CN112779270B - Functional gene for enhancing iron deficiency tolerance and iron accumulation of plants and application - Google Patents
Functional gene for enhancing iron deficiency tolerance and iron accumulation of plants and application Download PDFInfo
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Abstract
The invention relates to a functional gene for enhancing iron deficiency tolerance and iron accumulation of plants and application thereof, belonging to the technical field of bioengineering. The functional gene sequence for enhancing the iron deficiency tolerance and iron accumulation of the plants is shown in a sequence table SEQ ID No: l is shown. The invention relates to a method for preparing a polypeptide shown in a sequence table SEQ ID No: l, knocking out the functional gene for enhancing the tolerance of the plant to the iron deficiency stress in the plant, so that the functional gene is not expressed in the plant, and the plant shows the iron deficiency tolerance character. The functional gene for enhancing iron deficiency tolerance and iron accumulation of the plant can provide new gene resources and technical guarantee for stress resistance genetic improvement of crops, and plant seeds with enhanced stress resistance and stress tolerance can be cultivated by applying the functional gene.
Description
Technical Field
The invention belongs to the technical field of bioengineering, and particularly relates to a method for enhancing the tolerance of plants to iron deficiency stress and improving the iron content in the plants by using the gene as a negative regulatory factor.
Background
Iron (Fe) is one of the essential nutrient elements for animals and plants. It has important effect in maintaining normal physiological function of animal body, and can participate in hemopoiesis, growth and development maintenance, infectious disease resistance, etc. Similarly, Fe has an important role in plant bodies, and is a component of cytochrome and non-heme ferritin involved in photosynthesis, biological nitrogen fixation, and respiration, which are important substances in plant energy metabolism. The synthesis of chloroplast needs the participation of Fe, so under the stress of Fe deficiency, the plant has the phenomenon that the leaf veins are in green, and the green of the leaf is usually found in young leaves. Although the soil is rich in a large amount of Fe, the availability ratio is extremely low, the high pH value and the high carbonate content seriously reduce the effectiveness of Fe in the soil, so that the plants are difficult to absorb Fe from the soil, the Fe deficiency stress is caused, the plant growth is seriously influenced, and the crop yield is reduced. Therefore, the improvement of the tolerance of crops to iron deficiency stress and the improvement of the absorption of iron become great matters related to the civilian life, the important point of further research on agricultural production and food safety is formed, the attention of political and academic circles of various countries in the world is paid, and the hot spot of the current life science research is formed.
Arabidopsis thaliana, as a classical model plant, is widely used in the research fields of plant genetics, crop biology, developmental biology, molecular biology, and the like. The whole genome of the arabidopsis thaliana is sequenced, and the arabidopsis thaliana has the characteristics of simple structure, short growth period, high propagation coefficient, strong vitality, self-pollination and easiness in transformation, can more quickly and better reach the expected target of an experiment by taking the arabidopsis thaliana as a research object, can shorten the experiment time to a great extent and simplify the experiment conditions, and shows incomparable superiority of other organisms. The research on the mechanism of plant response to iron deficiency stress by using model organism arabidopsis thaliana provides a new gene resource for the stress resistance genetic improvement of crops. The search and discovery of new functional genes with proprietary intellectual property rights based on the arabidopsis sequencing database (www.arabidopsis.org) is one of the hot spots in the international research field of botany and is the focus of technological competition among different countries. Arabidopsis thaliana shares about 1.3 hundred million base pairs, 2.9 ten thousand genes. At present, the function research of most genes is not clear, the research of gene functions by using a mutant screening technology becomes an effective method, at present, CRISPR/Case9 mediated gene editing technical methods can be used, T-DNA insertion, transposon insertion, EMS mutagenesis, RNAi interference and other methods can be used, the research functional genes are knocked out finally by using the genetic engineering technical methods, and transformed plant tissues are cultivated into mutant plants.Through research on arabidopsis mutants, some functional genes that regulate iron deficiency response have been discovered, such as:FIT、bHLH38、bHLH39、bHLH100、bHLH101、bHLH34、bHLH104、bHLH105、bHLH115、 PYE、MYB10、MYB72and the like.
