Phytosiderophore release does not relate well with zinc efficiency in different bread wheat genotypes

Using six bread wheat genotypes (Triticum aesttvum L. cvs. Dagdas‐94, Gerek‐79, BDME‐10, SBVD 1–21, SBVD 2–22 and Partizanka Niska) and one durum wheat genotype (Triticum durum L. cv. Kunduru‐1149) experiments were carried out to study the relationship between the rate of phytosiderophore release and susceptibility of genotypes to zinc (Zn) deficiency during 15 days of growth in nutrient solution with (1 μM Zn) and without Zn supply. Among the genotypes, Dagdas‐94 and Gerek‐79 are Zn efficient, while the others are highly susceptible to Zn deficiency, when grown on severely Zn deficient calcareous soils in Turkey. Similar to the field observations, visual Zn deficiency symptoms, such as whitish‐brown lesions on leaf blades occurred first and severely in durum wheat Kunduru‐1149 and bread wheats Partizanka Niska, BDME‐10, SBVD 1–21 and SBVD 2–22. Visual Zn deficiency symptoms were less severe in the bread wheats Gerek‐79 and particularly Dagdas‐94. These genotypic differences in susceptibility to Zn deficiency were not related to the concentrations of Zn in shoots or roots. All bread wheat genotypes contained similar Zn concentration in the dry matter. In all genotypes supplied adequately with Zn, the rate of phytosiderophore release was very low and did not exceed 0.5 μmol/48 plants/ 3 h. However, under Zn deficiency the release of phytosiderophores increased in all bread wheat genotypes, but not in the durum wheat genotype. The corresponding rates of phytosiderophore release in Zn deficient durum wheat genotype were 1.2 umol and in Zn deficient bread wheat genotypes ranged between 8.6 μmol for Partizanka Niska to 17.4 umol for SBVD 2–22. In Dagdas‐94, the most Zn efficient genotype, the highest rate of phytosiderophore release was 14.8 umol. The results indicate that the release rate of phytosiderophores does not relate well with the susceptibility of bread wheat genotypes to Zn deficiency. Root uptake and root‐to‐shoot transport of Zn and particularly internal utilization of Zn may be more important mechanisms involved in expression of Zn efficiency in bread wheat genotypes than release of phytosiderophores.     

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.     

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 response of various c‐3 and c‐4 grain crop genotypes: Strategy II mechanism evaluated

The relative amount of phytosiderophore produced by various Strategy II plants has been categorized as follows: barley (Hordeum vulgare L.) > wheat (Triticum aestivum L.) > oat (Avena byzantina C. Koch.) > rye (Secale cereale L.) >> corn (Zea mays L.) >> sorghum (Sorghum bicolor (L.) Moench) > rice (Oryza sativa L.). With the exception of rice, these plants developed under oxidized soil conditions, and the C‐3 species produce more phytosiderophore than C‐4 species under Fe‐deficiency stress. Iron‐efficient Coker 227 oat produced phytosiderophore in response to Fe‐deficiency stress, while Fe‐inefficient TAM 0–312 oat did not. Although Fe‐efficient WF9 corn and Fe‐inefficient ys₁ corn differed in their ability to obtain Fe, neither produced sufficient quantities of phytosiderophore to explain these differences. The objectives of this research were to determine: (a) if phytosiderophore production of Fe‐deficiency stressed C‐4 species millet (Panicum miliaceum L.) and corn is low or absent compared to identically stressed C‐3 species oat and barley, and (b) if native, inbred and hybrid corn cultivars differ in ability to produce and utilize phytosiderophores.

Although release of phytosiderophore for Fe‐stressed corn and millet was generally lower than oat, quantity of release was not always related to obtaining Fe and maintaining green plants. Barley maintained high leaf Fe and low chlorosis with a minor release of phytosiderophore. Oat with increased release acted similarly to barley, whereas a relatively high release of phytosiderophore from White maize did not effect Fe uptake or greening. Likewise, small amounts of phytosiderophore were produced by all corn types, but corn was generally unable to obtain adequate Fe from the growth medium. Two of the native corns, Coneso and Tepecintle, maintained relatively low chlorosis, but they differed in phytosiderophore release. Thus, it appears that the C‐4 plants studied herein generally release a lower amount of phytosiderophore than do C‐3 species, but overcoming Fe‐deficiency chlorosis is not guaranteed by such release. The Strategy II mechanism of mere release of phytosiderophore and consequential Fe acquisition appears simplistic. There is a need for understanding what other factors are involved.     

