Genetics of tolerance to iron chlorosis in rice

Genetics of tolerance to iron chlorosis was investigated in eight crosses involving parents distinctly different in their level of tolerance. The segregating populations with parents and F₁s were screened under actual stress conditions in the field. Also, selected crosses were studied for Fe³⁺ uptake capacity. Tolerance/moderate tolerance to Fe chlorosis was dominant over susceptibility and it was controlled by two sets of nonallelic genes with complementary interaction. Gene Ic ₁ has been found to be basic and in complementation with Ic ₃ it confers tolerance. Likewise, Ic ₂ with Ic ₄ confers tolerance. The basic genes Ic ₁ and Ic ₂ are nonallelic and, in the absence of their respective complementary genes Ic ₃ and ₄ , ineffective, which results in susceptibility. Of tolerant cultivars, ARC 10372 and Cauvery have been tentatively assigned the genotype of Ic ₁ , Ic ₂ , Ic ₃ , Ic ₄ , and moderately tolerant IET 7613, Prasanna and Akashi Ic ₁ , ₂ Ic ₃ Ic ₄ . The susceptible ARC 5723 has been assigned Ic ₁ , ₂ , Ic ₃ , Ic ₄ , and IET 9829, Ic ₁ , ₂ Ic ₃ Ic ₄ . IET 7614 is susceptible, due to the presence of inhibitory genes I-Ic ₁ , I-Ic ₂ together with ic ₁ pt>, ic ₂ , Ic ₃ , Ic ₄ . Further, the gene Pc for purple coleoptile shows linkage with one of the complementary genes with a crossover value of 15.26%, while the gene(s) for seedling height Ts with Ic ₁ with a crossover value of 1.7%. It is possible that the gene(s) for iron chlorosis tolerance might belong to the second linkage group, where genes for purple leaf were located.     

Selection of olive varieties for tolerance to iron chlorosis

Under certain conditions, olive trees grown on calcareous soils suffer from iron chlorosis. In the present study several olive varieties and scion-rootstock combinations were evaluated for their tolerance to iron chlorosis. Plants were grown over several months in pots with a calcareous soil, under two fertilization treatments. These consisted of periodic applications of nutrient solutions containing either, 30 μmol/L FeEDDHA or not Fe. Tolerance was assessed by the chlorosis and growth parameters of plants grown without Fe, compared to those plants grown with Fe. Results show that there are differences in tolerance among olive varieties and that tolerance is mainly determined by the genotype of the rootstock. These results open the way to use tolerant varieties for those conditions where iron chlorosis could become a problem.     

Fe-EDDHA对矫治秦美猕猴桃叶片失绿和营养元素组成的影响

在陕西石灰性土壤秦美猕猴桃果园施用不同剂量叶绿灵 (Fe- EDDHA) ,结果表明 ,萌芽期株施 45 g叶绿灵对矫治秦美猕猴桃叶片失绿黄化效果显著 ,可一直维持到当年落叶。同黄化对照植株相比 ,叶片叶绿素含量增加 1 82 .5 % ,活性铁含量提高 5 3 .2 % ,果实品质得到明显改善。施用叶绿灵还改善了叶片营养元素的含量     

管道输液滴干对苹果缺铁失绿症的影响

为探索防止果树缺铁失绿症方案,该研究以Fe-N为铁肥品种,以管道输液滴干的方式对中秋王苹果树输入不同浓度的溶液,测定叶片的叶绿素相对含量(SPAD值)、百叶重、百叶厚、全铁和活性铁含量及荧光参数等指标。结果表明:Fe-N管道输液滴干处理显著提高了苹果叶片的SPAD值、百叶重、百叶厚、全铁以及活性铁的含量,发现叶片SPAD值与活性铁含量之间呈现良好的线性关系(相关系数为0.899),表明叶片SPAD值可以作为苹果缺铁诊断指标,同时也说明了苹果树体叶片黄化主要是由于活性铁含量低所致。16.4×10~(-3)mol·L~(-1)Fe-N溶液处理的叶片SPAD值、百叶重、百叶厚、全铁及活性铁含量显著高于其它处理,分别比对照提高了89.66%、20.42%、9.26%、158.0%、277.4%,该处理荧光参数也达到了较优水平。该研究结果为管道输液滴干防止苹果缺铁失绿症的应用奠定了基础。     