According to the published genomic sequences of the arabidopsis database,ATMYC1(AT4G00480) is a member of the Arabidopsis thaliana bHLH transcription factor family, and bHLH is a specific transcription regulator newly found in plants in recent years, and belongs to the bHLH family. It has been found that a part of bHLH transcription factors in Arabidopsis play a crucial role in responding to iron deficiency stress in plants. For theATMYC1Only research on genes has found thatATMYC1The gene is involved in the biosynthesis of grape flavone and the regulation of plant root hair and hair cell differentiation, however, it is not clear whether it plays a regulatory role in plant abiotic stress response.
In the face of the increasingly severe soil problems of high pH and high carbonate, the finding of functional genes for regulating plant iron homeostasis and enhancing iron accumulation and the clarification of the functions of the functional genes have important theoretical and practical significance.
Disclosure of Invention
The invention aims to provide a functional gene for enhancing the iron deficiency tolerance and iron accumulation of plants, and the second aim of the invention is to provide the application of the functional gene as a negative regulatory factor in regulating the iron deficiency tolerance and iron accumulation of plants.
The DNA sequence of a functional gene for enhancing the iron deficiency tolerance and accumulation of plants is shown as SEQ ID No. 1.
The sequence table SEQ ID No: l, knocking out the functional gene for enhancing the tolerance of the plant to iron deficiency stress and iron accumulation in the plant, namely not expressing in the plant, wherein the plant shows the iron deficiency tolerance character, and the plant is arabidopsis thaliana.
The sequence table SEQ ID No: l, knocking out the negative regulatory protein ATMYC1 gene in the arabidopsis thaliana by a T-DNA insertion method of the functional gene for enhancing the iron deficiency tolerance and accumulation of the plant. So as to improve the Fe absorption of the arabidopsis thaliana, lead the Fe content in the arabidopsis thaliana to be increased, and further show the tolerance character to iron deficiency stress.
The beneficial technical effects of the invention are embodied in the following aspects:
1. according to a genome sequence published by an arabidopsis database, ATMYC1 is a member of an arabidopsis bHLH transcription factor family, and the applicant finds that a plant is tolerant to iron deficiency stress under the treatment of iron deficiency stress after ATMYC1 gene mutation, which indicates that the ATMYC1 gene responds to the regulation of the iron deficiency stress. Therefore, the function of the gene is researched, and further research results show that the Fe content in the ATMYC1 gene mutant plant is higher than that in the wild plant, which shows that the knockout gene ATMYC1 can improve the Fe absorption of arabidopsis thaliana, so that the Fe content in the arabidopsis thaliana is increased, and the arabidopsis thaliana is further tolerant to iron deficiency stress.
2. The ATMYC1 gene can enhance the iron deficiency tolerance and accumulation of plants after mutation, provides new gene resources and technical guarantee for crop stress resistance genetic improvement, and can be used for cultivating plant seeds with enhanced stress resistance and stress tolerance.
Drawings
FIG. 1 is a schematic diagram of a T-DNA insertion site.
FIG. 2 isatmyc1In the mutantATMYC1And (4) identifying the transcription level of the gene.
FIG. 3 is a drawing showingATMYC1The gene is induced and expressed by iron deficiency stress in wild arabidopsis thaliana.
FIG. 4 is a drawing showingatmyc1Comparative photographs of mutants and wild type plants (WT) were cultured vertically on petri dishes and seeded directly on MS and-Fe medium, respectively, for 2 weeks in normal light conditions, with MS medium as control.
FIG. 5 shows the cloning of ATMYC1 gene, the identification of ATMYC1 overexpression vector and the screening of ATMYC1 overexpression plant.
FIG. 6 shows the detection of the expression level of ATMYC1 gene in ATMYC1 over-expressed plants.
FIG. 7 is a photograph comparing ATMYC1 overexpressing plants (OE 1 and OE 5) and wild type plants (WT) cultured vertically on petri dishes, spotted directly onto MS, -Fe medium, respectively, for 2 weeks under normal lighting conditions, with MS medium as control.
FIG. 8Is wild type andatmyc1comparing the iron content in the roots and stems of the mutant plants.