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.     

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.     

Phytosiderophore release from manganese‐induced iron deficiency in barley

An experiment was conducted in the phytotron with barley (Hordeum vulgare L. cv. Minorimugi) grown in nutrient solution to compare iron (Fe) deficiency caused by the lack of Fe with manganese (Mn)‐induced Fe deficiency. Dark brown spots on older leaves and stems, and interveinal chlorosis on younger leaves were common symptoms of plants grown in either Mn‐toxic or Fe‐deficient treatments. Dry matter yield was affected similarly by Fe deficiency and Mn toxicity. The Mn toxicity significantly decreased the translocation of Fe from roots to shoots, caused root browning, and inhibited Fe absorption. The rate of Fe translocated from roots to shoots in the 25.0 μM Mn (toxic) treatment was similar to the Fe‐deficient treatment. Manganese toxicity, based on the release of phytosiderophore (PS) from roots, decreased from 25.0>250>2.50 uM Mn. The highest release of PS from roots occurred 7 and 14 days after transplanting (DAT) to Mn‐toxic and Fe‐deficient treatments, respectively; but was always higher in the Fe‐deficient treatment than the Mn‐toxic treatments. The release of PS from roots decreased gradually with plant age and with severity of the Mn toxicity symptoms. The PS content in roots followed the PS release pattern.     

EFFECT OF CADMIUM ON PHYSIOLOGICAL RESPONSES OF WHEAT AND CORN TO IRON DEFICIENCY

This work evaluated the effect of cadmium (Cd) on the physiological responses of corn (Zea Mays L.) and wheat (Triticum aestivum L.) to iron (Fe) deficiency. For this purpose, seedlings of corn and wheat were cultivated under controlled conditions, plants were grown in different strength Hoagland's solutions for one month. In the fifth week, some seedlings were still in full strength Hoagland's solution (+Fe) and others were in full strength Hoagland's solutions without iron (?Fe). The plants were exposed to different cadmium (Cd) concentrations for four days. The plant chlorophyll content of young leaves, Fe and Cd content in shoots and roots, biomass production, and phytosiderophores (PS) release by roots were assessed. Results showed that Cd decreased the chlorophyll content of young leaves, accompanied by a significant shoot and root biomass reduction for Fe-deficient and Fe-sufficient wheat and corn across all Cd treatments. However, chlorophyll content and shoot and root biomass of Fe-deficient wheat and corn were lower than Fe-sufficient plants at different Cd concentrations. Iron-deficiency induced Cd accumulation compared to Fe-sufficient in wheat and corn; however, a depressive effect of Cd on iron acquisition in shoots and roots of Fe-deficient and Fe-sufficient wheat and corn across all Cd treatments was observed. Cadmium also inhibited PS release in Fe-deficient and Fe-sufficient wheat and corn. Iron-deficient PS release was higher than Fe-sufficient corn and wheat across all Cd treatments. These results suggested that Cd might reduce capacity of plants to acquire iron from solution by inhibiting PS release.     

MAIZE HYBRIDS DIFFER IN THEIR 24-H PATTERNS OF PHYTOSIDEROPHORE RELEASE

Iron (Fe) chlorosis tolerant gramineous species respond to Fe-deficiency stress by releasing phytosiderophores. The objective of this study was to characterize the diurnal pattern of phytosiderophore release from Fe-chlorosis tolerant and susceptible maize (Zea mays L.) hybrids. Phytosiderophore was collected from individual, Fe-deficient maize plants in 10 mg L^?1 Micropur solution during 4-h collection periods. The diurnal pattern of phytosiderophore release differed among maize hybrids. Similar levels of phytosiderophore were released during light and dark conditions for three hybrids. Greater phytosiderophore release was measured from a fourth hybrid when the collection procedure began 2-h after the initiation of light compared to 2-h before the initiation of dark. When a single plant was used for six consecutive collection periods, phytosiderophore release declined during the final 8- to 12-h of collection. The decline in phytosiderophore release was attributed to dissolution of apoplastic Fe and deactivation of the Fe-stress response mechanism.     

