微生物在植物铁营养中的潜在作用

根据近十多年来相关研究成果讨论了土壤微生物在植物根系吸收铁中可能的作用机理。这种机理可能包括缺铁植物根系分泌小分子有机化合物,如酚类和黄素类等化合物,这些化合物作为抑菌剂和(或)作为微生物生长的碳源物质来影响根际(Rhizosphere)微生物的群落结构,并在植物根际诱导形成特异性微生物种群,此类特异性微生物转而通过分泌高铁载体(Siderophore),增加土壤中铁的生物有效性,从而提高了根系对铁的吸收。此外,与植物根系共生的一些微生物也会改善植物的铁营养,这种作用可能包括:根瘤菌(Rhizobium)的结瘤作用,增强植物耐缺铁的生理响应;根系感染的菌根真菌通过增加植物根系的养分吸收面积和分泌对铁具有螯合作用的物质来改善植物的铁营养。本文在讨论这种可能的微生物作用机制的基础上,指出今后的研究方向和有待解决的问题。     

根际动态过程与植物营养

养分的有效性是由土壤物理、化学和生物学特性,特别是根系主导的根际动态过程所决定的。根系引起根际pH值和氧化还原电位、根分泌物以及由此引起微生物种群、数量和活性的改变,从根本上决定着根际养分的动态。而根系主导的根际动态又具有明显的基因型差异,并受植物营养状况的诱导和调控。因此,根际动态变化的方向和强度对植物适应土壤化学和物理逆境具有重要意义。本文从根际的一般概念入手,综述了近十余年来国内外根际动态研究的新成果,重点讨论了根际动态与植物营养的关系,并运用根际微生态系统的概念,把根际动态与植物对养分胁迫的适应性及其调控机理紧密联系起来,使根际动态和植物矿质营养基因型差异机理两个基础性研究热点融为一体,为解决重大的全球性环境、生态以及农业持续发展问题提供新的途径和理论依据。     

施肥对婆婆纳、离子芥、小花糖芥3种麦田杂草根际pH的影响

根际是受根系影响而在物理、化学和生物学性质等方面均不同于土体的极小部分土壤区域.研究表明:植物生长过程中,由于根系的代谢作用而向根外排泄H 或HCO3-,使根际土壤pH发生较大变化.而根际土壤的pH直接影响到土壤养分的活性和生物有效性.故不同植物根际土壤的pH状况可反映出它们对土壤氮、磷、钾养分吸收利用能力的相对大小.     

植物根系分泌物与根际营养关系评述

根系分泌物（ｒｏｏｔ ｅｘｕｄａｔｅｓ,ＲＥ）主要有粘胶、外酶、有机酸、糖、酚及各种氨基酸。不同营养基因型的植物,ＲＥ级分明显不同,存在养分和环境胁迫时,植物通过增加粘胶、酶及某些有机酸的分泌量以适应变化的环境。ＲＥ也是植物改善根际营养环境的重要手段。ＲＥ可改善土壤物理结构,促进矿物风化、提高土壤ＣＥＣ,影响土壤ｐＨ、土壤矿物表面吸附性能及土壤生物学性质。ＲＥ还存化根际土壤养分,促进植物对养分吸收     

机理Ⅰ植物铁营养的吸收转运及信号调控机制研究进展

铁是植物正常生长发育必需的微量元素之一。在通气良好的碱性或石灰性土壤中,常常因铁有效性低而难以满足植物生长发育所需,缺铁已成为继缺氮和缺磷之后农业生产所面临的又一重要的营养障碍因子。与机理Ⅱ植物相比,机理Ⅰ植物更易缺铁,因此全面了解机理Ⅰ植物的铁吸收及利用机制是分子育种改良此类植物铁营养的重要基础。基于国内外的相关研究进展,从机理Ⅰ植物的根际铁活化、根系铁吸收、木质部和韧皮部中的铁运输以及胞外和胞内铁的再利用等几方面进行综述;在此基础上,从bHLH和MYB转录因子调控网络、蛋白的泛素化修饰以及小分子化学信号调控途径等几方面,重点阐述机理Ⅰ植物铁营养吸收、转运及稳态平衡过程的调控机制;同时,对研究中存在的部分问题及未来研究方向进行简要的讨论分析。     