Foliar fertilization to control iron chlorosis in pear (Pyrus communis L.) trees

The effectiveness of foliar fertilization to re-green chlorotic leaves in iron-deficient pear trees has been studied. Trials were made to assess the influence of (i) the level of Fe deficiency, (ii) the leaf surface treated (adaxial or abaxial), and (iii) two different surfactants, L-77 and Mistol. Treatments were ferrous sulphate alone, ascorbic, citric and sulphuric acids, applied either alone or in combination with ferrous sulphate, Fe-DTPA and water as a control. Solutions were applied with a brush and leaves were treated twice each year. None of the treatments caused a full recovery from Fe deficiency chlorosis. Treatments containing Fe caused the largest re-greening effects, and FeSO₄ had a similar re-greening effect to Fe(III)-DTPA. Increases in leaf Chl were more pronounced with abaxial leaf surface applications and in severely deficient leaves. Using Fe(III)-DTPA in foliar sprays does not seem to be justified, since their effects are not better than those of FeSO₄. The joint use of Fe(III)-DTPA and L-77 and that of FeSO₄ and citric acid do not seem to be suitable. With a single foliar application, FeSO₄ combined with acids gave slightly better results than FeSO₄ alone. Acidic solution applications without Fe may be effective in alleviating chlorosis in some cases, especially in the case of citric acid. In the current state of knowledge, foliar fertilization cannot offer yet a good alternative for full control of Fe chlorosis, although its low environmental impact and cost make this technique a good complementary measure to soil Fe-chelate applications and other chlorosis alleviation management techniques. Abbreviations: Chl – chlorophyll; EDDCHA – ethylenediamine di(5-carboxy-2-hydroxyphenylacetic) acid; EDDHA – ethylenediamine di(o-hydroxyphenylacetic) acid; EDDHMA – ethylenediamine di(o-hydroxy-p-methylphenylacetic) acid; EDDHSA – ethylenediamine di(2-hydroxy-5-sulfophenylacetic) acid     

Varietal response to iron chlorosis in upland rice

Summary In a study involving twenty upland rice genotypes, the induction of iron chlorosis was found to be more by superphosphate application than due to lime incorporation into an alfisol soil under greenhouse conditions in GI tray-grown seedlings as quantified by two parameters viz., total chlorphyll content and Fe²⁺ content. Of the two indices of iron chlorosis, Fe²⁺ was more sensitive than chlorophyll content. Genotypes were grouped into efficient and inefficient categories both in terms of absorption and utilization of iron based on the degree of reduction in response to added superphosphate.     

A comparison of ferric-chelate reductase and chlorophyll and growth ratios as indices of selection of quince,pear and olive genotypes under iron deficiency stress

Quince (Cydonia oblonga Mill.), pear (Pyrus communis L.) and olive (Olea europaea L.) genotypes were evaluated for their tolerance to iron deficiency stress by growing young plants in three types of aerated nutrient solutions: (1) with iron, (2) without iron or (3) low in iron and with 10 mM bicarbonate. Plants were obtained either from rooted softwood cuttings or from germination of seeds. The degree of tolerance was evaluated with several indices: (1) the chlorophyll content, (2) the root Fe³⁺ reducing capacity and (3) the whole plant relative growth. Fifteen hours before Fe³⁺ reducing capacity determination, iron was applied to the roots of plants with iron-stress, since this method resulted in increasing the reductase activity. All quince and pear genotypes increased the root Fe³⁺ reducing capacity when grown in the treatments for iron-stress, in relation to control plants of the same genotypes. In olive cultivars, the Fe³⁺ reducing capacity was lower in the iron-stress treatments than in the control one. Studying the relationship between relative growth and chlorophyll content for each genotype under iron-stress, in relation to both indices in control plants, a classification of species and genotypes was established. According to that, most olive cultivars and some pear rootstocks and cultivars appear more iron-efficient than quince rootstocks. Our study shows that in some woody species, determining root Fe³⁺ reducing capacity is not the best method to establish tolerance to iron deficiency stress.     

