CN111961676B9 - Mutant gene OsCOPT7 of copper low-accumulation mutant lc1 of rice and application thereof - Google Patents
Mutant gene OsCOPT7 of copper low-accumulation mutant lc1 of rice and application thereof Download PDFInfo
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Abstract
The invention discloses a low-copper-accumulation mutant of paddy ricelc1Mutant gene of (2)OsCOPT7And applications thereof. Low-accumulation mutant of copper in ricelc1Mutant gene of (2)OsCOPT7Wild speciesOsCOPT7The coding region of the gene has base mutation, namely the 344 th base at the downstream of the ATG is mutated from G to A, namely the 115 th codon of the coding region is changed from GGG to GAG, so that the 115 th amino acid in the protein sequence is mutated from glycine (Gly/G) to glutamic acid (Glu/E). The mutant geneOsCOPT7The coding nucleotide sequence of (A) is shown as SEQ ID No.1, and the coding amino acid sequence is shown as SEQ ID No. 2. Mutantslc1The copper content in the rice grain is obviously lower than that of the wild rice grain. The application of the mutant comprises one or more of transgenosis, hybridization, backcrossing or asexual propagation; the mutant geneOsCOPT7The rice plant comprises the mutant geneOsCOPT7Or rice comprising said mutant geneOsCOPT7The encoded amino acid of (1).
Description
Technical Field
The invention belongs to the field of crop genetic breeding, and particularly relates to a low-copper-accumulation mutant of ricelc1Mutant gene of (2)OsCOPT7And applications thereof.
Background
Copper (Copper, Cu) is one of the main elements of the cultivated land soil polluted by heavy metal in China. Many studies have shown that excessive copper accumulation inhibits plant growth and affects the associated physiological processes. In addition, copper, as a coenzyme factor, participates in various biochemical reaction processes, exists in the human body in the form of ceruloplasmin and cuprase, and is a trace element necessary for life health. And the intake of proper amount of copper is vital to the health of residents. Rice is the main grain of residents in China, and has great significance in cultivating copper low-accumulation rice varieties suitable for copper-polluted soil. Because the copper accumulation capacity difference among different varieties of rice is obvious, the method is an economic and effective way for planting by selecting the rice variety with low copper accumulation in grains.
The copper content in rice grains is influenced by the absorption, transportation and redistribution processes of copper by root systems, and the process is genetically controlled. In recent years, several references have been madeGenes and transporters associated with copper uptake, transport and phloem transport in rice have been discovered in succession. After copper element enters plant root system, heavy metal ATP enzyme (P) is mainly used1B-ATPase, HMA) family transporter is involved in its transport process, corresponding to the transporter OsHMA4-9 in rice. In addition, genes such as iron transport proteins OsYSL1 and OsYSL3, zinc transport proteins OsZIP2 and OsZIP4 may also participate in the copper transport process of plants. OsYSL16 localized in phloem can participate in the redistribution process of copper by transporting Cu-NA. The CTR high-affinity copper transport protein (COPT) is a main participant in the copper absorption process of rice root systems, however, at present, few researches on rice COPT family genes and protein functions are reported, and no research is carried out on the functions of the genes and the proteinsOsCOPTAnd (3) mutating the family member gene to obtain a mutant with low copper accumulation in the grains.
Disclosure of Invention
In order to overcome the defects of the prior art, the invention aims to provide a low-accumulation mutant of copper in ricelc1Mutant gene of (2)OsCOPT7And applications thereof.
Low-copper-accumulation mutant of ricelc1Mutant gene of (2)OsCOPT7Wild speciesOsCOPT7The coding region of the gene has base mutation, namely the 344 th base at the downstream of the ATG is mutated from G to A, the 115 th codon of the coding region is changed from GGG to GAG, and the coding nucleotide sequence is shown as SEQ ID No. 1; the rice copper low-accumulation mutantlc1The copper accumulation in the seeds is obviously reduced to be less than 20 percent of the wild type.
