Silicon dynamics in the rhizosphere: Connections with iron mobilization

Silicon (Si) is a beneficial element for plants as it increases their resistance to several biotic and abiotic stresses. In the rhizosphere, root exudates, especially when released by nutritionally stressed plants, promote the mineral weathering and, consequently, influence Si biogeochemistry. This study aims at evaluating the mineralogical alterations in the rhizosphere of Fe‐deficient or Fe‐sufficient barley plants grown either in a natural or in an artificial calcareous soil, focusing on the dynamics of both Fe and Si. After 6 d of soil–plant contact, X‐ray diffraction (XRD) analysis of rhizosphere soil samples of Fe‐deficient plants revealed, for both natural and artificial soil, a decrease of amorphous phases and an increase of smectite compared to the unplanted soil. Root exudates released by Fe‐deficient plants were most likely the main responsible for the weathering of the amorphous phases by a ligand controlled dissolution mechanism. When the soil–plant contact was prolonged up to 12 d, plants overcame Fe nutritional stress and their effect on soil mineralogy completely changed, as proved by the considerable increase of amorphous and decrease of smectite. Smectite decrease might evidence the effort of plant to mobilize Si and micronutrients other than Fe from the soil through the exudation of organic ligands. When the artificial soil was treated with Fe‐sufficient barley plants, the mineral weathering trend appeared reversed compared to the experiments with Fe‐deficient plants. Plant nutritional status regulates the root exudation pattern and, consequently, drives mineral weathering processes in the rhizosphere. Barley has shown to be able to mobilize Si from smectite, yet depending on its Fe supply and proving the strict connection between Si and Fe dynamics in the rhizosphere.     

麦根酸的分泌及其对土壤中铁的有效性的影响

Large amounts of phytosiderophore are detected from both the solution and the rhizosphere soil when cereal crops are under Fe deficiency stress.The extension of phytosiderophore in the rhizosphere soil is found only within 1 mm apart from the root surface.The rate of phytosiderophore secretion is negatively related to chlorophyll content in young leaves and positively related to the Fe-solubilizing capacity.Results from in vitro experiments whow 10 μmoles mugineic acid can dissolve 501 μg Fe from iron hydroxide and 146 ug from strengite.Thus,phytosiderophore can considerably enhance the soil iron availability by increasing the solubility of amorphous iron hydroxide and iron phosphate,and active Fe is consequently accumulated in the plant rhizosphere,43% higher than in the bulk soils There is evidence to support that mugineic acid chelates with Fe%3 at a rate of 1:1 in the acid condition.In addition,we observe mugineic acid has certain effects on mobilization of P as well as Fe by dissolving the insoluble iron phosphate,And phytosiderophore seems to be an effective remedy for the chlorosis of dicotyledonous plants.     

Factors Affecting the Formation,Nature, and Properties of Iron Precipitation Products at the Soil–Root Interface

Abstract

A number of iron oxides (hematite, goethite, lepidocrocite, maghemite, and magnetite) or short‐range ordered precipitates (ferrihydrite) may be found in soil environments, but in the rhizosphere the presence of organic ligands released by plants (exudates) or microorganisms promote the formation of ferrihydrite. Iron ions are liberated into soil solution by acidic weathering of minerals and then precipitated either locally or after translocation in soil environments. Humic and fulvic acids as well as organic substances produced by plants and microorganisms are involved in the weathering of primary minerals. Organic compounds play a very important role in the hydrolytic reactions of iron and on the formation, nature, surface properties, reactivity, and transformation of Fe oxides. Organic substances present in the rhizosphere interact with Fe promoting the formation of ferrihydrite and organo‐mineral complexes. The solubility of Fe precipitation products is usually low. However, the formation of soluble complexes of Fe(II) or Fe(III) with organic ligands, usually present in the rhizosphere increases the solubility of Fe‐oxides. Mobilization of Fe from Fe oxides by siderophores is of great importance in natural systems. They can form stable Fe(III) complexes (pK up to 32) and thus mobilize Fe from Fe(III) compounds. These higher Fe concentrations are important for the supply of Fe to plant roots which excrete organic acids at the soil–root interface. Iron oxides adsorb a wide variety of organic and inorganic anions and cations, which include natural organics, nutrients, and xenobiotics. There is competition between anions and cations for the surfaces of Fe‐oxides. Root exudates suppress phosphate or sulfate adsorption on Fe‐oxides. This is a mechanism by which plant roots mobilize adsorbed phosphate and improve their phosphate supply. Anions adsorption on iron oxides modify their dispersion/flocculation behavior and thus their mobility in the soil system. That can increase or decrease the possibility of contact between Fe‐oxides and organics or organisms able to dissolve them.     

