CN113801885B - Rice large grain gene LG1 and application thereof - Google Patents
Rice large grain gene LG1 and application thereof Download PDFInfo
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Abstract
The invention belongs to the field of plant genetic engineering, and particularly relates to a gene LG1 for controlling rice grain development, which is separated, cloned and functionally verified, and an application thereof in rice production practice. The invention discloses a rice large-grain gene LG1 and a protein coded by the same, and also discloses an application of the rice large-grain gene LG 1: increasing the grain size of gramineous plants and increasing the yield.
Description
Technical Field
The invention belongs to the field of plant genetic engineering, and particularly relates to a gene LG1 for controlling rice grain development, which is separated, cloned and functionally verified, and an application thereof in rice production practice.
Background
Rice is an important food crop for human beings, is mainly distributed in asia, the south of europe, the americas, africa and the like, and lives more than one third of the world population. With the increase of population year by year, the available cultivated land area is reduced, natural disasters are frequent, the trend of land circulation and non-grain transformation is aggravated, the grain safety becomes a major problem which troubles many countries, and the stable yield and the high yield of the grain are kept to form the primary target of crop breeding.
The rice yield mainly comprises three factors of effective spike number, solid grain number of each spike and thousand grain weight. In high-yield breeding of rice, thousand grain weight is one of three factors determining yield, and grain shape is an important agronomic trait determining thousand grain weight. Wherein, the indexes reflecting the particle shape mainly comprise the particle length, the particle width, the particle thickness and the length-width ratio. These indicators are closely related to the rice yield, and also affect the appearance quality of rice, and determine the final commodity value. So far, a plurality of genes for controlling the grain shape have been cloned, and the genes are mainly involved in the regulation and control of molecular mechanisms such as cell proliferation and expansion, phytohormone, ubiquitination, grouting, small RNA and the like, and lay an important foundation for improving the yield of rice and the quality of rice.
The grain length, the grain width, the grain thickness and the length-width ratio of the rice are quantitative traits controlled by multiple genes, most related Quantitative Trait Loci (QTL) of the rice mainly regulate grain shape by influencing the division of glume cells, and a molecular regulation mechanism of the grain shape is preliminarily disclosed. GS3, GW6a and GL7/SLG7 are mainly involved in regulation and control of rice grain length and grain weight. GS3 is located near the centromere of chromosome 3 and is a major QTL affecting grain length and grain weight (Fan et al, 2006). GS3 encodes a transmembrane protein containing 3 domains, of which the OSR domain is an essential element affecting grain length, while the VWFC domain has an inhibitory effect on the OSR domain (Mao et al, 2010). After the GS3 function is lost, grains are lengthened, narrowed and the grain weight is increased; after the GS3 is over-expressed, grains become small, plants are dwarfed, and growth and development are inhibited, which indicates that the GS3 is a negative regulation factor of grain length. Compared with the short-grain variety, the long-grain variety has nonsense mutation of the gene in the second exon, and the protein translation is terminated early. Genome association analysis of rice cultivars indicates that this nonsense mutation located in the second exon of GS3 is a significant cause of grain size differences in the natural population of rice (Fan et al, 2009). GW6a is located in chromosome 6, encodes a GNAT-like protein OsglHAT1 with histone acetyltransferase activity, and positively regulates grain length, grain weight and plant biomass. After overexpression of GW6a, the seed becomes longer, the seed becomes shorter after expression is inhibited, and the seed length is increased after overexpression of the gene in Arabidopsis (Song et al, 2015). GL7/SLG7 is located on chromosome 7, encodes a homologous protein of the arabidopsis longifola protein, positively regulating the longitudinal elongation of glume cells (Wang Y et al, 2015. It was found that a 17.1Kb tandem repeat in long grain japonica rice in the united states resulted in up-regulation of GL7 expression levels and down-regulation of GL7 expression in the vicinity of negative regulators, thereby increasing grain length and significantly improving rice appearance quality (Wang Y et al, 2015). GW2, GW5/qSW5 and GS5 are mainly involved in regulating and controlling the grain width and the grain weight of rice. GW2 is located on chromosome 2It is a major QTL for grain width and grain weight that encodes a RING protein with E3 ubiquitin ligase activity that is degraded by anchoring its substrates to the proteasome, thereby negatively regulating cell division. GW2 increases glume size mainly by increasing glume cell number, and also increases rice filling rate, thereby changing grain size and yield (Song et al, 2007). Besides influencing the development of grains, the GW2 gene also regulates the development of the whole plant. Therefore, it belongs to the pleiotropic gene as well as the GS3 gene. GW5/qSW5 is a major gene on chromosome 5 that controls grain width and grain weight and encodes a nuclear localization protein with an arginine-rich region. Yeast double-hybrid experiments show that GW5/qSW5 can participate in ubiquitin protease degradation pathway of rice, and suggest that GW5/qSW5 regulates glume cell division through ubiquitin-protease complex pathway in seed development process, thereby influencing seed width and grain weight, and GW5/qSW5 and rice functionGW2There is a similar effect (Shomura et al, 2008. The GS5 gene is located on chromosome 5 and encodes a serine carboxypeptidase that positively regulates rice grain width and grain weight (Li et al, 2011). The GS5 gene influences the width of grains by influencing the number and the size of cells, and the increase of the expression quantity of the GS5 gene promotes the division of glume cells so as to increase the grain width, otherwise, the grain width is increased. Studies have shown that both broad and narrow varieties encode the entire GS5 protein, and thus the difference in grain width may be due to the difference in the amount of expression of this gene. To date, GS3, GW5/qSW5, GS5, etc. have been selected by humans and widely used in rice breeding practice.
