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有色稻米及其相关基因研究进展
Colored Rice and Its Related Genes: A Review
为了解有色稻米及其相关基因的研究进展，归纳了有色稻米的功能，分析了Ra、Rc、Rd、OsC1和OsB2基因对有色稻米性状的影响。这些基因通过影响花青素和原花青素的积累，决定了米粒的红、黑、紫等颜色。研究指出了相关稻米颜色基因在水稻育种中的局限性，目前的研究还不够系统，导致无法充分利用这些基因的多样性，也不能证明这些基因在水稻其他性状上的作用。笔者认为可以采用全基因组关联分析和群体遗传学方法，系统分析这些基因在不同水稻品种中的变异和表达模式，为水稻遗传改良提供数据支持，也可以利用遗传学方法和现代生物技术手段探索这些基因在水稻其他性状中的功能，为水稻的多性状改良提供理论依据。
To understand the research progress of colored rice and its related genes, the functions of colored rice were summarized, and the effects of Ra, Rc, Rd, OsC1 and OsB2 genes on the traits of colored rice were analyzed. These genes determine the red, black, and purple colors of rice grains by influencing the accumulation of anthocyanins and proanthocyanidins. In this paper, we point out the limitations of related rice color genes in rice breeding. And the current research is not systematic enough to make full use of the diversity of these genes and prove the role of these genes in other rice traits. It is believed that genome-wide association analysis and population genetics method can be used to systematically analyze the variation and expression patterns of these genes in different rice varieties, and provide data support for rice genetic improvement. We can also use genetic methods and modern biotechnology methods to explore the functions of these genes in other traits of rice, to provide theoretical basis for the improvement of multiple traits of rice.
有色稻米 / 基因 / 形成机制 / 调控机制 / 分子标记 {{custom_keyword}} /
colored rice / gene / formation mechanism / regulation mechanism / molecular marker {{custom_keyword}} /
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Red rice contains high levels of proanthocyanidins and anthocyanins, which have been recognized as health-promoting nutrients. The red coloration of rice grains is controlled by two complementary genes, Rc and Rd. The RcRd genotype produces red pericarp in wild species Oryza rufipogon, whereas most cultivated rice varieties produce white grains resulted from a 14-bp frame-shift deletion in the seventh exon of the Rc gene. In the present study, we developed a CRISPR/Cas9-mediated method to functionally restore the recessive rc allele through reverting the 14-bp frame-shift deletion to in-frame mutations in which the deletions were in multiples of three bases, and successfully converted three elite white pericarp rice varieties into red ones. Rice seeds from T in-frame Rc lines were measured for proanthocyanidins and anthocyanidins, and high accumulation levels of proanthocyanidins and anthocyanidins were observed in red grains from the mutants. Moreover, there was no significant difference between wild-type and in-frame Rc mutants in major agronomic traits, indicating that restoration of Rc function had no negative effect on important agronomic traits in rice. Given that most white pericarp rice varieties are resulted from the 14-bp deletion in Rc, it is conceivable that our method could be applied to most white pericarp rice varieties and would greatly accelerate the breeding of new red rice varieties with elite agronomic traits. In addition, our study demonstrates an effective approach to restore recessive frame-shift alleles for crop improvement.© 2019 The Authors. Plant Biotechnology Journal published by Society for Experimental Biology and The Association of Applied Biologists and John Wiley & Sons Ltd.
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Anthocyanins cause purple, brown or red colors in various tissues of rice plants, but the specific determinant factors and regulatory systems for anthocyanin biosynthesis in almost all tissues remain largely unknown. In the present study, we mapped and isolated two complementary genes, OsC1 encoding a R2R3-MYB transcriptional factor and OsDFR encoding a dihydroflavonol 4-reductase, which are responsible for the purple coloration of apiculi and stigmas in indica cultivar Xieqingzao by the map-based cloning strategy. We also identified two tissue-specific pigmentation genes, OsPa for apiculi and OsPs for stigmas, by phylogenetic analysis of all anthocyanin biosynthesis-associated bHLH transcriptional factors in maize and rice, CRISPR/Cas9 knockout and transcriptional expression analysis. The OsC1, OsPa and OsPs proteins are all localized in the nucleus while the OsDFR protein is localized in the nucleus and cytoplasm, and the OsC1 and OsDFR genes are preferentially strongly expressed in both purple-colored tissues while the OsPa and OsPs genes are preferentially strongly expressed in apiculi and stigmas, respectively. OsC1 specifically interacts with OsPa or OsPs to activate OsDFR and other anthocyanin biosynthesis genes, resulting in purple-colored apiculi or stigmas. OsC1 itself does not produce color but can produce brown apiculi when functioning together with OsPa. Loss of function of OsDFR alone leads to brown apiculi and straw-white stigmas. Genotyping and phenotyping of a panel of 176 rice accessions revealed diverse genotypic combinations of OsC1, OsDFR, OsPa and OsPs that enable accurate prediction of their apiculus and stigma pigmentation phenotypes, thus validating the general applicability of the OsC1-OsDFR-OsPa and OsC1-OsDFR-OsPs models to natural populations. Our findings disclosed the biological functions of OsC1, OsPa and OsPs, and shed light on the specific regulatory systems of anthocyanin biosynthesis in apiculi and stigmas, a further step in understanding the regulatory network of anthocyanin biosynthesis in rice.
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Anthocyanin pigmentation provides an excellent system with which to study the regulation of gene expression in higher plants. In this study, OsPR1.1 promoter was isolated and the promoter activity was monitored using a reporter gene OSB2, which encodes a transcription factor for anthocyanin synthesis in rice plants. We introduced PR::OSB2 plasmid into an isogenic Taichung 65, no. 99-962 T-65 CBA B9F5 (T65 CBA), rice line (Oryza sativa L.) and found that the transgenic rice plants exhibited anthocyanin accumulation by the induced expression of OSB2 after chemical treatments with methyl jasmonate (MeJA) and 2,6-dichloroisonicotinic acid (DCINA). The shoots of the PR::OSB2 transgenic rice plants changed color to red after application of the chemicals accompanying with the increased anthocyanin content to approximately 5-fold by MeJA and 2-fold by DCINA, respectively. The anthocyanin accumulation was consistent with the increase of the expression of OSB2 and anthocyanidin synthase (ANS). This color change system could provide a useful and easy way to produce transgenic plants for monitoring of chemicals in the environment.
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水稻花青素的生物合成受很多因素的影响,最根本的影响因素是结构基因和调控基因的表达水平不同,导致色素积累的差异性。本文通过半定量PCR方法研究一些结构基因和调节基因的时空表达差异性。发现OsPAL, OsCHS和OsCHI为花青素的基本表达基因,在叶片、茎、花期及花后1~2周的种子中均有表达。OsF3H, OsF3’H和OsDFR不在茎中表达,在叶片、花期和花后1~2周内均有表达,且在花后1~2周内,其表达量与时间成正比。OsANS在黑米和紫叶水稻的叶片和花中有少量表达,在花后1~2周内,仅在黑米中表达,且表达量与时间成正比。OsB1和OsB2特异性地在紫叶水稻的叶片中表达,OsC1也特异地在花期时表达。说明水稻不同部位的花青素积累受控于不同的基因,具有一定的时空表达特异性。
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