大豆[Glycine max (L.) Merr.]是需磷量较大的作物，在整个生育期中均需要保持较高的磷营养水平，其中出苗到盛花期对磷的需求较为敏感。大豆植株在缺磷后表现出明显的缺素症状，例如植株变矮小、叶片发生萎缩、出现坏死的斑点等，甚至还会导致叶片脱落、花期推迟、结荚变少等症状，从而严重影响大豆产量和品质。大豆耐低磷的种质资源匮乏，遗传基础狭窄，成为大豆耐低磷育种中的瓶颈。野生大豆(Glycine soja Sieb.et Zucc.)是栽培大豆的近缘野生种，目前对野生大豆已经开展了包括生态学、品质化学、结构植物学、植物保护等基础的生物学研究。并且野生大豆具有抗病、抗旱、抗虫以及耐盐、耐贫瘠等多种抗逆性和环境适应能力。

本研究以 240 份野生大豆组成的自然群体为研究材料，通过水培试验进行表型鉴定与分析，结合高通量的SNP标记对耐低磷相关性状进行全基因组关联分析，对显著且稳定关联的SNP位点所在的区间内的候选基因进行功能预测和表达模式分析。主要研究结果如下：

1. 野生大豆苗期磷效率相关性状的表型变化显著：低磷胁迫处理后，总根长显著增加，总投影面积和总根表面积出现上升趋势，而平均根系直径、总根体积、根尖数和分支数的表型均值在正常磷水平下高于低磷水平均值，表明根系形态变化是大豆应对低磷胁迫的机制之一。而低磷胁迫处理后，地上部分鲜重、地上部分干重、地上部分磷浓度和地上部分磷积累量整体出现显著下降趋势，其中地上部分磷积累量在低磷胁迫处理下下降了约11倍。根冠比和地上部分磷利用效率则相反，鲜重根冠比和干重根冠比在低磷胁迫下分别增加了2.5倍和2倍，而地上部分磷利用效率在低磷胁迫下增加了5倍。此外，分析表明地上部分鲜重、地上部分干重、鲜重根冠比、干重根冠比、地上部分磷浓度、地上部分磷积累量和地上部分磷利用效率等均存在广泛的表型变异，变异系数在29.61％~116.01％，且大部分呈现出正态分布或接近于正态的分布，表现出数量性状的特点。

2. 分别利用两种高通量的SNP标记(以Williams82基因型为参考基因组，包括1,038,402个高质量SNPs和以W05野生基因型为参考基因组，包括1,293,045个SNPs)对野生大豆磷效率相关的16个性状进行全基因组关联分析。在以W05野生基因型为参考基因组时，GWAS发现同一性状在多环境重复定位的稳定关联位点147个；在以Williams82基因型为参考基因组时,GWAS结果中发现同一性状在多环境重复定位的稳定关联位点35个，另外，发现4个一因多效的稳定位点：地上部鲜重和根干重在E2和E4环境下定位到rs527215和rs527234；地上部鲜重、干重和地上部总磷积累量在E2和E4环境下以及根干重在E1和E2环境下都定位到rs505515；地上部鲜重和干重在E1、E2和E3环境下以及根干重在E1和E4环境下定位到rs822515。并将这些位点用于进一步研究。

3. 候选基因的预测：在以Williams82基因型为参考基因组的1,038,402个SNPs文件检测到的关联SNP位点附近80 kb的范围内寻找候选基因，在以W05基因型为参考基因组的1,293,045个SNPs文件检测到的关联SNP位点附近40 kb的范围内寻找候选基因，结合生物学信息，最终选出可能与磷效率相关的13个野生豆特有基因，分别命名为磷酸化传导信号Gs01、磷酸转运因子Gs02、酸性磷酸酶Gs03、蛋白激酶Gs04、蛋白磷酸酶Gs05、蛋白激酶Gs06、膜联蛋白Gs07、类受体激酶Gs08、乙酰辅酶α羧化酶Gs09、F-box蛋白Gs10、蛋白磷酸酶Gs11、LRR类受体丝氨酸/苏氨酸蛋白激酶基因Gs12、LOB蛋白Gs13。对13个候选基因诱导表达实验结果中，Gs01、Gs06、Gs07、Gs09、Gs10受到低磷胁迫的诱导，表达量显著提升，而Gs02、Gs03、Gs04、Gs05在正常磷水平下表达量显著高于低磷胁迫条件下。进一步对Gs01和Gs02的功能分析，构建了过表达载体转化大豆毛状根，当Gs01在大豆毛状根中过表达时，与对照相比，低磷条件下的根鲜重显著增加，同时过表达该基因促进了低磷条件下根的伸长，这些结果说明Gs01可能通过促进低磷条件下根生长进而参与大豆的低磷胁迫应答响应。与Gs01相反，过表达Gs02显著降低了低磷条件下的根鲜重，且低磷条件下的根长与对照相比显著降低，初步说明Gs02可能负调控大豆低磷胁迫应答反应中根系的变化，这些结果表明以上候选基因可能与大豆耐低磷相关。