Phytosiderophore release by Sorghum,wheat, and corn under zinc deficiency 1

Zinc (Zn) deficiency is more common in corn (Zea mays L.) than in sorghum [Sorghum bicolor (L.) Moench] or wheat (Triticum sp.). The ability of wheat to withstand low soil Zn conditions is related to increased release of phytosiderophore from its roots. The reasons for sorghum's ability and corn's inability to utilize low levels of soil Zn have not been explored adequately. The objectives of this research were to 1) ascertain if Zn deficiency could be induced in sorghum, wheat, and corn grown in a chelator‐buffered nutrient solution and 2) determine relative releases of phytosiderophore from roots of sorghum, wheat, and/or corn under Zn‐deficiency conditions. Sorghum, wheat, and corn were grown hydroponically in the greenhouse with a chelator‐buffered nutrient solution designed to induce Zn deficiency, while supplying adequate amounts of other nutrients. Root exudates were collected over time to measure phytosiderophore release. Shoot Zn concentrations and shoot and root dry matter yields were determined also. The technique was effective for inducing Zn deficiency in sorghum, wheat, and corn, as evidenced by reduced shoot and root dry matter yields, shortened internodes, reduced shoot Zn concentrations, and plant Zn concentrations below the suggested critical values for these species. Sorghum and wheat plants increased the release of phytosiderophore in response to Zn deficiency, but com did not. The total amount of phytosiderophore released by the roots was in the order wheat>sorghum>corn. The absence of a “phytosiderophore”; response to Zn deficiency of corn, coupled with the evidence that this species requires, or at least accumulates, more Zn than wheat or sorghum, provides an explanation as to why Zn deficiencies are more prevalent for corn than wheat or sorghum under field 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.     

Zinc‐efficient wild grasses enhance release of phytosiderophores under zinc deficiency

The effect of the zinc (Zn) nutritional status on the rate of phyto‐siderophore release was studied in three wild grass species (Hordeum murinum, Agropyron orientale, and Secale cereale) grown in nutrient solution under co‐trolled environmental conditions. These wild grasses are highly “Zn‐efficient”; and grow well on severely Zn‐deficient calcareous soils in Turkey (DTPA‐extractable Zn was 0.12 mg/kg soil and CaCO₃ was 37%). In all wild grasses studied, Zn deficiency reduced shoot growth but had no effect on root growth. Low amounts of phytosiderophores were released from roots of all wild grasses adequately supplied with Zn. In plants grown without Zn, release of phytosiderophores progressively increased with the onset of visual Zn deficiency symptoms, such as inhibition of shoot elongation and appearance of chlorotic and necrotic patches on leaves. Compared to Zn‐sufficient plants, phytosiderophore release increased 18–20‐fold in deficient plants. HPLC analysis of root exudates showed that the dominating phytosiderophore in Zn‐deficient Agropyron and Hordeum was 3‐epi‐hydro‐xymugineic acid (epi‐HMA) and was 3‐hydroxy‐mugineic acid (HMA) in Secale. Besides HMA, epi‐HMA and mugineic acid (MA) were also detected in exudates of Zn‐deficient Secale. The results indicate the importance of phytosiderophores in adaptation of wild grasses to Zn‐deficient calcareous soils. Phytosiderophores might enhance mobilization of Zn from sparingly soluble Zn pools and from adsorption sites, both in the rhizosphere and within the plants.     

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.     

Diurnal rhythm of release of phytosiderophores and uptake rate of zinc in iron-deficient wheat

Abstract

The diurnal rhythm of release of phytosiderophores and uptake rate of zinc (Zn) was studied in iron (Fe) deficient wheat (Triticum aestivum L. cv. Ares) plants grown in nutrient solution under controlled environmental conditions. Different forms of Zn (e.g. ZnSO₄, ZnEDTA) were used to obtain different degrees of loading of the root apoplasmic pool with Zn.