小麦与花生间作改善花生铁营养的效应研究

采用砂-土联合培养根箱试验装置,模拟田间试验研究石灰性土壤小麦与花生间作改善花生Fe营养的效应结果表明,石灰性土壤高pH和高CaCO3是导致花生缺Fe黄化的主要原因。叶片已发生黄化的花生与小麦间作可明显改善花生缺Fe症状,间作16d后花生根际土壤有效铁含量、花生新叶叶绿素和活性Fe含量均显著提高。小麦与花生间作对改善花生Fe营养的效应可能与缺Fe小麦根分泌的Fe载体对土壤中Fe活化有关。     

施氮对小麦根-土界面磷迁移及根际磷素组分变化特征的影响

植物根系养分吸收性能与土壤养分、水分、pH值和温度等环境条件有着密切的关系,同时又影响着近根微区(根际)土壤中养分的迁移和有效性.土壤中磷素在土壤中的迁移和吸收直接与土壤性质有关.     

植物应答缺铁胁迫的分子生理机制及其调控

铁是植物生长发育中所必需的微量营养元素。虽然土壤中铁的丰度很高,但其生物有效性非常低,特别是在碱性石灰性土壤上,高pH和高重碳酸盐含量严重降低了土壤中铁的有效性。因此如何有效地提高植物对铁的利用效率及增强植物对缺铁胁迫的响应已成为目前该领域的研究热点。本文重点阐述了植物两种不同的铁吸收机制,以及对缺铁胁迫的应答反应;对目前所发现的植物中调控缺铁胁迫的相关基因进行了全面的综述,包括新发现的吞噬机理中所涉及的NRAMP基因;同时也介绍了感应铁缺乏的众多相关信号,包括植物激素、气体信号分子及microRNAs等;此外,还提出利用铁吸收相关基因的转导、控制铁吸收相关因子以及各种农艺措施的实施来提高植物铁的生物有效性从而有效缓解缺铁胁迫。最后对未来有关植物吞噬机制、铁缺乏感应信号及改善植物铁营养新途径等研究方向作了初步展望。     

酸性根际肥对石灰性土壤pH和铁有效性的影响研究

在无植物栽培的条件下通过肥料在土壤中的扩散试验研究酸性根际肥对石灰性土壤 pH值、有效铁含量的影响 ,利用盆栽试验验证对石灰性土壤上花生缺铁失绿黄化症的矫正效果。结果表明 ,酸性根际肥 (pH 1.0～ 2 .0 )中的酸在土壤中扩散的影响半径可达 6cm ,但对土壤pH降低作用最显著的是在距肥料 2cm内 ;在施肥 2 8d内 ,距肥料 2cm处 ,土壤 pH值降低了 0 .9个单位 ,土壤铁有效性 (DTPA浸提量 )增加了 5 .9mg kg ;施用酸性根际肥可使花生叶绿素SPAD值与叶片活性铁含量显著提高 ,克服了花生缺铁黄化症状 ,使施肥区 (肥料周围 2cm内 )土壤pH值显著降低 ,并显著提高了该区土壤铁的有效性和花生对土壤Fe的吸收量。     

根际养分活化与吸收过程的定量模拟研究进展

在论述植物根系作为养分的主动吸收槽，对根示养分生物有铲性产生重要作用的前提下，本文扼要讨论了根际ＰＨ值变化，根分泌螯合物与 性物及根际微生物在根际土壤养分活化中的作用过程，并重点评述了有关养分活化与吸收模拟模型的进展与发展趋势。     

Release of zinc mobilizing root exudates in different plant species as affected by zinc nutritional status

The effect of zinc nutritional status of the plant on the release of zinc mobilizing root exudates was studied in various dicotyledonous (apple, bean, cotton, sunflower, tomato) and graminaceous (barley, wheat) plant species grown in nutrient solutions. In all species, zinc deficiency increased root exudation of amino acids, sugars and phenolics. However, the root exudates of zinc deficient dicotyledonous species did not enhance zinc mobilization from a synthetic resin (Zn chelite), or a calcareous soil, although mobilization of iron from Fe^III hydroxide was increased. By contrast in the graminaceous species, root exudates from zinc deficient plants greatly increased mobilization of both zinc and iron from the various sources. These differences in capability of mobilization of zinc and iron between the plant species are the result of an enhanced release of phytosiderophores with zinc deficiency in the graminaceous species.     

