WO2020207510A2 - Application of gene in boosting rice grain yield - Google Patents
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- the invention relates to genes related to rice grain yield and applications thereof, and belongs to the field of crop molecular genetic breeding.
- Rice is an important food crop, and high-yield breeding is an effective guarantee to ensure my country's food security and sustainable agricultural development.
- the yield of rice is closely related to the grain weight of the grain. Kernel weight refers to the weight of each seed, which is largely determined by the size of the seed and the degree of filling. Therefore, discovering genes that can increase rice grain weight in rice has great application value for high-yield rice breeding.
- Rice yield is determined by the three elements of tiller number, grain number per panicle and grain weight.
- the so-called tiller number refers to the number of branches produced by each rice plant to the later stage, while the grain number per ear refers to the number of grains on each branch, and the grain weight refers to the weight of each seed.
- the grain weight is determined by the size of the seed and the degree of filling.
- the number of tillers, grains per panicle, and grain weight ultimately determine the yield of rice, in actual breeding work, there is a very strong negative correlation between the three. That is to say, the increase in the number of tillers often leads to a decrease in the number of grains per panicle. Grain becomes smaller and the number of grains per spike is increased, which often leads to a decrease in the number of tillers and grains, and a larger grain leads to a decrease in the number of tillers or grains per spike.
- the technical problem to be solved by the present invention is to provide a gene for promoting rice grain yield and its use.
- the present invention provides an application of a gene in promoting (increasing) rice grain yield, and the nucleotide sequence of the gene is shown in SEQ ID NO: 2 or SEQ ID NO: 3.
- mutant-1 SEQ ID NO: 2
- mutant-2 SEQ ID NO: 3
- the two mutants of the Os05g0462000 gene obtained above and the wild-type control Nipponbare were planted in rice fields, and it was found that the thousand-grain weight of the mutant plants increased significantly ( Figure 2). Therefore, the mutant gene can increase the grain yield of rice and has the potential for high-yield rice breeding applications. Transplanting the mutant gene into conventional cultivated rice or super hybrid rice varieties will increase the yield of rice.
- Figure 1 shows the mutation site sequence analysis of rice Os05g0462000 gene mutant.
- Figure 2 is a comparison of the flat area of 300 seeds of rice Os05g0462000 gene mutant and wild-type control.
- Figure 3 shows the thousand-grain weight comparison of rice Os05g0462000 gene mutant and wild-type control.
- Figure 4 is a comparison of yield per plant between rice Os05g0462000 gene mutant and wild-type control.
- sgRNAs were designed through the CRISPR Design program (http://crispr.mit.edu/).
- the sequence is sgRNA -1: 5'-GAACAAGGAGGAGTGCATGG-3', sgRNA-2: 5'-GATGTTGGCATGCTTCTCCAGGG-3'; and synthesize the sequence of sgRNA in a biotechnology company (such as Shanghai Invitrogen).
- Example 1 Rice tissue culture and vector transgene, according to the document "A protocol for Agrobacterium-mediated transformation in rice” (quoted from: Nat Protoc. 2006, 1(6): 2796-2802.), the method constructed in Example 1 The vector was genetically transformed into a wild-type rice variety Nipponbare (English name: Nipponbare) to obtain transgenic rice with Os05g0462000 gene mutation.
- the genomic DNA of genetically modified rice was extracted by the SDS method: 0.1g of rice leaves were taken, ground with liquid nitrogen, and 600 ⁇ l of extract (0.1mol/L Tris-Cl pH8.0, 500mmol/L NaCl, 1.25g/L SDS) was added, Incubate at 65°C for 30 min. Add 200 ⁇ l 5mol/L KAC, mix well, and ice bath for 30min. Then add 500 ⁇ l chloroform, mix well, centrifuge at 10000r/min for 5min, take the supernatant, add 2/3 of the supernatant volume of isopropanol, mix well, centrifuge at 12000r/min for 5min. Discard the supernatant, wash the pellet with 70% ethanol, invert to dry, and add 100 ⁇ l TE to dissolve the DNA.