FIG. 9 is a schematic view ofatmyc1The plant and the wild plant normally grow on the MS culture medium for 2 weeks, and the expression level of the iron deficiency related gene is detected after the plant and the wild plant are transferred to the Fe culture medium for 3 days.
Detailed Description
The present invention will be further described with reference to the following examples.
The experimental procedures in the following examples are conventional unless otherwise specified.
Example 1 obtaining of ATMYC1 mutant
We screened mutants responding to plant iron deficiency stress using Arabidopsis thaliana mutant seeds obtained from the American Arabidopsis thaliana germplasm resources pool. Screening an arabidopsis seed mutant library by using an MS culture medium without iron to obtain an iron-deficiency tolerant mutant, and screening and identifying to find that the mutant is an ATMYC1 gene function-deficient mutant named as ATMYC1-1, wherein the seed number of the mutant is SALK-056899. In order to further research the effect of the ATMYC1 gene in response to iron deficiency stress of plants, another ATMYC1 gene function deletion mutant named as ATMYC1-2 is obtained from an American Arabidopsis germplasm resource library, and the seed number of the mutant is SALK-057388. Because the seed is T-DNA insertion mutation, PCR amplification and sequencing are carried out through specific primers, the sequencing result is compared in NCBI database Blast, and T-DNA insertion site information is obtained through comprehensive analysis, and the figure 1 is shown. Using semi-quantitative PCR, onatmyc1-1Andatmyc1-2 transcript level characterization of the mutant material revealed that the expression level of ATMYC1 gene was significantly lower in both mutants than in Wild Type (WT), see fig. 2. In order to further determine whether the ATMYC1 gene participates in plant iron deficiency stress response, RNA of Wild Type (WT) Arabidopsis thaliana subjected to iron deficiency stress treatment at different time points is extracted and is subjected to reverse transcription to form cDNA, the transcription level of the ATMYC1 gene is analyzed by using a real-time quantitative PCR technology, the result shows that the expression level of the ATMYC1 gene is remarkably inhibited by iron deficiency stress relative to a control group, and the result further shows that the ATMYC1 is used as a negative regulatory factor to respond to the iron deficiency stress, and the reference is made to FIG. 3.
Wild Type (WT) was seeded simultaneously with atmyc1-1 and atmyc1-2 in a 90 mm diameter petri dish in solid media with and without iron and cultured vertically in a 22 ℃ thermostatically light incubator with a 16 h photoperiod, 8 h dark. After two weeks, it can be observed: no significant difference was observed in the root length and chlorophyll content of WT, atmyc1-1, atmyc1-2 grown on MS medium, as shown in B in FIG. 4 and C in FIG. 4. Directly dibbling the plant and culturing on a culture medium without iron,atmyc1-1andatmyc1-2the trait exhibited marked iron deficiency tolerance, as shown in a in fig. 4. Under iron deficiency stress, the root length and chlorophyll content of atmyc1-1 and atmyc1-2 were significantly higher than that of WT, see B in FIG. 4 and C in FIG. 4. The above results indicate that atmyc1-1, atmyc1-2 showed significant tolerance to iron deficiency stress as compared to WT.
