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稻米品质研究进展
Research Progress on Rice Quality
笔者简述了国内外稻米品质的研究现状和发展趋势,总结了稻米品质形成的遗传基础、调控分子机制以及优化稻米品质的策略,揭示了淀粉、蛋白质、脂质、香气及颜色相关基因与稻米品质之间的关联,阐述了基因编辑技术、分子标记辅助技术2种提升稻米品质的具体对策。指出稻米品质研究过程中面临的稻米品质与产量不平衡、稻米品质改良技术转化与推广难、育种过程中水稻存在地区适应性、育种成本高周期长以及消费者对改良稻米的接受程度等挑战,并从多方合作和资源共享、强化基础研究和创新技术、高效筛选和评价方法、考虑多样化需求以及推动产学研结合与技术转化等方面提出应对措施,以期为优化水稻品质和培育高品质稻米品种提供理论参考。
In this paper, the research status and development trend of rice quality at home and abroad were reviewed, the genetic basis of rice quality formation, regulatory molecular mechanisms and strategies for optimizing rice quality were summarized, and the correlation between rice quality and related genes such as starch, protein, lipid, aroma and color was revealed. Two specific countermeasures to improve rice quality, gene editing technology and molecular marker-assisted technology, were described. The challenges in rice quality research, such as imbalance between rice quality and yield, difficulty in transforming and popularizing rice quality improvement technology, regional adaptability of rice in breeding process, high breeding cost and long breeding cycle, and consumer acceptance of improved rice, were pointed out. Countermeasures were put forward from the aspects of multi-party cooperation and resource sharing, strengthening basic research and innovative technology, efficient screening and evaluation methods, considering diversified needs, and promoting the integration of industry-university-research and technology transformation, so as to provide theoretical guidance for optimizing rice quality and cultivating high-quality rice varieties.
水稻 / 稻米品质 / 育种 / 遗传基础 / 分子机制 {{custom_keyword}} /
rice / rice quality / breeding / genetic basis / molecular mechanism {{custom_keyword}} /
表1 不同地膜覆盖对大棚草莓植株农艺性状的影响 cm |
处理 | 始花期 | 采收末期 | 株高增长量 | 冠幅增长量 | |||
---|---|---|---|---|---|---|---|
株高 | 冠幅 | 株高 | 冠幅 | ||||
黑膜(CK) | 20.8 | 40.9 | 35.1 | 46.4 | 14.3 | 5.5 | |
银黑双色膜 | 19.6 | 39.9 | 28.3 | 46.5 | 8.7 | 6.6 |
表2 不同地膜覆盖对大棚草莓品质的影响 |
处理 | 早期 | 后期 | |||||||
---|---|---|---|---|---|---|---|---|---|
可溶性固形物 SSC% | 酸度 | 糖酸比 | 硬度 | 可溶性固形物 SSC% | 酸度 | 糖酸比 | 硬度 | ||
黑膜(CK) | 11.4 | 0.9 | 12.6 | 2.94 | 9.6 | 0.8 | 11.7 | 1.84 | |
银黑双色膜 | 12.5* | 0.9 | 13.7* | 3.25 | 9.8 | 0.7* | 14.8* | 2.07* |
注*表示通过0.05信度的显著性检验。 |
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【目的】挖掘Wx新等位变异,明确Wx410新等位基因对稻米品质性状的影响。【方法】以Wx<sup>lv</sup>、Wx<sup>a</sup>和Wx<sup>b</sup>等位基因为模板,利用PCR进行第10外显子第101位碱基的A-G单点突变,分别构建了不同Wx等位背景下的Wx410定点突变植物表达载体pEGFC-Wx<sup>lv</sup>410、pEGFC-Wx<sup>a</sup>410和pEGFC-Wx<sup>b</sup>410,阳性对照组载体分别为pEGFC-Wx<sup>lv</sup>、pEGFC-Wx<sup>a</sup>和pEGFC-Wx<sup>b</sup>。通过转化糯稻品种苏御糯,分析该位点的变异对稻米品质的遗传效应。【结果】花后7 d和14 d,转基因植株pEGFC-Wx<sup>lv</sup>410,pEGFC-Wx<sup>a</sup>410及pEGFC-Wx<sup>b</sup>410的胚乳Wx基因表达量较各自的阳性对照材料无显著变化,而颗粒结合淀粉合酶活性极显著降低;转基因植株直链淀粉含量较野生型显著降低,而糊化温度无明显变化;pEGFC-Wx<sup>a</sup>410及pEGFC-Wx<sup>b</sup>410的胶稠度较各自的阳性对照材料显著升高,而pEGFC-Wx<sup>lv</sup>410的胶稠度较其阳性对照材料显著降低。【结论】pEGFC-Wx410为水稻淀粉合成的一个新的功能等位基因,控制的直链淀粉含量为4%~6%,刚好弥补了目前所鉴定的复等位基因所调控的直链淀粉含量在这个范围内的空缺,为稻米食味和加工相关品质改良提供更丰富的遗传资源。