- extract 0.1mol/L Tris-Cl pH8.0, 500mmol/L NaCl, 1.25g/L SDS
- the primer sequences F:5'-AGATAGATAAGTAAGCAGTGAG-3' and R:5'-AGCTAGCTCTCGTCGTCGTC-3' of the Os05g0462000 gene were amplified by PCR, and the primer DNA was synthesized in a biotechnology company (such as Shanghai Invitrogen). 20 ⁇ l system for PCR amplification, containing 25-50ng template DNA, 100 ⁇ mol/L each dNTP, 2 ⁇ l 10 ⁇ Buffer, 0.2 ⁇ mol/L primer F and R, 1UintTaq DNA polymerase.
- the amplification procedure is: pre-denaturation at 94°C for 5 min; denaturation at 94°C for 30 sec, annealing at 55°C for 30 sec, extension at 72°C for 60 sec, 35 cycles; extension at 72°C for 5 min.
- the PCR products were electrophoresed on a 1% agarose gel containing ethidium bromide (EB) for 30 min.
- the PCR product was sequenced and analyzed in a biotechnology company (such as Shanghai Invitrogen) to obtain the Os05g0462000 gene sequence of the transgenic rice, and compared and analyzed with SeqMan software, and two homozygous mutants of the Os05g0462000 gene mutant-1 (from sgRNA-1) were identified And mutant-2 (from sgRNA-2), both lead to a frameshift mutation in the Os05g0462000 gene ( Figure 1), resulting in a loss of function of the gene.
- the nucleotide sequence of mutant-1 is shown in SEQ ID NO:2, and the nucleotide sequence of mutant-2 is shown in SEQ ID NO:3.
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Abstract
The present invention pertains to a rice grain yield-related gene and an application thereof, belonging to the field of crop molecular genetic breeding. Specifically, disclosed is an application of a gene in boosting rice grain yield, the nucleotide sequence of the gene being as shown in SEQ ID NO: 2 or SEQ ID NO: 3. The gene can be used for promoting an increase in rice grain yield, and can also be used for promoting an increase in grain size.
Description
本发明涉及水稻籽粒产量相关基因及其应用，属于作物分子遗传育种领域。The invention relates to genes related to rice grain yield and applications thereof, and belongs to the field of crop molecular genetic breeding.
水稻是重要的粮食作物，高产育种是确保我国粮食安全和农业可持续发展的有效保障。水稻的产量与籽粒的粒重密切相关。粒重是指每粒种子的重量，很大程度上粒重由种子的大小和灌浆程度所决定。因此，在水稻中发掘能够提高水稻粒重的基因，对水稻高产育种有很大的应用价值。Rice is an important food crop, and high-yield breeding is an effective guarantee to ensure my country's food security and sustainable agricultural development. The yield of rice is closely related to the grain weight of the grain. Kernel weight refers to the weight of each seed, which is largely determined by the size of the seed and the degree of filling. Therefore, discovering genes that can increase rice grain weight in rice has great application value for high-yield rice breeding.
水稻产量由分蘖数、穗粒数和粒重三要素所决定。所谓分蘖数是指每一株水稻发育到后期所产生的分枝数，而穗粒数是指每一分枝上拥有的籽粒数，粒重则是指每粒种子的重量，很大程度上粒重由种子的大小和灌浆程度所决定。尽管分蘖数、穗粒数和粒重最终决定了水稻产量，然而，在实际育种工作中，这三者间却存在非常强的负相关，也就是说，分蘖数增加往往导致穗粒数减少或籽粒变小，穗粒数增加常常会导致分蘖数减少、籽粒变小，籽粒变大导致分蘖数减少或穗粒数减少等。Rice yield is determined by the three elements of tiller number, grain number per panicle and grain weight. The so-called tiller number refers to the number of branches produced by each rice plant to the later stage, while the grain number per ear refers to the number of grains on each branch, and the grain weight refers to the weight of each seed. The grain weight is determined by the size of the seed and the degree of filling. Although the number of tillers, grains per panicle, and grain weight ultimately determine the yield of rice, in actual breeding work, there is a very strong negative correlation between the three. That is to say, the increase in the number of tillers often leads to a decrease in the number of grains per panicle. Grain becomes smaller and the number of grains per spike is increased, which often leads to a decrease in the number of tillers and grains, and a larger grain leads to a decrease in the number of tillers or grains per spike.
