Shigeyama Takuma, Watanabe Asuka, Tokuchi Konatsu, Toh Shigeo, Sakurai Naoki, Shibuya Naoto, Kawakami Naoto
Department of Life Sciences, School of Agriculture, Meiji University, Higashimita 1-1-1, Tama-ku, Kawasaki 214-8571, Japan.
Graduate School of Biosphere Science, Hiroshima University, Kagamiyama 1-3-2, Higashihiroshima 739-8528, Japan.
J Exp Bot. 2016 Oct;67(19):5615-5629. doi: 10.1093/jxb/erw321. Epub 2016 Sep 7.
Regulation and maintenance of cell wall physical properties are crucial for plant growth and environmental response. In the germination process, hypocotyl cell expansion and endosperm weakening are prerequisites for dicot seeds to complete germination. We have identified the Arabidopsis mutant thermoinhibition-resistant germination 1 (trg1), which has reduced seed dormancy and insensitivity to unfavourable conditions for germination owing to a loss-of-function mutation of TRG1/XYL1, which encodes an α-xylosidase. Compared to those of wild type, the elongating stem of trg1 showed significantly lower viscoelasticity, and the fruit epidermal cells were longitudinally shorter and horizontally enlarged. Actively growing tissues of trg1 over-accumulated free xyloglucan oligosaccharides (XGOs), and the seed cell wall had xyloglucan with a greatly reduced molecular weight. These observations suggest that XGOs reduce xyloglucan size by serving as an acceptor in transglycosylation and eventually enhancing cell wall loosening. TRG1/XYL1 gene expression was abundant in growing wild-type organs and tissues but relatively low in cells at most actively elongating part of the tissues, suggesting that α-xylosidase contributes to maintaining the mechanical integrity of the primary cell wall in the growing and pre-growing tissues. In germinating seeds of trg1, expression of genes encoding specific abscisic acid and gibberellin metabolism enzymes was altered in accordance with the aberrant germination phenotype. Thus, cell wall integrity could affect seed germination not only directly through the physical properties of the cell wall but also indirectly through the regulation of hormone gene expression.
细胞壁物理性质的调控与维持对植物生长和环境响应至关重要。在萌发过程中,下胚轴细胞扩张和胚乳弱化是双子叶植物种子完成萌发的先决条件。我们鉴定出了拟南芥突变体抗热抑制萌发1(trg1),由于编码α-木糖苷酶的TRG1/XYL1功能缺失突变,该突变体种子休眠降低且对不利于萌发的条件不敏感。与野生型相比,trg1伸长的茎表现出显著更低的粘弹性,果实表皮细胞纵向更短且横向扩大。trg1活跃生长的组织中游离木葡聚糖寡糖(XGOs)过度积累,种子细胞壁中的木葡聚糖分子量大幅降低。这些观察结果表明,XGOs通过作为转糖基化的受体来减小木葡聚糖的大小,并最终增强细胞壁的松弛。TRG1/XYL1基因在野生型生长的器官和组织中表达丰富,但在组织最活跃伸长部分的细胞中相对较低,这表明α-木糖苷酶有助于维持生长中和生长前组织中初生细胞壁的机械完整性。在trg萌发的种子中,编码特定脱落酸和赤霉素代谢酶的基因表达根据异常的萌发表型发生了改变。因此,细胞壁完整性不仅可以通过细胞壁的物理性质直接影响种子萌发,还可以通过激素基因表达的调控间接影响种子萌发。