Disclosure of Invention
The invention aims to solve the technical problem of providing a rice large-grain gene LG1 and application thereof in rice breeding.
In order to solve the technical problems, the invention provides a rice large-grain gene LG1, wherein the nucleotide sequence of the gene LG1 is shown as SEQ ID NO:2, the preparation method is as follows.
The invention also provides the protein coded by the rice large-grain gene LG1, and the amino acid sequence of the protein is shown as SEQ ID NO:4, the preparation method is described in the specification.
The invention also provides a recombinant vector and a transformant containing the gene.
The invention also provides the application of the rice large-grain mutant gene LG 1: increasing the grain size of graminaceous plants and increasing the yield.
Improvement of the use of the large rice grain gene LG1 of the invention: the gramineous plants are rice, the grain length, the grain width and the grain thickness of rice grains are increased, and the rice yield is improved.
The invention also provides a method for regulating and controlling the grain development of the gramineous plants, which comprises the following steps: comprises the steps of transforming gramineous plant (such as rice) cells by using the gene LG1, and then culturing the transformed gramineous plant cells into plants, wherein the grains of the plants are enlarged, and the thousand kernel weight of the plants is increased.
The improvement of the method for regulating and controlling the development of the gramineous plant kernel comprises the following steps: the grain length, the grain width and the grain thickness of the grains are increased.
The invention also provides a plasmid containing the gene, and engineering bacteria or host cells containing the gene or the vector.
The engineered bacteria and host cells are understood to be those used by the skilled worker in the context of transgenesis. However, with the development of science and technology, the selection of the engineering bacteria and the host cells may be changed, or in the application field of non-transgenic purpose, the utilization of the vector and the engineering bacteria is also related, but the invention is within the protection scope as long as the gene or the vector of the invention is contained.
Further, the invention also provides a host cell, which contains the gene sequence and is an Escherichia coli cell, an Agrobacterium cell or a plant cell.
Another object of the present invention is to provide the use of the above gene for transgenic crop improvement.
The preparation of the transgenic rice is a conventional technical means in the field, the invention is not limited, and the technical scheme of utilizing the gene to perform rice transgenosis is within the protection scope of the invention.
The specific technical steps for realizing the invention are as follows:
1. isolation and genetic analysis of mutant lg 1:
the large-grain rice mutant lg1 of the invention is derived from a natural mutant library of japonica rice cultivar Zhonghua 11 (ZH 11) (FIG. 1). The mutant is proved to be controlled by recessive monogene through positive and negative cross experiments with wild type (ZH 11).
2. Comparison of lg1 with wild-type spikelets
Grains of lg1 were observed during the maturation phase. The lg1 mutant can obviously increase the grain length, the grain width and the grain thickness, particularly the grain width and the grain thickness, and obviously increase the rice yield (figure 1).
3. Map-based cloning of LG1 Gene
In order to isolate LG1 gene, the invention firstly constructs a positioning population, and the F is formed by hybridizing LG1 and indica rice variety NJ6 2 And (3) positioning the population, and preliminarily positioning the LG1 locus on the 2 nd chromosome between the R9 marker and the R16 marker by using a plurality of molecular markers through a map location cloning method. By analyzing the sequence between these two markers, a new polymorphic marker was developed to finely map the LG1 gene to a region of about 33kb between the markers S19 and S24, and sequencing alignment analysis revealed that the DNA sequence of LOC _ Os02g09490 gene of LG1 mutant underwent base substitution compared with ZH11, resulting in premature translation termination (FIG. 2). Blast analysis found that LG1 gene (LOC _ Os02g 09490) is allelic with GH2 previously reported. The lg1 mutation site is inconsistent with the mutation site of the GH2 allele previously reported, and the base substitution of the mutant DNA sequence in lg1 results in premature translation termination. Analysis of LG1 gene expression showed that it was expressed in each tissue or organ, but was highly expressed in the ear (fig. 3).
The large-grain rice mutant lg1 has obvious phenotype difference with the existing gh2 mutant. The method specifically comprises the following steps: the main characteristic of the gh2 mutant is golden grain glume, while the lg1 mutant of the invention is mainly characterized in that grains are obviously increased and thousand kernel weight is increased. Therefore, the invention deeply expounds the genetic mechanism and the action mechanism of rice grain regulation and control through the cloning and the deep functional interpretation of the LG1 gene. Therefore, the invention provides important theoretical significance for further improvement of the yield of rice and molecular design breeding.
Rice (Oryza sativa) is basic food which nearly half of the population lives in the world, and has a large planting area in China and even Asia, and the large-particle control gene LG1 cloned by the invention and the protein coded by the gene have important production and application values in increasing the rice yield.