Example 2 cultivation of iron deficiency phenotype sensitive Arabidopsis thaliana
1、ATMYC1Gene overexpression transgenic lineATMYC1Obtaining of OE1, OE5
To further verify the function of the gene in regulation and control of iron deficiency stress of plants, an ATMYC1 gene overexpression vector (35S: ATMYC 1) is constructed. First, the target fragment is amplified. The wild arabidopsis thaliana is normally cultured on an MS culture medium for two weeks, total RNA of a plant is extracted, reverse transcription is carried out to synthesize cDNA, the synthesized cDNA is taken as a template, PCR is carried out, and a sufficient amount of target products are amplified, wherein the target products are shown in A in figure 5. And performing second amplification by using the PCR product as a template so as to introduce a restriction enzyme site. The PCR product and the vector pCAMBIA1301 are subjected to enzyme digestion and recovered. The recovered and purified target DNA fragment and the vector are then ligated overnight with T4 DNA ligase. Transferring the ligation solution into DH5a, detecting and screening positive clones, and sequencing. After the sequencing result is correct, the agrobacterium GV3101 is transferred by an electric shock transformation method. The agrobacterium GV3101 after electric shock transformation is coated on LB culture medium plate containing double antibody (Kan, Gen) after activation. Single colonies were picked at random, expanded in LB medium containing double antibody (kan, Gen) and identified by PCR using vector primers, see B in FIG. 5. After the size of the PCR amplified fragment is consistent with that of the target gene, an arabidopsis wild type plant is transformed by adopting a floral dip method, so that an ATMYC1 gene overexpression transgenic strain is obtained, and the expression is shown as C in figure 5. Wherein the content of the first and second substances,
primer 1: f5 'NNNGGTACCATGTCTTTGACAATGGCTGATG 3';
primer 2: r5 'NNNAAGCTTTTAATGAAAGATACAAATCGCCC 3' is provided.
2. Transcription level identification of ATMYC1 overexpression transgenic plant and comparison with iron deficiency of wild plant
Using semi-quantitative PCR we identified the transcription level of ATMYC1 overexpressing transgenic plants, and finally selected OE1 and OE5 for further experiments, see FIG. 6. Wild Type (WT) was seeded simultaneously with ATMYC1-OE1 and OE5 on a 90 mm diameter petri dish in solid media with and without iron, and cultured vertically in a 22 ℃ constant temperature light incubator (16 hours light cycle, 8 hours dark). After two weeks, it can be observed: WT and ATMYC1-OE1, OE5 grown on MS medium showed no significant difference in root length, chlorophyll, etc., as shown in B in FIG. 7 and C in FIG. 7; the ATMYC1-OE1 and OE5 cultured on a medium without iron directly inoculated on the plant all show obvious iron deficiency sensitive characters, and the characters are shown in A in figure 7. Under iron deficiency stress, ATMYC1-OE1, OE5 had significantly lower root length and chlorophyll content than WT, see B in FIG. 7 and C in FIG. 7. The results show that ATMYC1-OE1 and OE5 show more obvious sensitive traits to iron deficiency stress than WT shows, and further prove that ATMYC1 gene is used as a negative regulatory factor to respond to the regulation of iron deficiency stress.
3. Analysis of iron content in roots and stems of wild type and atmyc1 mutant plants
atmyc1The mutant and wild type plants (WT) were cultured vertically on petri dishes and were seeded directly on iron-supplemented and iron-deficient medium, respectively, and cultured vertically for 2 weeks under normal light conditions. The iron content of the roots and stems of WT, atmyc1-1, atmyc1-2 plants was determined under MS and-Fe treatment conditions, respectively. As a result, it was found that, under normal conditions,atmyc1-1、atmyc1-2the iron content in the stem and root was not significantly different from that of wild-type WT, as shown in FIG. 8A and FIG. 8B; under the condition of iron deficiency, the iron-rich alloy is mixed with the iron,atmyc1-1、 atmyc1-2the iron content in both the stem and root was higher than that of wild-type WT, as shown in FIG. 8A and FIG. 8B, and the above results were further obtainedThe gene and the coding protein thereof for enhancing the iron deficiency tolerance and accumulation of the plants by using ATMYC1 as a negative regulatory factor are shown.
4. Comparison of expression levels of iron deficiency-related genes in atmyc1 mutant and wild type WT plant under iron deficiency stress
Taking atmyc1 and wild WT plants under the condition of 3 days of iron deficiency stress treatment, extracting RNA and performing reverse transcription to form cDNA, and then determining the expression levels of related genes (FIT, IRT1 and FRO 2) by utilizing qRT-PCR technology, and finding that the expression levels of FIT, IRT1 and FRO2 in atmyc1 plants are obviously higher than that of WT, as shown in A in figure 9, B in figure 9 and C in figure 9, which indicates that the tolerance of atmyc1 plants to iron deficiency may be related to the obvious induction of the expression of FIT, IRT1 and FRO2 genes.
