理想株型是合理密集种植、提高作物产量和节省劳动力成本的生物学基础，对于提高农业生产力至关重要。地上部分株型由叶、茎、花器官和果实的形态共同决定。叶型，包括叶片位置和叶片形态，是植物株型的关键组成部分，直接决定植物外观、光合作用和最终生产力。叶片位置由叶序、叶角/叶柄角和叶柄长度的综合影响决定。适当的叶角/叶柄角能够增加种植密度，较大幅度提高作物产量，同时有利于在高密度种植条件下机械化收割，降低人工成本，提升经济效益。叶形态在光合作用、蒸腾作用和呼吸作用中发挥着重要作用。叶形态是由叶片的大小、数量、形态、厚度和曲率以及边缘配置和叶脉模式等组分共同决定的。叶片平整性和叶边缘锯齿形态是决定叶片形态多样性的关键指标。

黄瓜（Cucumis sativus L.），葫芦科中一种重要的蔬菜经济作物。较小叶柄角和适宜叶片形态的黄瓜品种，可以改善栽培模式、有利于合理密植和提高经济效益。对叶型目标性状调控基因的发掘与功能解析已成为黄瓜分子育种研究的重点。目前关于控制黄瓜叶柄角调控基因的研究寥寥无几，亟待开展。关于黄瓜叶片形态的研究相对较多，但针对叶片平展性和叶边缘的遗传机制和调控网络尚不明晰，有待完善。挖掘和克隆叶型形成关键基因，对于揭示黄瓜叶型的调控机理和加快黄瓜株型育种进程具有重要意义。本实验室从EMS（甲基磺酸乙酯）诱变华北型黄瓜CCMC的群体中鉴定出两个黄瓜叶型突变体，直立紧凑突变体erect and compact leaf architecture mutant（ecla）和光滑叶边缘突变体smooth leaf margin mutant（slm），可作为研究叶柄角度和叶片形态的理想材料。本研究通过遗传图谱构建、转录组分析以及蛋白互作验证等实验解析了两个突变体形成的遗传机制和调控网络，不仅为黄瓜叶柄角和叶片形态调控基因网络提供了新的基因资源，也为生长素调控黄瓜叶型提供了新的桥梁。另外，研究结果对葫芦科分子标记辅助育种及现有品种遗传改良具有一定的指导意义。

1. 黄瓜直立紧凑突变体ecla的基因克隆和功能分析

1.1从黄瓜EMS诱变群体中鉴定出的直立紧凑突变体ecla在营养器官和生殖器官表现出多效性表型，包括植株紧凑，高度降低，叶柄角明显变小，叶片和花器官褶皱和种子极小；通过扫描电镜和石蜡切片进行叶柄和叶片细胞大小和形态观察，结果显示突变体ecla的叶柄角变小与叶柄基部近-远轴的不对称细胞扩增减弱有关，而叶片褶皱与叶片不同部位的细胞扩增不平衡有关。

1.2基于分子标记构建连锁图谱将ecla位点定位在黄瓜第5号染色体上65 kb的基因组区间里。MutMap测序分析和dCAPS标记验证确定导致ecla突变表型的候选基因为CsaV3_5G037960，编码ATP结合盒式（ATP-binding cassette，ABC）转运蛋白ABCB19。突变体中CsABCB19基因的第十个外显子的非同义突变导致蛋白翻译提前终止；CsABCB19的异位表达完全恢复了拟南芥突变体Atabcb19中叶柄角减小和叶片皱褶的表型；亚细胞定位显示CsABCB19蛋白定位在烟草叶表皮细胞的质膜上，与拟南芥AtABCB19的结果一致。这些结果说明ABCB19作为生长素转运蛋白在调控叶片定位和平展性上功能保守。

1.3基因定量分析显示，与野生型（WT）相比，突变体ecla中CsABCB19在所有组织中的表达量均显著降低；转录组分析显示，生长素极性运输、生长素响应因子、生长素合成及极性调控相关基因在突变体ecla的叶柄和叶片中的表达与WT相比具有显著差异，说明叶柄角的发育和叶片平整性的维持需要生长素的稳定和极性的建立；叶柄和叶片中内源生长素测定和外源激素喷施效应进一步证明，CsABCB19通过介导生长素积累和运输来调节黄瓜叶柄角大小和叶片形态发育。

