植物适应酸铝胁迫机理的研究进展

酸铝胁迫是限制植物正常生长发育的重要非生物胁迫因子,严重制约了我国酸性土壤地区的农业生产水平。植物抵御酸铝胁迫的形式复杂多样,如分泌有机酸、提高根际pH、分泌黏液、细胞壁对Al³⁺的固定、有机酸对细胞溶质中Al³⁺的螯合与液泡区隔化等。现有研究多集中于常规生理特征分析,缺乏深入的分子生物学解析。基于此,本文对国内外植物适应酸铝胁迫机理的相关研究进行了归纳和总结,从酸铝胁迫对植物生长与生理代谢的影响、植物适应酸铝胁迫最主要的两种生理机制(Al排除机制、Al耐受机制)以及分子水平上调控相关耐铝基因进行了综述。最后针对现有研究的不足提出了展望,以期为深入揭示植物适应酸铝胁迫的机理以及挖掘适于酸土生长的优质作物资源提供理论依据。     

Zinc status does not affect aluminum deposition in tissues of rats

To examine whether zinc deficiency would increase the toxicity of dietary aluminum, weanling, male Sprague-Dawley rats were fed purified diets containing either 2 or 30 mg Zn/kg diet, with or without 500 mg Al/kg diet for 28 d. Individually pair-fed rats were fed the 30 mg Zn/kg diet with or without added aluminum to control for inanition secondary to zinc deficiency. Rats fed the 2 μg Zn/kg diet showed evidence of zinc deficiency, including anorexia, growth retardation, and depressed concentrations of zinc in tibias and livers. Zinc deficiency did not significantly increase the concentrations of aluminum in the tibias, livers, kidneys, or regions of the brain examined (cerebrum, cerebellum, midbrain, and hippocampus). Inclusion of aluminum in the diet did not alter aluminum concentrations in the various tissues. Under the conditions of this study, zinc deficiency did not result in greater sensitivity to dietary aluminum exposure.     

Renal aluminum excretion

Urinary aluminum (Al) excretion was studied in humans with normal and impaired renal function. Al was measured by atomic absorption spectrometry. In healthy volunteers (n=50), renal Al excretion was 12.2±8.5 μg/24 h. Two patients on plasma exchange therapy with normal renal function and an inadvertent load of 870 and 388 μg Al/treatment showed a 23 and 14% positive balance until next treatment.     

微生物铝毒害和耐铝毒机制研究进展

在酸性土壤中,铝毒是限制农作物生产的关键问题之一,铝同样对微生物产生毒害作用。研究微生物的铝毒害和耐铝毒机制可以为植物耐铝毒机制的研究提供一种新视角。目前的研究表明,铝作用于微生物细胞的细胞壁、细胞膜、细胞核和细胞器,影响微生物的物质和能量代谢,抑制微生物的生长和发育。针对这些毒害作用,铝毒耐受微生物通过多途径全方位的机制适应外界的铝毒环境。该文结合作者的研究工作,综述了微生物的铝毒害和耐铝毒机制。     

Effects of low-molecular-weight aluminum complexes on brain tissue calcium homeostasis

The in vitro effects of low-molecular-weight aluminum complexes (citrate, lactate, and ATP complex) on the Ca²⁺ uptake and aluminum-induced lipid peroxidation of brain tissue show that the modification of the calcium homeostasis is determined by the nature of the ligand and that there is no correlation between the aluminum-induced lipid peroxidation and the Ca²⁺ uptake. The same characteristics have been shown by a similar study performed with Ehrlich carcinoma cells. The electrophoretic analyses of the aluminum lactate-albumin and aluminum lactate-ATP interactions indicate an aluminum transfer from the lactate to the albumin and ATP ligands. The increased Ca²⁺ uptake when ATP is present in the incubation medium with aluminum citrate and aluminum lactate corroborates the suggested mediator role of ATP in cellular calcium homeostasis modification induced by iron.     