Drawings
The following describes embodiments of the present invention in further detail with reference to the accompanying drawings.
FIG. 1 is a comparison of grain phenotypes of wild type (ZH 11) and lg1 mutants;
in fig. 1:
a is the length comparison of ZH11 and lg1 mutant grains;
b is the width comparison of ZH11 and lg1 mutant grains;
c is length statistical data of ZH11 and lg1 mutant grains;
d is the width statistical data of ZH11 and lg1 mutant grains;
e is the thickness statistical data of ZH11 and lg1 mutant grains;
f is the thousand kernel weight statistical data of ZH11 and lg1 mutant grains;
FIG. 2 shows the phenotype of transgenic rice tested for gene localization and functional complementation;
in FIG. 2:
a is the location of LG1 gene;
b is LG1 gene coding cinnamyl alcohol dehydrogenase, and the mutation thereof causes the premature termination of protein translation;
c is genetic complementation experiment; wt represents the kernel of the wild type plant, lg1 represents the kernel of the mutant plant, and com represents the kernel of the complementary plant.
FIG. 3 shows the expression of LG1 gene.
Detailed Description
The following examples illustrate the invention in detail: the present embodiment is implemented on the premise of the technical solution of the present invention, and a detailed implementation manner and a specific operation process are given, but the protection scope of the present invention is not limited to the following embodiments.
Example 1:
1. rice material
Rice (Oryza sativa l.) mutant lg1 (large grain 1), original wild-type material was japonica rice cultivar midflower 11 (ZH 11) (fig. 1). lg1 mutants are natural mutagen material from ZH 11.
Example 2:
1. analyzing and locating populations
The positive and negative cross experiments of lg1 mutant and indica rice variety NJ6 show that the mutant is controlled by nuclear genes. lg1 mutant and indica rice variety NJ6, F 1 Selfing and breeding F 2 1006 individuals with large grain phenotype (i.e., the grain phenotype is consistent with the lg1 mutant) were selected from the population as a targeted population. At the same time, F 2 Segregation ratios of normal phenotype plants and mutant phenotype plants in the population were close to 3 by chi-square test.
TABLE 1 genetic analysis of mutant lg1
2. DNA extraction
Approximately 1g of leaves were taken from each plant during heading to extract total DNA. The method comprises the following specific steps:
(1) 1g of rice leaf was weighed, ground into powder with liquid nitrogen, and then added with 600. Mu.l of DNA extraction buffer prepared from CTAB solution (2% (m/V) CTAB,100mmol/L Tris-Cl,20mmol/L EDTA,1.4mol/L NaCl; pH 8.0), and water-bathed at 65 ℃ for 40 minutes. Then 600. Mu.l of chloroform/isoamyl alcohol (24. Centrifuge at 10,000rpm for 5 minutes and transfer the supernatant to a new centrifuge tube.
(2) And (2) adding 2/3-1 times volume of precooled (to 4 ℃) ethanol into the supernatant obtained after the centrifugation in the step (1), and gently mixing the mixture until DNA precipitation. Centrifuge at 12,000rpm for 10 minutes and pour the supernatant.
(3) Then, the DNA precipitate obtained in the above step (2) was washed with 500. Mu.l of 75% (vol.) ethanol.
(4) The washed DNA was air-dried and dissolved in 500. Mu.l of purified water.
(5) The integrity of the DNA was checked by electrophoresis on a 1% agarose gel. The intact DNA was used for PCR amplification and the incomplete DNA was re-extracted until the intact DNA was obtained.
Example 3:
1. preliminary mapping of LG1 Gene
F from lg1 mutant in combination with NJ6 hybrid 2 94 recessive individuals are randomly selected from the population, polymorphism analysis is carried out on the small population, and PCR amplification is carried out according to known reaction conditions for linkage analysis according to primers which are uniformly distributed on 12 chromosomes and stored in a laboratory.
The PCR amplification conditions were as follows: the total PCR reaction was 10. Mu.l: wherein 100 ng/. Mu.l of rice genomic DNA 1. Mu.l, 10 XPCR Buffer 1. Mu.l, 2mM dNTP 1. Mu.l, 10uM primer 2. Mu.l, 5U/. Mu.l rTaq 0.05. Mu.l, ddH 2 O4.95. Mu.l. The PCR amplification conditions are specifically: pre-denaturation at 94 ℃ for 5 min; denaturation at 94 ℃ for 30 sec, annealing at 55 ℃ for 30 sec, extension at 72 ℃ for 30 sec, 35 cycles, separation by 4% agarose gel electrophoresis and staining with Gelred nucleic acid dye, detection of polymorphism of PCR product, and preliminary mapping of LG1 gene between R9 and R16 markers on chromosome 2 (FIG. 2).
