水稻（Oryza sativa L.）作为全球主要的粮食作物之一，其产量和品质对保障粮食安全至关重要。株型是影响水稻产量的关键因素，包括株高、分蘖角度和叶片形态等性状，直接决定了光能利用效率和群体结构。近年来，水稻株型相关基因研究已经取得了一定的进展，这些基因通过调控细胞分裂、伸长和分化，影响水稻的生长发育过程。然而，尽管已有大量研究揭示了部分基因的功能，水稻形态建成的分子调控网络仍不完全清楚，尤其是基因与环境互作、激素信号通路之间的复杂关系仍需进一步探索。因此，深入研究水稻株型基因的功能及其调控机制，不仅有助于解析水稻发育的生物学基础，也为培育高产水稻新品种提供重要的理论依据。

1. FYVE结构域是一种保守的蛋白质结构域，能够特异性识别并结合磷脂酰肌醇-3-磷酸（phosphatidylinositol-3-phosphate, PI3P），广泛参与内体、液泡膜等细胞器膜动态调控及囊泡运输过程。在动物中被大量研究，但在植物中的研究较少。本实验室前期以宁粳7号为遗传背景，通过CRISPR/Cas9技术对水稻FYVE结构域编码基因OsFYVE2和OsFYVE10进行双基因敲除突变体创制，成功获得株型发育缺陷突变体osfyve2osfyve10（以下简称f2f10）。通过对突变体f2f10株型进行表型考察、细胞学观察及分子功能解析，初步探究了OsFYVE2和OsFYVE10在水稻株型调控中的作用，主要结果如下：

（1）利用CRISPR/Cas9技术在粳稻宁粳7号中创制了两种碱基编辑类型的f2f10敲除突变体。突变体在苗期表现出叶鞘弯曲和生长迟滞。随着发育，叶鞘弯曲表型逐渐恢复，但部分叶片呈现区域性高度螺旋卷曲，表现为叶片中部边缘向中脉卷曲形成筒状结构，包裹住叶片正反面。抽穗后，突变体呈现株高矮化、育性降低（花药弯曲但碘-碘化钾染色未显示花粉淀粉填充缺陷）。成熟籽粒表现为粒长显著减小、粒宽略增，且千粒重、结实率、穗粒数和穗长均显著低于野生型。

（2）对开花期螺旋卷曲的叶片和成熟籽粒颖壳进行细胞学观察，结果表明籽粒粒长变短和粒宽增加是由细胞扩张异常引起，而叶片螺旋卷曲表型则是由细胞分化和维管束系统发育异常导致的。

（3）OsFYVE2、OsFYVE10均是组成型表达基因，其表达水平存在组织特异性差异，根部表达量显著高于其他组织。亚细胞定位分析显示，OsFYVE2、OsFYVE10蛋白产物在细胞质中呈现点状分布。细胞器共定位实验显示，OsFYVE2、OsFYVE10均与网格蛋白（clathrin）标记的网格蛋白有被小泡（clathrin-coated vesicles, CCVs）和前液泡区室蛋白（prevacuolar compartment, PVC）共定位。

（4）Western blot与RT-qPCR分析表明，f2f10突变体中肌动蛋白基因OsActin的转录及蛋白水平均显著低于野生型。酵母双杂交检测到OsFYVE10与微丝蛋白存在弱互作。Alexa-488-鬼笔环肽染色显示，突变体微丝束结构完整性受损（断裂增多、空间弥散）。FM4-64标记实验证实其胞吞速率减缓，暗示OsFYVE2/OsFYVE10功能缺失可能影响肌动蛋白依赖的囊泡运输过程。

（5）f2f10突变体呈现与生长素极性运输缺陷型相似的叶鞘/叶片异常弯曲表型。重力反应实验显示其叶鞘向重力性反应增强。LC-MS检测表明突变体地上部IAA含量升高50.86%，RT-qPCR分析进一步揭示色氨酸途径生长素合成关键基因显著上调。

2. 本研究以水稻品种广陆矮为受体亲本，T65为供体亲本构建的高世代回交群体中自发产生的矮秆小粒突变家系GLA-c1为研究对象，系统分析突变家系株高和粒型等表型特征，并通过图位克隆鉴定到候选基因RGA1。

（1）将矮秆小粒家系GLA-c1与受体亲本GLA杂交，构建F₂分离群体，对F₂群体进行遗传分析，利用BSA-seq技术和基于分子标记的基因定位将候选基因定位到5号染色体长臂端约1.2 Mb的区间内。

（2）候选区间内存在已报导的控制矮秆小粒性状的基因LOC_Os05g26890（RGA1），对该基因进行全序列测序分析发现，其第十个外显子产生了单碱基突变，导致其第272位赖氨酸（K）变成了精氨酸（R）。通过多物种序列比对显示，该赖氨酸残基在不同物种中都较为保守。因此，暂将RGA1作为GLA-c1矮秆家系的候选基因，后续将通过转基因互补实验进一步证实。

