CN112553242B - Application of promoter GmLCLb2 in regulating and controlling near-day rhythmic expression level of gene in response to environmental light quality change - Google Patents
Application of promoter GmLCLb2 in regulating and controlling near-day rhythmic expression level of gene in response to environmental light quality change Download PDFInfo
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Abstract
The invention relates to the technical field of plant genetic engineering, in particular to application of a promoter GmLCLb2 in regulating and controlling the near-day rhythmicity expression level of genes in response to environmental light quality change. The invention provides a soybean promoter GmLClb2 capable of responding to red light and blue light qualities to change and regulate the near-day rhythmicity expression level of genes, wherein the activity of the promoter responds to ambient red light and blue light signals, can drive target genes to differentially express under red light and blue light conditions, enables the expression peak value of the target genes to appear in the early morning, and has high application value in plant gene expression regulation and transgenic plant construction and breeding with excellent properties.
Description
Technical Field
The invention relates to the technical field of plant genetic engineering, in particular to application of a promoter GmLCLb2 in regulating and controlling the near-day rhythmicity expression level of genes responding to environmental light quality change.
Background
In order to adapt to the environment, the living bodies on the earth generate a biological clock regulation mechanism in the evolution process. Plant biological clocks have the following basic characteristics: the biological clock is endogenously generated by an organism and can be autonomously maintained, and the nearly 24-hour periodic rhythm can be still maintained under the constant condition of an external environment; the biological clock has a temperature compensation mechanism, and the cycle length of the biological clock can not be obviously changed along with the temperature change in a certain temperature range. The plant biological clock core oscillator is composed of a plurality of transcription factors and transcription accessory factors.
The light is the most important environmental signal of the plant, the light intensity, the light quality and the light cycle all regulate and control the growth and development of the plant, and the red light, the blue light and the far-red light play an important role in the growth and development of the plant. The red light inhibits the elongation of plant stems, promotes branching and tillering, and enhances the accumulation of anthocyanin chlorophyll and carotenoid. Blue light inhibits the extension area of leaves and the growth rate of plants, enhances the carbon-nitrogen ratio, and regulates and controls chlorophyll synthesis, the ratio of chlorophyll a to chlorophyll b and chloroplast formation. The development of a promoter capable of regulating the near-day rhythmicity expression of a gene in response to light quality change has important significance for plant gene expression regulation and near-day rhythmicity regulation.
Disclosure of Invention
The invention aims to provide a soybean promoter capable of regulating and controlling the expression level of the gene in the near-day rhythmicity in response to the change of the light quality of the environment.
In order to achieve the purpose, the invention carries out a large amount of screening on the upstream sequence (promoter region) of the gene which is involved in the biological clock regulation and presents the rhythmic expression in the soybean. The invention discovers that the upstream sequence of the GmLCLb2 gene can respond to the light quality change of red light and blue light and regulate the near-day rhythmicity expression level of the gene. The invention also finds that the sequence fragments with different lengths at the upstream of the GmLCLb2 gene have obviously different regulation and control effects on red light and blue light response and rhythmic expression level, wherein the activity of the promoter shown as SEQ ID NO.1 can respond to ambient red light and blue light signals, the target gene is efficiently driven to be differentially expressed under red light and blue light conditions, the expression level under the blue light condition is obviously higher than that under the red light condition, and the expression peak value of the target gene appears in the early morning.
Specifically, the invention provides the following technical scheme:
in a first aspect, the invention provides application of a promoter GmLCLb2 or an expression cassette, a vector or a microorganism containing the promoter GmLCLb2 in regulating and controlling differential expression of genes in plants in response to environmental light quality change.
The expression level of the GmLCLb2 promoter driving gene under the blue light condition is obviously higher than that under the red light condition.
Preferably, the application is to up-regulate the expression level of the gene by giving a blue light irradiation treatment, or to down-regulate the expression level of the gene by giving a red light irradiation treatment.
In a second aspect, the invention provides application of a promoter GmLCLb2 or an expression cassette, a vector or a microorganism containing the promoter GmLCLb2 in regulating the expression level of a near-day rhythmicity in response to environmental light quality change in a plant.
Specifically, the application is to up-regulate the near-day rhythmic expression level of the gene by giving blue light irradiation treatment, or to down-regulate the near-day rhythmic expression level of the gene by giving red light irradiation treatment.
In a third aspect, the present invention provides the use of the promoter GmLCLb2 for driving the near-day rhythmic expression of a target gene in plants under constant-temperature, continuous red light conditions, with the peak of expression in the early morning.