201711469003X的发明《水稻高光合效率的基因及其应用》虽然告知了敲除Os05g0462000基因能增加水稻叶片的叶绿素含量，即，敲除Os05g0462000基因能有效促进水稻光合效率。但是，并没有告知敲除Os05g0462000基因与水稻籽粒的粒径大小以及与水稻籽粒产量的关系。Although the invention of 201711469003X "Genes for High Photosynthetic Efficiency in Rice and Their Applications" informs that knocking out the Os05g0462000 gene can increase the chlorophyll content of rice leaves, that is, knocking out the Os05g0462000 gene can effectively promote the photosynthetic efficiency of rice. However, it did not inform the relationship between the knockout of Os05g0462000 gene and rice grain size and rice grain yield.
发明内容Summary of the invention
本发明要解决的技术问题是提供一种促进水稻籽粒产量基因及其用途。The technical problem to be solved by the present invention is to provide a gene for promoting rice grain yield and its use.
为了解决上述技术问题，本发明提供一种基因在促进(提升)水稻籽粒产量上的应用，该基因的核苷酸序列如SEQ ID NO：2或者SEQ ID NO：3所示。In order to solve the above technical problems, the present invention provides an application of a gene in promoting (increasing) rice grain yield, and the nucleotide sequence of the gene is shown in SEQ ID NO: 2 or SEQ ID NO: 3.
作为本发明的基因在促进水稻籽粒产量上的应用的改进：促进水稻籽粒产量增加；促进籽粒的粒径增大。As an improvement in the application of the gene of the present invention in promoting rice grain yield: promoting an increase in rice grain yield; promoting an increase in grain size.
本发明的技术方案具体如下：The technical scheme of the present invention is specifically as follows:
利用CRISPR/Cas9技术，设计特异性靶向Os05g0462000基因(SEQ ID NO:1)的sgRNA，对水稻品种日本晴的Os05g0462000基因进行敲除突变，获得了该基因的2个敲除突变体mutant-1与mutant-2。对这些突变体的PCR扩增及测序分析，表明突变位点发生在 Os05g0462000基因的不同位置(图1)，但是都导致了该基因发生移码突变，而造成该基因的功能缺失。Using CRISPR/Cas9 technology, we designed an sgRNA that specifically targets the Os05g0462000 gene (SEQ ID NO:1), and performed a knockout mutation on the Os05g0462000 gene of the rice variety Nipponbare, and obtained two knockout mutants of this gene, mutant-1 and mutant-2. The PCR amplification and sequencing analysis of these mutants showed that the mutation sites occurred in different positions of the Os05g0462000 gene (Figure 1), but they all caused the frameshift mutation of the gene, which resulted in the loss of function of the gene.
上述2个敲除突变体的序列分别为：mutant-1(SEQ ID NO:2)，mutant-2(SEQ ID NO:3)。The sequences of the above two knockout mutants are: mutant-1 (SEQ ID NO: 2) and mutant-2 (SEQ ID NO: 3).
将以上获得的Os05g0462000基因两种突变体与野生型对照日本晴种植在水稻大田，发现突变体的植株的千粒重显著性增加(图2)。因此，该突变基因可增加水稻的籽粒产量，具有水稻高产育种应用的潜力，将该突变基因转育到常规栽培稻或超级杂交稻品种中，将能提高水稻的产量。The two mutants of the Os05g0462000 gene obtained above and the wild-type control Nipponbare were planted in rice fields, and it was found that the thousand-grain weight of the mutant plants increased significantly (Figure 2). Therefore, the mutant gene can increase the grain yield of rice and has the potential for high-yield rice breeding applications. Transplanting the mutant gene into conventional cultivated rice or super hybrid rice varieties will increase the yield of rice.