Manganese‐iron interactions in the plant‐soil system

Dicotyledonous plants had significantly higher Mn and Fe intake rates on a near neutral soil, had a significantly higher Mn intake rate on a slightly calcareous soil, but had lower Mn and Fe intake rates on a calcareous soil, than monocotyledonous plants. This dependency on soil reaction suggests that dicots utilize primarily a chemical reduction mechanism, whereas monocots utilize some less pH‐dependent mechanism (possibly Mn(III)‐, Fe(III)‐organic complexation) to mobilize soil Mn and Fe. Soluble soil Mn and Fe fractions in the rhizosphere were consistently positively correlated with each other, as were Mn and Fe intake rates. These results suggest that for soil‐grown plants, Mn and Fe uptake was positively interrelated because both Mn and Fe were mobilized by similar root processes.     

Water- and pyrophosphate-extractable humic substances fractions as a source of iron for Fe-deficient cucumber plants

The capacity of Fe-deficient cucumber plants to utilise water-extractable and pyrophosphate-extractable humic substances as a source of Fe was investigated. Plants were grown for 13 days in nutrient solution in the presence or absence of Fe and during the last 7 days water-extractable and pyrophosphate-extractable humic substances were added to the solution at a final concentration of 5 μg organic C ml^–1. The water-extractable humic fraction did not significantly modify leaf area and dry matter accumulation, leaf total Fe or chlorophyll content of cucumber plants adequately supplied with Fe. In contrast, pyrophosphate-extractable humic substances caused a slight but significant decrease of all the leaf parameters considered, with the exception of the chlorophyll content. Root Fe content of Fe-sufficient plants was decreased by more than 50% in the presence of each humified fraction. Addition of each humic fraction to Fe-deficient plants led to a partial disappearance of leaf chlorosis symptoms with a significant increase in chlorophyll and leaf Fe content. Fe content of roots was also significantly increased in Fe-deficient plants by the addition of humic substances to the nutrient solution. These results show that Fe-deficient cucumber plants can utilise Fe contained in the two fractions of humified organic matter. However, by calculating the amount of total Fe accumulated per plant in the presence of water-extractable or pyrophosphate-extractable humic substances, it could be seen that Fe contained in the water-extractable humic fraction was almost totally used by Fe-deficient cucumber plants, while that present in the pyrophosphate-extractable fraction could only be partially absorbed. The results strongly support a role of humified organic matter in Fe nutrition of plants and are discussed in terms of a possible interaction between soil humic substances and the biochemical mechanisms involved in the plant response to Fe deficiency. Received: 6 November 1996     

Root exudation of sugars,amino acids,and organic acids by maize as affected by nitrogen,phosphorus, potassium,and iron deficiency

Root exudates play a major role in the mobilization of sparingly soluble nutrients in the rhizosphere. Since the amount and composition of major metabolites in root exudates from one plant species have not yet been systematically compared under different nutrient deficiencies, relations between exudation patterns and the type of nutrient being deficient remain poorly understood. Comparing root exudates from axenically grown maize plants exposed to N, K, P, or Fe deficiency showed a higher release of glutamate, glucose, ribitol, and citrate from Fe‐deficient plants, while P deficiency stimulated the release of γ‐aminobutyric acid and carbohydrates. Potassium‐starved plants released less sugars, in particular glycerol, ribitol, fructose, and maltose, while under N deficiency lower amounts of amino acids were found in root exudates. Principal‐component analysis revealed a clear separation in the variation of the root‐exudate composition between Fe or P deficiency versus N or K deficiency in the first principal component, which explained 46% of the variation in the data. In addition, a negative correlation was found between the amounts of sugars, organic and amino acids released under deficiency of a certain nutrient and the diffusion coefficient of the respective nutrient in soils. We thus hypothesize that the release of dominant root exudates such as sugars, amino acids, and organic acids by roots may reflect an ancient strategy to cope with limiting nutrient supply.     