2. 黄瓜光滑叶边缘突变体slm的基因克隆和功能分析

2.1从EMS诱变黄瓜群体中鉴定出突变体smooth leaf margin mutant（slm），在营养阶段和生殖阶段都表现出明显表型变异，主要包括叶片圆形，叶边缘没有锯齿，花瓣没有尖端和果实没有种子。石蜡切片显示子房中胚珠启动的失败导致雌性不育。遗传作图结合MutMap测序确定候选基因CsaV3_2G032490，编码拟南芥NAKED PINS IN YUC MUTANTS1（NPY1）的同源蛋白，命名为CsNPY1。CsNPY1编码BTB-NPH3蛋白，突变导致NPH3结构域的缺失，说明NPH3是NPY1调控叶边缘发育所必须的。qRT-PCR结果显示CsNPY1基因在突变体slm所有被检测的组织中的表达量均明显低于WT，尤其是在叶边缘。转录组分析显示生长素相关基因及调控叶片极性和锯齿发育的转录因子在WT和突变体slm叶片中显著差异表达。

2.2突变体slm与CsPID突变导致的圆叶突变体rl具有相似的表型，都缺失明显叶尖端呈现圆叶叶形。但是叶边缘锯齿表现具有明显区别，突变体slm表现出完全缺失锯齿的光滑叶边缘，突变体rl的叶边缘存在均匀锯齿。本研究通过对两个突变体杂交后代的表型进行分析证明了rl和slm在黄瓜叶边缘锯齿发育中发挥协调作用。转录组分析显示突变体slm中PID基因的表达明显上调，说明PID和NPY1基因在转录水平上存在联系。另外，Pull-down实验证明PID和NPY1可以在体外直接互作，说明PID和NPY1在蛋白水平存在联系。因此，NPY1和PID在控制黄瓜生长发育上存在功能联系。

2.3转录组联合分析显示突变体slm和rl中差异表达基因明显重叠，证明NPY1和PID可能在黄瓜叶边缘发育同一调控网络中。内源生长素测定结果显示slm叶边缘中生长素IAA的含量相对低于WT，但是突变体rl中IAA的含量明显高于WT。因此，NPY1和PID在生长素调控黄瓜叶边缘发育同一个调控网络中，但具有不同调控机制。另外，使用极性运输抑制剂NPA和生长素IAA持续喷施野生型都产生均匀锯齿的心型叶，类似两个突变体杂交获得的中间型，说明生长素可以通过生长素极性运输和积累来调控黄瓜叶边缘锯齿发育。

Optimal plant architecture is the biological basis of dense planting, high crop yield and labour cost savings, and is thus critical for improving agricultural productivity. The above-ground parts are collectively determined by the morphology of leaves, stems, flowering organs, and fruit morphology. Leaf architecture, including leaf position and leaf morphology, are critical components of plant architecture that directly determines plant appearance, photosynthetic utilization, and ultimate productivity. Leaf positioning is determined by the combined effects of leaf order, leaf angle/leaf petiole angle, and petiole length. Proper leaf angle/petiole angle can substantially increase crop yield by increasing planting density and facilitate mechanized harvesting under high density planting conditions, reducing labor costs and improving economic efficiency. Leaf morphology plays an important role in photosynthesis, respiration and light perception. The overwhelming differences in leaf morphology among plant species are determined by a combination of leaf shape, thickness, curvature, leaf margin configuration, and leaf vein pattern. Leaf flatness and leaf margin pattern are key indicators of leaf morphological diversity.

Cucumber (Cucumis sativus L.) is an important vegetable cash crop in the Cucurbitaceae. The breeding of cucumber varieties with smaller petiole angles and suitable leaf morphology can improve cultivation patterns, rationalize dense planting and increase economic efficiency. The discovery and functional elucidation of genes regulating leaf architecture target traits has become the focus of molecular breeding research in cucumber. There are few studies on the genes controlling the regulation of petiole angle in cucumber, which need to be carried out urgently. There are relatively more studies on cucumber leaf morphology, but studies on genetic mechanisms and regulatory networks targeting leaf flattening and leaf margins need to be improved. The excavation and cloning of key genes for leaf architecture formation is important for revealing the regulatory mechanism of cucumber leaf architecture and accelerating the plant architecture breeding process of cucumber. Two leaf architecture mutants, erect and compact leaf architecture mutant (ecla) and smooth leaf margin mutant (slm), were identified from an EMS-mutagenized population of Changchunmici (CCMC, as wild type, WT) cucumber, which are ideal materials for studying petiole angle and leaf morphology. In this study, the genetic mechanism and regulatory network of the formation of two leaf architecture mutants were resolved through genetic map construction, transcriptome analysis and protein interaction verification experiments. This not only provides new genetic resources for the regulatory gene network of cucumber petiole angle and leaf morphology, but also provides a new bridge for auxin regulation of cucumber leaf shape. In addition, the results of the study have some guiding significance for molecular marker-assisted breeding and genetic improvement of existing varieties of Cucurbitaceae.

1. Gene cloning and functional analysis of ecla in cucumber

1.1 The mutant ecla additionally exhibited a pleiotropic phenotype, including reduced plant height, significantly smaller petiole angles, crinkled leaves and floral organs, and extremely small seeds. The size and morphology of petiole and leaf cells were observed by scanning electron microscopy and paraffin sectioning, and the results showed that the reduced petiole angle in the mutant ecla was associated with diminished asymmetric cell expansion in the adaxial-abaxial axis at the base of the petiole, while the leaf crinkling was associated with unbalanced cell expansion in different parts of the leaf.