下面结合附图对本发明的具体实施方式作进一步详细说明。The specific embodiments of the present invention will be described in further detail below with reference to the accompanying drawings.
图1为水稻Os05g0462000基因突变体的突变位点序列分析。Figure 1 shows the mutation site sequence analysis of rice Os05g0462000 gene mutant.
图2为水稻Os05g0462000基因突变体与野生型对照的300粒种子平铺面积比较。Figure 2 is a comparison of the flat area of 300 seeds of rice Os05g0462000 gene mutant and wild-type control.
图3为水稻Os05g0462000基因突变体与野生型对照的千粒重比较。Figure 3 shows the thousand-grain weight comparison of rice Os05g0462000 gene mutant and wild-type control.
图4为水稻Os05g0462000基因突变体与野生型对照的单株产量比较。Figure 4 is a comparison of yield per plant between rice Os05g0462000 gene mutant and wild-type control.
图1-4中，日本晴：野生型对照品种；mutant-1与mutant-2：水稻Os05g0462000基因敲除突变体；图3和图4中的数值为平均值±标准差，**表示在水稻Os05g0462000基因突变体与野生型对照日本晴比较中，t检验存在极显著性(P<0.01)差异。In Figure 1-4, Nipponbare: wild-type control variety; mutant-1 and mutant-2: rice Os05g0462000 knockout mutants; the values in Figures 3 and 4 are the mean ± standard deviation, and ** indicates the rice Os05g0462000 Compared with the wild-type control Nipponbare, there was a significant difference in t test (P<0.01).
实施例1、Os05g0462000基因的CRISPR/Cas9载体构建Example 1. CRISPR/Cas9 vector construction of Os05g0462000 gene
根据Os05g0462000基因的核苷酸序列(SEQ ID NO:1)，通过CRISPR Design程序(http://crispr.mit.edu/)设计了两个CRISPR/Cas9编辑的靶点序列sgRNA，其序列为sgRNA-1：5'-GAACAAGGAGGAGTGCATGG-3'，sgRNA-2：5'-GATGTTGGCATGCTTCTCCAGGG-3'；并在生物技术公司(如上海Invitrogen)合成sgRNA的序列。According to the nucleotide sequence of the Os05g0462000 gene (SEQ ID NO: 1), two CRISPR/Cas9 editing target sequence sgRNAs were designed through the CRISPR Design program (http://crispr.mit.edu/). The sequence is sgRNA -1: 5'-GAACAAGGAGGAGTGCATGG-3', sgRNA-2: 5'-GATGTTGGCATGCTTCTCCAGGG-3'; and synthesize the sequence of sgRNA in a biotechnology company (such as Shanghai Invitrogen).
采用CRISPR/Cas9试剂盒(Biogle，Cat#BGK03)分别构建了2个相应的载体，方法按照产品说明。Two corresponding vectors were constructed using CRISPR/Cas9 kit (Biogle, Cat#BGK03), and the method was in accordance with the product instructions.
实施例2、CRISPR/Cas9载体的水稻遗传转化Example 2. Rice genetic transformation of CRISPR/Cas9 vector
水稻的组织培养与载体转基因，根据文献《A protocol for Agrobacterium-mediated transformation in rice》(引自：Nat Protoc.2006,1(6):2796-2802.)的方法，将实施例1中构建的载体遗传转化到野生型水稻品种日本晴(英文名：Nipponbare)中，获得Os05g0462000基因突变的转基因水稻。Rice tissue culture and vector transgene, according to the document "A protocol for Agrobacterium-mediated transformation in rice" (quoted from: Nat Protoc. 2006, 1(6): 2796-2802.), the method constructed in Example 1 The vector was genetically transformed into a wild-type rice variety Nipponbare (English name: Nipponbare) to obtain transgenic rice with Os05g0462000 gene mutation.