In a fourth aspect, the present invention provides the use of the promoter GmLCLb2 for driving the near-day rhythmic expression of a target gene in plants under constant-temperature, persistent blue-light conditions, with the peak of expression in the early morning.
The plant of the invention is preferably a dicotyledonous plant, more preferably a leguminous plant, most preferably a soybean.
The nucleotide sequence of the promoter GmLCLb2 is shown in SEQ ID NO. 1.
The expression cassette containing the promoter GmLCLb2 can be an expression unit obtained by operably connecting any target gene sequence at the downstream of the promoter GmLCLb 2.
The vector containing the promoter GmLCLb2 can be any vector known in the art, such as a cloning vector, an expression vector, an integration vector or a transposon.
The microorganism of the present invention includes, but is not limited to, Escherichia coli, Agrobacterium, and the like.
The gene of the invention can be a functional gene, an antisense gene of the functional gene or a small RNA gene capable of interfering the expression of the functional gene.
In a fifth aspect, the present invention provides a method for regulating differential expression of genes in plants in response to changes in the light qualities of ambient red light and blue light, the method comprising: the gene is operably connected to the downstream of a promoter GmLCLb2, and the promoter GmLCLb2 is used for driving the differential expression of the gene in response to the change of the light quality of the ambient red light and the blue light in plants.
Specifically, the method comprises the following steps: operably linking the gene downstream of a promoter GmLCLb2 to drive the expression of the gene by the promoter GmLCLb 2; an expression cassette or vector containing the promoter GmLCLb2 was introduced into plants.
In the above-described method, the plant is preferably a dicot, more preferably a leguminous plant, and most preferably soybean.
In the method, the nucleotide sequence of the promoter GmLCLb2 is shown as SEQ ID NO. 1.
The beneficial effects of the invention at least comprise: the invention provides a soybean promoter GmLCLb2 capable of responding to red light and blue light qualities to change and regulate the near-daily rhythmicity expression level of genes, wherein the activity of the promoter responds to ambient red light and blue light signals, can drive target genes to differentially express under red light and blue light conditions, enables the expression peak value of the target genes to appear in the early morning, and has higher application value in plant gene expression regulation and control and transgenic plant construction and breeding with excellent properties.
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FIG. 1 shows bioluminescence signal detection results of expression of LUC gene in soybean hairy roots in soybean transformed with pH2GW7 delta-GmLCLb 2, LUC in example 2 of the present invention; wherein, A is hair root field imaging; b is the result of bioluminescence imaging of hairy roots.
FIG. 2 is a graph showing the results of the analysis of the level of the node-regulated expression of LUC gene in soybean hairy roots in soybean transformed to pH2GW 7. delta. -GmLCLb2: LUC under Red and Blue light conditions in example 2 of the present invention, wherein Red represents the Red light condition and Blue represents the Blue light condition.
Detailed Description
Preferred embodiments of the present invention will be described in detail with reference to the following examples. It is to be understood that the following examples are given for illustrative purposes only and are not intended to limit the scope of the present invention. Various modifications and alterations of this invention will become apparent to those skilled in the art without departing from the spirit and scope of this invention.
The experimental procedures used in the following examples are all conventional procedures unless otherwise specified.
Materials, reagents and the like used in the following examples are commercially available unless otherwise specified.
Example 1 cloning of the Soybean GmLCLb2 promoter
Using forward primer 5'-cgggatcCATGTGTTATACAAGAGAAGTTGA ACCG-3' (SEQ ID NO.2) and reverse primer: 5'-ggggtaccGAGAAGCGAC GGAATTCATTTG-3' (SEQ ID NO.3) obtains a GmLCLb2 promoter with the length of 3172bp from soybean genome by PCR amplification, and the nucleotide sequence of the GmLCLb2 promoter is shown in SEQ ID NO.1 by sequencing verification. In order to facilitate the subsequent connection with the vector, the two ends of the PCR product are respectively provided with enzyme cutting sites of BamH I and Kpn I and protective basic groups.
The PCR amplification system (total volume 20. mu.l) was:
PCR amplification procedure: 5min at 95 ℃; 30s at 95 ℃, 30s at 55 ℃ and 3min at 72 ℃ for 28 cycles; 10min at 72 ℃.