虹吸输铁对黄化苹果叶片光合参数及细胞器结构的影响

该研究以红富士苹果品种为材料,通过虹吸输入铁肥的方式,设置输铁液(600倍FeS04·7H20)处理并以输离子水和不输液为对照,观察了处理后苹果树的复绿情况,测定了叶绿素含量、光合参数、进行了叶绿体、线粒体超微结构的电镜扫描。结果表明:输铁后10 d,树体复绿情况明显,复绿等级由2.01变为0.53,复绿的叶片叶绿素含量有显著增加,增幅达到200%。同时,净光合速率提高了68%、蒸腾速率提高了21%、气孔导度提高了49%,胞间CO2浓度降低了100%。输铁处理后的单个细胞叶绿体个数比对照有显著增加,且个体更饱满,淀粉粒和嗜锇颗粒有明显减少;基粒片层结构规律整齐,基质清晰,被膜结构完整,线粒体内嵴清晰度高,数量增多。在输铁处理后,苹果树体复绿明显,叶绿素合成能力恢复,叶绿体、线粒体被膜结构均得到修复,表明用虹吸的方法将铁肥输入树体对苹果缺铁黄化病有很好的疗效。     

Relationship between iron chlorosis and alkalinity in Zea mays

Mengel, K. and Geurtzen, G. 1988. Relationship between iron chlorosis and alkalinity in Zea mays . - Physiol. Plant. 72: 460–465.
Maize ( Zea mays L. cv. Anjou 21) grown in nutrient solution with Fe-EDTA and with nitrate as the sole nitrogen source showed typical Fe-chlorosis symptoms after a growth period of 14–21 days. Alkalinity in roots, stems and leaves of the chlorotic plants was high. Transferring the chlorotic plants from the nitrate-containing nutrient solution to a solution of (NH₄)₂SO₄ resulted in a regreening of leaves within 2–3 days which was associated with a decrease in solution pH, a decrease in alkalinity of plant parts, a translocation of Fe from roots to tops and a release of Fe into the outer solution. Similar effects were obtained when Fe chlorotic plants were transferred to a dilute HO solution with pH 3.5.
Spraying chlorotic leaves with indoleacetic acid or with fusicoccin led also to a regreening of leaves without having a major effect on leaf alkalinity.
Interpretation of the experimental results is based on the assumption that nitrate as sole N source leads to a high pH level in the apoplast resulting in the precipitation of Fe compounds, probably Fe oxide hydrate. Ammonium nutrition has the reverse effect since it lowers the apoplast pH and this can result in the dissolution of Fe compounds. Application of indoleacetic acid as well as fusicoccin supposedly stimulates the proton pumps in the plasmalemma of the leaf tissue. The resulting decrease in apoplast leaf pH in the microenvironment also leads to a dissolution of Fe compounds in the apoplast and thus promotes the uptake of Fe by the symplasm.     

Chlorosis correction and agronomic biofortification in field peas through foliar application of iron fertilizers under Fe deficiency

Effectiveness of different iron (Fe) foliar sprays for leaf chlorosis correction and grain Fe boosting was studied in field peas under Fe deficiency. No chlorophyll reduction was observed in Fe deficient plants treated with foliar sprays. EDDHA [ethylenediamine-N,N′-bis(2-hydroxyphenylacetic acid)] followed by FeSO₄ (73.7?mg/l Fe) treated at the start of flowering was most responsive in correcting chlorosis and increasing shoot dry biomass in peas. Inductively coupled plasma-atomic emission spectroscopy data showed significant increase of Fe in grains while treated with all foliar sprays at the time of grain filling in Fe-deficient plants. Among them, FeSO₄ (73.7?mg/l Fe) was the most efficient in biofortifying Fe in mature grain under Fe deficiency in peas. Results also pinpoint that flowering is a suitable time for applying foliar sprays to boost Fe in mature grains. Taken together, application of Fe foliar sprays facilitated both chlorosis correction and Fe boosting in peas and can be further used by breeders and farmers.     