Sequence listing
<110> university of fertilizer industry
<120> functional gene for enhancing iron deficiency tolerance and iron accumulation of plants and application thereof
<130> 2021
<160> 1
<170> SIPOSequenceListing 1.0
<210> 1
<211> 1743
<212> DNA
<213> Arabidopsis thaliana (Arabidopsis thaliana)
<400> 1
atgtctttga caatggctga tggtgtagaa gctgcagcag gaagaagtaa aagacaaaac 60
agcttattaa gaaaacaact tgctttagct gtaagaagtg ttcaatggag ctacgcaatc 120
ttctggtcgt cttcacttac tcaacctggg gttttggagt ggggagaagg atgttacaat 180
ggagatatga agaagaggaa gaagagttat gaatctcatt ataaatatgg gttgcaaaaa 240
agcaaggagc ttcggaaact ttatttgtct atgcttgaag gagacagtgg tactactgtt 300
agtactactc atgataatct caatgatgat gatgataatt gtcacagtac aagtatgatg 360
ctgtcaccag atgacctctc tgatgaagag tggtactatt tagtctccat gtcctatgtc 420
ttctctcctt cacaatgttt gcctggaaga gcttcagcga cgggtgagac catatggctc 480
tgcaacgctc aatatgcgga gaacaagctc ttctctcgtt ctttgttagc aagaagcgca 540
tcaattcaga ctgttgtgtg tttcccttac ttgggcggag tcattgagct gggcgtcact 600
gaattgattt cagaagacca taacctgctt cgaaacatca aatcttgctt gatggaaata 660
tctgcacacc aagacaacga tgacgagaag aagatggaga ttaagatcag tgaagagaag 720
catcagcttc cattaggtat ttctgatgaa gacttgcatt acaaaagaac catttcaaca 780
gtactcaact actccgcaga tagatcaggt aagaacgata agaacattcg tcatcgtcag 840
ccaaatattg ttacttctga acctggctca agtttcttgc ggtggaagca atgtgagcag 900
caagtctcgg gttttgttca gaaaaaaaag tcacagaatg tgttgcggaa gatattgcat 960
gatgtccctt tgatgcacac aaagagaatg ttcccaagtc agaactctgg tctgaatcaa 1020
gatgatcctt cagatagaag aaaagagaac gaaaagttca gtgtccttag aactatggtt 1080
cccactgtca acgaggttga taaagaatcg atactaaaca acacaatcaa gtacctgcaa 1140
gaactggagg caagagtaga agagctagaa tcttgtatgg gatcagttaa ttttgtagaa 1200
agacaaagaa agacgacaga gaaccttaac gactctgtgt tgatcgaaga gacatcaggg 1260
aactacgatg atagcacgaa gatcgatgac aattcaggag aaaccgaaca agtcactgtt 1320
ttcagagata agacacattt gagagttaaa ctcaaagaaa cagaagttgt gatcgaagta 1380
agatgttctt acagagacta catagttgcg gacatcatgg aaactctgag caatcttcac 1440
atggatgctt tctctgttag atctcacacg ctcaataagt tcctcacatt gaatctcaag 1500
gccaagtttc gcggggctgc agttgcgtcc gtaggaatga ttaagcgaga gctgagaaga 1560
gtcattgact ttcgtgaacc gatatgcgat gtgccattat ctttacatca agttttcagg 1620
gtttttgtat gtaaagtttg ccaaagtttg gttggaattt tcgacaacgt tgtctcttcc 1680
tcttctacaa aaccaagatc tatacttatt cataattcat gggcgatttg tatctttcat 1740
taa 1743
Claims (2)
1. As shown in a sequence table SEQ ID No: the application of the functional gene shown by the general formula I in enhancing the tolerance of plants to iron deficiency and iron accumulation is characterized in that: the sequence table SEQ ID No: l, knocking out the functional gene shown in the formula I in the plant, namely not expressing the functional gene in the plant, wherein the plant shows the iron deficiency tolerance character and is arabidopsis thaliana.
2. Use according to claim 1, characterized in that: the sequence table SEQ ID No: l, knocking out the negative regulatory protein ATMYC1 gene in the arabidopsis thaliana by a T-DNA insertion method.
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