In the Fe-deficient plants the release of phytosiderophores from the roots followed a distinct diurnal rhythm with a steep peak about 4 h after the onset of the light period. These plants also showed a similar pattern in the rates of Zn uptake over the 24 h day-night cycle. During the light period there was a steep transient peak (factor 3.8) in Zn uptake rate in the Fe-deficient plants supplied with ZnSO₄. This transient peak was much less distinct in plants supplied with ZnEDTA (factor 1.8) and absent in plants supplied with ZnEDTA plus free chelator (+ NaEDTA) in excess. The peak in Zn uptake coincided with the maximum rate of phytosiderophore release in the Fe-deficient plants. In the Fe-sufficient plants the release of phytosiderophores was very low and no such peak in Zn uptake rates could be observed.

These results demonstrate that phytosiderophores mobilize Zn not only in the rhizosphere, but also from the root apoplast. Thus, the apoplasmic pool of micronutrient cations has to be taken into account as potential source for both uptake and diurnal variation in uptake rates of Micronutrient cations.     

Phytosiderophore release in relation to multiple micronutrient metal deficiency in wheat

Phytosiderophore (PS) release, which occurs mainly under iron deficiencies in the representative Poaceae, has been speculated to be a general adaptive response to enhance the acquisition of micronutrient metals. However, it is very common to encounter deficiency of micronutrients other than iron (Fe) in soils and interactions with respect of multi-micronutrient deficiency to effect on PS release are not known. Further, the diurnal rhythm for the release of PS may also be affected under multiple micronutrient deficiency. PS release capacity and PS content of roots and the diurnal rhythm of PS release was measured in selected efficient and inefficient wheat genotypes varied on individual and combined deficiency of Fe, zinc (Zn), copper (Cu) and manganese (Mn) in nutrient solution culture. A nutrient sufficient treatment was also taken as experimental control. Lack of Fe in the nutrient medium caused a significantly higher release of PSs followed by Zn, Mn and Cu in the same order. The diurnal rhythm of PS release was similar in the absence of either of the micronutrients or under their combined deficiency. Micronutrient sufficient control did not release any PS. Fe-use-efficient cultivars produced and released a larger amount of PS and differed from the inefficient cultivars in terms of the PS release but not in the PS biosynthesis in the roots. Thus, indicating that the limitation at the level of release of the PS is responsible for low Fe use efficiency of the Fe deficiency susceptible cultivars. Further, the diurnal variation in the PS release was similar for all the investigated wheat cultivars and did not influence the variation in the Fe use efficiency.     

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.     

根分泌物与禾本科植物对缺铁胁迫的适应机理

本文系统地总结了自然基金重点项目根分泌物在根际微生态系统中的营养机理的研究进展和部分主要成果。研究表明,养分专一性根分泌物是植物营养遗传特性控制基因的标记物,它受某一养分缺乏的诱导,是在植物体内合成并可通过主动分泌作用进入根际的代谢产物。它的合成和分泌只受该养分胁迫的专一诱导和控制,只要改善这一营养状况就能抑制或阻止其合成和分泌。当植物缺乏这一养分时,植物体可通过自身的调节能力,合成专一性物质并自根分泌到根际,促进该养分的活化并提高植物对其吸收利用效率,从而达到克服或缓解该营养胁迫的目的。用单基因突变材料进行的研究表明,植物铁载体的生物合成和吸收利用是受单基因控制的过程。这一发现不仅使人类有可能运用生物学研究技术来解决营养缺乏问题,而且也为有效地利用自然资源、降低生产成本、减少环境污染提供了可能性。麦根酸类(mugineic.acids)植物铁载体(phytosiderophores)在缺铁禾本科植物体内的生物合成,从植物根内向根际的分泌、在根际环境中对铁的活化及植物对F63+植物铁载体螯合体的吸收四个过程组成了禾本科植物对缺铁胁迫的适应机理。植物铁载体只在早晨日出后2～6h内大量分泌的节律性增加了它们在根际土壤中的相对浓度,减少了它们与土壤颗粒的接触和被吸附;分泌部位集中在微生物尚未侵染的根尖避开了微生物的破坏和分解,同时也增加了它们在土壤微区中的相对浓度。分泌作用和螯合作用不受介质pH值和Ca2+离子浓度影响的特性使该机理在经常出现缺铁现象的石灰性土壤上具有特殊意义。研究结果还表明,缺铁可以诱导激活根细胞原生质膜上可能存在的Fe3+植物铁载体复合体的专一性吸收和运载蛋白,高pH值和CaCO3对这一蛋白的载体功能只有很小的抑制作用,即使在pH值和CaCO3含量都较高的石灰性土壤上,也能有效地发挥作用。这一结论揭示了小麦等禾本科植物适应铁胁迫的实质及其生态学意义。     