Iron-deficiency response and expression of genes related to iron homeostasis in poplars

Iron (Fe) deficiency is a serious agricultural problem, especially in calcareous soils, which are distributed worldwide. Poplar trees are an important biomass plant, and overcoming Fe deficiency in poplars will increase biomass productivity worldwide. The poplar Fe-deficiency response and the genes involved in poplar Fe homeostasis remain largely unknown. To identify these genes and processes, we cultivated poplar plants under Fe-deficient conditions, both in calcareous soil and hydroponically, and analyzed their growth rates, leaf Soil and Plant Analyzer Development (SPAD) values, and metal concentrations. The data clearly showed that poplars have notable growth defects in both calcareous soil and a Fe-deficient hydroponic culture. They exhibited serious chlorosis of young leaves after 3 weeks of Fe-deficient hydroponic culture. The Fe concentrations in old leaves with high SPAD values were markedly lower in Fe-deficient poplars, suggesting that poplars may have good translocation capability from old to new leaves. The Zn concentration in new leaves increased in Fe-deficient poplars. The pH of the hydroponic solution decreased in the Fe-deficient culture compared to the Fe-sufficient culture. This finding shows that poplars may be able to adjust the pH of a culture solution to better take up Fe. We also analyzed the expression of Fe homeostasis-related genes in the roots and leaves of Fe-sufficient and Fe-deficient poplars. Our results demonstrate that PtIRT1, PtNAS2, PtFRO2, PtFRO5, and PtFIT were induced in Fe-deficient roots. PtYSL2 and PtNAS4 were induced in Fe-deficient leaves. PtYSL3 was induced in both Fe-deficient leaves and roots. These genes may be involved in the Fe uptake and/or translocation mechanisms in poplars under Fe-deficient conditions. Our results will increase a better understanding of the Fe-deficiency response of poplars and hence improve the breeding of Fe-deficiency-tolerant poplars for improved biomass production, the greening of high pH soils, and combatting global warming.     

Bioenergy grass [Erianthus ravennae (L.) Beauv.] secretes two members of mugineic acid family phytosiderophores which involved in their tolerance to Fe deficiency

Ravenna grass, Erianthus ravennae (L.) Beauv. (E. ravennae) is a potential high biomass-energy crop with low input requirements. Iron (Fe) deficiency in calcareous soils is a widespread agronomic problem which reduces crop yields. Fe is sparingly soluble under aerobic conditions at high soil pH, such as in calcareous soils; therefore, plants cannot take up enough Fe. Increasing crop productivity of giant grasses, such as Ravenna grass in calcareous soil, has a positive effect by alleviating environmental problems. However, the growth character in calcareous soil and Fe homeostatic trait of Ravenna grass are largely unknown. In this study, we analyzed characteristics of Ravenna grass. The growth of E. ravennae plants were impaired in calcareous soil compared to those in the normal soil. In calcareous soil, the growth of E. ravennae plants differ among the water and fertilizer conditions; E. ravennae plants were grown better in the submerged condition adding micronutrient among conditions. These results suggested that impaired growth of E. ravennae in calcareous soil might be micronutrient shortage. We found that E. ravennae roots possess Fe reductase activities which were upregulated under Fe-deficient conditions. E. ravennae produced and secreted mugineic acid (MA) and deoxymugineic acid (DMA) to acquire Fe from the soil. The amount of MA was higher than that of DMA. Thus, E. ravennae might have both partial Strategy-I and Strategy-II Fe uptake systems. E. ravennae intercropped with transgenic rice plants producing and secreting MA through the introduction of the barley MA synthase gene showed improved growth compared to monocropped E. ravennae plants, suggesting that the increased amounts of MA enhanced their tolerance to Fe deficiency. Our results suggest that there is a considerable potential to improve the growth of E. ravennae plants in calcareous soils by enhancement of their Fe uptake systems through increase of MA production.     