铝对豌豆生长的影响

研究酸性条件下不同浓度Al^3 对豌豆根芽生长的影响。结果表明,Al对豌豆根芽的生长有显著的抑制作用,对豌豆根尖细胞分裂产生危害的初始浓度为1mg.L^-1,Al诱导豌豆根尖染色体发生多种类型畸变,初始浓度为0．5mg.L^-1,低浓度下Al对豌豆根芽生长的抑制与植物对Ca、Mg的吸收没有明显的相关性。     

Actions of aluminum on voltage-activated calcium channel currents

Summary 1. Extracellular and intracellular effects of aluminum (Al) on voltage-activated calcium channel currents (VACCCs) of cultured rat dorsal root ganglion (DRG) neurons were investigated. Al (0.54 to 5.4 µg/ml=20–200 µM) applied extracellularly reduces VACCCs in a concentration-dependent manner. The IC₅₀ was calculated to be 2.3 µg/ml (83 µM). All types of VACCCs were similarly reduced by Al treatment. A slight shift of the current-voltage relation to depolarized potentials was observed for higher Al concentrations (>2 µg/ml). The action of Al was found to be use dependent, with little recovery (max. 20%) after wash.2. The effect of Al was highly pH dependent in the investigated range (pH 6.4 to 7.8). We observed a rightward shift of the concentration-response curve at pH 7.7 (IC₅₀:3.1 µg/ml) and a leftward shift at pH 6.4 (IC₅₀:0.56 µg/ml) compared to the concentration-response curve at pH 7.3.3. The VACCC declined when 2.7 µg/ml Al was added to the internal solution. A steady state was reached within a few minutes. Additional extracellular application of the same concentration lead to an additional decrease of the current. These observations strongly suggest the existence of both intracellular and extracellular accessible binding sites for Al on voltage-activated calcium channels (VACCs).4. The special characteristics of the action of Al on VACCCs, i.e., the irreversibility, use dependence, and pH dependence, as well as the additional internal binding site may contribute to its neurotoxicity.     

Hippocampal tin,aluminum and zinc in Alzheimer's disease

The use of inductively coupled plasma source mass spectrometry (ICP-MS) for multi-element analysis has led to the observation, in two separate studies, of increased blood tin in Alzheimer's disease (AD). We have therefore applied the technique of ICP-MS to hippocampal tissues obtained post-mortem from patients with AD and from controls. There was no significant difference in tin concentrations in AD. There were increased concentrations of aluminum and silicon, and reduced concentrations of zinc and selenium. It is postulated that displacement of hippocampal zinc by heavy metals may be important in producing clinical memory disturbance. However, analysis of the CA1 region, rather than of the dentate gyrus, would have been preferable.     

Interaction of aluminum ion with ATP. Mechanism of the aluminum inhibition of glycerol kinase and its reversal by spermine

Aluminum ion inhibited yeast glycerol kinase competitively with respect to the substrate MgATP. The K value of the enzyme for aluminum ion was about 3 M. Spermine at physiological concentrations prevented glycerol kinase from the inhibition by aluminum ion. Nuclear magnetic resonance spectroscopy showed the specific elimination by spermine of aluminum from the metal-ATP complex, but no dissociation of MgATP complex by spermine. Inhibition by aluminum ion of glycerol kinase as well as hexokinase can reduce the utilization of energy fuel in yeast. Change in polyamine concentration may control energy production in vivo, and is responsible for the development of age-related aluminum toxicity. © Rapid Science 1998.     

Continuous flow rhizostat for the study of aluminum toxicity

Abstract. Wide variations in total aluminum (AL) concentration reported to have a toxic effect on plants arc attributed to imprecise definition of the chemical environment at the root interface. Because AL can complex with natural ligands and form time-dependent metastable species in water, the requirements of a fixed AL speciation in the nutrient medium can only be met by simultaneously holding constant the pH, the AL/OH ratio and the renewal time of the solution. It is also important that the root be constantly perfused by a fresh solution so that the root interface is as chemically close to the bulk solution as possible. The rhizostat presented here uses a process controller to constantly replenish four tanks containing different combinations of nutrients and AL. The solution is pumped from each tank to a tray where it is continuously delivered to seedlings growing on artificial medium. This rhizostat provides an accurate control for AL speciation. Preliminary experiments on white spruce seedlings show that levels of 10 micromolal or less total AL are sufficient to cause toxic effects at pH of 4.5 and without the formation of AL complexes.     

