Thomas A L, Guerreiro S M C, Sodek L
Departamento de Fisiologia Vegetal, Brazil.
Ann Bot. 2005 Dec;96(7):1191-8. doi: 10.1093/aob/mci272. Epub 2005 Sep 30.
Flooding results in hypoxia of the root system to which N2 fixation of nodulated roots can be especially sensitive. Morphological adaptions, such as aerenchyma formation, can facilitate the diffusion of oxygen to the hypoxic tissues. Using soybean, the aim of the study was to characterize the morphological response of the nodulated root system to flooding and obtain evidence for the recovery of N metabolism.
Sections from submerged tissues were observed by light microscopy, while sap bleeding from the xylem was analysed for nitrogenous components.
Flooding resulted in the rapid formation of adventitious roots and aerenchyma between the stem (immediately above the water line), roots and nodules. In the submerged stem, taproot, lateral roots and adventitious roots, lysigenous aerenchyma arose initially in the cortex and was gradually substituted by secondary aerenchyma arising from cells derived from the pericycle. Nodules developed aerenchyma from cells originating in the phellogen but nodules situated at depths greater than 7-8 cm showed little or no aerenchyma formation. As a result of aerenchyma formation, porosity of the taproot increased substantially between the 4th and 7th days of flooding, coinciding with the recovery of certain nitrogenous products of N metabolism of roots and nodules transported in the xylem. Thus, on the first day of flooding there was a sharp decline in xylem ureides and glutamine (products of N2 fixation), together with a sharp rise in alanine (product of anaerobic metabolism). Between days 7 and 10, recovery of ureides and glutamine to near initial levels was recorded while recovery of alanine was partial.
N metabolism of the nodulated soybean root system can recover at least partially during a prolonged period of flooding, a process associated with aerenchyma formation.
洪水会导致根系缺氧,而结瘤根的固氮作用对其尤为敏感。形态学适应,如通气组织的形成,有助于氧气向缺氧组织扩散。本研究以大豆为材料,旨在描述结瘤根系对洪水的形态学响应,并获取氮代谢恢复的证据。
通过光学显微镜观察淹没组织的切片,同时分析木质部汁液渗出物中的含氮成分。
洪水导致不定根迅速形成,且在茎(紧挨着水面上方)、根和根瘤之间形成通气组织。在淹没的茎、主根、侧根和不定根中,溶生性通气组织最初在皮层中产生,并逐渐被源自中柱鞘细胞的次生通气组织所取代。根瘤由起源于木栓形成层的细胞发育出通气组织,但位于深度大于7 - 8厘米处的根瘤几乎没有或没有通气组织形成。由于通气组织的形成,主根的孔隙率在洪水淹没的第4天至第7天大幅增加,这与根和根瘤中氮代谢的某些含氮产物在木质部中运输的恢复相吻合。因此,在洪水淹没的第一天,木质部中的脲类和谷氨酰胺(固氮产物)急剧下降,同时丙氨酸(无氧代谢产物)急剧上升。在第7天至第10天期间,脲类和谷氨酰胺恢复到接近初始水平,而丙氨酸仅部分恢复。
结瘤大豆根系的氮代谢在长时间洪水淹没期间至少可以部分恢复,这一过程与通气组织的形成有关。