De Zio Elena, Trupiano Dalila, Montagnoli Antonio, Terzaghi Mattia, Chiatante Donato, Grosso Alessandro, Marra Mauro, Scaloni Andrea, Scippa Gabriella S
Dipartimento di Bioscienze e Territorio, University of Molise, 86090 Pesche (IS), Italy.
Dipartimento di Biotecnologie e Scienze della Vita, University of Insubria, 21100 Varese, Italy.
Ann Bot. 2016 Oct 1;118(4):865-883. doi: 10.1093/aob/mcw159.
Background and Aims Progress has been made in understanding the physiological and molecular basis of root response to mechanical stress, especially in the model plant Arabidopsis thaliana, in which bending causes the initiation of lateral root primordia toward the convex side of the bent root. In the case of woody roots, it has been reported that mechanical stress induces an asymmetric distribution of lateral roots and reaction wood formation, but the mechanisms underlying these responses are largely unknown. In the present work, the hypothesis was tested that bending could determine an asymmetric response in the two sides of the main root axis as cells are stretched on the convex side and compressed on the concave side. Methods Woody taproots of 20 seedlings were bent to an angle of 90° using a steel net. Changes in the anatomy, lignin and phytohormone content and proteome expression in the two sides of the bent root were analysed; anatomical changes, including dissimilarities and similarities to those found in poplar bent woody stem, were also considered. Key Results Compression forces at the concave side of poplar root induced the formation of reaction wood which presented a high lignin content and was associated with the induction of cambium cell activity. Auxin seemed to be the main hormone triggering lignin deposition and cell wall strengthening in the concave sides. Abscisic acid appeared to function in the water stress response induced by xylem structures and/or osmotic alterations in the compression sides, whereas gibberellins may control cell elongation and gravitropisms. Conclusions Poplar root reaction wood showed characteristics different from those produced in bent stem. Besides providing biomechanical functions, a bent root ensures water uptake and transport in the deforming condition induced by tension and compression forces by two different strategies: an increase in xylem thickness in the compressed side, and lateral root formation in the tension side.
背景与目的 在理解根系对机械应力的生理和分子基础方面已取得进展,尤其是在模式植物拟南芥中,弯曲会导致侧根原基向弯曲根的凸侧起始。对于木本根系,据报道机械应力会诱导侧根不对称分布和反应木形成,但其潜在机制大多未知。在本研究中,对弯曲能否在主根轴两侧确定不对称反应这一假设进行了验证,因为细胞在凸侧被拉伸而在凹侧被压缩。
方法 使用钢丝网将20株幼苗的木本主根弯曲至90°角。分析弯曲根两侧的解剖结构、木质素和植物激素含量以及蛋白质组表达的变化;还考虑了包括与杨树弯曲木本茎中发现的异同在内的解剖学变化。
关键结果 杨树根凹侧的压力诱导了反应木的形成,其木质素含量高,且与形成层细胞活性的诱导有关。生长素似乎是触发凹侧木质素沉积和细胞壁强化的主要激素。脱落酸似乎在由压缩侧木质部结构和/或渗透变化诱导的水分胁迫反应中起作用,而赤霉素可能控制细胞伸长和向地性。
结论 杨树根反应木表现出与弯曲茎中产生的反应木不同的特征。除了提供生物力学功能外,弯曲的根通过两种不同策略确保在张力和压力诱导的变形条件下的水分吸收和运输:压缩侧木质部厚度增加,以及张力侧形成侧根。