CN116751815B - Application of OsEnS-73 gene in regulation of rice quality - Google Patents
Application of OsEnS-73 gene in regulation of rice quality Download PDFInfo
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- Y—GENERAL TAGGING OF NEW TECHNOLOGICAL DEVELOPMENTS; GENERAL TAGGING OF CROSS-SECTIONAL TECHNOLOGIES SPANNING OVER SEVERAL SECTIONS OF THE IPC; TECHNICAL SUBJECTS COVERED BY FORMER USPC CROSS-REFERENCE ART COLLECTIONS [XRACs] AND DIGESTS
- Y02—TECHNOLOGIES OR APPLICATIONS FOR MITIGATION OR ADAPTATION AGAINST CLIMATE CHANGE
- Y02A—TECHNOLOGIES FOR ADAPTATION TO CLIMATE CHANGE
- Y02A40/00—Adaptation technologies in agriculture, forestry, livestock or agroalimentary production
- Y02A40/10—Adaptation technologies in agriculture, forestry, livestock or agroalimentary production in agriculture
- Y02A40/146—Genetically Modified [GMO] plants, e.g. transgenic plants
Abstract
The invention discloses an application of an OsEnS-73 gene in regulating and controlling rice quality. The OsEnS-73 gene over-expression strain has the advantages of reduced whole polished rice rate, reduced thousand grain weight, increased chalkiness rate and chalkiness degree. The expression level of the OsEnS-73 gene in the two-week spike is inversely related to the quality. Therefore, the expression level of the rice OsEnS-73 gene is involved in regulating and controlling the rice quality, and can be used for genetic improvement of the rice quality.
Description
Technical field:
the invention belongs to the field of rice breeding, and relates to application of an OsEnS-73 gene in regulation and control of rice quality.
The background technology is as follows:
with the development of the economy in China and the continuous improvement of the living standard of people, the further improvement of the rice quality becomes the main aspect of rice breeding on the premise of current stable yield and increase of rice. In the market, the bad quality rice is little and the price of the high quality rice is several times, even tens times, that of the general rice. The strong development of high-quality rice is the need of market development and is also an important measure for realizing structural reform and country vibration strategy of agricultural supply side in China.
Rice quality includes processing quality, appearance quality, cooking taste quality, and nutritional quality. In the new revised national standard of high-quality rice, GB/T17891-2017, whole polished rice rate, chalkiness and taste are three indicators of high-quality rice grading. The polished rice rate means that after the rice is dehulled and milled into polished rice, the whole polished rice (the length of which reaches three-fourths or more than the average length of the whole rice) accounts for the percentage of the mass of the whole rice, and the final yield of the rice is affected, so that the commodity value and economic benefit of the rice are directly determined. The whole polished rice rate of the rice is 10-72%, and the whole polished rice rate of most high-quality rice varieties bred in China is 50-60%. Taking the first-class high-quality variety "Yuzhixiang" obtained by "national high-quality rice variety taste quality evaluation golden prize" for two consecutive years as an example, the whole polished rice rate is 55.8%. The average mu yield of the rice is 800 jin, the selling price of the rice is 35 yuan per jin, if the whole polished rice rate is improved by 10%, the broken rice is calculated according to half price, and the economic benefit can be improved by 1400 yuan per mu. Therefore, the improvement of the whole polished rice rate of the rice can bring great economic benefit. In addition, quality traits of rice examined varieties are gradually improved in 1977 to 2018 in China, wherein genetic improvement of chalkiness and chalkiness rates is significantly improved, but chalkiness indexes of most cultivars are still higher at present.
In view of this, the present invention has been made.
The invention comprises the following steps:
the invention aims to provide the function of the OsEnS-73 gene in regulating and controlling the rice quality, and provides technical support for improving the rice quality by utilizing the OsEnS-73 gene.
The first object of the present invention is to provide the function of the OsEnS-73 gene in regulating the whole polished rice rate of rice: the whole polished rice rate of the OsEnS-73 gene overexpression line is reduced.
The second object of the present invention is to provide the function of the OsEnS-73 gene in regulating grain weight: thousand kernel weight of the OsEnS-73 gene overexpression line is reduced.
The third object of the present invention is to provide the function of the OsEnS-73 gene in regulating the chalky grain rate of rice: the chalk rate of the OsEnS-73 gene over-expression line is increased.