Distribution of exudates of lupin roots in the rhizosphere under phosphorus deficient conditions

Abstract

The distribution of secretory acid phosphatase and organic acids enhanced by phosphorus deficiency in lupin rhizosphere was investigated using a rhizobox system which separated the rhizosphere soil into 0.5 mm fractions. In the soil fraction closest to the root surface, the lupin exudates displayed an acid phosphatase activity of 0.73 u g^?1 dry soil and citrate concentration of 85.2 μmol g^?1 dry soil, respectively. The increase of the acid phosphatase activity-induced an appreciable depletion of organic P in the rhizosphere, indicating that lupin efficiently utilized the organic P from soil through the enzyme activitye The sterile treatments demonstrated that the acid phosphatase in the rhizosphere was mainly derived from lupin root secretions. The secretory organic acids enhanced considerably the solubility of the inorganic P in three types of soil and a sludge. However, the secretory acid phosphatase and organic acids from lupin roots were only detected in a considerable amount in 0-2.5 mm soil fractions from root surface.     

Rhizosphere calcareous soil P-extraction at the expense of organic carbon from root-exuded organic acids induced by phosphorus deficiency in several plant species

The amount of organic acids in root exudates rapidly increases under phosphorus (P) deficiency. Loss of carbon from root-exuded organic acids, which are derived from plant net photosynthetic products, is generally considered negligible. The present study aimed to study the characteristics of root-exuded organic acids, extraction of phosphorus (P extraction) in calcareous soil and the expression of organic carbon from root-exuded organic acids in two woody Moraceae plants (Broussonetia papyrifera L. Vent and Morus alba L.) and two herbaceous cruciferous plants (Orychophragmus violaceus L. Schulz and Brassica napus L.) under two P levels (P-normal and P-deficient). P extraction and the amount of root-exuded organic acids simultaneously and disproportionately increased in the four plant species tested under P deficiency. The maximum P-extracting capability of the four plant species was observed after 40 days of treatment. Additionally, the response of root-exuded organic acids induced by P deficiency was species-specific. B. papyrifera extracted more P in calcareous soil, and expended less organic acid for the same P-extraction than M. alba. Similarly, O. violaceus extracted more P in calcareous soil, and consumed less organic acid for the same level of P-extraction than B. napus. Root-exuded oxalic and malic acids accounted for most of the increment of P extraction in woody Moraceae plants, while root-exuded citric acid accounted for most of the increment in P extraction in herbaceous cruciferous plants. B. papyrifera and O. violaceus exhibited the strongest P-extracting capability at lower expense of organic carbon over the treatment duration in the four plant species. O. violaceus had the most rapid response of root-exuded organic acids to P deficiency, while B. napus had the slowest response. Thus, rapid response with low organic carbon cost and high efficiency of extraction on P in calcareous soil may underlie the strong adaptability of B. papyrifera and O. violaceus to a Karst environment.     

Organic acid behaviour in a calcareous soil implications for rhizosphere nutrient cycling

Calcareous soils are frequently characterized by the low bioavailability of plant nutrients. Consequently, many vascular plant species are unable to successfully colonize calcareous sites and the floristic composition of calcareous and acid silicate soils has been shown to differ markedly. The root exudation of oxalate and citrate has been suggested to play a pivotal role in same nutrient acquisition mechanisms operating in calcareous soils. The aim of this study was therefore to investigate the nutrient extraction efficiency of three individual organic acids commonly identified in root exudates, i.e. citric, malic and oxalic acid. Our results clearly demonstrate the context dependent nature of nutrient release by organic acids. The degree of P extraction was highly dependent on which organic acid was added, their concentration and pH, and their contact time with the soil. P is generally more efficiently extracted by organic acids at a high pH and follows the series oxalate>citrate>malate. The opposite relationship between pH and extraction efficiency was apparent for most other cations examined (e.g. Zn, Fe), which are more efficiently extracted by organic acids at low pH. A serious constraint to the ecological importance of organic acid exudation in response to P deficiency is, however, their very low P mobilization efficiency. For every mol of soil P mobilized, 1000 mol of organic acid has to be added. It can, however, be speculated that in a calcareous soil with extremely low P concentrations it is still beneficial to the plants to exude organic acids in spite of the seemingly high costs in terms of carbon.     