Example 2 use of the soybean GmLCLb2 promoter to drive expression of the LUC Gene
The gel-cut recovered product of the GmLCLb2 promoter cloned by PCR in example 1 was digested with BamH I and Kpn I. Mixing the vector pENTR-1A-LUC + (Xie Q, et al (2014) LNK1 and LNK2 are transferable activators in the Arabidopsis circular activator. plant Cell 26(7) (2843) 2857) with BamHI and Kpn I were digested simultaneously, and the recovered fragments were ligated with the promoter of GmLClb2 by using T4 DNA ligase (Thermo Co., Ltd., product No. EL 0014). Obtaining an intermediate vector pENTR-GmLCLb2: LUC, recombining the intermediate vector into a plant expression vector pH2GW7 delta (Xie Q, et al (2014)) LNK1 and LNK2 are transformed and transformed into Escherichia coli DH5 alpha for propagation by utilizing an LR reaction kit (Thermo company, Cat. 11791019), and obtaining a recombinant plant expression vector pH2GW7 delta-GmLCLb 2: LUC.
The specific procedures for transformation of competent cells of E.coli and identification of vectors are as follows:
(1) preparing an LB solid culture medium containing antibiotics;
(2) melting competent cells stored in an ultra-low temperature refrigerator in an ice bath, adding 5 mu l of a connection product or an LR reaction product, gently mixing uniformly, and standing in the ice bath for 25 minutes;
(3) heat shock is carried out in a water bath kettle at 42 ℃ for 90 seconds, and then the mixture is immediately kept stand in an ice bath for 5 minutes;
(4) adding 500. mu.l of LB liquid medium, shaking-culturing at 37 ℃ for 1 hour (rotation speed of 150 rpm);
(5) uniformly coating 100 mu l of bacteria recovery culture solution on a screening culture medium, carrying out inverted culture at 37 ℃ for about 15 hours, and picking 3 single colonies for shake culture;
(6) and extracting plasmids, carrying out enzyme digestion identification and sequencing.
And (3) transforming the correctly identified recombinant plant expression vector pH2GW7 delta-GmLCLb 2: LUC into agrobacterium rhizogenes K599 to obtain positive transformed agrobacterium rhizogenes K599.
The pH2GW7 delta-GmLCLb 2: LUC is transferred into soybean WS82 by utilizing an agrobacterium rhizogenes mediated transformation method, and hairy roots with the pH2GW7 delta-GmLCLb 2: LUC are obtained through bioluminescence signal screening (figure 1).
The specific method for agrobacterium rhizogenes mediated soybean transformation is as follows:
(1) taking out soybean seeds sterilized by a chlorine fumigation method for 12 hours, placing the soybean seeds on a super clean workbench to blow off residual chlorine, and soaking the soybean seeds for about 16 hours by using sterile ultrapure water for later use;
(2) picking beltA single clone of Agrobacterium rhizogenes K599 with pH2GW7 delta-GmLCLb 2: LUC was subjected to small shaking in a liquid YEP medium in a test tube, and then transferred to a conical flask for shaking until the bacterial liquid OD 600 About 1.0. The cells were collected by centrifugation at 4000rpm for 10 minutes and resuspended in transformation medium (1/10X Gamborg B) 5 Salt, 30g/L sucrose, 3.9g/L MES, pH 5.4, added to 40mg/L acetosyringone after sterilization);
(3) cutting off plumule of imbibed soybean, taking hypocotyl as explant, soaking the explant in heavy suspension for 30 min, completely infecting, blotting the infection solution on filter paper, and placing the infected soybean explant in co-culture medium (1/10X Gamborg B) 5 Salt, 30g/L of sucrose, 3.9g/L of MES, 4.25g/L of agar and pH 5.4, and is added with 400 mg/L of Cysteine and 40mg/L of acetosyringone) after sterilization to be cultured for 3 days in dark place;
(4) inserting hypocotyl of co-cultured soybean explant into hairy root induction medium (1X Gamborg B) 5 Salt, sucrose 30g/L, MES 0.59g/L, agar 7g/L, pH 5.7, sterilized, Cefotaxime 100mg/L), and cultured at 25 ℃ under 12L/12D conditions for 14 days to induce rooting.
Hairy roots of the screened transformed pH2GW 7. delta. -GmLCLb2: LUC were cut into about 1 cm pieces and placed in 96-well plates with solid medium, to each well of which 40. mu.l firefly luciferase substrate (1.25mM) was added. Transfer to blue light (20. mu. mol/m) at 25 ℃ 2 S) or red light (20. mu. mol/m) 2 /s) continuous lighting conditions, and detecting bioluminescent signals every 3 hours using a bioluminescent detector.