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【目的】分析在同一主效基因(Wx<sup>mp</sup>)背景下可溶性淀粉合成酶基因SSⅡa和去分支酶基因PUL对稻米蒸煮食味品质的影响,以期为水稻品质遗传改良提供依据。【方法】选择在SSⅡa和PUL存在多态性而其他淀粉合成酶相关基因没有多态性的半糯品系宁0145和粳稻品种武运粳21进行杂交,获得F<sub>2</sub>群体与F<sub>3</sub>株系。利用分子标记,选择含有Wx<sup>mp</sup>基因的F<sub>2</sub>单株与F<sub>3</sub>株系,将这些F<sub>2</sub>单株与F<sub>3</sub>株系分成SSⅡa<sup>n</sup>PUL<sup>n</sup>、SSⅡa<sup>n</sup>PUL<sup>w</sup>、SSⅡa<sup>w</sup>PUL<sup>n</sup>和SSⅡa<sup>w</sup>PUL<sup>w</sup>4种基因型(<sup>n</sup>和<sup>w</sup>分别表示该基因来源于宁0145和武运粳21),分析不同基因型蒸煮食味品质性状的差异,探讨同一Wx<sup>mp</sup>基因背景下不同SSⅡa和PUL等位基因对蒸煮食味品质性状的影响。【结果】不同基因型间蒸煮食味品质性状均存在显著差异,来源于武运粳21的SSⅡa<sup>w</sup>基因和PUL<sup>w</sup>基因分别使直链淀粉含量增加0.29%~1.00%和0.62%~1.18%,且PUL的效应大于SSⅡa,两者间存在互作效应。SSⅡa<sup>w</sup>基因和PUL<sup>w</sup>基因降低胶稠度和崩解值,提高了热浆黏度、冷胶黏度、消减值和回复值,对糊化温度、峰值黏度和峰值时间的作用较小。【结论】明确了Wx<sup>mp</sup>背景下SSⅡa和PUL基因对稻米蒸煮食味品质的遗传效应,该研究结果为SSⅡa和PUL基因的分子标记辅助选择改良稻米品质提供了理论依据。
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【目的】探讨优良食味水稻品种的籽粒蛋白质积累特征及其对氮素水平的响应。【方法】以食味值不同的常规粳稻和杂交稻为材料,在结实期设置不同氮素施用水平处理,分析不同类型品种在不同氮素水平下的蒸煮食味品质及其与稻米蛋白质及其组分含量的关系。进一步分析各品种在不同氮素水平下稻穗不同部位的氨基酸含量及籽粒蛋白质含量在结实期的动态变化,总结优良食味水稻品种的籽粒蛋白质积累特征及其对氮素水平的响应特征。【结果】优良食味水稻品种籽粒蛋白质含量较低,且随着氮素水平的增加而上升;优良食味水稻崩解值较高,消减值较低;蒸煮食味品质受氮素水平影响较小。优良食味水稻品种蛋白组分含量较低,且稻米蛋白质含量与食味品质呈显著负相关。在常规粳稻中,稻米食味值与清蛋白、球蛋白和醇溶蛋白含量均显著负相关;在杂交稻中,稻米食味值与醇溶蛋白和谷蛋白含量显著负相关。优良食味水稻品种籽粒充实过程中游离氨基酸含量较低,并呈现较低水平的蛋白质积累。而食味较差品种灌浆期籽粒的氨基酸含量较高,成熟籽粒蛋白质含量也较高,且氮素供应水平提升其籽粒蛋白质含量的效应更为显著。【结论】优良食味水稻品种的籽粒蛋白质含量较低,与其充实过程中蛋白质积累水平较低有紧密关系,且受氮素水平影响较小。
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Aromatic rice belongs to a small but important sub-group of rice, which is highly regarded for its excellent aroma and superior grain quality. Aromatic rice, especially Basmati- and Jasmine-type rice, is being traded at a high price in the local and global markets. Genetically, rice aroma is a phenotypical expression of spontaneous recessive mutations of the OsBadh2 gene (also known as fgr / badh2 / osbadh2 / os2AP gene). These mutations inhibit the flow of γ-aminobutyraldehyde (GAB-ald) to γ-aminobutyric acid (GABA), and consequently, the accumulated GAB-ald is diverted to a potent flavour component 2-acetyl-1-pyrroline (2AP) by a non-enzymatic reaction with methylglyoxal. The natural incidence of non-functional osbadh2 mutation along with selection and nursing by the farmer from the ancient time makes rice aroma as a prominent natural gift. As GABA and methylglyoxal play significant roles in stress tolerance, and their biosynthesis is strictly regulated in rice plants, the accumulation of 2AP in aromatic