The fourth object of the invention is to provide the function of the OsEnS-73 gene in regulating the chalkiness of rice: the OsEnS-73 gene over-expression line has increased chalkiness.
A fifth object of the present invention is to provide the second object of the present invention as described above, which provides the use of the OsEnS-73 gene in improving rice quality.
Preferably, the application in rice breeding is in improving whole polished rice rate, grain weight, chalkiness rate and chalkiness degree.
Compared with the prior art, the invention has the following beneficial effects:
(1) The invention provides the function of the OsEnS-73 gene in regulating and controlling the quality of rice. The transgenic gene proves that the expression level of the OsEnS-73 gene can regulate the rice quality.
(2) By analyzing 200 parts of materials with extreme expression level of the OsEnS-73 gene, the expression level of the OsEnS-73 gene is inversely related to the quality, and further the expression level of the OsEnS-73 gene can be applied to the improvement of rice quality.
Description of the drawings:
fig. 1: whole genome association analysis (GWAS) results of 300 parts indica rice. A and B represent manhattan plots of GWAS results at the Yangjiang test base in 2016, guangzhou test base and 2018, respectively. The OsEnS-73 gene in the invention is a candidate gene identified by GWAS to a QTL (named qHMRR 4-2), and the qHMRR4-2 is marked by an arrow in the figure.
Fig. 2: qHMRR4-2 candidate gene analysis. Panel A and B represent a phenotypic comparison of the whole rice rates between different haplotype materials in 2016 Guangzhou and 2018 Yangjiang qHMRR4-2, respectively. Haplotypes a and C refer to variations on chromosome 4 29582749. Statistical comparisons used a one-sided t-test. Panel C shows the local Manhattan plot of GWAS and the qHMRR4-2 linkage disequilibrium heat map. Panel D shows the expression level of LOC_Os04g49500 in high HMRR germplasm (Nos. 539, 645 and 659) and low HMRR germplasm (Nos. 452, 523 and 651) by RNA-seq measurement. Panel E shows qRT-PCR to verify LOC_Os04g49500 expression.
Fig. 3: an overexpression vector of the OsEnS-73 gene.
Fig. 4: functional verification of the OsEnS-73 gene. A represents the detection of overexpression of the OsEnS-73 gene line (OE-7, OE-9). B represents a polished rice picture of the control (659) and OsEnS-73 gene overexpression lines (OE-7 and OE-9), head polished rice represents whole polished rice, and Broken rice represents crushed rice. Panels C-F show statistical comparisons of control (659) and OsEnS-73 gene over-expression lines (OE-7 and OE-9) in terms of whole polished rice rate (HMRR), thousand Grain Weight (TGW), chalky grain rate (PGWC) and chalky Degree (DEC), respectively. Statistical comparisons used a one-sided t-test, which represents a significant difference in p < 0.05 in t-test, and p < 0.01 in t-test.
Fig. 5: influence of the expression level of the OsEnS-73 gene on the rice quality. Panel A shows a comparison of 100 parts of OsEnS-73 gene highly expressed and 100 parts of OsEnS-73 gene lowly expressed material. Panel B, panel C and panel D show comparisons of polished rice rate, chalkiness rate and chalkiness between materials with high and low expression of the OsEnS-73 gene, respectively. Statistical comparisons used a one-sided t-test.
The specific embodiment is as follows:
the following examples are further illustrative of the invention and are not intended to be limiting thereof. The specific experimental conditions and methods are not noted in the following examples, and the techniques employed are conventional and well known to those skilled in the art.
Example 1: whole genome association analysis of whole polished rice rate
The 300 parts of diversity indica rice germplasm introduced from international rice is taken as a material, the 300 parts of indica rice are respectively planted in a Guangzhou Dafeng test base and a Yangjiang test base in 2016 and 2018 late-time planting, and after seeds are harvested, the whole polished rice rate of the rice is measured according to national standard GB T21719-2008. After obtaining the phenotype data, a whole genome association analysis (GWAS) was performed using 70 ten thousand SNPs data from the 300 whole genome of indica rice. Whole genome association analysis utilizationThe software package GAPIT 2.0 performs, in combination with the kineship matrix and principal component PC of the population, calculations using a hybrid linear model. Screening criteria based on conventional whole genome correlation analysis results, i.e. 2P values less than 10 over a continuous 200kb interval of the genome -4 The interval serves as a QTL site. Two years of GWAS results identified a large effect QTL on chromosome 4 with a peak position of 29582749, designated qHMRR4-2 (fig. 1).