Iron fertilizers applied to calcareous soil on the growth of peanut in a pot experiment

A greenhouse pot experiment was conducted with peanuts (Arachis hypogaea L., Fabceae) to evaluate iron compound fertilizers for improving within-plant iron content and correcting chlorosis caused by iron deficiency. Peanuts were planted in containers with calcareous soil fertilized with three different granular iron nitrogen, phosphorus and potassium (NPK) fertilizers (ferrous sulphate (FeSO₄)–NPK, Fe–ethylendiamine di (o-hydroxyphenylacetic) (EDDHA)–NPK and Fe–citrate–NPK). Iron nutrition, plant biomass, seed yield and quality of peanuts were significantly affected by the application of Fe–citrate–NPK and Fe–EDDHA–NPK to the soil. Iron concentrations in tissues were significantly greater for plants grown with Fe–citrate–NPK and Fe–EDDHA–NPK. The active iron concentration in the youngest leaves of peanuts was linearly related to the leaf chlorophyll (via soil and plant analyzer development measurements) recorded 50 and 80 days after planting. However, no significant differences between Fe–citrate–NPK and Fe–EDDHA–NPK were observed. Despite the large amount of total iron bound and dry matter, FeSO₄–NPK was less effective than Fe–citrate–NPK and Fe–EDDHA–NPK to improve iron uptake. The results showed that application of Fe–citrate–NPK was as effective as application of Fe–EDDHA–NPK in remediating leaf iron chlorosis in peanut pot-grown in calcareous soil. The study suggested that Fe–citrate–NPK should be considered as a potential tool for correcting peanut iron deficiency in calcareous soil.     

Interactive effects of organic acids in the rhizosphere

Organic acids released into the rhizosphere may perform many beneficial functions to the plant including metal detoxification and enhancement of nutrient acquisition. Typically, these organic acids are studied in isolation; however, roots simultaneously exude a cocktail of organic acids and other substances, and their combined impact on rhizosphere processes may be quite different. It has been hypothesized that some exudates may play secondary roles (e.g. inhibitors of microbial activity, blockage of sorption sites), which might enhance the longevity and nutrient-mobilization capacity of others. Here we investigated how the decomposition, sorption and P-solubilizing effects of citrate, malate and oxalate are affected by the presence of malonate and shikimate. We found that in a range of agricultural soils the decomposition of citrate, malate and oxalate was rapid, but not influenced by the presence of large quantities of shikimate or malonate. This suggests that the individual organic acids are taken up by different transport mechanisms or components of the microbial community. At large concentrations, malonate decreased sorption of citrate, malate and oxalate on the soil, whilst shikimate had little effect. The capacity of citrate, malate and oxalate to desorb P was significantly greater in cocktails containing malonate compared with the single organic acid; no effect was seen with shikimate. We conclude that neither malonate nor shikimate at realistic concentrations will significantly affect the biodegradation of citrate, malate or oxalate in the rhizosphere, and while malonate did enhance P desorption, this effect is additive rather than synergistic. Overall, we found little evidence that malonate and shikimate act as secondary regulators of citrate, malate and oxalate behavior in soil.     

The effects of Fe deficiency on low molecular weight organic acid exudation and cadmium uptake by Solanum nigrum L.