石灰性土壤上HCO3-诱导花生缺铁失绿机制

采用土壤-营养液结合的分根培养方法,研究了部分根系供应HCO- 3或铁对花生铁营养的调控及其作用机制。结果表明,对花生部分根系供应HCO- 3或铁可以调控花生的铁营养,仅供HCO- 3可以诱导缺铁,而只供铁能矫正失绿,同时供应HCO- 3和铁时则不引起失绿。在花生新生叶失绿和复绿的过程中,其中的活性铁含量和全铁含量也有相应的消长。当花生表现缺铁失绿症状时,地上各部分的全铁含量显著降低,而土中根的全铁含量不降低、质外体铁含量升高。在HCO- 3存在的条件下,不同部分根系的铁( )还原酶活性因其生长介质而不同,营养液中根系的铁( )还原酶活性降低而土中根的铁( )还原酶活性不受影响。当花生表现缺铁失绿症状时,土壤中HCO- 3含量升高,有效铁含量不高,p H值无变化。因此,本试验证实了石灰性土壤上的高HCO- 3含量,主要是降低了花生地上部的铁含量而引起失绿,而且花生缺铁失绿又导致土壤HCO- 3含量升高     

锰胁迫对甘蔗幼苗缺铁和失绿的影响

近年来,甘蔗主产区的甘蔗幼苗出现严重的缺铁失绿问题,影响了我国甘蔗生产及食糖安全。为了揭示锰诱导甘蔗幼苗缺铁失绿机制,该研究采用水培试验法,对过多锰诱导的甘蔗幼叶失绿及其与铁素营养的关系进行了探讨。结果表明:过多锰胁迫下随着甘蔗中锰含量的增加,幼叶明显失绿。250、500、750μmol·L-1处理10 d后,幼叶中的叶绿素含量分别从对照处理的1.71 mg·g-1FW下降至0.86、0.85、0.64 mg·g-1FW。过多锰抑制甘蔗对铁的吸收,每株植株对铁吸收量(3.22~4.40 mg)显著减少。幼叶中铁含量(116.8~128.6 mg·kg-1DW)也随着锰处理浓度的增加而显著降低。250~750μmol·L-1处理的甘蔗幼叶中铁的含量仅相当于对照处理的89.4%~81.2%。相反,锰处理后根和茎中铁的含量却显著增加。锰胁迫下幼叶中活性铁含量和活性铁与全铁比值(0.14~0.21)也显著降低。高锰胁迫下,幼叶中的活性铁含量(4.1~6.9 mg·kg-1FW)相当于对照处理的25.5%~55.2%。相关分析结果显示,锰胁迫下的甘蔗叶片中活性铁含量与叶绿素含量呈显著的正相关;锰处理后幼叶中活性铁与锰含量的比值从对照的0.71下降至0.04~0.01。这说明过多的锰可诱导甘蔗幼叶失绿,而失绿与过多的锰胁迫下甘蔗对铁的吸收、运输受阻及铁的钝化有关。     

Plant growth and nutrient uptake characteristics of Fe-deficiency chlorosis susceptible and resistant subclovers