Example 2: identification of qHMRR4-2 candidate Gene
There was a significant difference in overall polished rice rate between materials of different haplotypes of qHMRR4-2 (fig. 2A and 2B). Based on linkage disequilibrium analysis, qHMRR4-2 was divided into regions of about 260kb (from 29.37Mb to 29.63Mb on chromosome 4) (FIG. 2C). According to MSU rice genome annotation 7 th edition, qHMRR4-2 region total 39 annotated genes. According to the haplotype analysis of qHMRR4-2, we selected 3 high whole-rice haplotypes and 3 low whole-rice haplotypes for gene differential expression analysis. The RNA-seq results and qRT-PCR verification show that only the expression pattern of LOC_Os04g49500 has expression differences between different haplotype materials. The LOC_Os04g49500 expression level in the low whole rice material is higher than that in the high whole rice material (FIGS. 2D and 2E). LOC_Os04g49500 is endosperm specific expression gene, and the gene is named OsEnS-73.
Example 3: functional verification of OsEnS-73 gene
OsEnS-73 is an endosperm-specific expressed gene whose nucleotide sequence is shown in SEQ ID No.1 and is expressed primarily during endosperm development (Nie, D.M., ouyang, Y.D., wang, X., zhou, W., hu, C.G., yao, J. (2013), genome-wide analysis of endosperm-specific genes in rice. Gene,530 (2), 236-247.). We constructed an overexpression vector for the OsEnS-73 gene using the endosperm-specific expression promoter DXCP35 (Yan Y, lin YJ. Cloning and functional identificat ion of an endosperm-specific promoter DXCP from Rice. Journal of Huazhong Agricultu ral university.2014 (5): 15-20 (in Chinese)), FIG. 3. The carrier construction process is as follows: (1) target fragment amplification and recovery: the gene sequence of OsEnS-73 (its core) was obtained using the RGAP (http:// price. Plan biology. Msu. Edu /) databaseThe nucleotide sequence is shown as SEQ ID NO. 1), and an amplification primer F is designed: 5'-Acta ggtctcG CGAT GGCGCCGCCGCCGTCGTCATT-3', R:5'-Acta g gtctcT TCAT CAACTATTTCCCTTCTGTGA-3' cDNA obtained by RNA reverse transcription of the spike of indica rice material 659 in the filling stage is used as a template for amplification. The promoter DXCP35 was amplified from 659 material leaf DNA using primers DXCP35-1F:5'-Acta ggtctcG CTCG CTAACGCTAT TCAACCGTGA-3' and DXCP35-1R:5'-Acta ggtctcT ATCG ATCGATCCCAAACTACGAGA-3'. NOS was amplified using NOS-3F:5'-Acta ggtctcG ATGA TCCT CCCGATCGTTCAAACA-3' and NOS-3R:5'-Acta ggtctcT ACCG GTGCTCCCGATCTAGTAA C-3' with pYLCRISPR/Cas9Pubi-N as templates. PCR reaction system: 2 XPCR buffer 25. Mu.l, 2mM dNTPs 10. Mu.l, F primer 1. Mu.l, R primer 1. Mu.l, KOD-FX 1. Mu.l, template DNA 1. Mu.l, and H 2 O to 50. Mu.l. PCR reaction conditions: 94 ℃ for 2min;98℃10sec,55℃30sec,68℃2min (32 cycles); and at 68℃for 5min. The three fragments of DXCP35 (1357 bp)/OsEnS-73 (2145 bp)/NOS (270 bp) were cut out and the target fragment was recovered using agarose gel DNA recovery kit (DP 209-03) from Tiangen Biochemical Co., ltd. (2) cleavage-ligation reaction: 10 XCutSmart Buffer 1.5. Mu.l, 10mM ATP 1.5. Mu.l, PEG plasmid (backbone of PEG plasmid as in FIG. 3, modified with pc1300 plasmid) 100ng, target fragment (DXCP 35, osEnS-73, NOS) 50 ng/each, bsa I-HF 10U,T4 DNA ligase 35U, H 2 O to a total volume of 15. Mu.l, the reaction parameters were 5min at 37℃and 5min at 20 ℃. (3) recombinant plasmid transformation and bacterial detection: transferring the constructed vector into DH5a escherichia coli competent cells, and performing colony PCR detection by using EnS (det+) (5'-GTGCCCAATTACAGGTCAAG-3')/RB-R (5'-AAGTTG GGTAACGCCAGGGT-3'), wherein the band size is about 894bp; and (5) detecting positive bacterial plaques, and picking and shaking the bacterial plaques. And (3) carrying out Sanger sequencing on the plasmid extracted from the positive bacterial plaque, and detecting by using a sequencing primer Lac-F (5'-TC ACTCATTAGGCACCCCAG-3')/RB-R, wherein the sequence of a sequencing result is consistent with the sequence of a target fragment, so that the construction of the overexpression vector is successful. After the constructed over-expression vector is transformed into agrobacterium EHA105, infection transformation is carried out on callus of Japanese sunny by using the agrobacterium, and the obtained transgenic line is screened by hygromycin and further passes through a primer F:5'-CAGCAGATGCAGAAAGCGTG-3' R:5'-TCGTTCAGATGCCTCGTTGG-3' the expression level of the OsEnS-73 gene was examined to obtain a transgenic line in which OsEnS-73 was expressed in an increased amount (FIG. 4A).