2. Fine localization of LG1 Gene
F is formed by the hybridization of lg1 and indica rice variety NJ6 2 And (3) positioning the remaining 912 recessive individuals of the population, continuously designing a molecular marker on the basis of initial positioning, and finally accurately positioning the LG1 gene in a region of about 33kb between the markers S19 and S24. Sequencing alignment analysis revealed that the DNA sequence of LOC _ Os02g09490 gene of lg1 mutant underwent base substitution, leading to premature translation termination, compared with ZH11 (FIG. 2). The primer sequences are shown in table 2:
TABLE 2 location marker sequence of LG1 Gene
Name of label | Front primer (5 '-3') | Rear primer (5 '-3') | Size of fragment |
R9 | AACCTACCACTGCCATTGC | GGCATTATCCATACCAGCAG | 253bp |
R16 | TGCTGATGTTAATCTGTGGGTG | TGGCGGCTTGAGAGTGTTTGTAG | 105bp |
S19 | GGAGGTAATGCTTATAGTGGG | CATTGCTGCGGGAAATTTAG | 680bp |
S24 | GGCCAACTGGAGCACCTTG | GACCTAGCGTCAATTCCGAC | 457bp |
3. Gene prediction and comparative analysis:
from the fine localization results, a total of 4 candidate genes were found in this interval, predicted by Rice Genome Annotation Project http:// Rice. Through designing a sequencing primer, a PCR method is adopted to amplify candidate genes from lg1 mutant genomes and wild type ZH11 genomes respectively for sequencing analysis. The DNA sequence of LOC _ Os02g09490 gene of lg1 mutant underwent base substitution, resulting in premature termination of translation. The Blast analysis found that LG1 gene (LOC _ Os02g 09490) was allelic with GH2 previously reported. The lg1 mutation site is inconsistent with the mutation site of the GH2 allele previously reported, and the base substitution of the mutant DNA sequence in lg1 results in premature translation termination.
The results are verified repeatedly for 3 times respectively, the same result is obtained, and the accuracy of the result is proved. Based on the gene annotation information (NCBI), the LG1 gene was predicted to encode a cinnamyl alcohol dehydrogenase.
The PCR reaction system is as follows: 2. Mu.l of 100 ng/. Mu.l rice genomic DNA,GXL Buffer 10. Mu.l, 2.5mM dNTP 4. Mu.l, 10. Mu.M primer 4. Mu.l> GXL DNA Polymerase 4μl,ddH 2 O26. Mu.l, 50. Mu.l in total. The PCR amplification conditions are specifically: pre-denaturation at 94 ℃ for 2 min; denaturation at 98 ℃ for 10 seconds, annealing at 60 ℃ for 15 seconds, extension at 68 ℃ for 70 seconds, 30 cycles, gel cutting and recovery after 1% agarose gel electrophoresis, transferring to escherichia coli, and selecting positive monoclonal sequencing. The LG1 gene sequencing primers were as follows: LG1 CX-1F.
The nucleotide sequence of the rice large-grain gene LG1 is shown as SEQ ID NO:2, the amino acid sequence of the encoded protein is shown as SEQ ID NO:4 is shown in the specification;
the nucleotide sequence of flower 11 (ZH 11) in the original wild type material japonica rice variety is SEQ ID NO:1, the amino acid sequence of the protein coded by ZH11 is shown as SEQ ID NO:3, the preparation method is as follows.
Description of the drawings:
SEQ ID NO:1 comprises a 5-end DNA sequence, an exon, an intron and a 3-end DNA sequence;
SEQ ID NO:2 comprises a 5-terminal DNA sequence, an exon, an intron and a 3-terminal DNA sequence.
Example 4, plant transformation:
the DNA sequence of ZH11 LG1 (containing SEQ ID NO: 1) was amplified using homologous recombination and PCR techniques. And recovering and purifying DNA fragments after electrophoresis, connecting products with the correct sizes of the recovered and purified fragments to a vector after enzyme digestion for escherichia coli transformation, and selecting positive monoclonal sequencing. Obtaining a correct transformation vector pCA1301-LG1, and transforming the vector into a rice mutant LG1 callus in an Agrobacterium tumefaciens strain LBA4404 by an electric shock method. The callus induced by the mutant seeds is cultured for 3 weeks by an induction culture medium, and the vigorous growing callus is selected to be used as a transformation receptor. The rice calli were infected with the LBA4404 strain containing the recombinant plasmid vector, and after co-cultivation in the dark at 25 ℃ for 3 days, they were cultured on a selection medium containing 300mg/LG 418. Selection of resistant calli was cultured on pre-differentiation medium containing 250mg/L G418 for about 10 days. The pre-differentiated calli were transferred to differentiation medium and cultured under light at 25 ℃. Obtaining resistant transgenic plants in about two months, and transplanting the resistant transgenic plants to a field for continuous growth. The plant is subjected to kernel phenotype identification and observation in the mature period, and the lg1 kernel size is recovered to be normal (figure 2), and the specific data are shown in the following table 3. It was shown that the present invention obtained transgenic rice that restored the lg1 mutant to the normal phenotype (FIG. 2).
TABLE 3
Length of grain | Grain width | Particle thickness | Thousand seed weight | |
wt | 7.16±0.16 | 3.21±0.17 | 2.46±0.22 | 26.42±0.45 |
lg1 mutant | 7.50±0.19 | 3.73±0.21 | 2.78±0.21 | 30.25±0.59 |
Resistant transgenic plants | 7.11±0.20 | 3.22±0.13 | 2.47±0.18 | 26.65±0.62 |
Finally, it is also noted that the above-mentioned list is only a few specific embodiments of the present invention. It is obvious that the invention is not limited to the above embodiments, but that many variations are possible. All modifications which can be derived or suggested by a person skilled in the art from the disclosure of the present invention are to be considered within the scope of the invention.