Soybean [Glycine Max (L.) Merr. ] is a crop that needs a large amount of phosphorus, and it needs a high phosphorus nutrient level throughout its growth period, especially from seedling emergence to flowering stage. Soybean plants with phosphorus deficiency will show obvious deficiency symptoms,such as shorter plants,smaller leaves, necrotic spots,etc. If plants with phosphorus deficiency in pod-setting stage,it will also lead to leaf shedding,delayed flowering,less pod-setting and other symptoms,thus seriously affecting soybean yield and quality. The lack of germplasm resources and narrow genetic basis of soybean low phosphorus tolerance has become the bottleneck of soybean low phosphorus tolerance breeding. Wild soybean (Glycine soja Sieb. Et Zucc.) is a relative wild species of cultivated soybean. The basic biology of wild soybeans has been studied,including ecology,structure botany,quality chemistry and plant protection. The results showed that wild soybeans had special chemical quality and strong resistance to disease,insects,drought and barren.

In this study,240 wild soybean populations were used as research materials. Phenotypic identification and analysis were carried out through hydroponic tests. Genome-wide association analysis was conducted for traits related to low phosphorus tolerance by combining high throughput SNP markers. The candidate genes were predicted and analyzed according to the interval and functional annotation of the identified stable associated SNP loci.The main research contents are as follows：

1. Phenotypic variation of wild soybean root related traits at seedling stage: Phosphorus stress after processing,the total root length,total projection area,plant total root surface area show an upward trend.The mean value of traits including average root diameter, total root volume,tips and forks of phenotype were slightly higher at normal P level than at low P level.Phenotypic variation of phosphorus efficiency related traits in wild soybean seedlings is significant: under P stress treatment, shoot dry weight, shoot fresh weight, P concentration of shoot and total P of shoot showed a significant downward trend,and total P of the shoot decreased by about 11 times under low P stress. The root-shoot ratio and P utilization efficiency of shoot showed the opposite.The root-shoot ratio by fresh weight increased by 2.5 times and the root-shoot ratio by dry weight increased 2 times under low P.while the P utilization efficiency of shoot increased 5 times.The analysis under low P stress. In addition, the analysis shows that the shoot fresh weight, shoot dry weight, root-shoot ratio by dry weight , root-shoot ratio by fresh weight,P concentration of shoot ,total P of shoot and P utilization efficiency of shoot show a wide range of phenotypic variation, the variation coefficient was 29.61% ~ 116.01%, and most of this traits present normal distribution or close to normal distribution, It shows the characteristics of quantitative characters.

2. Genome-wide association analysis of 16 traits related to phosphorus efficiency in wild soybean was conducted using two high-throughput SNP markers (1,038,402 high-quality SNPs with Williams82 genotype as the reference genome and 1,293,045 SNPs with W05 genotype as the reference genome). Genome-wide association analysis of 16 traits related to phosphorus efficiency in wild soybean was conducted. When using W05 wild genotype as the reference genome, we found 147 stable SNPs for the same trait repeatedly located in multiple environments.When using Williams82 genotype as the reference genome, 35 stable SNPS were identified, In addition, 4 stable SNPs tended to have pleiotropism: rs527215 and rs527234 were located in shoot fresh weight and root dry weight in E2 and E4 environments.we located rs505515 in shoot fresh weight, dry weight and total P of shoot in E2 and E4 and root dry weight in E1 and E2.rs822515 was located in the fresh weight and dry weight of shoot in E1, E2 and E3,and the dry weight of root in E1 and E4. These SNPs will be used for further study.

3. Prediction of candidate genes: Candidate genes are searched within a range of 80 KB near the SNPS with stable association detected when using W82 Genotype file, and 40KB near the SNPS with stable association detected when using W05 Genotype file.several genes were found,and 13 genes that might be related to phosphorus efficiency were finally selected based on literature review and biological information. They are Phosphorylated transduction signal Gs01,phosphate transporter Gs02, acid phosphatase Gs03, protein kinase Gs04,protein phosphatase Gs05, protein kinase Gs06,and Annexin-like protein Gs07,receptor-like kinase Gs08,acetyl-CoA carboxylase Gs09,F-box protein Gs10,protein phosphatase Gs11, LRR receptor-like serine/threonine protein kinase gene Gs12,LOB protein Gs13.Among the 13 candidate genes，Gs01，Gs06，Gs07，Gs09 and Gs10 were significantly induced by low phosphorus.Gs02, Gs03, Gs04 and Gs05 were significantly higher under normal P than under low P stress.By further functional analysis of Gs01 and Gs02, an overexpression vector was constructed to transform hairy roots of soybean. When Gs01 was overexpressed in hairy roots of soybean, the root fresh weight significantly increased under low P condition compared with the control. Meanwhile, overexpression of Gs01 promoted root elongation under low P condition. These results suggest that Gs01 may be involved in soybean response to low P stress by promoting root growth under low P stress. In contrast to Gs01, overexpression of Gs02 significantly reduced the root fresh weight and root length under low P conditions compared with the control, suggesting that Gs02 may negatively regulate the changes of the root system in response to low P stress. These results suggest that the above candidate genes may be related to low P tolerance of soybean.

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