Nitrate induced iron deficiency chlorosis in Juncus acutiflorus

Chlorosis caused by iron deficiency is commonly associated with high bicarbonate levels in the soil. However, in rare cases such chlorosis has been observed in soils with high nitrate levels. In a dutch rich-fen, chlorosis has been noted in stands of Juncus acutiflorus at locations where groundwater containing high levels of nitrate reached the surface. Experiments revealed that the chlorosis could be attributed to iron deficiency although iron levels in the shoots were well above the known physiological threshold values for iron deficiency. It is postulated that increased nitrate assimilation leads to an increased apoplastic pH and to a concomitant immobilisation of iron and/or lower iron (III) reduction. Moreover free amino acid levels were markedly higher in the iron deficient plants in the field. It was found, however, that the percentage of nitrogen present as free amino acids was not influenced directly by low iron levels but mainly by the C/N ratios in the shoots. Nowadays, nitrate concentrations in ground water as high 1000 µM are no longer an exception in the Netherlands. We propose that strongly increased nitrate inputs may cause iron stress in natural vegetations, especially in wet habitats.     

Physiological and biochemical adjustment of iron chlorosis affected low-chill peach cultivars supplied with different iron sources

A pot experiment was carried out to investigate the effect of iron supplementation on physiological and biochemical status of the low-chill peach cultivars (Saharanpur Prabhat, Shan-e-Punjab and Pratap) suffered from iron chlorosis in artificially created calcareous soil. Three most commonly used iron sources viz. Fe-sulphate (1.0 % and 0.5 %), Fe-citrate (1.0 % and 0.5 %) and FeEDTA (0.1 % and 0.2 %) were sprayed on the 4th and 5th leaves from the apex of the twig. And after 1 week of spraying, observation on various physiological and biochemical parameters in leaves were recorded. Improvement in plant physiological parameters like chlorophyll content index (CCI), photosynthetic rate (P_n), stomatal conductance (g_s) and intercellular CO₂ conc. (C_i) were recorded best with the application of 1.0 % Fe-sulphate both in treated and untreated upper leaves. The maximum recovery in biochemical parameters such as total leaf chlorophyll content, superoxide dismutase (SOD) and peroxidase (POD) activity was also noted with the application of 1.0 % Fe-sulphate. However, application of 1.0 % Fe-sulphate and 0.5 % Fe-sulphate had similar effect for most of the parameters under study. The ability of iron sources to induce physiological and biochemical responses in iron deficient low-chill peach plants were in the following order Fe-sulphate>Fe-citrate>FeEDTA. Differential responses in plant physiological and biochemical parameters were also exhibited by the low-chill peach cultivars with regard to supplementation of various iron sources. Among the low-chill peach cultivars, Saharanpur Prabhat responded best with the application of iron sources followed by Shan-e-Punjab and Pratap.     

Effects of iron chlorosis and iron resupply on leaf xylem architecture, water relations, gas exchange and stomatal performance of field-grown peach (Prunus persica)

There is increasing evidence suggesting that iron (Fe) deficiency induces not only leaf chlorosis and a decline of photosynthesis, but also structural changes in leaf morphology, which might affect the functionality of leaves. In this study, we investigated the effects of Fe deficiency on the water relations of peach ( Prunus persica (L.) Batsch.) leaves and the responses of previously chlorotic leaves to Fe resupply via the root or the leaf. Iron deficiency induced a decline of maximum potential photosystem II (PSII) efficiency (F _V/F _M), of rates of net photosynthesis and transpiration and of water use efficiency. Iron chlorosis was associated with a reduction of leaf xylem vessel size and of leaf hydraulic conductance. In the course of the day, water potentials in chlorotic leaves remained higher (less negative) than in green leaves. In chlorotic leaves, normal stomatal functioning was disturbed, as evidenced by the lack of opening upon withdrawal of external CO₂ and stomatal closure after sudden illumination of previously darkened leaves. We conclude that the Fe deficiency induced limitations of xylem conductivity elicited a water saving strategy, which poses an additional challenge to plant growth on high pH, calcareous soils. Fertilisation with Fe improved photosynthetic performance but the proper xylem structure and water relations of leaves were not fully restored, indicating that Fe must be available at the first stages of leaf growth and development.     