Sequence listing
<110> institute of Rice research in China
<120> large rice grain gene LG1 and application thereof
<160> 4
<170> SIPOSequenceListing 1.0
<210> 1
<211> 3941
<212> DNA
<213> Rice (Oryza sativa)
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ctttaattat atatactcgc tgcgggacgc accaccccct ctccagaatt cagcgagaga 60
gagcgaatca gccactgcac ttcttgttct tgttctcttc ctcctcccga atccgatcca 120
tccattggcc gccgccgccg ccgccgcctc gtgcgtcgcg tgttcgtgtc cagagacgat 180
gggcagcctc gccgccgaga agaccgtcac cgggtgggcc gccagggacg cctccggcca 240
cctcaccccc tacaactaca ccctcaggta tgatatgata tgatcgatca ccctctcctt 300
ctcgatcttg ttcttgcaag ttttgcttaa ggtcggtaga aacggaggga ttaggtttta 360
gtttctcttt tgtctcaatt ctgaaactct ggtacatgcg ttctactaca gtttagttgt 420
tcagatggtt cgtttctgaa tttctgaaaa ctgatggcgt tccagtgctt tagttgttta 480
gatgcccccg tgttttaatt tgtttgtgtt catgtcatct gcctcgagta caactcagat 540
tgtttctgcc tcttggccaa agtgtgaaga ggtgaagaga gcaattggat ttcttttttc 600
ttttttgggc tcctccattc gatccattcc acacttccca cgtgagtttg actctatttt 660
gtattcctgc tcctctccta gtataagtcc aacatgaaga gtatagtggt gtagtactag 720
tgtactacca aaactgcttc acctaccgcc aattggctat tgcaatcatc gtcacgtctt 780
ggttttgttg ggtgaaagga acagaatccc tgaatccgga gcagttgtgg caaacagaca 840
gaatgcatgg agttcttttt ggctagttgc ctggctccac atatatatat tttcagaagc 900
attctttata attttctaat atatatattt gctatgcaat tctcgtgcca tgcctcctta 960
aagaaaagga ggattatata tatatcaata atttccttga gttgtcttct tgtagaatgg 1020
aataactgaa gaagcatctt atgcaacaga tcaactaatg ttcttttttt ttcctgtatg 1080
tggtggtacc aggaagactg ggcctgaaga tgtggtggtg aaggttttgt attgtggtat 1140
ctgccatact gacatccacc aggccaagaa ccaccttggt gcttccaagt accccatggt 1200
ccctgggtaa gattattcac agttatctag atggatcatg gctcatgtca atggagatct 1260
tctactagca attgatttga tggtgtattc tatagtagac tgaagataaa cttattaact 1320
gattgggata acaaacatat cagaaaagag taggcactag taactagcaa atctggtcag 1380
tgcttttttc agatttacaa aaggcataga tgcacatgca ccactatatc acactccttt 1440
ggaacagaaa gataagttag tgcatcgctt ttagcatttt tcttttttga tcaacacctt 1500
tcatgacagg tatagtacag attagatgca ccatcagttt cagagacacg tgtaaagaaa 1560
acctcagcga gaaccttccc tgatctgatg ggctttaaat aagcacacga tagatcccca 1620
acaccaggct gaaaaatgat gcctaattag tagtaattta gtactactac accatctgca 1680
tgtcgtacta agtatgcaac aataattggt tggattaatt agttggtgac caagaattcc 1740
tttttccagt acacttccat tgtcgtacca atattcagtc atgtgttgga ttttgtcggt 1800
gaccaagaaa attctctcct cttctcgtct gcagccatga ggtggtcggc gaggtggtgg 1860
aggtcgggcc ggaggtgacc aagtacagcg ccggcgacgt cgtcggcgtc ggcgtcatcg 1920
tcggctgctg ccgcgagtgc catccgtgca aggccaatgt ggagcagtac tgcaacaaga 1980
ggatttggtc ctacaacgac gtctacaccg acggccggcc aacccagggc ggcttcgcct 2040
ccgccatggt cgtcgaccag aagtgagtgc taaacacagc tttactccaa caagtaatca 2100
ccaaccaact gatcacgata tcattatcat attatcatat tgtcagtgtt ctaggtagag 2160
aaaactagac tggacttttc cgaacatcca aagtaaagtc cagtctgtta gaaaattttg 2220
tggagaaaat gtcatctgag actgaaagag atgtcgagaa gtcggtactg tgttgagtgt 2280
aaagtgagaa gtcaccaaag ttaggtataa agtgttgatt aactgataag tcctcaccta 2340
ccccggtcca ttagtccatt tgttgttgga agaattgctt cttgaacaga gcaaaagttg 2400
gattgttgca tagcgtcgtt gtagtcagtt ggtaaactag cagagacagt gaaatgttgg 2460
gctcattttg tttggtgcgc taccaaatca ttggtagcgt caaatcatgg tcaagatttg 2520
gcactaccaa tattttagta tagttggttt ataagagcaa gttcaatagt aaagttaact 2580