rice depends on the interaction of various genetic and environmental factors, and its production may come at some costs of sacrificing tolerance. This review focused on some potential underlying genes in the 2AP and GABA biosynthesis pathways, and analyzed most aspects of aroma formation in rice, and summarized the molecular mechanism of aroma production together with its genetic and non-genetic influencing factors. The present review also stated approaches to produce high-quality aromatic rice via developing novel cultivars and with good agronomic knowledge-based practice. {{custom_citation.content}}
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特种稻是一类具有特殊遗传性状和用途的水稻品种,目前已成为杂交水稻育种的一个重要方向,选择和利用不同来源特种稻种质资源并对其特殊性状基因型进行鉴定和遗传特性分析,对进一步促进特种稻的遗传改良及新品种的选育具有重要意义。本研究利用特异性分子标记InDel-E2、FMbadh2-E7、CAPSRa和CAPS-Ra以及自主开发设计的特异性功能标记FMbadh2-E2、FMbadh2-E7A、FMbadh2-E7B、Ra-CAPS1和Ra-CAPS2并结合基因克隆和测序技术对来自不同地区的32份优质香型水稻亲本材料的香味基因和8份黑米水稻品种的黑色种皮基因进行了基因型鉴定和分析。研究发现,在32份香型水稻材料中,有28份香味性状来源于水稻第8号染色体上编码甜菜碱脱氢酶基因Badh2的功能缺失突变,且为常见的badh2-E7等位突变类型,其在Badh2基因第7外显子处发生8-bp缺失和3个单核苷酸(SNPs)位点多态性突变;2份(板仓香糯和八桂香)表现为香味基因杂合型(Badh2/badh2);2份(溪香和莉香占)在Badh2基因第2和第7外显子处均未发生碱基缺失突变,且具有完整的Badh2基因位点编码序列,其香味性状可能不受Badh2等位基因控制。另外,研究发现,8份黑米水稻品种的黑色种皮均由位于水稻第4号染色体上的Ra/Pb基因控制,其在第7外显子处均表现为2-bp(GT)的缺失突变。同时本研究通过构建多个杂交F2群体对Badh2和Ra基因进行了遗传特性分析,发现其香味和黑色种皮性状分别受1对细胞核隐性和1对细胞核显性主效基因控制。本研究通过对不同来源优质香型水稻亲本材料的香味基因和黑米水稻品种的黑色种皮基因进行鉴定、特异性功能分子标记的开发以及遗传特性分析,为进一步通过分子标记辅助选择育种技术,选育优质、高产且具有香味的杂交水稻新品种及黑米品种提供了重要的理论基础,可明显提高特种稻的选育效率,同时在进一步为发现和挖掘新的香味控制基因及黑色种皮基因的研究上具有重要意义。
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The area between the upper part of the leaf sheath and the basal portion of the leaf blade contains several specialized organs, such as the laminar joint, auricle and ligule. Here we report the identification of T-DNA insertional mutant lines that lack all of these organs. The gene knocked out in the mutant lines encodes a protein that contains a SBP (SQUAMOSA promoter Binding Protein)-domain and is highly homologous to the maize LIGULELESS1 (LG1) gene. At the amino acid sequence level, the OsLG1 protein is 69% identical to maize LG1 and 78% identical to barley LG1. We named the rice gene OsLIGULELESS1 (OsLG1). Transient expression of an OsLG1:RFP (Red Fluorescent Protein) fusion protein indicated that the protein is localized to the nucleus. Transgenic plants harboring the OsLG1 promoter:GUS (beta-glucuronidase) reporter gene construct display preferential expression in developing laminar joint regions and meristemic regions. The gene is also weakly expressed in the ligule, auricles, and leaf sheaths at the basal region. These results indicate that OsLG1 is a transcriptional factor that plays an important role in building the laminar joint between leaf blade and leaf sheath boundary, thereby controlling ligule and auricle development.