The bioluminescence detection result is shown in fig. 2, and the result shows that the activity of the soybean GmLCLb2 promoter under red light is obviously lower than that of blue light, and the near-day rhythmic expression level of a target gene under the red light condition is obviously lower than that of the blue light condition; under the blue light condition, the activity of the GmLCLb2 promoter is 1.8-2.5 times higher in peak value and valley value than under the red light condition. The results show that the GmLCLb2 promoter can be used to drive differential expression of a gene of interest in plants under red or blue light conditions.
Although the invention has been described in detail hereinabove with respect to a general description and specific embodiments thereof, it will be apparent to those skilled in the art that modifications or improvements may be made thereto based on the invention. Accordingly, such modifications and improvements are intended to be within the scope of the invention as claimed.
Sequence listing
<110> university of Henan
Application of <120> promoter GmLCLb2 in regulating and controlling near-day rhythmic expression level of gene in response to environmental light quality change
<130> KHP201118724.6
<160> 3
<170> SIPOSequenceListing 1.0
<210> 1
<211> 3172
<212> DNA
<213> Artificial Sequence (Artificial Sequence)
<400> 1
cagaacactc ttatacttgt cattgcttat catctgaatt tgatcaccca cattcactgt 60
gaaggagtat ggttatcacc gttggatcaa aattaaggtg gcctaccctt taattttata 120
tatgctatac cctactctta cctgaaagat ggagggttgg agggccattc ctttatgtaa 180
tcttcgatag ggtgacagtg aagtctcaag aagtccctct agttggaaac cttcttacta 240
ttcacgttga aactagtgga caccctgctt gccttgaatg gatcgttgga gtaactcttt 300
aatctctcac tctctggtaa accgaaaaac tcccttgaca ccttcatcat cttttcaatc 360
acaccctctg gaaccatgct ttgtcacctt cattcatttc aaattttaca caaacgtaag 420
gttaatcacc aattctttcg ctctgtgtca attgaatggc actagctacc tgcatctgtg 480
aagatgcatg catgtgtgag ctaggacaga ccacgtatca tatctatctt aaaacataaa 540
agagcgtatt taaggaagag taagtttttt ctttatactt tgttttgttc tctatttttt 600
caatcatgct aagagagaat ttttctcttt tttttttttc atagacaacc taaaaatcta 660
ttttttttca atcatgtaaa tagagcattt tactttgtat tcttcacagt ataatcaatc 720
gtaggcagca atgaataaag aatttaactg acttgggaaa atcttaaaca taccaatatt 780
tgggatattt aagtgcaaga aggagagata aagaggaaaa aaaaaataag agatgaagta 840
atagatataa taaataatgt aatagaaaat aaagagtggg ataaagaaaa ataactcata 900
agctaaaaca aaattttagt tctttattta tatttttgtt ttaatttagt acttaaattt 960
tatgtgttac attatactag ttctttcgtc acacgtgata aattgtgaaa tatgacatgt 1020
ggtatggaat agtaatcatt aatgaaaatt gatattattt aagagataaa aataaaaatt 1080
taaaaactta agagattata ttgtcacaaa aaatataata attaatgaag tttacatagg 1140
tgaaaaaatc cataattttg gcaagcttgg gcaatctgtt ggatgatgtg ggagcgattg 1200
gggccatgaa ggtcttgaag atgaatgaga ggattggacg gtcagcaagt ggcctgatga 1260
aattagaggg aacctacttc aaagttcaaa gcagtactgg cttttgggtc atgttattgc 1320
tgtccattga ttcaaccact ttgctaatgc tacctctcta gctatagaat acaaataccc 1380
tcctctacct ctagctttta tttataggaa ccgagtaacg tgaaagaaat tcaattttcc 1440
tatattacta ataacaggta ttggttcgca taagaggctt tataaactct ttttaaaata 1500
ggttaaatta gttcattatt tcctataata caaatattgt tttttaggtt aaattagtaa 1560
attttaaaat cgattcaatt tgatataatt atataaacta aatcaatagt tttaaatcac 1620
tacgtcaaga ctgctcaact caaattaatt tataagacga gatcaaattg aataaatttt 1680
aaaattagag gaccaaatta ataaaaaaaa aactaaaaga ctcaatttaa ttaattttta 1740
aaattaaaag attaaattga aaagaaaaaa aatcaaatca aacttaaaaa ataaattaaa 1800
gtaataataa actaatttaa ccatatgtct tataaacaaa atctctcaaa taataaatat 1860
caaaaggaat atattattac aaaaaattga ccgtccattg gttggcagat agagcttcga 1920
agttggtaca atttcctact gtgggaatta gttccattct tcgtaaccgt ccaataacat 1980