实施例3、Os05g0462000基因突变株的测序分析Example 3 Sequencing analysis of Os05g0462000 gene mutant strain
采用SDS法提取转基因水稻的基因组DNA：取水稻叶片0.1g，用液氮研磨后，加600μl提取液(0.1mol/L Tris-Cl pH8.0，500mmol/L NaCl，1.25g/L SDS)，65℃温育30min。加200μl 5mol/L KAC，混匀后，冰浴30min。再加500μl氯仿，混匀，10000r/min离心5min，取上清，加2/3上清体积的异丙醇，混匀，12000r/min离心5min。弃去上清，用70％乙醇洗涤沉淀，倒置晾干，加100μl TE溶解DNA。The genomic DNA of genetically modified rice was extracted by the SDS method: 0.1g of rice leaves were taken, ground with liquid nitrogen, and 600μl of extract (0.1mol/L Tris-Cl pH8.0, 500mmol/L NaCl, 1.25g/L SDS) was added, Incubate at 65°C for 30 min. Add 200μl 5mol/L KAC, mix well, and ice bath for 30min. Then add 500μl chloroform, mix well, centrifuge at 10000r/min for 5min, take the supernatant, add 2/3 of the supernatant volume of isopropanol, mix well, centrifuge at 12000r/min for 5min. Discard the supernatant, wash the pellet with 70% ethanol, invert to dry, and add 100 μl TE to dissolve the DNA.
用PCR扩增Os05g0462000基因的引物序列F:5’-AGATAGATAAGTAAGCAGTGAG-3’和R:5’-AGCTAGCTCTCGTCGTCGTC-3’，在生物技术公司(如上海Invitrogen)合成引物的DNA。PCR扩增用20μl体系，含25～50ng模板DNA、100μmol/L each dNTP、2μl 10×Buffer、0.2μmol/L引物F与R、1Uint Taq DNA聚合酶。扩增程序为：94℃预变性5min；94℃变性30sec，55℃退火30sec，72℃延伸60sec，35个循环；72℃延伸5min。PCR产物在含溴化乙锭(EB)的1％琼脂糖凝胶中电泳30min。The primer sequences F:5'-AGATAGATAAGTAAGCAGTGAG-3' and R:5'-AGCTAGCTCTCGTCGTCGTC-3' of the Os05g0462000 gene were amplified by PCR, and the primer DNA was synthesized in a biotechnology company (such as Shanghai Invitrogen). 20μl system for PCR amplification, containing 25-50ng template DNA, 100μmol/L each dNTP, 2μl 10×Buffer, 0.2μmol/L primer F and R, 1UintTaq DNA polymerase. The amplification procedure is: pre-denaturation at 94°C for 5 min; denaturation at 94°C for 30 sec, annealing at 55°C for 30 sec, extension at 72°C for 60 sec, 35 cycles; extension at 72°C for 5 min. The PCR products were electrophoresed on a 1% agarose gel containing ethidium bromide (EB) for 30 min.
PCR产物在生物技术公司(如上海Invitrogen)进行测序分析，获得转基因水稻的Os05g0462000基因序列，用SeqMan软件进行比对分析，鉴定了Os05g0462000基因2种纯合突变体mutant-1(来自sgRNA-1)与mutant-2(来自sgRNA-2)，都导致了Os05g0462000基因发生移码突变(图1)，而造成该基因的功能缺失。突变体mutant-1的核苷酸序列如SEQ ID NO:2所述，突变体mutant-2的核苷酸序列如SEQ ID NO:3所述。The PCR product was sequenced and analyzed in a biotechnology company (such as Shanghai Invitrogen) to obtain the Os05g0462000 gene sequence of the transgenic rice, and compared and analyzed with SeqMan software, and two homozygous mutants of the Os05g0462000 gene mutant-1 (from sgRNA-1) were identified And mutant-2 (from sgRNA-2), both lead to a frameshift mutation in the Os05g0462000 gene (Figure 1), resulting in a loss of function of the gene. The nucleotide sequence of mutant-1 is shown in SEQ ID NO:2, and the nucleotide sequence of mutant-2 is shown in SEQ ID NO:3.