The transgenic lines (OE-7 and OE-9) with the OsEnS-73 expressed in an increasing mode and a control material (659) are simultaneously sown, germinated and raised, and when the rice seedlings grow to the four-leaf period, the rice seedlings are transferred to a transgenic greenhouse. And (5) conventional water and fertilizer management. After the harvested mature seeds are naturally dried in the sun, husking rice by a rice huller to obtain brown rice, grinding the brown rice into polished rice by a rice polisher, manually separating whole polished rice and broken rice, and measuring the whole polished rice rate according to the method of national standard 'rice whole polished rice rate test method' (GB/T21719-2008). The chalky grain rate and chalky whiteness of whole polished rice were measured by using a rice appearance quality tester (SC-E, hangzhou). The whole polished rice rate of the OsEnS-73 gene-overexpressing strain was reduced, while the thousand seed weight of the OsEnS-73 gene-overexpressing strain was found to be reduced, and the chalkiness rate and chalkiness were increased (FIGS. 4B to 4F).
Example 4: influence of OsEnS-73 Gene expression on Rice quality
Transcriptome data of 350 rice varieties and 2 weeks of the grouting period of resource materials are obtained in the early stage of the research laboratory, and the influence of OsEnS-73 gene expression on the quality of seeds is further analyzed by using materials with extreme OsEnS-73 gene expression quantity, including 100 high-expression materials and 100 low-expression materials (figure 5A). Statistical comparisons found that materials highly expressing the OsEnS-73 gene had lower whole polished rice rates, higher chalky and chalky levels (fig. 5B, 5C and 5D). This suggests that lower levels of expression of the Os EnS-73 gene can improve grain quality.
The OsEnS-73 gene sequence is as follows (SEQ ID NO. 1):
ATGGCGCCGCCGCCGTCGTCATTGCTTCGCGACCTCCTCGCCGCTGATGGCTTCAGAAACCGTCGC
AAGCCGCCTGACAGTAACCCTCCGGCCGCCCCAAGGACCACGAGCATGCCCCTCCAGCACCGGCG
GCCGAGTAGGCCAGCACGGTCGCAGTCCGACGTCCTCACCCGCAGCCGCCTCAGAGAAACAAACA
ACGTCGGTACTAGCGATGGCGACGGAGTCGACGCCGGCGAGGAGCAGCGGACCGCCACACGGAG
GTCATCGGCGTCGCTGATGAGCGCGAGGAGCTACAACAACAACAAGGACAGCGGCGGCGGCGCCA
TGAGGGGCGGCTCAGCTGCCGCGGTACCGGCTCTCGACGAGTCCGTGCTCACCGCGTTGATCTCCC
TGGTCGCGGGAGCCGTGAAGCGGTTCGTCAGGGACGAGGGCTTCCGCGCGTCGCTCCGCGGCGGG
TGCATGTCGTGCCTCGGCGGCGAGTCCAACCACCGGGCGGTCCTGGACCTCCGCGTGATCGTGCAC