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Nuclear factor Y (NF-Y) is a heterotrimeric transcription factor that consists of three subunits, NF-YA, NF-YB, and NF-YC. Gene functions of NF-Ys during endosperm development are not well understood. In this study, we identified eight rice NF-Y-encoding genes, namely OsNF-YA8, OsNF-YB1,9, and OsNF-YC8,9,10,11,12, that are predominantly expressed in the endosperm. Interestingly, the close homologs of these OsNF-Ys are present only in monocot species and are also preferentially expressed in the endosperm, suggesting that they have roles in the regulation of endosperm development. A systemic analysis of interactions between rice endosperm-preferential NF-Ys in yeast revealed that OsNF-YBs and OsNF-YCs could interact with each other. We also found that the endosperm-preferential OsNF-YBs and OsNF-YCs could interact with some ethylene response factors (ERFs) of rice. Unlike OsNF-YC8,9,10, the members of OsNF-YB1,9 or OsNF-YC 11,12 showed no transcriptional activation when present alone. However, they displayed functional activity while in dimer form. In addition, OsNF-YB1-knockout lines showed significant changes in seed morphology, further confirming its role in endosperm development. Our findings provide evidence that a group of phylogenetically conserved NF-Ys is probably differentiated in monocots to regulate endosperm development.
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任俊, 曹跃炫, 黄勇, 等. 基因编辑技术及其水稻中的发展和应用[J]. 中国稻米, 2021, 27(4):92-100.
CRISPR/Cas系统作为一种新兴的基因编辑系统,以其简单、高效、特异性高等特点已广泛应用于植物功能基因组研究和品种改良。在本文中,首先,我们系统总结了CRISPR/Cas技术及其衍生技术在植物中的开发和优化;其次,重点介绍了基于基因组编辑技术的水稻种质和品种改良的最新进展,并描述了基于基因编辑技术的水稻育种新策略,这是传统育种很难实现的;最后,还讨论了基因编辑技术在未来作物和粮食生产中所要面对的挑战。
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为了获得经济价值更高的香稻品种,利用成熟的CRISPR/Cas9技术,在水稻Badh2基因上设计sgRNA-E2和sgRNA-E3 2个靶位点,对超级稻品种龙粳31的香味品质进行改良,其中靶位点sgRNA-E2跨第2个内含子和第2个外显子,靶位点sgRNA-E3位于第3外显子区。经检测共获得有碱基突变的转基因植株6株,其中2个单株为纯合突变,4个单株为双等位突变。突变类型包括碱基缺失、插入及碱基变异,其中突变单株Badh2-E2-13有大片段缺失,为双等位突变,分别缺失64,108 bp。在转录水平上发现6个突变株系的Badh2基因表达量均比野生型显著降低(P0</sub>突变体种子的香味物质2AP含量,发现5个突变单株的2AP含量相对野生型显著提高(P1</sub>主要农艺性状的调查发现,3个株系在香味物质含量提高的同时主要农艺性状和野生型没有显著差异。进一步对T<sub>1</sub>植株进行筛选,共获得了无转基因序列的突变单株10个。综上,利用CRISPR/Cas9技术成功对超级稻品种龙粳31的香味品质进行了遗传改良,实现了香味物质含量的显著提高,为香稻和超级稻的育种提供了基础材料。
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为促进长粒型粳稻品种的选育,以粳稻品种东富139、龙粳31和东农427为试验材料,利用CRISPR/Cas9基因编辑技术构建了pYLCRISPR/Cas9-GS3-RNA和pYLCRISPR/Cas9-GS3-GS9-RNA 2个敲除载体,通过农杆菌转化法侵染受体材料的愈伤组织,对GS3和GS9基因进行定点编辑。最终,3个品种在T<sub>2</sub>都获得了GS3单基因突变、GS9单基因突变和GS3、GS9双基因突变,且无T-DNA元件的纯合植株。在成熟期对T<sub>2</sub>突变体及其野生型的农艺性状进行考察分析,结果表明,与野生型相比,3个品种的gs3突变植株的粒长、千粒质量均显著增加,粒宽、结实率和穗粒数无显著变化;gs9突变体粒长显著增加,粒宽显著减少,千粒质量、结实率和穗粒数无显著变化;gs3gs9突变体粒长增加,且增幅大于gs3和gs9,同时粒宽显著减少,千粒质量显著增加,而结实率和穗粒数无显著变化。综上,利用CRISPR/Cas9技术对东富139、龙粳31和东农427等3个粳稻品种的粒型进行改良,加快了长粒型粳稻新品种的选育进程。
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