Department of Environmental Management Engineering, Faculty of Environmental Science and Technology, The Graduate School of Environmental Science, Okayama University, Okayama 700-8530, Japan.
J Integr Plant Biol. 2009 Nov;51(11):1024-39. doi: 10.1111/j.1744-7909.2009.00873.x.
Non-mycorrhizal Brassica does not produce specialized root structures such as cluster or dauciform roots but is an effective user of P compared with other crops. In addition to P-uptake, utilization and remobilization activity, acquisition of orthophosphate (Pi) from extracellular sparingly P-sources or unavailable bound P-forms can be enhanced by biochemical rescue mechanisms such copious H(+)-efflux and/or carboxylates exudation into rhizosphere by roots via plasmalemma H(+) ATPase and anion channels triggered by P-starvation. To visualize the dissolution of sparingly soluble Ca-phosphate (Ca-P), newly formed Ca-P was suspended in agar containing other essential nutrients. With NH(4)(+) applied as the N source, the precipitate dissolved in the root vicinity can be ascribed to rhizosphere acidification, whereas no dissolution occurred with nitrate nutrition. To observe in situ rhizospheric pH changes, images were recorded after embedding the roots in agar containing bromocresol purple as a pH indicator. P-tolerant cultivar showed a greater decrease in pH than the sensitive cultivar in the culture media (the appearance of typical patterns of various colors of pH indicator in the root vicinity), and at stress P-level this acidification was more prominent. In experiment 2, low P-tolerant class-I cultivars (Oscar and Con-II) showed a greater decrease in solution media pH than low P-sensitive class-II (Gold Rush and RL-18) cultivars, and P-contents of the cultivars was inversely related to decrease in culture media pH. To elucidate P-stress-induced remodeling and redesigning in a root architectural system, cultivars were grown in rhizoboxes in experiment 3. The elongation rates of primary roots increased as P-supply increased, but the elongation rates of the branched zones of primary roots decreased. The length of the lateral roots and topological index values increased when cultivars were exposed to a P-stress environment. To elucidate Pi-uptake kinetics, parameters related to P influx: maximal transport rate (V(max)), the Michaelis-Menten constant (K(m)), and the external concentration when net uptake is zero (C(min)) were tested in experiment 4. Lower K(m) and C(min) values were better indicative of the P-uptake ability of the class-I cultivars, evidencing their adaptability to P-starved environmental cues. In experiment 5, class-I cultivars exuded two- to threefold more carboxylates than class-II cultivars under the P-stress environment. The amount and types of carboxylates exuded from the roots of P-starved plants differed from those of plants grown under P-sufficient conditions. Nevertheless, the exudation rate of both class-I and class-II cultivars decreased with time, and the highest exudation rate was found after the first 4 h of carboxylates collection. Higher P uptake by class-I cultivars was significantly related to the drop in root medium pH, which can be ascribed to H(+)-efflux from the roots supplied with sparingly soluble rock-P and Ca(3)(PO(4))(2). These classical rescue strategies provided the basis of P-solubilization and acquisition from sparingly soluble P-sources by Brassica cultivars to thrive in a typically stressful environment.
非菌根 Brassica 不会产生专门的根结构,如簇状或 dauciform 根,但与其他作物相比,它是 P 的有效使用者。除了 P 吸收、利用和再利用活性外,通过生化拯救机制,如大量 H(+)流出和/或根系通过质膜 H(+)ATP 酶和阴离子通道从细胞外 P 来源或不可用的结合 P 形式中分泌羧酸,可以增强对正磷酸盐 (Pi) 的获取。为了观察难溶性 Ca-磷酸盐 (Ca-P) 的溶解,新形成的 Ca-P 悬浮在含有其他必需养分的琼脂中。当用 NH(4)(+) 作为 N 源时,沉淀可以归因于根际酸化而溶解,而硝酸盐营养则不会发生溶解。为了观察根际 pH 值的原位变化,在将根嵌入含有溴甲酚紫的琼脂中后记录图像,作为 pH 指示剂。在培养基中(根附近出现各种颜色 pH 指示剂的典型图案),耐 P 品种的 pH 值下降幅度大于敏感品种,在胁迫 P 水平下,这种酸化更为明显。在实验 2 中,低耐 P 品种(Oscar 和 Con-II)在低 P 敏感品种(Gold Rush 和 RL-18)的溶液培养基 pH 值下降幅度更大,并且品种的 P 含量与培养基 pH 值下降呈负相关。为了阐明 P 胁迫诱导的根构型系统的重塑和重新设计,在实验 3 中在根盒中种植了品种。随着 P 供应的增加,主根的伸长率增加,但主根分支区的伸长率下降。当品种暴露于 P 胁迫环境时,侧根的长度和拓扑指数值增加。为了阐明 Pi 吸收动力学,在实验 4 中测试了与 P 内流相关的参数:最大转运速率 (V(max))、米氏常数 (K(m)) 和净吸收时的外部浓度 (C(min))。较低的 K(m)和 C(min)值更好地表明了 I 类品种的 P 吸收能力,证明了它们对 P 饥饿环境线索的适应性。在实验 5 中,I 类品种在 P 胁迫环境下分泌的羧酸是 II 类品种的两到三倍。从 P 饥饿植物根系分泌的羧酸的数量和类型与在 P 充足条件下生长的植物不同。然而,I 类和 II 类品种的分泌率随时间下降,并且在收集羧酸后的前 4 小时发现分泌率最高。I 类品种对 P 的高吸收与根介质 pH 值下降显著相关,这可归因于从供应难溶性岩石 P 和 Ca(3)(PO(4))(2)的根中流出的 H(+)。这些经典的拯救策略为 Brassica 品种从难溶性 P 来源中溶解和获取 P 提供了基础,使其在典型的胁迫环境中茁壮成长。