gccggctcca aaattcacca cattatttaa gagataacta aacaatggta gatacagtaa 2640
ttggctacac ccatgcttta accaaggcat cctgtcaggt gttggtaact tctgcatgtt 2700
caagttctaa ttttcgattg gttgtcccga gctgcagggg gccagagcgc ctaggcgctc 2760
gtgcttgagc cggtgactgg attgtcgtcc gctttgctct ctctgctttc cctctctcta 2820
tcatatatgt taaaatatta ggtaagcagg ctattagcct gctattgtac ctgctctaag 2880
agttgtatta cctttggtaa attttgacat cattacaaac atatacatgg tactatttaa 2940
agtgccaact atttgatagg gcaaattctg gcatcaaacc aaaacatccc gtgatttttg 3000
atccacctgc ctaatctgaa agttgtcacg tcttaaaaag attaaaatat ttggttagtg 3060
agatggtaag aaactagtaa aaaaagtaat gaatttgtgt gcaggtttgt ggtgaagatc 3120
ccggcggggc tagcgccgga gcaggcggcg ccgctgctgt gcgccgggct gacggtgtac 3180
agcccactga agcacttcgg gctgatgtcg ccaggtctcc gcggcggcgt cctggggctc 3240
ggcggcgtgg ggcacatggg cgtgaaggtg gccaagtcga tggggcacca cgtgacggtg 3300
atcagctcgt cggcgaggaa gcgcggcgag gccatggacg acctgggcgc cgacgcctac 3360
ctcgtcagct ccgacgcggc ggcgatggcg gccgccggcg actcgctgga ctacatcatc 3420
gacaccgtgc cggtgcacca cccgctggag ccgtacctgg cgctgctgaa gctggacggg 3480
aagctgatcc tgatgggggt gatcaaccag ccgctgagct tcatctcgcc catggtgatg 3540
ctcggccgga aggccatcac cggcagcttc atcgggagca tggccgagac ggaggaggtg 3600
ctcaacttct gcgtcgacaa ggggctcacc tcccagatcg aggtcgtcaa gatggactac 3660
gtcaaccagg ccctcgagcg cctcgagcgc aacgacgtcc gctaccgctt cgtcgtcgac 3720
gtcgccggca gcaacatcga cgacgccgac gcgccgcccg cctgatcgcc gccgtgcacc 3780
ctcatggcag catccgtatg atccgttctt gaacttgttt gtgtgagact ctgacgactt 3840
gtccatgtaa cgtccgtgtg cctttccggt ataaattcgg cctcgcaata tatggaccat 3900
tttgcatttc cgatcaaaat catgcttgcg tttcacttgc a 3941
<210> 2
<211> 3941
<212> DNA
<213> Rice (Oryza sativa)
<400> 2
ctttaattat atatactcgc tgcgggacgc accaccccct ctccagaatt cagcgagaga 60
gagcgaatca gccactgcac ttcttgttct tgttctcttc ctcctcccga atccgatcca 120
tccattggcc gccgccgccg ccgccgcctc gtgcgtcgcg tgttcgtgtc cagagacgat 180
gggcagcctc gccgccgaga agaccgtcac cgggtgggcc gccagggacg cctccggcca 240
cctcaccccc tacaactaca ccctcaggta tgatatgata tgatcgatca ccctctcctt 300
ctcgatcttg ttcttgcaag ttttgcttaa ggtcggtaga aacggaggga ttaggtttta 360
gtttctcttt tgtctcaatt ctgaaactct ggtacatgcg ttctactaca gtttagttgt 420
tcagatggtt cgtttctgaa tttctgaaaa ctgatggcgt tccagtgctt tagttgttta 480
gatgcccccg tgttttaatt tgtttgtgtt catgtcatct gcctcgagta caactcagat 540
tgtttctgcc tcttggccaa agtgtgaaga ggtgaagaga gcaattggat ttcttttttc 600
ttttttgggc tcctccattc gatccattcc acacttccca cgtgagtttg actctatttt 660
gtattcctgc tcctctccta gtataagtcc aacatgaaga gtatagtggt gtagtactag 720
tgtactacca aaactgcttc acctaccgcc aattggctat tgcaatcatc gtcacgtctt 780
ggttttgttg ggtgaaagga acagaatccc tgaatccgga gcagttgtgg caaacagaca 840
gaatgcatgg agttcttttt ggctagttgc ctggctccac atatatatat tttcagaagc 900
attctttata attttctaat atatatattt gctatgcaat tctcgtgcca tgcctcctta 960
aagaaaagga ggattatata tatatcaata atttccttga gttgtcttct tgtagaatgg 1020
aataactgaa gaagcatctt atgcaacaga tcaactaatg ttcttttttt ttcctgtatg 1080
tggtggtacc aggaagactg ggcctgaaga tgtggtggtg aaggttttgt attgtggtat 1140
ctgccatact gacatccacc aggccaagaa ccaccttggt gcttccaagt accccatggt 1200
ccctgggtaa gattattcac agttatctag atggatcatg gctcatgtca atggagatct 1260
tctactagca attgatttga tggtgtattc tatagtagac tgaagataaa cttattaact 1320
gattgggata acaaacatat cagaaaagag taggcactag taactagcaa atctggtcag 1380