1.2 Molecular marker-based construction of a linkage map localized the ecla locus to a 65 kb genomic interval on cucumber chromosome 5. MutMap sequencing analysis and dCAPS marker validation identified the candidate gene responsible for the ecla mutant phenotype as CsaV3_5G037960, encoding the ATP-binding cassette (ABC) transporter protein ABCB19. A nonsynonymous mutation in the tenth exon of the CsABCB19 gene in the mutant resulted in premature termination of protein premature termination of translation. Ectopic expression of CsABCB19 completely rescued the phenotype of reduced petiole angle and leaf crinkling in the Arabidopsis mutant Atabcb19. Subcellular localization experiments localized the CsABCB19 protein to the plasma membrane of tobacco leaf epidermal cells, consistent with the results in the Arabidopsis AtABCB19 protein. These results confirm that ABCB19 is functionally conserved as an auxin transporter protein in the regulation of leaf positioning and flattening.

1.3 Analysis of qRT-PCR results showed that the expression levels of CsABCB19 gene were significantly reduced in all tissues in the mutant ecla compared with the WT. Transcriptome analysis revealed that the expression of auxin polarity transport, auxin response genes, auxin synthesis/metabolism and polarity-related genes were significantly different in the mutant petioles and leaves compared to those in WT. This indicates that the development of petiole angle and maintenance of leaf flatness require auxin stabilization and polarity establishment. The results of auxin endogenous assay and the effects of exogenous auxin inhibitor NPA and auxin IAA application in the petioles and leaves of mutant and WT further demonstrate that CsABCB19 regulates cucumber petiole angle size and leaf morphological development by mediating auxin accumulation and transport.

2. Gene cloning and functional analysis of mutant slm in cucumber

2.1 The smooth leaf margin mutant (slm) mutant exhibited significant phenotypic variation in both the nutritional and reproductive stages, mainly including the absence of serrations on the leaf margins, petals lacking tips, and fruits without seeds. Paraffin sections revealed a failure of ovule initiation in the ovary resulting in seedless fruits. Genetic mapping combined with MutMap sequencing identified the candidate gene CsaV3_2G032490, encoding a homolog of Arabidopsis NAKED PINS IN YUC MUTANTS1 (NPY1), named CsNPY1. CsNPY1 encodes the BTB-NPH3 protein, and the mutation in the mutant slm resulted in the early production of the stop codon, leading to early termination of translation resulting in the deletion of the entire NPH3 structural domain, indicating that NPH3 is required for NPY1 to regulate leaf margin development. The qRT-PCR results showed that the expression level of CsNPY1 gene was significantly lower in all tissues examined in mutant slm than in WT, especially in the leaf margins. Transcriptome analysis revealed that auxin-related genes and transcription factors regulating leaf polarity and serration development were significantly differentially expressed in WT and mutant slm leaves.

2.2 The mutant slm resulting from the CsNPY1 mutation and the mutant rl resulting from the CsPID mutation exhibited a similar phenotype with rounded leaves lacking a distinct leaf tip. However, the phenotypes of leaf margin serrations were significantly different, with the mutant slm showing smooth leaf margins lacking serrations, while the mutant rl had uniform serrations on the leaf margins. Phenotypic analysis of the progeny of the two mutant crosses demonstrated that rl and slm play a coordinated role in the development of cucumber leaf margin serrations. Transcriptome data analysis showed that the expression of PID gene was significantly up-regulated in mutant slm, indicating that PID and NPY1 genes interacted at the transcriptional level. In addition, Pull-down experiments demonstrated that PID and NPY1 proteins directly interact with each other in vitro, suggesting that PID and NPY1 are linked at the protein level. Therefore, there is a functional link between NPY1 and PID to control cucumber growth and development.

2.3 Transcriptome association analysis revealed significant overlap of differentially expressed genes in mutants slm and rl, demonstrating that NPY1 and PID may be in the same pathway of auxin-regulated cucumber leaf margin development. The results of endogenous auxin content measurement in leaf margins showed that the content of IAA in slm was relatively lower than that in WT, but the content of IAA in rl was relatively higher than that in WT. Thus, NPY1 and PID are in the same regulatory network of auxin-regulated cucumber leaf margin development, but with different regulatory mechanisms. Furthermore, continuous treatment of the WT with the auxin polarity transport inhibitor NPA and the auxin IAA produced uniformly serrated heart-shaped leaves, similar to the intermediate type obtained by crossing the two mutants. This suggests that auxin can regulate cucumber leaf margin serration development via auxin polar transport and accumulation.

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