Abstract

The influence of Fe-deficiency on the root exudation of low molecular weight organic acids (LMWOAs), pH alteration and cadmium (Cd) accumulation and translocation were investigated in morel (Solanum nigrum L.) in hydroponic culture experiments. Tartaric, citric, malic and acetic acids were monitored because these acids were abundant and often detected as root exudates. Results showed that Fe-deficient plants excreted large amounts of LMWOAs in comparison with Fe-sufficient plants across all Cd treatments (p <0.05). In both cases the concentrations of the four organic acids were tartaric > citric > malic > acetic. The results showed that the Fe-deficient plants with higher concentrations of LMWOAs accumulated more Cd (p <0.05) and induced a decrease in solution pH compared with the Fe-sufficient plants. Cadmium accumulation in the Fe-deficient and Fe-sufficient plants had significant positive correlations with the exudation of malic and acetic acids (p <0.05 and p<0.01). Cadmium accumulation in the Fe-sufficient plants had a significant (p<0.01) positive correlation with the exudation of tartaric acid, whereas there was a negative correlation (p<0.01) between Cd accumulation and the exudation of tartaric acid in the Fe-deficient plants. No significant correlation between the exudation of citric acid and Cd accumulation was obtained. Our results indicate that Fe-deficiency could induce Cd accumulation and translocation through an increase of LMWOAs exudation and pH alteration, both of which enhance Cd bioavailability.     

低分子量有机酸对石灰性潮土无机磷形态转化的影响

无菌培养条件下,模拟缺磷胁迫时植物根系分泌的低分子量有机酸种类和数量,采用石灰性土壤无机磷分级方法,研究了低分子量有机酸对石灰性潮土无机磷形态转化的影响。结果表明:①在石灰性潮土中无机磷主要以有效性较低的磷酸钙盐(Ca10-P等)形式存在,而有效性较高的形式(Ca8-P等)含量较少,Ca2-P就更少。②有机酸通过结合土壤中可以使磷固定的Ca、Fe及Al,致使根际范围酸化,促进磷酸盐的形态转换,增加磷的有效性。这种促进能力因有机酸种类和性质的不同而异,其促进能力大小顺序为草酸>柠檬酸>酒石酸。     

有机酸对活化土壤中镉和小麦吸收镉的影响

向土壤中加入外源有机酸,研究有机酸对活化土壤中镉的作用和小麦吸收镉的影响。结果表明：有机酸对土壤中镉有一定的活化能力,对镉活化能力强弱顺序为ＥＤＴＡ〉缺铁小麦根分泌物〉柠檬酸〉苹果酸〉水。但ＥＤＴＡ却降低了小麦地上部镉的含量,缺铁小麦根分泌物明显增加了小麦地上部镉含量。与对照相比,柠檬酸和苹果酸对小麦地上部的镉含量有一定的促进作用,也增加了小麦地上部镉含量。     

Sorption of organic acids in acid soils and its implications in the rhizosphere

Organic acids have been implicated in many soil-forming and rhizosphere processes, but their fate in soil is poorly understood. We examined the sorption of four simple short-chain organic acids (citric, oxalic, malic and acetic) in five acid soils and on synthetic iron hydroxide (ferrihydrite). The results for both soils and ferrihydrite indicated that the sorption depended on concentration in the following order of strength: phosphate >> oxalate > citrate > malate >> acetate. The sorption reactions in soil were shown to be little influenced by pH, whereas for ferrihydrite, sorption of all ligands increased strongly with decreasing pH. The sorption of organic anions onto ferrihydrite was influenced to a lesser extent by the presence of metal cations in solution. From the results we calculated that when organic acids enter solution they rapidly become sorbed onto the soil's exchange complex (> 80% within 10 min), and we believe that this sorption will greatly diminish their effectiveness to mobilize nutrients from the rhizosphere.     

Amino acid sequence of proteins induced in Fe-deficient stressed alfalfa (Medicago sativa L.) roots