综上，本研究初步研究了水稻OsFYVE2和OsFYVE10的功能和在水稻株型调控的作用；通过矮秆小粒突变家系GLA-c1定位，鉴定到RGA1可能的新型变异形式，是GLA-c1矮秆家系的潜在候选基因。

Rice (Oryza sativa L.) serves as a global staple crop, where yield and grain quality are critical for food security. Plant architecture—comprising traits such as plant height, tiller angle, and leaf morphology—directly determines photosynthetic efficiency and population structure, thereby governing yield potential. Although genes regulating rice architecture via cell division, elongation, and differentiation have been identified, the molecular networks underlying morphogenesis, particularly gene-environment interactions and hormonal signaling integration, remain incompletely elucidated.

The FYVE domain is a conserved protein module that specifically binds phosphatidylinositol-3-phosphate (PI3P), orchestrating endosomal/vacuolar membrane dynamics and vesicular trafficking. While extensively studied in animals, its functions in plants are poorly characterized. Using the japonica cultivar Ningjing 7 as the genetic background, we generated dual-gene knockout mutants of OsFYVE2 and OsFYVE10 via CRISPR/Cas9-mediated genome editing, designated osfyve2osfyve10 (f2f10). Phenotypic, cytological, and molecular analyses revealed their roles in rice architecture regulation:

Two base-edited f2f10 mutants exhibited seedling-stage phenotypes: leaf sheath curvature and growth retardation. Sheath curvature gradually recovered during development, but leaves displayed regional spiral curling, manifesting as tube-like structures formed by adaxial rolling of mid-lamina margins, enclosing both leaf surfaces. Post-heading mutants showed reduced plant height, impaired fertility (with curved anthers but normal pollen starch filling by I₂-KI staining), and mature grains with significantly shortened length, slightly increased width, and reduced 1,000-grain weight, seed-setting rate, spikelet number per panicle, and panicle length compared to wild-type (WT).

Cytological examination of spirally curled leaves and grain husks indicated that shortened grain length and increased width resulted from aberrant cell expansion, while leaf curling arose from defective cell differentiation and vascular bundle development.

(3) OsFYVE2 and OsFYVE10 are constitutively expressed with tissue-specific variation (highest in roots). Subcellular localization revealed punctate cytoplasmic distribution. Co-localization assays confirmed their association with clathrin-coated vesicles (CCVs) and prevacuolar compartments (PVCs).

(4) Western blot and RT-qPCR demonstrated significantly reduced transcript and protein levels of actin gene OsActin in f2f10. Yeast two-hybrid (Y2H) detected weak interaction between OsFYVE10 and microfilaments. Alexa-488-phalloidin staining showed disrupted microfilament bundling (increased fragmentation and diffuse spatial distribution). FM4-64 endocytosis assays revealed slowed endocytic trafficking, suggesting that loss of OsFYVE2/OsFYVE10 function compromises actin-dependent vesicular transport.

(5) f2f10 phenocopied auxin polar transport mutants in leaf/sheath curvature. Gravitropism assays indicated enhanced negative gravitropic response in sheaths. LC-MS quantified a 50.86% increase in aerial tissue indole-3-acetic acid (IAA) content, while RT-qPCR showed upregulated expression of tryptophan-dependent auxin biosynthesis genes.

2. In this study, we focused on the dwarf and small-grain mutant line GLA-c1, which spontaneously arose in an advanced backcross population constructed using the rice cultivar Guangluai as the recurrent parent and T65 as the donor parent. We systematically analyzed phenotypic traits of the mutant, including plant height and grain morphology, and identified the candidate gene RGA1 through map-based cloning.

(1) The dwarf and small-grain mutant line GLA-c1 was backcrossed with its recurrent parent GLA to generate an F₂ segregating population. Genetic analysis of the F₂ population, combined with BSA-seq (Bulked Segregant Analysis sequencing) and molecular marker-based gene mapping, localized the candidate gene to a 1.2 Mb genomic interval on the long arm of chromosome 5.

(2) Within the candidate interval, the previously reported gene LOC_Os05g26890 (RGA1), known to regulate dwarf and small-grain traits, was identified. Full-length sequencing of this gene revealed a single nucleotide mutation in its tenth exon, leading to a substitution of lysine (K) to arginine (R) at residue position 272. Multiple species sequence alignment demonstrated that this lysine residue is highly conserved across diverse taxa. Based on these findings, RGA1 is provisionally proposed as the candidate gene underlying the dwarf phenotype in the GLA-c1 mutant line, with further validation planned via transgenic complementation assays.

In summary, this study provides a preliminary investigation into the functions of OsFYVE2 and OsFYVE10 in rice and their roles in regulating plant architecture. Additionally, through genetic mapping of the dwarf and small-grain mutant line GLA-c1, we identified a novel allelic variant of RGA1, which is proposed as the potential candidate gene underlying the dwarf phenotype in GLA-c1.

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