Effect of zinc deficiency in wheat on the release of zinc and iron mobilizing root exudates

The effect of Zn deficiency in wheat (Triticum aestivum L. cv. Ares) on the release of Zn mobilizing root exudates was studied in nutrient solution. Compared to Zn sufficient plants, Zn deficient plants had higher root and lower shoot dry weights. After visual Zn deficiency symptoms in leaves appeared (15–17 day old plants) there was a severalfold increase in the release of root exudates efficient at mobilizing Zn from either a selective cation exchanger (Zn-chelite) or a calcareous soil. The release of these root exudates by Zn deficient plants followed a distinct diurnal rhythm with a maximum between 2 and 8 h after the onset of light. Re-supply of Zn to deficient plants depressed the release of Zn mobilizing root exudates within 12 h to about 50%-, and after 72 h to the level of the control plants (Zn sufficient plants). The root exudates of Zn deficient wheat plants were equally effective at mobilizing Fe from freshly precipitated Fe^III hydroxide as Zn from Zn-chelite. Furthermore, root exudates from Fe deficient wheat plants mobilized Zn from Zn-chelite, as well as Fe from Fe^III hydroxide. Purification of the root exudates and identification by HPLC indicated that under Zn as well as under Fe deficiency, wheat roots of the cv. Ares released the phytosiderophore 2′-deoxymugineic acid. Additional experiments with barley (Hordeum vulgare L. cv. Europa) showed that in this species another phytosiderophore (epi-3-hydroxymugineic acid) was released under both Zn and Fe deficiencies. These results demonstrate that the enhanced release of phytosiderophores by roots of grasses is not a response mechanism specific for Fe deficiency, but also occurs under Zn deficiency. The ecological relevance of enhanced release of phytosiderophore also under Zn deficiency is discussed.     

Iron deficiency‐induced zinc uptake by bread wheat

The present study aimed to test the contribution of the iron (Fe) deficiency‐induced uptake system to zinc (Zn) and copper (Cu) uptake by using bread wheat (Triticum aestivum cv. Bezostaja). For this purpose, two different uptake experiments, long‐term and short‐term, were set up in a nutrient solution culture under controlled growth chamber conditions. For the long‐term experiment, wheat cv. plants were grown with different concentrations of Fe or Zn. Results show that there was an uptake system induced under Fe‐limiting conditions which also contributed to Zn and Cu uptake. However, the Zn deficiency‐induced uptake mechanism affected neither Fe nor Cu uptake by wheat. Short‐term uptake experiments indicate that Fe deficiency‐induced Zn²⁺ uptake was more enhanced than the absorption of Zn‐phytosiderophore (PS) complexes. In addition, the Fe‐deficient plants absorbed more Zn in comparison to those plants supplied with sufficient Fe. Similar tendencies in Zn uptake under Fe deficiency in both short‐ and long‐term experiments suggest that there may be a specific Fe uptake system induced under Fe‐limiting conditions for non‐chelated metals in bread wheat. Moreover, this system also contributes to the transport of inorganic forms of some other metals, such as Zn and Cu. Although evidence is still needed involving the use of molecular biological techniques, it is hypothesized that IRT‐like proteins are responsible for this uptake system. Moreover, the release of Fe deficiency‐induced phytosiderophores and uptake of Fe(III)‐phytosiderophore complexes may not be the only mechanisms involved in the adaptation of wheat to Fe‐limiting conditions.     

Alien cytoplasm effects on phytosiderophore release in two spring wheats (Triticum aestivum L.)