ACCGTCGAGAGGGCCGCGAGCGAGGGGCTCGACGACCCGCGCGACCTGAAGCGCGCGTCGCTCA
GGCTGCACGCCATGGCGTCCCTCGACGCGAAGGAAGCGGACGCGGTGAGCGCGGCCGGCGTGCCC
TACCAGCGCCTCGCCGCGTGCGCGCACCTGTACATGTCCGTCATCTCCAAGCTCCAGGAGAAGGAC
CACTCCTCCGCGGTGCACGTGCTGGAGGCCTTCTGCTTGGCGCCGCACGAGGCCCGCACATCCCTG
CTTCCCGCGCTCTGGGACAGGCTCTTCCGCTCGGGCCTCTCGCACCTCAAAGCGTGGCGTGACCGC
GAGTCGGCGACCGCGGCGAGCTCGGATACGAGGGTGAAGGACGTGGAGAAGATGTTCGTAGAGGC
AGTGGACGATGGCACACGCGCGCTCGCATGCTACTACAGAGATTGGCTTCTGGGGCGCAGCCAAGC
CATGTCTCTCCCGGTTGTTCCTGCTCCTCCCAGCACCGTTCTTGCCAGCGCGCTGAGGTTCTCGACG
TCGACGTCCTATGACATTGGCTCGGATGTCGCTTGTAGCTCCGGGAGTTACAGTCCAGCTGTAAAGT
TTGCGCTCGACGAGACTCCATCACAATATGATCGAGAGATCGAAGAGGAAGAAGAAGCAGAGGTG
GACGAGAAGACAGCAGATGCAGAAAGCGTGTTCCATGAGTGCGATGGCACTGAACCGAAGAGCTA
CACTCACTCACTTCAGACAGAAGAAAACGAGCTAATGCCTGACAAGCTAGCCAACGAGGCATCTG
AACGAAAGAGTGAAGATGAACGGAGCAGACAACCAGATGAGTCAACGAGTTATGTGCCAATTTCT
GATATCACGGCTATTGACCTTCCTACACTCGAGTTCTGTGAAGGGCCTGATCTCCAGAGTGACACAG
ATGGCAGTCAAATATCCATTTTTGCCACCATTCCTAGTGATTTCCTCTGTCCACTGACGAGGCAGATC
TTCAACCGTCCGGTGACAATCGAGACAGGTCAGACGTTCGAGCGGCATGCTATAGTTCAGTGGTTC
GATAAAGGCATCAGAATGTGCCCAATTACAGGTCAAGAACTTGAAAGCCTGTCGATTCCAGACATA
AACCGTGTGTTAAAACGCTTGATCGATAACTGGAAGTCCGAACACTGCAAAACTCTGAACTCTGAA
AGTACATGTCCTGAAAAGGAGTTAACTGCAACATTCGTAGAGAATGTCCTTTCTTCTGAATGCGAGA
CGTCTGAAAAACTTGAGAAGGCAAGGCGTCTTATGGCAATCGGTGGCATAGACTTTCTTCTGCATAA
ATTTCATGGAGGAGGAGAAGATGAGAAGGCACAAGCAGCTGAGCTCCTTTTGCTCTGCATCAGGGC
AGAAGGAAGCTGCAGAAACTACATGGCCATAAGGATTTGTAACTCAAGTGTTGTTCAGCTTCTCCA
CAGTGAGGTGATTTCAGCAAGGAGTTCTGCAGTTCGTCTGCTTGTTGAACTGCTCTGCCTGAAAAG
GTTCTGTCTCCTGTTCGTAACACCAAATGCATCCAAATATGTCTTGTATGTACATGCGCTTCCATCTC
ACAAAATATTTCAATTGTCACAGAAGGGAAATAGTTGA。
Claims (2)
- the application of the OsEnS-73 gene expression level in the regulation of rice quality is characterized in that the quality of an OsEnS-73 gene over-expression strain is reduced, the nucleotide sequence of the OsEnS-73 gene is shown as SEQ ID NO.1, and the rice quality is the whole polished rice rate, thousand grain weight of rice, chalkiness rate of rice and chalkiness degree of rice.
- The application of the OsEnS-73 gene in rice breeding is characterized in that the low expression of the OsEnS-73 gene is applied to improving the whole rice yield, the chalkiness rate and the chalkiness degree of rice, and the nucleotide sequence of the OsEnS-73 gene is shown as SEQ ID NO. 1.
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