The rice grain low-copper accumulation mutantlc1Mutant gene of (2)OsCOPT7The mutant geneOsCOPT7The coded amino acid sequence is shown as SEQ ID No.2, and the 115 th amino acid in the wild type OsCOPT7 protein sequence is mutated from glycine to glutamic acid.
The low-copper-accumulation mutant of ricelc1Mutant gene of (2)OsCOPT7The rice plant comprises the mutant geneOsCOPT7(ii) a Or rice comprising the mutant geneOsCOPT7The encoded amino acid of (1).
The application mode comprises one or more of transgenosis, hybridization, backcross or asexual propagation.
An expression cassette or recombinant vector containing the mutant geneOsCOPT7。
The invention has the beneficial effects that:
mutant material obtained by the present inventionlc1The copper content in the kernel is reduced to below 20 percent. Thus, mutantslc1Is/are as followsOsCOPT7The gene mutation mode has obvious influence on the accumulation of the copper in the seeds and can play a remarkable role in subsequent breeding. And the mutantlc1Can be used for the subsequent construction and utilization of the dominant variety of the copper-polluted soil.
Drawings
FIG. 1 shows wild-type and copper-poor accumulating mutants (lc1) Basic parameter diagram of plants.
FIG. 2 shows wild type andlc1copper content in plant brown rice and rice ear is shown.
Detailed Description
The invention is further described below with reference to the following figures and examples.
The invention uses Indica rice 9311(Oryza _ Indica) as wild type seeds (M)0Generation) and performing EMS mutagenesis treatment, planting the treated seeds to obtain M1And (5) plant generation. M1Inbred seed production (M) from plant generations2Generation), planting M on the field2And (4) plant generation, namely after husking the harvested rice grains, measuring the copper content in the brown rice by using an inductively coupled plasma spectrometer (ICP), and screening the plants with low copper accumulation. Further selfing and screening the obtained low copper accumulation rice line, and planting and verifying in different rice fields to obtain a mutation homozygous single plant which has low copper accumulation amount and can be stably inherited, and is named aslc1（low copper mutant 1) And used for cross breeding and biotechnological research.
Low-accumulation mutant of copper in ricelc1The mutation site is located in riceOsCOPT7The 344 th base downstream of the ATG of the gene initiation codon is mutated from G to A, the 115 th codon in the CDS sequence is changed from GGG to GAG, the 115 th amino acid in the OsCOPT7 protein sequence is mutated from glycine (Gly/G) to glutamic acid (Glu/E), and the result is thatlc1The copper content in the mutant rice grain is significantly lower than that of the wild rice grain.
Low-accumulation mutant of copper in ricelc1Mutant gene thereofOsCOPT7The nucleotide sequence is shown as SEQ ID No.1, and the amino acid sequence is shown as SEQ ID No. 2.
Firstly, obtaining a low-copper-accumulation mutant of rice:
in 2013, 450g of conventional rice wild type seeds (M in this case)0) Soaking the seeds in clear water for 12 hours, soaking the seeds in 1.0% Ethyl Methanesulfonate (EMS) at room temperature for 12 hours, stirring the mixture during the soaking, discarding the EMS treatment solution, and washing the treated seeds with clear water for 4 hours. Seeds after mutagenesis treatment are soaked, germinated and sown, and are transplanted after 25d seedling age, and the seeds are managed according to conventional fertilizer and water (M)1). And (4) harvesting 1-3 ears per plant according to the seed setting rate during maturation, and harvesting 4140 plants in total. Except for single plant seed preservation, each plant is randomly mixed into M by taking 15 grains2And (4) seeds.