Detection of the regions of phytosiderophore release from barley roots

Abstract

We devised a method to detect regions of phytosiderophore release from barley (Hordeum vulgare cv. Minorimugi) roots. Plants were grown in Fe-sufficient (+Fe) or Fe-deficient (?-Fe) water cultures for 14 d in a phytotron. Intact or excised roots were sampled and put between two sheets of filter papers just after the onset of the light period. The filter papers with roots were wrapped with vinyl film to avoid drying out and covered with aluminum foil to shade. The roots between the filter paper were kept in the phytotron for 4 h and phytosiderophores (PS) released from roots were absorbed by the filter paper. Then, the shape of the roots was preserved by photocopy and PS released on the filter paper were visualized by the method employed for detecting PS on thin layer chromatography (TLC). Gelatinous Fe solubilizing activity was employed for detecting PS on the filter paper. Released PS were detected as white spots on the orange-colored background and the regions of PS release from the roots were visible on the filter paper. It was evident that the apical zones of roots were the main regions of PS release. It was apparent that a distinct primary root newly formed on the basal parts had higher activity to release PS than the other roots of ?Fe plants. It was also shown that apical root zones of +Fe plants released PS.     

Responses of aerobically grown iron chlorosis tolerant and susceptible rice (Oryza sativa L.) genotypes to soil iron management in an Inceptisol

Iron deficiency is a serious nutritional disorder in aerobic rice, causing chlorosis, poor yields and reduced grain nutritional quality. The problem can be managed by complementing the use of Fe-efficient plant type with a suitable Fe management strategy. In the present paper, we report the effect of eight iron management practices to resolve the problem of iron (Fe) chlorosis through the use of an iron deficiency tolerant (IDTR) and iron deficiency susceptible (IDSR) rice genotype, i.e. Pusa 33 and ADT 39, respectively. Fe deficiency tolerance of these genotypes was related to the root release of PS which enabled a higher uptake of Fe in the IDTR than the IDSR under Fe deficiency. In general, IDTR performed better than the IDSR as evident from a significant increase in total iron, active iron, chlorophyll content and grain and straw yield. IDSR produced the highest grain and straw yield under slow iron release nano clay complex source. Grain Fe content of the IDTR and IDSR increased by 18.9 and 13.4%, respectively, under recommended dose of Fe. The results identified the most effective soil management strategies for the alleviating Fe deficiency chlorosis and improving Fe nutrition of both IDTR and IDSR genotypes.     

Morpho‐physiological Approaches to Investigate Lime‐Induced Chlorosis in Deciduous Fruit Tree Species

Abstract

Several deciduous fruit tree species are affected by iron chlorosis when grown in calcareous soils and worldwide research on rootstock breeding and screening in relation to iron deficiency is carried out. For many years the goal of our department has been to investigate morphological and physiological aspects related to lime‐induced chlorosis, and to identify markers usable as tools for tolerance screening of clonal rootstocks. Experimental strategies using pot, hydroponic and in vitro culture have been used to understand if they are able to originate suitable indicators on developmental and biochemical changes resulting from iron deficiency. Results risen up from in vitro trials suggest that iron deficiency sensitive system should work in a divergent way from that of bicarbonate system. Physiological and morphological data highlight diverse syndromes and behaviors among the genotypes, probably reflecting properly perception, regulation, and activation of biological answer system to iron deficiency and bicarbonate. Recently, quince (Cydonia oblonga Mill.) minus‐ and plus‐variants have been obtained by in vitro somaclonal variation to clarify the agonistic relationship between bicarbonate and iron deficiency sensitive systems, that in turn can induce the activation of two specific gene expression pathways.