tgcttttttc agatttacaa aaggcataga tgcacatgca ccactatatc acactccttt 1440
ggaacagaaa gataagttag tgcatcgctt ttagcatttt tcttttttga tcaacacctt 1500
tcatgacagg tatagtacag attagatgca ccatcagttt cagagacacg tgtaaagaaa 1560
acctcagcga gaaccttccc tgatctgatg ggctttaaat aagcacacga tagatcccca 1620
acaccaggct gaaaaatgat gcctaattag tagtaattta gtactactac accatctgca 1680
tgtcgtacta agtatgcaac aataattggt tggattaatt agttggtgac caagaattcc 1740
tttttccagt acacttccat tgtcgtacca atattcagtc atgtgttgga ttttgtcggt 1800
gaccaagaaa attctctcct cttctcgtct gcagccatga ggtggtcggc gaggtggtgg 1860
aggtcgggcc ggaggtgacc aagtacagcg ccggcgacgt cgtcggcgtc ggcgtcatcg 1920
tcggctgctg ccgcgagtgc catccgtgca aggccaatgt ggagcagtac tgcaacaaga 1980
ggatttgatc ctacaacgac gtctacaccg acggccggcc aacccagggc ggcttcgcct 2040
ccgccatggt cgtcgaccag aagtgagtgc taaacacagc tttactccaa caagtaatca 2100
ccaaccaact gatcacgata tcattatcat attatcatat tgtcagtgtt ctaggtagag 2160
aaaactagac tggacttttc cgaacatcca aagtaaagtc cagtctgtta gaaaattttg 2220
tggagaaaat gtcatctgag actgaaagag atgtcgagaa gtcggtactg tgttgagtgt 2280
aaagtgagaa gtcaccaaag ttaggtataa agtgttgatt aactgataag tcctcaccta 2340
ccccggtcca ttagtccatt tgttgttgga agaattgctt cttgaacaga gcaaaagttg 2400
gattgttgca tagcgtcgtt gtagtcagtt ggtaaactag cagagacagt gaaatgttgg 2460
gctcattttg tttggtgcgc taccaaatca ttggtagcgt caaatcatgg tcaagatttg 2520
gcactaccaa tattttagta tagttggttt ataagagcaa gttcaatagt aaagttaact 2580
gccggctcca aaattcacca cattatttaa gagataacta aacaatggta gatacagtaa 2640
ttggctacac ccatgcttta accaaggcat cctgtcaggt gttggtaact tctgcatgtt 2700
caagttctaa ttttcgattg gttgtcccga gctgcagggg gccagagcgc ctaggcgctc 2760
gtgcttgagc cggtgactgg attgtcgtcc gctttgctct ctctgctttc cctctctcta 2820
tcatatatgt taaaatatta ggtaagcagg ctattagcct gctattgtac ctgctctaag 2880
agttgtatta cctttggtaa attttgacat cattacaaac atatacatgg tactatttaa 2940
agtgccaact atttgatagg gcaaattctg gcatcaaacc aaaacatccc gtgatttttg 3000
atccacctgc ctaatctgaa agttgtcacg tcttaaaaag attaaaatat ttggttagtg 3060
agatggtaag aaactagtaa aaaaagtaat gaatttgtgt gcaggtttgt ggtgaagatc 3120
ccggcggggc tagcgccgga gcaggcggcg ccgctgctgt gcgccgggct gacggtgtac 3180
agcccactga agcacttcgg gctgatgtcg ccaggtctcc gcggcggcgt cctggggctc 3240
ggcggcgtgg ggcacatggg cgtgaaggtg gccaagtcga tggggcacca cgtgacggtg 3300
atcagctcgt cggcgaggaa gcgcggcgag gccatggacg acctgggcgc cgacgcctac 3360
ctcgtcagct ccgacgcggc ggcgatggcg gccgccggcg actcgctgga ctacatcatc 3420
gacaccgtgc cggtgcacca cccgctggag ccgtacctgg cgctgctgaa gctggacggg 3480
aagctgatcc tgatgggggt gatcaaccag ccgctgagct tcatctcgcc catggtgatg 3540
ctcggccgga aggccatcac cggcagcttc atcgggagca tggccgagac ggaggaggtg 3600
ctcaacttct gcgtcgacaa ggggctcacc tcccagatcg aggtcgtcaa gatggactac 3660
gtcaaccagg ccctcgagcg cctcgagcgc aacgacgtcc gctaccgctt cgtcgtcgac 3720
gtcgccggca gcaacatcga cgacgccgac gcgccgcccg cctgatcgcc gccgtgcacc 3780
ctcatggcag catccgtatg atccgttctt gaacttgttt gtgtgagact ctgacgactt 3840
gtccatgtaa cgtccgtgtg cctttccggt ataaattcgg cctcgcaata tatggaccat 3900
tttgcatttc cgatcaaaat catgcttgcg tttcacttgc a 3941
<210> 3
<211> 363
<212> PRT
<213> Rice (Oryza sativa)
<400> 3
Met Gly Ser Leu Ala Ala Glu Lys Thr Val Thr Gly Trp Ala Ala Arg
1 5 10 15
Asp Ala Ser Gly His Leu Thr Pro Tyr Asn Tyr Thr Leu Arg Lys Thr
20 25 30
Gly Pro Glu Asp Val Val Val Lys Val Leu Tyr Cys Gly Ile Cys His
35 40 45