Interactive regulation of nitrogen and aluminum in rice

Despite many studies on the high aluminum (Al) tolerance of rice (Oryza sativa), its exact mechanisms remain largely unknown. It is also unclear why Al improves growth of some plants. Our research on interactions between nitrogen (N) and Al may help to understand these phenomena. Previously, we found that ammonium-supplemented rice was more Al tolerant than nitrate-supplemented rice. Furthermore, Al-tolerant rice varieties preferred ammonium, while Al-sensitive ones preferred nitrate; in fact, Al tolerance was significantly correlated with the ammonium/nitrate preference among rice varieties. Al even enhanced growth of ammonium-supplemented rice, while it inhibited growth of nitrate-supplemented rice. Based on our own and other reports on N-Al interactions, we propose that intermediate products of N metabolism may play a role in rice Al tolerance. Al-enhanced ammonium utilization may explain why Al promotes growth of some plants, since Al often coexists with higher levels of ammonium than nitrate in acid soils.     

Targeted expression of SbMATE in the root distal transition zone is responsible for sorghum aluminum resistance

Aluminum (Al) toxicity is one of the major limiting factors for crop production on acid soils that comprise significant portions of the world's lands. Aluminum resistance in the cereal crop Sorghum bicolor is mainly achieved by Al‐activated root apical citrate exudation, which is mediated by the plasma membrane localized citrate efflux transporter encoded by SbMATE. Here we precisely localize tissue‐ and cell‐specific Al toxicity responses as well as SbMATE gene and protein expression in root tips of an Al‐resistant near‐isogenic line (NIL). We found that Al induced the greatest cell damage and generation of reactive oxygen species specifically in the root distal transition zone (DTZ), a region 1–3 mm behind the root tip where transition from cell division to cell elongation occurs. These findings indicate that the root DTZ is the primary region of root Al stress. Furthermore, Al‐induced SbMATE gene and protein expression were specifically localized to the epidermal and outer cortical cell layers of the DTZ in the Al‐resistant NIL, and the process was precisely coincident with the time course of Al induction of SbMATE expression and the onset of the recovery of roots from Al‐induced damage. These findings show that SbMATE gene and protein expression are induced when and where the root cells experience the greatest Al stress. Hence, Al‐resistant sorghum plants have evolved an effective strategy to precisely localize root citrate exudation to the specific site of greatest Al‐induced root damage, which minimizes plant carbon loss while maximizing protection of the root cells most susceptible to Al damage.     

27Al-NMR studies of aluminum transport across yeast cell membranes

Aluminum (Al) transport across yeast cells was studied using Dy(NO₃)₃ as a shift reagent by ²⁷Al-NMR spectroscopy. The results showed that (a) Al enters the yeast cells at 15 min and over a period of time, within 4 h, an equilibrium sets in between outside and inside Al; (b) citrate does not favor Al going into the yeast cells at pH 5.0; and (c) EDTA brings out all the Al that has entered the yeast cells. (Mol Cell Biochem 175: 59–63, 1997)     

In vivo and in vitro effects of aluminum treatment on rat liver mitochondrial function

This study examines the effect on mitochondrial respiration and permeability of in vivo and in vitro aluminium (Al) exposure. Rats were treated intraperitoneally with AlCl₃ to achieve serum and liver Al concentrations comparable to those seen in Al-related disorders. Mitochondria isolated from Al-treated rats had higher (p<0.01) Al concentration, lower (p<0.05) state 3 respiration, respiratory control (RCR), and ADP/O ratio (succinate substrate), and greater passive swelling in 100 mM KCl or 200 mM NH₄NO₃ than controls. The in vitro addition of Al (0–180 μM) to mitochondria from normal rats also decreased (p<0.01) state 3 respiration, RCR, and ADP/O and stimulated passive swelling in KCl and NH₄NO₃ at 42–180 μM Al. These studies show that Al depresses mitochondrial energy metabolism and increases membrane permeability. The toxicity associated with Al may be related to its effect on mitochondria.     

Oxidative stress due to aluminum exposure induces eryptosis which is prevented by erythropoietin