Alfalfa (Medicago sativa L.) grows well in soils with a moderately high pH and dissolves insoluble iron in the rhizosphere. We have investigated active uptake mechanisms under Fe-deficient nutrient conditions and the effects of Fe-deficiency on plants. Previously, we observed that Fe-deficient alfalfa roots exuded many compounds (Masaoka et al. 1993) such as fiavonoids. We also identified a new compound “alfafuran” which is a phenol compound and is different from organic acids or phytosiderophore-type amino acid derivatives exuded by Fe-deficient plant roots. This compound is also very effective in dissolving ferric phosphate (Noguchi et al. 1994), suggesting that alfalfa may have developed several strategies against Fe-deficient stress including the exudation of organic compounds like alfafuran which accelerate the Fe³⁺-reducing activity on the root cell membrane to dissolve insoluble iron compounds. Suzuki et al. (1995, 1997) observed that in barley several peptide spots obtained by electrophoresis were induced under Fe-deficient stress when mugineic acid-family phytosiderophores were secreted from the roots. They suggested that these peptides control the mugineic acid synthesis and secretion. We examined the peptides induced in Fe-deficient alfalfa roots.     

Rhizospheric organic compounds in the soil–microorganism–plant system: their role in iron availability

Poor iron (Fe) availability in soil represents one of the most important limiting factors of agricultural production and is closely linked to physical, chemical and biological processes within the rhizosphere as a result of soil–microorganism–plant interactions. Iron shortage induces several mechanisms in soil organisms, resulting in an enhanced release of inorganic (such as protons) and organic (organic acids, carbohydrates, amino acids, phytosiderophores, siderophores, phenolics and enzymes) compounds to increase the solubility of poorly available Fe pools. However, rhizospheric organic compounds (ROCs) have short half‐lives because of the large microbial activity at the soil–root interface, which might limit their effects on Fe mobility and acquisition. In addition, ROCs also have a selective effect on the microbial community present in the rhizosphere. This review aims therefore to unravel these complex dynamics with the objective of providing an overview of the rhizosphere processes involved in Fe acquisition by soil organisms (plants and microorganisms). In particular, the review provides information on (i) Fe availability in soils, including mineral weathering and Fe mobilization from soil minerals, ligand and element competition and plant‐microbe competition; (ii) microbe–plant interactions, focusing on beneficial microbial communities and their association with plants, which in turn influences plant mineral nutrition; (iii) plant–soil interactions involving the metabolic changes triggered by Fe deficiency and the processes involved in exudate release from roots; and (iv) the influence of agrochemicals commonly used in agricultural production systems on rhizosphere processes related to Fe availability and acquisition by crops.     

油葵和苦荬菜根际土壤中镉和锌的活化机制及修复潜力比较

  目的  研究油葵和苦荬菜根际土壤固、液相对镉（Cd）和锌（Zn）的活化机制,比较两种植物在轻、中度复合污染农田的修复潜力。  方法  通过大田试验种油葵和苦荬菜,测定成熟期土壤的pH值、有机酸、重金属总量及其生物有效性;测定土壤溶液中的溶解性有机质（DOM）、主要离子、水溶态重金属及其形态分布;测定植物各部位中重金属的浓度及形态,通过计算重金属在植物中的富集系数（BCF）和转运系数（TF）,比较两种植物对土壤重金属污染的修复潜力。  结果  油葵和苦荬菜根系分泌的低分子有机酸均使根际土壤pH值下降明显,显著低于非根际土壤（P < 0.05）;苦荬菜根际土中低分子有机酸及DOM的浓度显著高于油葵根际土（P < 0.05）。两种植物根际土壤溶液中的Cd以离子态和DOM结合态为主,Zn以离子态为主;两种植物根际土壤中有效态的Cd差异不显著,油葵根际有效态Zn显著高于苦荬菜;两种植物根际土壤的Zn和Cd有效态与土壤溶液中Cd-DOM和Zn-DOM呈显著相关。苦荬菜根对重金属的富集能力较强,但油葵地上部分能吸收转运更多的Cd和Zn,并在叶中以毒性较低的不溶性磷酸盐结合态和草酸结合态富集。  结论  两种植物根际分泌的有机酸可以增加根际土壤中的Cd-DOM和Zn-DOM的浓度,提高土壤中的Cd和Zn的有效性,苦荬菜根际对重金属有较强的活化能力,但油葵地上部分对Cd和Zn的吸收转运能力更强。两种植物都具有较强的土壤重金属修复潜力,但从经济角度出发,油葵更适合现阶段我国农田重金属污染的修复。