2014, M2Soaking seeds, accelerating germination, sowing, transplanting the single seeds with the age of 25d into the same field (the pH of soil = 7.5), and managing according to the conventional fertilizer and water. After maturation, the single plant is harvested, and 35689 strains are harvested together. Selecting 433 rice plants with low fruiting rate and 12527 rice plants with normal fruiting rate, shelling corresponding plant seeds after harvesting, and measuring the content of elements such as copper in the brown rice by using an inductively coupled plasma spectrometer (ICP). The copper content of the brown rice is 1.4mg/kg, and the copper content of the corresponding wild variety brown rice is 3.6-4.1 mg/kg.
The pH value of the previous soil is high, so that the copper absorption of the rice is influenced. In 2015, the obtained copper accumulation material seeds of the rice are soaked and germinated in the next year, then sown, and 18 individual seedlings with the age of 25d are transplanted in a field with the pH value of 5.4, and are managed according to conventional fertilizer and water. After the mature, the single plant is harvested, and 10 plants are harvested in total. Husking the harvested rice seeds, and measuring the copper content in the brown rice by using ICP (inductively coupled plasma), wherein the copper content in the brown rice of 9 rice plants is obviously lower than that in the wild type rice, the concentration distribution is 1.7-2.1mg/kg, and the copper content in the brown rice of the wild type variety is 7.0-8.2 mg/kg.
In 2016, 9 low-copper-accumulation rice material seeds harvested in the previous year are soaked and germinated in the next year, then sown, 18 single seedlings are transplanted in a field with the soil pH value of 5.4 at the age of 25d, and the field is managed according to the conventional fertilizer and water. After the rice is matured, the single plant is harvested, the seeds are hulled, and the copper content in the brown rice is measured by ICP. The result shows that the copper content in the brown rice is not separated and is about 60 percent lower than that of the wild plant rice. Wherein the brown rice copper content of the 1 line rice is obviously lower than that of the wild type rice, the concentrations are respectively 1.5-2.3mg/kg, and the brown rice copper content of the wild type variety is 5.4-6.8 mg/kg.
Considering that the low copper accumulation character of the seeds of the plant line is not separated, the plant line is numbered as K152 in 2017, and a column line with lower content is screened for further selfing purification, and is respectively planted and verified in the Zhejiang Fuyang two-field rice field. The results show that the copper content in the mutant strain seeds is obviously lower than that of the wild type seeds, is respectively 7% and 10% of that of the wild type seeds, and can be stably inherited. In addition, validation in 2018 in another background field showed that mutant lines still have significant low copper advantage. Therefore, K152 is taken as a mutant with low copper accumulation in grains for subsequent experiments, and the name islc1 (low copper mutant 1)。
Second, low copper accumulation mutantlc1Copper accumulation characteristics
Low copper accumulation mutantslc1The tillering number, the seed setting rate and the plant height of the plant are obviously lower than those of a wild type, and other main agronomic traits are not obviously different from those of the wild type (see figure 1).
2016. 2017, the mutants are planted in Chang Yang Changan rice field in Hangzhou and Zhonghuayuan rice field in Zhejiang Fuyang in 2018, and the results show that the mutants are all shownlc1The copper content in the brown rice is obviously lower than that of the wild type brown rice; the rice field planting result of Zhejiang Fuyang Zhonghui Huayuan in 2019 shows that the mutantlc1The copper content in the rice ear and brown rice of the plant is 50% and 16% of that of the wild type respectively, and the detailed results are shown in figure 2.
III, copper low accumulation mutantlc1Determination of mutant genes
Wild type and copper low-accumulation rice mutantlc1The variety is crossed, and all F1 plantsLow copper accumulation characteristics were not exhibited. In the same year, F is treated in Hainan in winter1Selfing for the first time to obtain F2 segregating population. Randomly selecting about 500F particles2And (3) seed generation, seed soaking and germination accelerating, planting in a Hainan propagation base, harvesting a single plant, and measuring the copper content in the brown rice by using ICP (inductively coupled plasma). F2The low-accumulation and normal-accumulation strains of copper in the brown rice in the segregation population are typically 1: 3 separation, indicating that the rice is ricelc1The special character of the mutant is controlled by recessive monogene.