Thr Asp Ile His Gln Ala Lys Asn His Leu Gly Ala Ser Lys Tyr Pro
50 55 60
Met Val Pro Gly His Glu Val Val Gly Glu Val Val Glu Val Gly Pro
65 70 75 80
Glu Val Thr Lys Tyr Ser Ala Gly Asp Val Val Gly Val Gly Val Ile
85 90 95
Val Gly Cys Cys Arg Glu Cys His Pro Cys Lys Ala Asn Val Glu Gln
100 105 110
Tyr Cys Asn Lys Arg Ile Trp Ser Tyr Asn Asp Val Tyr Thr Asp Gly
115 120 125
Arg Pro Thr Gln Gly Gly Phe Ala Ser Ala Met Val Val Asp Gln Lys
130 135 140
Phe Val Val Lys Ile Pro Ala Gly Leu Ala Pro Glu Gln Ala Ala Pro
145 150 155 160
Leu Leu Cys Ala Gly Leu Thr Val Tyr Ser Pro Leu Lys His Phe Gly
165 170 175
Leu Met Ser Pro Gly Leu Arg Gly Gly Val Leu Gly Leu Gly Gly Val
180 185 190
Gly His Met Gly Val Lys Val Ala Lys Ser Met Gly His His Val Thr
195 200 205
Val Ile Ser Ser Ser Ala Arg Lys Arg Gly Glu Ala Met Asp Asp Leu
210 215 220
Gly Ala Asp Ala Tyr Leu Val Ser Ser Asp Ala Ala Ala Met Ala Ala
225 230 235 240
Ala Gly Asp Ser Leu Asp Tyr Ile Ile Asp Thr Val Pro Val His His
245 250 255
Pro Leu Glu Pro Tyr Leu Ala Leu Leu Lys Leu Asp Gly Lys Leu Ile
260 265 270
Leu Met Gly Val Ile Asn Gln Pro Leu Ser Phe Ile Ser Pro Met Val
275 280 285
Met Leu Gly Arg Lys Ala Ile Thr Gly Ser Phe Ile Gly Ser Met Ala
290 295 300
Glu Thr Glu Glu Val Leu Asn Phe Cys Val Asp Lys Gly Leu Thr Ser
305 310 315 320
Gln Ile Glu Val Val Lys Met Asp Tyr Val Asn Gln Ala Leu Glu Arg
325 330 335
Leu Glu Arg Asn Asp Val Arg Tyr Arg Phe Val Val Asp Val Ala Gly
340 345 350
Ser Asn Ile Asp Asp Ala Asp Ala Pro Pro Ala
355 360
<210> 4
<211> 118
<212> PRT
<213> Rice (Oryza sativa)
<400> 4
Met Gly Ser Leu Ala Ala Glu Lys Thr Val Thr Gly Trp Ala Ala Arg
1 5 10 15
Asp Ala Ser Gly His Leu Thr Pro Tyr Asn Tyr Thr Leu Arg Lys Thr
20 25 30
Gly Pro Glu Asp Val Val Val Lys Val Leu Tyr Cys Gly Ile Cys His
35 40 45
Thr Asp Ile His Gln Ala Lys Asn His Leu Gly Ala Ser Lys Tyr Pro
50 55 60
Met Val Pro Gly His Glu Val Val Gly Glu Val Val Glu Val Gly Pro
65 70 75 80
Glu Val Thr Lys Tyr Ser Ala Gly Asp Val Val Gly Val Gly Val Ile
85 90 95
Val Gly Cys Cys Arg Glu Cys His Pro Cys Lys Ala Asn Val Glu Gln
100 105 110
Tyr Cys Asn Lys Arg Ile
115
Claims (4)
1. Rice large grain geneLG1The application of the compound fertilizer in increasing seeds and yield of gramineous plants is characterized in that: rice large grain geneLG1The nucleotide sequence of (a) is shown as SEQ ID NO:2, rice large grain geneLG1The amino acid sequence of the encoded protein is shown as SEQ ID NO:4 is shown in the specification; the gramineous plant is rice.
2. Use according to claim 1, characterized in that: increase the grain length, the grain width and the grain thickness of rice grains and improve the rice yield.
3. A method for regulating and controlling the development of gramineous plant kernels is characterized in that: comprising the use of SEQ ID NO:2, or a pharmaceutically acceptable salt thereofLG1Transformation of gramineous plantsCultivating the transformed gramineous plant cells into plants, wherein the seeds of the plants are increased and the thousand seed weight is increased; the gramineous plant is rice.
4. A method of modulating grain development in a graminaceous plant as claimed in claim 3, wherein: the grain length, the grain width and the grain thickness of the grains are increased.
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