2 empty vector root pigeon pea) and root transgenic plant drought treatment phenotype were observed. Control and root transgenic plants were 5 pots each, as biological replicates. As shown in FIG. 4, the growth condition of the pigeon pea in which the CIPK14 gene is overexpressed in roots under drought stress is obviously better than that of the control pigeon pea. The root transgenic pigeon pea with the CIPK14 gene over-expressed has stronger drought resistance than the control pigeon pea.
Example 4
CIPK14 gene root overexpression pigeon pea drought treatment survival rate detection
The drought treatment method was as described in example 3. After 14 days of drought stress treatment, adding water into the tray to enable each small basin to fully absorb water, and observing survival rate of the pigeon pea after rehydration, wherein the results are shown in figure 5, CK is 7.059%,8.202% and 8.739 respectively; CIPK14-OE was 20.252,%,23.215%,28.533%, respectively.
The foregoing is merely a preferred embodiment of the present invention and is not intended to limit the present invention in any way. It should be noted that modifications and adaptations to the present invention may occur to one skilled in the art without departing from the principles of the present invention and are intended to be comprehended within the scope of the present invention.
Sequence listing
<110> Beijing university of forestry
Application of <120> CIPK14 gene in improving drought resistance of pigeon pea
<160> 4
<170> SIPOSequenceListing 1.0
<210> 1
<211> 25
<212> DNA
<213> Artificial sequence (Artificial Sequence)
<400> 1
atgaatgggc aaaaaattaa gcgcc 25
<210> 2
<211> 25
<212> DNA
<213> Artificial sequence (Artificial Sequence)
<400> 2
caggaaacca ataccaaaat catat 25
<210> 3
<211> 29
<212> DNA
<213> Artificial sequence (Artificial Sequence)
<400> 3
ggattgatgt gatatctcca ctgacgtaa 29
<210> 4
<211> 20
<212> DNA
<213> Artificial sequence (Artificial Sequence)
<400> 4
cgattaagtt gggtaacgcc 20
Claims (7)
- Application of CIPK14 gene in promoting growth of pigeon pea under drought condition and improving survival rate of pigeon pea under drought condition;the CIPK14 gene is derived from pigeon pea;the CIPK14 gene is numbered XM_020370625.1 in the pigeon pea genome database.
- 2. The use according to claim 1, characterized by the steps of:1) Connecting cDNA of CIPK14 gene into a vector to obtain an expression vector;2) Transforming the expression vector obtained in the step 1) into competent agrobacterium to obtain transformed bacteria;3) And (3) infecting pigeon pea with the transformed bacteria obtained in the step (2).
- 3. The use according to claim 2, wherein the step 1) vector comprises a pROK2 vector.
- 4. The use according to claim 2, wherein the step 1) is characterized in that the cDNA of CIPK14 gene and the vector are ligated by using T4 ligase after double digestion with XbaI and KpnI.
- 5. The use according to claim 2, wherein said step 2) agrobacterium comprises agrobacterium K599.
- 6. The use according to claim 2, wherein the expression vector is transformed into competent agrobacterium by heat shock.
- 7. The use according to claim 2, wherein the transformed bacteria of step 3) infest 1cm above the base of pigeon pea.
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| 李莉 ; 李毅 ; 王长春 ; 王锋青 ; 杨玲 ; .番茄LeCIPK3的克隆及非生物胁迫诱导的表达分析.植物生理学通讯.2010,第46卷(第7期),第659-663页. * |
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