Further selection of mutantslc1And (3) taking the F2 population hybridized with the wild type variety as a positioning population, respectively selecting 30 individuals from the separated brown rice with extremely low copper content and extremely high phenotype, respectively extracting DNA, then equivalently mixing 30 sample DNAs to construct a DNA pool, marking as F2-L and F2-H, and performing whole genome re-sequencing.
The DNA extraction was carried out as follows: taking rice leaves with the length of about 2cm, and placing the rice leaves into a 2ml centrifugal tube filled with steel balls; after freezing by liquid nitrogen, crushing the leaves by using a crusher, taking out and adding 500ml of 1.5 × CTAB, and shaking the test tube at room temperature until the leaf powder is uniformly mixed with the extractant; water bath at 65 deg.C for 20-30min, and mixing by reversing every 5min for 1 time; adding equal volume of chloroform, turning upside down, mixing, and keeping for 10 min; centrifuging at 10000rpm for 10 min; sucking 400ml of supernatant into a new centrifuge tube, adding 200ml of isopropanol, turning upside down, and mixing uniformly for 10 min; centrifuging at 10000rpm for 10 min; centrifuging, removing supernatant, adding 500ml 75% ethanol, rinsing by inversion, and centrifuging at 12000rpm for 5 min; discarding supernatant, drying in a superclean bench or naturally drying in the air, adding 100ml ddH2And dissolving the DNA, and detecting the quality of the DNA by electrophoresis.
The genome re-sequencing steps were as follows: after obtaining the DNA, the two parents of the population and the two offspring extreme phenotype pool DNA samples are randomly broken into fragments with the length of 350 bp. The Library is built by adopting a TruSeq Library Construction Kit, the preparation of the whole Library is completed by the steps of end repair, ployA tail addition, sequencing joint addition, purification, PCR amplification and the like of a DNA fragment, and the built Library is sequenced by the illumina HiSeq.
The Raw data 36.381G is generated in the sequencing process, the filtered Clean data is 36.321G, the sequencing quality is high (Q20 is more than or equal to 96.83%, Q30 is more than or equal to 91.69%), and the GC content is 42.98% -43.98%. In conclusion, the data quantity of all samples is enough, the sequencing quality is qualified, the GC distribution is normal, and the library construction sequencing is successful.
Valid sequencing data are aligned to a reference genome Oryza _ Indica through BWA software, the alignment result is subjected to SAMTOOLS to remove duplicates, and the alignment rate of all samples is 96.35% -99.18%. The comparison result is normal, and can be used for subsequent variation detection and related analysis.
A Unifield Genotyper module of GATK3.3 software is adopted to detect a plurality of sample SNPs, and Filtration is carried out by using Variant Filtration, so that 254,730 total SNPs are obtained. The number of synonymous variants on exon was 7435 and the number of non-synonymous variants was 11698. Based on the results of genotyping, markers of homozygous differences between the two parents were screened, and a total of 3743 polymorphic markers were selected. And according to the distribution of the filial generation SNP-index on the chromosome, finding that the chromosome 9 is the chromosome where the candidate gene is located.
For candidate polymorphic marker sites, annotation results of ANNOVAR are extracted, and a candidate gene BGIOSGA030839 is screened, wherein the CDS corresponding coding sequence of the candidate gene is Copper transport protein OsCOPT7 (Copper Transporter 7) and the mutant thereof is subjected to mutationlc1The coding sequence is shown in SEQ ID No. 1. The candidate gene is located between 14863852-14864301bp of chromosome 9,lc1the base at 344bp downstream of ATG in the mutant is mutated from G to A, so that the 115 th amino acid in the amino acid sequence of OsCOPT7 is mutated from glycine to glutamic acid (G115E), and the amino acid sequence is shown as SEQ ID No. 2.