Iron (Fe) deficiency is a difficult nutrient problem particularly in crop plants grown on calcareous soils. Recently, phytosiderophore (PS) release has been linked to the ability of graminaceous species and genotypes to withstand Fe-deficiency chlorosis. So enhancing PS release is a critical step to improve iron efficiency of plants grown on iron stressed soils. The effects of alien cytoplasm on PS release in spring wheat were studied by analyzing PS release from twenty wheat genotypes, including two spring wheat 881 and 352-35, and their 18 alloplasmic lines with the participation of cytoplasms from the Aegilops and Triticum species. Different genotypes were grown in iron sufficient and deficient nutrient solution under controlled environmental conditions. PS release rates were determined at two or three days intervals after onset of iron deficiency symptoms by the measurement of iron mobilizing capacity of root exudates from freshly precipitated Fe^III hydroxide. High amounts of phytosiderophores were released from roots of all wheat genotypes without iron supplied, and the amount progressively increased with the development of iron deficiency chlorosis. The results revealed that (1) the release rate of phytosiderophores from roots of common wheat could be considerably influenced by alien cytoplasms. Some alien cytoplasms exerted positive effects, some ones did negative effects, and the other ones had no significant effects. (2) the same alien cytoplasm could affect similarly or oppositely the phytosiderophores release from different wheat. (3) some alien cytoplasms, such as Chinese Spring, Ae. speltoides Tausch and Ae. cylindrica Host showed promising and potential in improving the rate of phytosiderophore release in common wheat. These cytoplasms which showed the desired effect should be given priority in interspecific and intergeneric hybridization to develop and reconstruct the needed wheat cultivars.     

EFFECT OF REDUCED SEED AND APPLIED ZINC ON ZINC EFFICIENCY OF WHEAT GENOTYPES UNDER ZINC DEFICIENCY IN NUTRIENT SOLUTION CULTURE

Improving zinc efficiency of cereals especially wheat under zinc deficiency is a priority area of research to determine and manipulate the plant factors that govern zinc uptake and utilization. Experiments conducted to ascertain variability and also the contribution of seed zinc (complete seed and half seed) to zinc efficiency in diverse wheat genotypes raised on zinc sufficient and deficient nutrient solution, showed wide genetic variability and general deficiency response such as, increase in root to shoot ratio and decrease in leaf chlorophyll, superoxide dismutase activity, total plant zinc concentration, and uptake. Since difference in zinc deficiency response were more distinct at the initial growth stage, seed zinc could be crucial for determining plant establishment, vigor, and yield. A few exceptional genotypes, where cutting seed zinc supply infect triggered growth, root characteristics, zinc uptake and use efficiency were identified and their zinc uptake capacity on low zinc soil was measured. The difference in zinc efficiency was attributable to phytosiderophore release capacity of the genotypes.     

Phytosiderophore release as a criterion for genotypic evaluation of iron efficiency in oat

Iron (Fe) deficiency in small grains grown on calcareous soils results in reduced yields, is difficult and expensive to treat with fertilizer, and is complicated to overcome by genetic field screening due to heterogeneous soil and environmental conditions. Recently, phytosiderophore release has been linked to ability of species and genotypes to resist Fe‐deficiency chlorosis. We propose a laboratory technique to measure phytosiderophore release by Fe‐deficient oat (Avena sativa L.) genotypes as a selection method for Fe‐deficiency chlorosis resistance in oat. Plants were grown in Fe‐limiting nutrient solution and phytosiderophore release was measured on 11 days. Summations of daily phytosiderophore release by 17 oat genotypes correlate well with Fe‐deficiency chlorosis scores in the field (r = ‐0.70, p = 0.01). The proposed method consistently identified the genotypes most susceptible to Fe deficiency but did not clearly separate the moderately susceptible genotypes. In these latter genotypes, other factors such as active uptake sites, root growth rate, utilization of acquired Fe, or soil interactions may be modifying factors to phytosiderophore in Fe efficiency. Quantification of phytosiderophore provides a useful selection criterion for oat by eliminating the most inefficient types and with refinement, may become a powerful tool for identifying Fe efficiency in grass crops.     

Physiological basis of iron chlorosis tolerance in rice (Oryza sativa) in relation to the root exudation capacity

Micronutrient deficiency in cultivable soil, particularly that of iron (Fe) and zinc (Zn), is a major productivity constraint in the world. Low Fe availability due to the low solubility of the oxidized ferric forms is a challenge. An experiment was, thus, executed to assess the performance of eight genetically diverse rice genotypes on Fe-sufficient (100 µM) and Fe-deficient (1 µM) nutrient solution, and their ability to recover from Fe deficiency was measured. Fe efficiency under Fe deficiency in terms of biomass production showed a significant positive correlation with the root release of phytosiderophore (PS) (R² = 0.62*). This study shows that the Fe deficiency tolerance of Pusa 33 was related to both a high release of PS by the root and an efficient translocation of Fe from the root to the shoot as the Fe–PS complex, which could be useful for improving the Fe nutrition of rice particularly under aerobic conditions.     