tacaaactat atttattaaa tataatatat agtatgtagt ataattatta acaataaaaa 2040
ttactaagat ataagctatt actattaata tatatttccg tgttctgttt ttaaatatat 2100
tttcaactaa ttatttttta aaaaacaacg aaggttttaa tttataaaaa tagcattaga 2160
tttaatatat atatatatat atatatatat atatatatat atatatatat atatataatg 2220
tttaataaat gttgtaatca atatattaaa ggtagcattg actgttctac atcattaaat 2280
agtgtattta ttaaatatta tgtatggttt cttctaaatt tttttataga atatgtttta 2340
ggccaaccca atttaagtgt attcatttat ttaaagacag ccggtcatta ctacacttat 2400
aatttaagac ataatttaat tttaacttga aatagaataa gtttcatatt ggactaacag 2460
ttaaaaaatc gtttttgagt taaatgtatt tgctccaaaa aagaagaaaa agaagttaaa 2520
tgtatttaat tatattatat tataattgtt ttcgtaatta tttaaaaaaa acattctttt 2580
atttaaaatt taaatttgat cagattataa ttaaattggt tgaaattaga ttgtagattt 2640
aaattaaaca aaaagtgtta caattgtatg tctggtgctt agattattat catcgagaaa 2700
caaatctaaa ttgagccatg gaacgcacag caagtaacga accgtgaaga gagacgctcc 2760
gtgcggcggt gtgattggac gaagccgttg ccgcgcaact tgagggcccc acgtatgaca 2820
gctggcggcg cttttacgga ctgaaaacaa tggcgtcgga gtttgtggat gagatttcag 2880
ttgaggggag aaggccccaa aaaacaatat taaaggattc aggaaaagtg cattaaccac 2940
taaattgatg ccacttgtca caatcccttg cacacttagc agcctccaag taaaacgaca 3000
ccttcgtttt catcccagga aattctagtg gctgagattg cttctcactg ttgaacagtt 3060
ggctgtgctt gcttctgttc cttgtagttg aagtcgctcc ccctcattct tccttttcta 3120
cttgccactg tttcctattc gtcattgccg caaatgaatt ccgtcgcttc tc 3172
<210> 2
<211> 35
<212> DNA
<213> Artificial Sequence (Artificial Sequence)
<400> 2
cgggatccat gtgttataca agagaagttg aaccg 35
<210> 3
<211> 30
<212> DNA
<213> Artificial Sequence (Artificial Sequence)
<400> 3
ggggtaccga gaagcgacgg aattcatttg 30
Claims (3)
1. The application of the promoter GmLCLb2 or an expression cassette, a vector or a microorganism containing the promoter GmLCLb2 in the differential expression of regulatory genes in soybean hairy roots in response to the change of environmental light quality is characterized in that the expression level of the genes is up-regulated by giving blue light illumination treatment, or the expression level of the genes is down-regulated by giving red light illumination treatment;
the nucleotide sequence of the promoter GmLCLb2 is shown in SEQ ID NO. 1.
2. The application of a promoter GmLClb2 or an expression cassette, a vector or a microorganism containing the promoter GmLClb2 in regulating the expression level of a gene in the soybean hairy roots in response to the change of environmental light quality to regulate the expression level of the near-day rhythmicity, and is characterized in that the expression level of the near-day rhythmicity of the gene is up-regulated by giving blue light illumination treatment or the expression level of the near-day rhythmicity of the gene is down-regulated by giving red light illumination treatment;
the nucleotide sequence of the promoter GmLCLb2 is shown in SEQ ID NO. 1.
3. A method for regulating differential expression of genes in response to light quality changes of environmental red light and blue light in soybean hairy roots is characterized in that the genes are operably connected to the downstream of a promoter GmLCLb2, and the promoter GmLCLb2 is used for driving the differential expression of the genes in response to the light quality changes of the environmental red light and the blue light in the soybean hairy roots; up-regulating the expression level of the gene by giving a blue light irradiation treatment, or down-regulating the expression level of the gene by giving a red light irradiation treatment;
the nucleotide sequence of the promoter GmLCLb2 is shown in SEQ ID NO. 1.
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