实施例4、转基因水稻的农艺性状鉴定Example 4 Identification of Agronomic Characters of Transgenic Rice
在相同的实验条件下(水稻生长最适宜的季节)，将Os05g0462000基因两个突变体mutant-1与mutant-2及野生型对照日本晴种植在大田中，成熟后每个品种随机取30株，按照文献《中国稻种资源》(引自：中国农业科技出版社，1993)评价标准，测量水稻的产量等性状，进行3次生物学重复。采用t-Test对突变体与野生型的测量数据进行显著性差异的分析。结果表明，Os05g0462000基因的两个突变体mutant-1与mutant-2的籽粒大小显著性大于野生型对照日本晴(图2)；野生型对照的千粒重为23.0±0.28g，而Os05g0462000基因突变体mutant-1与mutant-2的千粒重分别为24.9±0.29g和24.7±0.41g，极显著(P<0.01)高出野生型对照8.3％和7.4％(图3)；野生型对照的单株产量为22.9±0.98g，而Os05g0462000基因突变体mutant-1与mutant-2的千粒重分别为25.9±0.78g和26.5±0.86g，极显著(P<0.01)高出野生型对照13.1％和25.7％(图4)。因此，得知：Os05g0462000基因突变不但能促进籽粒增大，还能促进水稻籽粒的产量。Under the same experimental conditions (the most suitable season for rice growth), two mutants of Os05g0462000 gene mutant-1 and mutant-2 and wild-type control Nipponbare were planted in the field. After maturity, 30 plants of each variety were randomly selected, according to The document "Chinese Rice Seed Resources" (quoted from: China Agricultural Science and Technology Press, 1993) evaluates the standard, measures rice yield and other traits, and performs 3 biological replicates. Use t-Test to analyze the significant difference between the mutant and wild-type measurement data. The results showed that the grain size of the two mutants of Os05g0462000 gene mutant-1 and mutant-2 was significantly larger than that of the wild-type control Nipponbare (Figure 2); the thousand-grain weight of the wild-type control was 23.0±0.28g, while the Os05g0462000 gene mutant mutant- The thousand-grain weights of 1 and mutant-2 were 24.9±0.29g and 24.7±0.41g, respectively, which were extremely significantly (P<0.01) higher than the wild-type control by 8.3% and 7.4% (Figure 3); the wild-type control yield per plant was 22.9 ±0.98g, while the thousand-grain weights of mutant-1 and mutant-2 of Os05g0462000 gene were 25.9±0.78g and 26.5±0.86g, which were significantly higher (P<0.01) than the wild-type control by 13.1% and 25.7% (Figure 4) ). Therefore, it is known that: Os05g0462000 gene mutation can not only promote grain enlargement, but also promote rice grain yield.
最后，还需要注意的是，以上列举的仅是本发明的若干个具体实施例。显然，本发明不 限于以上实施例，还可以有许多变形。本领域的普通技术人员能从本发明公开的内容直接导出或联想到的所有变形，均应认为是本发明的保护范围。Finally, it should be noted that the above-listed are only a few specific embodiments of the present invention. Obviously, the present invention is not limited to the above embodiments, and many variations are possible. All modifications that can be directly derived or associated by those of ordinary skill in the art from the disclosure of the present invention should be considered as the protection scope of the present invention.
Claims (3)
- 基因在促进水稻籽粒产量上的应用，其特征是：所述基因的核苷酸序列如SEQ ID NO：2或者SEQ ID NO：3所示。The application of the gene in promoting rice grain yield is characterized in that the nucleotide sequence of the gene is shown in SEQ ID NO: 2 or SEQ ID NO: 3.
- 根据权利要求1所述的基因在促进水稻籽粒产量上的应用，其特征是：促进水稻籽粒产量增加。The application of the gene in promoting rice grain yield according to claim 1, characterized in that it promotes an increase in rice grain yield.
- 根据权利要求1或2所述的基因在促进水稻籽粒产量上的应用，其特征是：促进籽粒的粒径增大。The use of the gene according to claim 1 or 2 in promoting rice grain yield, characterized in that it promotes the increase of grain size.
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