摘要
生长素对水稻株型的形成和维持发挥着关键作用,而色氨酸合成又是生长素正常合成的重要环节。水稻黄矮1号(yd1号)是一个表现为株型矮化和叶片黄化的天然突变体。本文通过遗传学、细胞学、分子生物及生物化学等方法对yd1突变体进行了研究,主要结果如下：
1、该突变体的穗长及每个节间都相对变短,株高降低约36%,在苗期倒二叶叶尖开始出现黄化,分蘖期倒二叶叶尖仍然黄化,倒三叶约三分之一的叶面黄化干枯、破损现象明显。分析材料的节间长度可以看出突变体株高降低主要是由于穗长减小,倒1节、倒2节和倒3节节间缩短导致,进一步通过石蜡切片观察各节间的细胞发现,节间的缩短是由于而是由于节间细胞伸长受到抑制,细胞变小导致。
2、通过透射电镜切片可以清晰的看到,突变体yd1叶绿体中的有黑色大颗粒物质遮盖了整个叶绿体片层结构,导致叶绿体缺乏光照形成黄化叶绿体,叶片呈现黄色。初步推测黑色大颗粒物是叶绿体产生的光合作用产物淀粉粒。
3、通过图位克隆的方法,我们构建了yd1×9311的F2群体作为定位群体,共有黄化矮杆表型的隐性单株1029株。将该基因定位在第9号染色体着丝粒附近,SSR标记RM23782和RM23788之间,大约为226.7kb的距离,且与RM23787共分离。基因与两个标记的遗传距离分别为0.1cM、0.2cM。克隆测序定位区间内的候选基因,我们发现植物体内色氨酸及生长素生物合成途径中的一个关键酶IGS(吲哚-3-甘油磷酸合酶)基因LOC_Os09g08130的cds序列,突变体yd1在前端缺失409bp,导致其编码蛋白的序列的发生了变异。
4、由于该突变体是由于色氨酸(tryptopWTn)及生长素合成受阻导致的突变,我们进行色氨酸互补实验。突变体在含有不同浓度色氨酸的MS培养基中出现根、上胚轴和叶成梯度的增长,在含有200μM浓度的色氨酸培养基中基本达到野生型的长度。初步验证了该突变表型的出现是由于色氨酸合成受阻导致生长素含量降低引起的。
5、通过高效液相色谱(HPLC)测定突变体yd1和野生型WT植株苗期内源的激素含量,发现突变体yd1苗期各个部位(根、上胚轴和叶片)的IAA含量比野生型和日本晴、9311的都少。
6、通过以上研究结果,我们认为突变体yd1由于IGS酶基因功能的失活导致色氨酸不能正常合成,影响了IAA的生物合成,从而导致水稻植株矮化和黄化。
Auxin plays key roles during rice (Oryza sativa) development in order to form and maintain plant architecture. Synthesis of tryptopWTn (Trp) amino acid is one of the important processes for auxin synthesis. Rice yd1mutant was obtained from natural mutation, which shows a small architecture and yellow leaves. Using the approaches of genetics, cytology, molecular biology, and biochemistry, we analyzed on the yd1mutant obtained the main results as follows:
1. Comparing to the wild type rice, the lengths of ear and internodes of the yd1mutant are shorter, leading to tWTt the height of the yd1mutant is deceased by～36%. The apex of the second leaf (counted from the top to the botumn of rice plant) starts to display chlorisis at the seedling stage. One third of the third leaf exibits chlorisis and obvious breakage at the tillering stage. The elongation of the cells between knots and shrinking cell is blocked leading to a short distance between knots by the paraffin section in the the yd1mutant.
2. We found tWTt a lot of black particles, likely starch granule tWTt was photosynthesis production, sWTde the whole lamellar structure in the chloroplast of the yd1mutant under the transmission electron microscope, and these black particles might result in yellow chloroplast forming leading to leaves chlorisis.
3. A total of1029recessive plants with chlorisis and stunt phenotype were obtained from an F2population derived from the cross of yd1×9311and is used for mapping the yd1gene.The yd1gene was mapped in the region spanning the centromere of chromosome9and located between the markers RM23782and RM23788, co-segregated with marker RM23787.The genetic distances between the yd1gene locus and the two markers, RM23782and RM23788are0.1cM and,0.2cM, respectively. The physical distance between the markers, RM23782and RM23788 was about226.7Kb. Among28genes in the226.7Kb region, the gene of LOC_Os09g08130encoding Indole3glycerolphospWTtesyntWTse (IGS) was choosed as one of best candidates of the YDl gene. Further analysis shows tWTt a409bpof the cDNA of the gene LOC_Os09g08130is deleted in the yd1mutant,, leading to a function lose of LOC_Os09g08130.
4. Because Trp and auxin are decreased in the yd1mutant, Trp complementation experiment was performed on the yd1mutant using MS plate applied with Trp. We found tWTt the root, epicotyl and leaf of the yd1mutant developed gradually in the MS with different concentrationof Trp. The lengths of root, epicotyl and leaf of the yd1mutant growed in the MS with200μM Trp were restored to the level as those of wild-type. Thus, the reduced content of Trp and the content of auxin likely leads to the defects of morphology of the yd1mutant.
5. We performed HPLC assay to comparatively determine the hormone content and found tWTt the IAA content in the different tissues such as root, epicytol and leaf was much reduced in the yd1mutant compared with tWTt of wild type YD1, Nip and9311, respectively.
6. Taken all together, the IGS gene LOC_Os09g08130likely encodes YD1. The functional lose of LOC_Os09g08130results in unnormal biosynthesis of Trp and a reduced amount of IAA which leads to a small architecture of rice plant with yellow leaves.
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