Sequence listing
<110> institute of Rice research in China
<120> mutant gene OsCOPT7 of copper low accumulation mutant lc1 of paddy rice and application thereof
<141> 2020-09-01
<160> 2
<170> SIPOSequenceListing 1.0
<210> 1
<211> 450
<212> DNA
<213> Indica rice 9311(Oryza Indica)
<400> 1
atgatgcaca tgaccttcta ctgggggaag gacgtgacga tcctgtttga cggctggcgg 60
acggccacgt ggaccgggta cctcctctcc ctcgtcgcgc tcctcctcgc ctccgccttc 120
taccagtacc tcgaggcgtt ccggatccgc gtcaagctcc tcgcgggcgc caagccggcc 180
tccatcccgc ccccggcgtc ctccgacgcc gcgcgggccc cgctcctcct cccctcgtcc 240
gccgccggac gctggccggc gcggctggcg acggccgggc tgttcggggt gaactccggc 300
ctgggctacc tcctcatgct cgccgtcatg tcgttcaacg gcgaggtgtt cgtcgccgtg 360
gtcgtcgggc tcgcggcggg ctacttggcg ttccgcagca gcgacggcga ggacctcgtc 420
gtggtcgaca acccctgcgc ctgcgcctag 450
<210> 2
<211> 149
<212> PRT
<213> Indica rice 9311(Oryza Indica)
<400> 2
Met Met His Met Thr Phe Tyr Trp Gly Lys Asp Val Thr Ile Leu Phe
1 5 10 15
Asp Gly Trp Arg Thr Ala Thr Trp Thr Gly Tyr Leu Leu Ser Leu Val
20 25 30
Ala Leu Leu Leu Ala Ser Ala Phe Tyr Gln Tyr Leu Glu Ala Phe Arg
35 40 45
Ile Arg Val Lys Leu Leu Ala Gly Ala Lys Pro Ala Ser Ile Pro Pro
50 55 60
Pro Ala Ser Ser Asp Ala Ala Arg Ala Pro Leu Leu Leu Pro Ser Ser
65 70 75 80
Ala Ala Gly Arg Trp Pro Ala Arg Leu Ala Thr Ala Gly Leu Phe Gly
85 90 95
Val Asn Ser Gly Leu Gly Tyr Leu Leu Met Leu Ala Val Met Ser Phe
100 105 110
Asn Gly Glu Val Phe Val Ala Val Val Val Gly Leu Ala Ala Gly Tyr
115 120 125
Leu Ala Phe Arg Ser Ser Asp Gly Glu Asp Leu Val Val Val Asp Asn
130 135 140
Pro Cys Ala Cys Ala
145
Claims (5)
1. Low-copper-accumulation mutant of ricelc1Mutant gene of (2)OsCOPT7Characterized by being wildOsCOPT7The coding region of the gene has base mutation, namely the 344 th base of the ATG is mutated from G to A, the 115 th codon of the coding region is changed from GGG to GAG, and the coding nucleotide sequence is shown in SEQ ID No. 1.
2. The mutant gene according to claim 1OsCOPT7Characterized in that the mutant geneOsCOPT7The coded amino acid sequence is shown as SEQ ID No.2, namely the 115 th amino acid in the wild type OsCOPT7 protein sequence is mutated from glycine to glutamic acid.
3. The low copper accumulation mutant of rice as claimed in claim 1lc1Mutant gene of (2)OsCOPT7Use of a mutant gene according to claim 1 in rice for reducing copper accumulation in riceOsCOPT7(ii) a Or rice comprising the mutant gene as claimed in claim 2OsCOPT7The encoded amino acid of (1).
4. Use according to claim 3, wherein the mode of application comprises one or more of transgenesis, crossing, backcrossing or asexual propagation.
5. An expression cassette or recombinant vector comprising the mutant gene according to claim 1OsCOPT7。
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Correction item: Claims Correct: Claims 1-5 submitted on July 30, 2021 False: Claims 1-5 submitted on June 29, 2021 Number: 38-02 Page: ?? Volume: 37 |
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