The widespread use of aluminum (Al) provides easy exposure of humans to the metal and its accumulation remains a potential problem. In vivo and in vitro assays have associated Al overload with anemia. To better understand the mechanisms by which Al affects human erythrocytes, morphological and biochemical changes were analyzed after long-term treatment using an in vitro model. The appearance of erythrocytes with abnormal shapes suggested metal interaction with cell surface, supported by the fact that high amounts of Al attached to cell membrane. Long-term incubation of human erythrocytes with Al induced signs of premature erythrocyte death (eryptosis), such as phosphatidylserine externalization, increased intracellular calcium, and band 3 degradation. Signs of oxidative stress, such as significant increase in reactive oxygen species in parallel with decrease in the amount of reduced glutathione, were also observed. These oxidative effects were completely prevented by the antioxidant N-acetylcysteine. Interestingly, erythrocytes were also protected from the prooxidative action of Al by the presence of erythropoietin (EPO). In conclusion, results provide evidence that chronic Al exposure may lead to biochemical and morphological alterations similar to those shown in eryptosis induced by oxidant compounds in human erythrocytes. The antieryptotic effect of EPO may contribute to enhance the knowledge of its physiological role on erythroid cells. Irrespective of the antioxidant mechanism, this property of EPO, shown in this model of Al exposure, let us suggest potential benefits by EPO treatment of patients with anemia associated to altered redox environment.     

Spectrofluorimetric determination of aluminum ions via complexation with luteolin in absolute ethanol

An optimized and validated spectrofluorimetric method has been developed for the rapid determination of aluminum in absolute ethanol. The method is based on the chelation of aluminum and luteolin which results in a complex exhibiting an intense emission signal. The characterization of Al–luteolin complex was studied using ultraviolet–visible spectrometry, infrared spectrometry, fluorescence and mass spectrometry. The complex stoichiometry ratio of aluminum:luteolin was 1:2. The fluorescence of the complex was monitored at an emission wavelength of 545 nm with excitation at 518 nm. The linear concentration range was 6.5 × 10^‐7 to 4.0 × 10^‐5 M with a correlation coefficient of r = 0.998. The detection limit was 5.0 × 10^‐7 M. The method was appropriately validated and yielded relative standard deviations of < 2.3% (n = 5), which was considered acceptable. The method was successfully applied in the determination of aluminum in river water, skin care products and pharmaceutical samples. Copyright © 2013 John Wiley & Sons, Ltd.     

Rapid evaluation of sorghum for aluminum tolerance

Two screening methods tested in this study were based on the observation that roots of freshly germinated seeds exhibit their relative tolerance to Al during the first 48 hours of growth. Sorghum (Sorghum bicolor L. Moench) varieties were evaluated using toxic/nontoxic soil pairs in petri dishes and toxic/nontoxic solution pairs in a flowing aqueous system. The soils had 0, 61, and 72% Al saturation and the solutions had 0, 1.85 and 3.70 M l^–1 Al from AlCl₃ and 0.25 mM l^–1 Ca from Ca(NO₃)₂. Relative root lengths in both systems correctly differentiated 13 genotypes of known Al tolerance into tolerant and intolerant groups. Twenty six other sorghums genotypes were also screened using genotypes of known Al reactions as checks. The soil with 72% Al saturation and the 1.85 M l^–1 Al solution gave the highest correlation between the two systems and both effectively arrayed sorghums of known and unknown tolerance. For routine screening the authors prefer the soil system for its simplicity, efficiency, and use of a natural growth medium.Journal paper 11637. Purdue University Agricultural Experiment Station, Lafayette, Indiana. Contribution from the Agronomy Department.     

小麦根系对铝毒的反应及其与根细胞壁组分和细胞壁对铝的吸附-解吸性能的关系

研究了小麦根系对铝毒的反应与不同根段细胞壁的组分及细胞壁对铝的吸附解吸性能的关系。结果表明,30μmol/LAlCl3可迅速抑制根系伸长,在铝处理30h时其根长仅为对照的30.2%;小麦根系相对伸长率随着铝浓度的提高而急剧降低,30μmol/LAlCl3处理24h对根系伸长的抑制率高达70.9%。小麦根系中距根尖0~10mm根段的铝含量和细胞壁中果胶糖醛酸含量明显高于距根尖10~20mm根段;距根尖0~10mm根段细胞壁对铝的吸附量明显大于距根尖10~20mm根段,而前者吸附态铝的解吸率低于后者;铝浓度从10μmol/L提高到20μmol/L时细胞壁对铝的吸附量增加,但对铝的解吸没有明显影响。采用1.0mol/LNH3·H2O对细胞壁预处理2h降低果胶甲基酯化程度后,铝吸附量降低了20.9%,但对铝解吸率没有影响。由此可见,小麦根尖是铝毒的主要位点,细胞壁果胶含量和果胶甲基酯化程度对小麦不同根段细胞壁对铝的吸附、积累具有重要作用,铝与细胞壁的结合是根系对铝毒胁迫反应的重要原因。