The objective of this study was to evaluate the growth and nutrient-uptake characteristics of Fe-deficiency resistant and susceptible subclover (Trifolium subterraneum L., T. yanninicum Katzn. and Morley, T. brachcalycinum Katzn. and Morley) cultivars on a calcareous soil. Ten subclover cultivars showing varying susceptibilities to Fe-deficiency chlorosis (Karridale, Nangeela, Geraldton, Mt. Barker, Woogenellup, Larisa, Trikkala, Rosedale, Koala and Clare) were grown on a low-Fe, calcareous soil (Petrocalcic Paleustoll) under moist (18% water content, 85% of water holding capacity) and water-saturated conditions using a Cone-tainer^® culture system. Chlorosis and its correlation with growth traits and mineral nutrition of the 10 cultivars were examined. The Fe-deficiency susceptibilities of the 10 cultivars decreased in the above order under the moist condition, but in slightly different order under the saturated condition. Shoot and root dry weights, total dry weight, and root-to-shoot ratio were each negatively correlated with chlorosis under both soil-moisture conditions, as was total shoot content of P, Ca, Fe, Mn and Zn. Shoot P and Fe concentrations were each positively correlated with chlorosis under the moist soil condition. Iron and Cu utilization efficiencies (biomass per unit weight of nutrient) in the shoot were each negatively correlated with chlorosis under the moist soil condition. These results suggest that there may be several characteristics of Fe-deficiency chlorosis resistance in subclovers, such as a more effective soil-Fe mobilizing mechanism(s), more balanced nutrition, lower required Fe concentration in the shoot, higher shoot-Fe utilization efficiency, and higher root/shoot ratio under Fe-deficiency stress conditions.     

Evaluation of early generations for iron chlorosis in relation to productivity in groundnut (Arachis hypogaea L.)

Five popular but iron-inefficient cultivars were crossed with three efficient genotypes and both parents and F₁s were evaluated for iron-efficiency in potted calcareous and noncalcareous soil. The iron-efficient genotypes were dark green or green in both noncalcareous and calcareous soils whereas inefficient types were light green to yellow in calcareous soil. The chlorophyll and active iron (Fe²⁺) concentration of leaves was less in iron-efficient genotypes compared to efficient types in calcareous soil and reduction of both the parameters from noncalcareous to calcareous soil was considerably high in iron-inefficient lines. There was significant correlation between visual scores, chlorophyll and active iron content. There were no differences among F₁s for iron chlorosis and they were all iron-inefficient. The frequency of iron-inefficient plants was higher than the efficient plants in all F₂ populations. But most of the productive plants came from iron-efficient segregants indicating strong association between iron-efficiency and productivity. Based on the results selection for iron-efficiency in early generations and extensive evaluation for productivity in advanced generations is suggested for developing varieties for cultivation in calcareous soils.     

Effect of iron chlorosis on mineral nutrition and lipid composition of thylakoid biomembrane in Prunus persica (L.) Bastch.

The effect of iron chlorosis on mineral, thylakoid lipids and fatty acids composition of field grown peach tree leaves was studied. Significant differences were found in iron extracted by using , -dipyridyl (active iron), total iron, P, K, Cu and the P/Fe and Fe/Mn ratios. The levels of total chlorophyll, total glycolipids and phospholipids were reduced under iron chlorosis. A slight iron deficiency does not modify the fatty acid composition of thylakoid membranes, while a strong deficiency changes the proportion of some fatty acids.Abbreviations Chl chlorophyll - DGDG digalactosyldiglycerol - MGDG mono-galactosyldiglycerol - PC phosphatidycholine - PE phophatidylethanolamine - PG phophatidylglycerol - TLC thin layer chromatography - 16:0 palmitic acid - 16:1 palmitoleic acid - 16:lt trans-hexadecenoic - 18:0 steric acid - 18:1 oleic acid - 18:2 linoleic acid - 18:3 linolenic acid     

The effects of nutrient supply,predominantly addition of iron,and rhizobial inoculation on the tolerance of Lupinus pilosus genotypes to a calcareous soil

