Department of Bioscience, Tokyo University of Agriculture, 1-1-1 Sakuragaoka, Setagaya-ku, Tokyo 156-8502, Japan.
Mol Plant. 2011 Jan;4(1):17-24. doi: 10.1093/mp/ssq063. Epub 2010 Oct 13.
While an increase in the number of xyloglucan tethers between the cellulose microfibrils in plant cell walls increases the walls' rigidity, the degradation of these tethers causes the walls to loosen. Degradation can occur either through the integration of xyloglucan oligosaccharides due to the action of xyloglucan endotransglucosylase or through direct hydrolysis due to the action of xyloglucanase. This is why the addition of xyloglucan and its fragment oligosaccharides causes plant tissue tension to increase and decrease so dramatically. Experiments involving the overexpression of xyloglucanase and cellulase have revealed the roles of xyloglucans in the walls. The degradation of wall xyloglucan in poplar by the transgenic expression of xyloglucanase, for example, not only accelerated stem elongation in the primary wall, but also blocked upright-stem gravitropism in the secondary wall. Overexpression of cellulase also reduced xyloglucan content in the walls as cellulose microfibrils were trimmed at their amorphous region, resulting in increased cell volume in Arabidopsis leaves and in sengon with disturbed leaf movements. The hemicellulose xyloglucan, in its function as a tether, plays a key role in the loosening and tightening of cellulose microfibrils: it enables the cell to change its shape in growth and differentiation zones and to retain its final shape after cell maturation.
虽然植物细胞壁中木葡聚糖连接物的数量增加会增加细胞壁的刚性,但这些连接物的降解会导致细胞壁松动。降解可以通过木葡聚糖寡糖的整合发生,这是由于木葡聚糖内切葡聚糖酶的作用,或者通过木葡聚糖酶的直接水解发生。这就是为什么添加木葡聚糖及其片段寡糖会导致植物组织张力如此剧烈地增加和减少的原因。涉及木葡聚糖酶和纤维素酶过表达的实验揭示了木葡聚糖在细胞壁中的作用。例如,通过过表达木葡聚糖酶在杨树中降解细胞壁木葡聚糖,不仅加速了初生细胞壁的伸长,而且阻断了次生细胞壁的直立茎向地性。纤维素酶的过表达也减少了细胞壁中的木葡聚糖含量,因为纤维素微纤维在无定形区域被修剪,导致拟南芥叶片和破坏运动的银荆的细胞体积增加。半纤维素木葡聚糖作为一种连接物,在纤维素微纤维的松动和紧固中起着关键作用:它使细胞能够在生长和分化区改变形状,并在细胞成熟后保持其最终形状。