IRON EFFICIENCY IN KENTUCKY BLUEGRASS NOT RELATED TO PHYTOSIDEROPHORE RELEASE

Some Kentucky bluegrass (KBG; Poa pratensis L.) is susceptible to iron (Fe)-deficiency chlorosis. Under Fe-deficiency stress, phytosiderophore is produced and released by the roots of many grasses to solubilize soil Fe and enhance uptake. In other species, quantifying phytosiderophore screens for Fe-deficiency resistant cultivars. A hydroponic study was conducted at 1 and 10 μM solution Fe to variously stress ‘Baron’, ‘Award’, ‘Limousine’, and ‘Rugby II’ KBG cultivars. One μM Fe solution produced more Fe-deficiency stress in all cultivars compared to 10 μM, resulting in greater chlorosis and phytosiderophore release but reduced shoot and root Fe concentrations and shoot weight. Of the four cultivars, Baron was the most susceptible to Fe deficiency and exhibited severe Fe chlorosis and low shoot Fe but, surprisingly, produced the most phytosiderophore. These results imply that Fe-deficiency susceptibility in KBG may be less related to phytosiderophore release and more related to inefficient uptake or utilization mechanisms.     

Release of phytosiderophores from roots of wheat and triticale under nickel‐deficient conditions

Despite numerous studies on phytosiderophores (PS) there is still an open question whether nickel (Ni) deficiency induces release of PS from graminaceous plant roots. Seedlings of two wheat cultivars (Triticum aestivum L. cvs. Rushan and Kavir) and a triticale cultivar (X. triticosecale) were grown in Ni‐free nutrient solution (Ni‐deficient, Ni–) and with 10 µM NiSO₄ (Ni‐sufficient, Ni+, control). Root exudates were collected weekly for 4 weeks and the amount of PS in the root exudates was measured. The response to Ni deficiency on the release of PS differed between species. Roots of Rushan and triticale exuded higher PS in response to Ni‐deficient conditions. Nickel deficiency significantly enhanced shoot Fe and Zn concentrations in wheat, while it decreased shoot Fe and Zn concentrations in triticale. In Kavir, PS exudation was decreased by Ni deficiency at weeks 3 and 4 and the reduced release of PS from roots of Kavir was accompanied by lower concentrations of Fe and Zn in plant roots but higher Fe and Zn concentrations in shoot tissue. The PS release by Kavir was triggered by a Ni‐induced Zn deficiency particularly in the shoots. According to the results, it is suggested that in the studies concerning the phytosiderophore release under Ni deficiency, special attention should be given to different responses among and within cereals and to the plant Zn or Fe nutritional status.     

Iron deficiency stress responses of five gram‐inaceous monocots

Plant growth, leaf chlorosis, root reductive capacity, rhizosphere pH, and phytosiderophore release capacity were used as indices to evaluate the responses of maize (Zea mays L. cv ‘clipper'), millet (Pennisetum glaucum L. cv. ‘Dwarf Gero'), sorghum (Sorghum bicolor L. cv. YG 5760), barnyard grass (Echinochloa crus galli L. cv: unknown), wheat (Triticum aestivum L. cv. ‘tonic'), and white lupin (Lupinus albus L. cv ‘lucky') to iron‐deficiency stress. Generally, root and shoot dry matter increased with iron treatment and leaves became less chlorotic. Neither the order nor the magnitude of the root reductive capacities of the monocots studied was affected by iron deprivation, but these reductive capacities and the changes in rhizosphere pH differed markedly. Significant iron stress‐induced phytosiderophore release was observed only in wheat and sorghum in which accompanying increases in rhizosphere pH were also evident. Such phytosiderophore release matched the severity of leaf chlorosis and iron uptake and depended on the form in which the element was supplied. These results, from experiments conducted in non‐axenic hydroponic cultures, indicate that in iron‐ deficiency stress mechanisms ‐ similar to those found in dicots ‐could account for iron uptake in some graminaceous monocots, and that strategy II‐type response proposed for all in this category of plants would be an over simplification.