Two glasshouse experiments were conducted to examine the effects of nutrient supply and rhizobial inoculation on the performance of Lupinus pilosus genotypes differing in tolerance to calcareous soils. In experiment 1, plants were grown for 84 days in a calcareous soil (50% CaCO₃; soil water content 90% of field capacity) at four nutrient treatments (no-added nutrients, added nutrients without Fe, added nutrients with soil applied FeEDDHA, added nutrients with foliar applied FeSO₄). In experiment 2, plants were grown for 28 days with supply of NH₄NO₃ without inoculation or inoculated with Bradyrhizobium sp. (Lupinus). Chlorosis in the youngest leaves was a good indicator of the relative tolerance of the genotypes to the calcareous soil in both experiments, except the treatment with FeEDDHA at 5 mg kg^–1 soil which was toxic to all genotypes. Chlorosis scores correlated with chlorophyll meter readings and chlorophyll concentrations. The foliar application of FeSO₄ did not fully alleviate chlorotic symptoms despite concentrations of active or total Fe in the youngest leaves being increased. Adding nutrients and chemical nitrogen did not change the severity of chlorosis or improve the growth of the plant. The nutrient supply did not alter the ranking of tolerance of genotypes to the calcareous soil. The results suggest that nutrient deficiency or poor nodulation was not a major cause of poor plant growth on calcareous soils and that bicarbonate may exert a direct effect on chlorophyll synthesis. The mechanism for tolerance is likely to be related to an ability to exclude bicarbonate or prevent its transport to the leaves.     

Correcting iron deficiencies in annual and perennial plants: Present technologies and future prospects

Correction of Fe chlorosis is done mainly by foliar sprays because soil applications generally are ineffective, especially for annual crops. Inorganic Fe sources applied to soils react rapidly to forms which are not as available to plants; ferrous Fe is oxidized to the ferric form in well-aerated soils, especially as soil pH increases. Several synthetic chelates and organic complexes have been used with varying success, depending upon Fe source and rate, application method, plant species, and weather and soil conditions. Use of Fe-efficient cultivars is one method of counteracting Fe deficiencies in some species. Future prospects for improving control of Fe chlorosis lie more with development of Fe-efficient cultivars of Fe-sensitive species than with development of improved Fe fertilizers and methods of application. The techniques of molecular biology should be applicable to help solve this important plant nutrition problem, but priority has not been given to conduct this research at this time.     

The mobile ferrous iron pool in Escherichia coli is bound to a phosphorylated sugar derivative

Based on in vivo Mössbauer spectroscopy it has previously been demonstrated that the intracellular iron pool of Escherichia coli, grown in iron deficient media supplemented with siderophores as the sole iron source, is dominated by a single Fe²⁺ and a single Fe³⁺ species. We have isolated the ferrous ion species and have purified it employing native column PAGE, chromatography and ultrafiltration. The purified compound displays an M _app of 2.2 kDa and an extremely low isoelectric point (pI) of 1.05. It is shown that this ferrous ion binding compound is neither a protein nor a nucleotide, rather it is composed mainly of phosphorylated sugar derivatives. This compound binds approximately 40% of the cytoplasmic iron. Therefore it is proposed that this oligomeric ferrous carbohydrate phosphate represents the long sought after mobile, low molecular mass iron pool.     

Effect of selected soil factors on chlorosis of peanut plants grown in calcareous soils in Cyprus

In 1986 and 1987 surveys were conducted of 34 (1986) and 35 (1987) peanut (Arachis hypogaea L.) fields in which the plants showed various degrees of chlorosis. In the areas concerned, plant appearance was classified according to a chlorotic index and corresponding soil samples were taken and analysed for CaCO₃, pH, NO₃–N and DTPA-extractable Fe in 1986 and for CaCO₃, NO₃–N and active lime in 1987.Regression analyses showed that CaCO₃, active lime and NO₃–N were positively correlated, while DTPA-extractable Fe was negatively correlated, with the chlorosis problem. The critical levels above which plants were chlorotic were 20 to 25% CaCO₃ and 10% active lime. Plants began to be chlorotic when DTPA-extractable Fe was below 2.5 mg·kg^–1. The soil factors examined explained about 60% of the variability in plant chlorosis.