Boron (B) alleviates aluminum (Al) toxicity in higher plants; however, the underlying mechanisms behind this phenomenon remain unknown. Here, we used bromocresol green pH indicator, noninvasive microtest, and microelectrode ion flux estimation techniques to demonstrate that B promotes root surface pH gradients in pea () roots, leading to alkalization in the root transition zone and acidification in the elongation zone, while Al inhibits these pH gradients. B significantly decreased Al accumulation in the transition zone (∼1.0-2.5 mm from the apex) of lateral roots, thereby alleviating Al-induced inhibition of root elongation. Net indole acetic acid (IAA) efflux detected by an IAA-sensitive platinum microelectrode showed that polar auxin transport, which peaked in the root transition zone, was inhibited by Al toxicity, while it was partially recovered by B. Electrophysiological experiments using the Arabidopsis () auxin transporter mutants (; []) and the specific polar auxin transporter inhibitor1-naphthylphthalamic acid showed that PIN2-based polar auxin transport is involved in root surface alkalization in the transition zone. Our results suggest that B promotes polar auxin transport driven by the auxin efflux transporter PIN2 and leads to the downstream regulation of the plasma membrane-H-ATPase, resulting in elevated root surface pH, which is essential to decrease Al accumulation in this Al-targeted apical root zone. These findings provide a mechanistic explanation for the role of exogenous B in alleviation of Al accumulation and toxicity in plants.
Engineered nanoparticles (NPs) are considered potential agents for agriculture as fertilizers and growth enhancers. However, their action spectrum differs strongly, depending on the type of NP, its concentrations, and plant species per se, ranging from growth stimulation to toxicity. This work aimed to investigate effects of iron oxide (Fe3O4) NPs on growth, photosynthesis, respiration, antioxidant activity, and leaf mineral content of wheat plants. Wheat seeds were treated with NP for 3 h and plants were grown in the soil at two light intensities, 120 and 300 μmol (photons) m−2·s−1, followed by physiological assessment at several time points. High NP treatment (200 and 500 mg·L−1) enhanced plant growth, photosynthesis and respiration, as well as increasing the content of photosynthetic pigments in leaves. This effect depended on both the light intensity during plant growth and the age of the plants. Regardless of concentration and light intensity, an effect of NPs on the primary photochemical processes was not observed. Seed treatment with NP also led to increased activity of ascorbate peroxidase and reduced malondialdehyde (MDA) content in roots and leaves. Treatment with Fe3O4 also led to noticeable increases in the leaf Fe, P, and K content. It is concluded that iron oxide (Fe3O4)-based NP could enhance plant growth by improving photosynthetic performance and the availability of Fe and P.
Root border cells, which form a cell layer around the root tip, seem to play multiple roles in the rhizosphere of the apical root. As these cells (species‐dependent dozens to several thousand per root tip) are rapidly sloughed off in water, studies in hydroponic culture fail to elucidate their role in most conventional physiological studies. The common method for harvesting these cells consists in germination of seeds in a humid atmosphere (usually a Petri dish), but labor and time constraints allow to yield only very limited amounts of uniform cells. We thus developed a low‐cost mist‐culture method, where intact border cells can be collected in the range of several grams. We applied this technique in a preliminary experiment where the influence of aluminum (Al) supply on calcium (Ca) release and viability of this cell type was studied. Purified detached border cells of pea were incubated with 0, 50, and 500 mmol m–3 AlCl3 solution (pH 4.5) for 90 min at a ratio of 3 × 105 cells (4 mL)–1. After incubation, cells contained 4.27 and 13.28 mg Al g–1 C at 50 mmol m–3 and 500 mmol m–3 AlCl3, respectively, while their total Ca content decreased correspondingly. Cell viability of border cells as tested by fluorescein diacetate‐propidium iodide (FDA‐PI) fluorescence yielded unexpected results: the test exhibited significantly lower vitality at 50, but not at 500 mmol m–3 AlCl3. Assessing mitochondrial activity by 3‐(4,5)‐dimethylthiazol‐2‐yl‐2,5 diphenyl‐tetrazolium‐bromide (MTT) reduction showed that viability decreased in a dose‐dependent manner with increasing Al concentrations. This apparent contradiction is attributed to the formation of dense mucilage around border cells at high Al concentrations, which likely inhibits the access of the dye PI or may chemically inactivate this compound and thus wrongly suggest higher viability. Mist culture allows harvesting selectively large amounts of homogeneous border cells quickly and to study their physiological reactions separated from the root tip.
Aluminum (Al) toxicity is the primary factor limiting crop growth in acidic soils. Boron (B) alleviates Al toxicity in plants, which is mainly considered to be due to the formation of Rhamnogalacturonan II-B (RGII-B) complexes, which helps to stabilize the cytoskeleton. It is unclear yet whether this is due to the increasing of net negative charges and/or further mechanisms. Kinetics of Al accumulation and adsorption were investigated using entire cells, cell wall and pectin of root border cells (RBCs) of pea (Pisum sativum), to reveal the mechanism of B in interacting with alkali-soluble and chelator-soluble pectin for an increased Al tolerance in RBCs. The results show that B could rescue RBCs from Al-induced cell death by accumulating more Al in the cell wall, predominately in alkali-soluble pectin. Boron also promotes Al3+ adsorption and inhibits Al3+ desorption from alkali-soluble pectin. Thus, more Al3+ is immobilized within the alkali-soluble pectin fraction and less in the chelator-soluble pectin, rendering Al3+ less mobile. Boron induces an increase of RG-II (KDO,2-keto-3-deoxyoctonic acid) content for forming more borate-RGII complexes, and the decrease of pectin methyl-esterification, thus creates more negative charges to immobilize Al3+ in cell wall pectin. The study provides evidence that abundant B supply enhances the immobilization of Al in alkali-soluble pectin, thus most likely reducing the entry of Al3+ into the symplast from the surroundings.
We investigated the hypothesis that a discrepancy of Al binding in cell wall constituents determines Al mobility in root border cells (RBCs) of pea (Pisum sativum), which provides protection for RBCs and root apices under Al toxicity. Plants of pea (P. sativum L. ‘Zhongwan no. 6’) were subjected to Al treatments under mist culture. The concentration of Al in RBCs was much higher than that in the root apex. The Al content in RBCs surrounding one root apex (104 RBCs) was approximately 24.5% of the total Al in the root apex (0–2.5 mm), indicating a shielding role of RBCs for the root apex under Al toxicity. Cell wall analysis showed that Al accumulated predominantly in alkali-soluble pectin (pectin 2) of RBCs. This could be attributed to a significant increase of uronic acids under Al toxicity, higher capacity of Al adsorption in pectin 2 [5.3-fold higher than that of chelate-soluble pectin (pectin 1)], and lower ratio of Al desorption from pectin 2 (8.5%) compared with pectin 1 (68.5%). These results indicate that pectin 2 is the primary target of Al immobilization in RBCs of pea, which impairs Al access to the intracellular space of RBCs and mobility to root apices, and therefore protects root apices and RBCs from Al toxicity.
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