Effects of branch solid Fe sulphate implants on xylem sap composition in field-grown peach and pear: changes in Fe, organic anions and pH

The effects of placing solid implants containing Fe sulfate in branches of Fe-deficient pear and peach trees on the composition of the xylem sap have been studied. Iron sulfate implants are commercially used in northeastern Spain to control iron chlorosis in fruit trees. Implants increased Fe concentrations and decreased organic acid concentrations in the xylem sap, whereas xylem sap pH was only moderately changed. The citrate to Fe ratios decreased markedly after implants, therefore improving the possibility that Fe could be reduced by the leaf plasma membrane enzyme reductase, known to be inhibited by high citrate/Fe ratios. In peach, the effects of the implants could be observed many months post treatment. In pear, some effects were still observed one year after the implants had taken place. Results obtained indicate that solid Fe sulfate implants were capable of significantly changing the chemical composition of the xylem sap in fruit trees.     

Mapping genetic loci for iron deficiency chlorosis in soybean

The objective of this study was to map genes controlling iron deficiency chlorosis in two intraspecific soybean [Glycine max (L.) Merrill] populations. Chlorosis symptoms were evaluated by visual scores and spectrometric chlorophyll determinations at the V4 stage (third trifoliolate leaf fully developed) in the field in 1993, and at V2 (first trifoliolate leaf fully developed) and V4 stages in 1994. A total of 89 RFLP and 10 SSR markers in the Pride B216 x A15 population, and 82 RFLP, 14 SSR and 1 morphological I (hilum color) markers in the Anoka x A7 population were used to map quantitative trait loci (QTL) affecting iron deficiency chlorosis. QTL with minor effects were detected on six linkage groups of the Pride B216 x A15 population, suggesting a typical polygene mechanism. In contrast, in the Anoka x A7 population, one QTL contributed an average of 72.7% of the visual score variation and 68.8% of the chlorophyll concentration variation and was mapped on linkage group N. Another QTL for visual score variation, and one for chlorophyll concentration variation were detected on linkage groups A1 and I, respectively. Due to the large LOD score and major genetic effect of the QTL on linkage group N, the quantitative data was reclassified into qualitative data fitting a one major gene model according to the means of the QTL genotypic classes. The major gene was mapped in the same interval of linkage group N using both visual scores and chlorophyll concentrations, thus verifying that one major gene is involved in segregation for iron chlorosis deficiency in the Anoka x A7 population. This study supported a previous hypothesis that two separate genetic mechanisms control iron deficiency in soybean.     

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     

Characterization of the tolerance to iron chlorosis in different peach rootstocks grown in nutrient solution

Iron chlorosis is an important problem in peach trees, but differences exist between peach rootstocks in their tolerance to Fe chlorosis in calcareous soils. The purpose of this investigation was to characterize the tolerance of different rootstocks to Fe chlorosis induced by bicarbonate in nutrient solution. The rootstocks studied included peach (Nemaguard), plums (Brompton, San Julian A and Puebla de Soto 101) and almond × peach hybrids (Adafuel and GF677). Young plants obtained from rooted cuttings or from in vitro culture techniques were grown individually, under controlled conditions, in flasks with 700 mL of aerated nutrient solution low in iron and with or without 10 mM bicarbonate or 10 mM phosphate. Susceptiblity to bicarbonate-induced chlorosis was inversely correlated with both the Fe content in young leaves and the reducing capacity of roots, but not with the phosphorus content in young leaves. The plum Puebla de Soto 101 and the hybrid GF677 showed the lowest degree of chlorosis and the highest reducing capacity. Phosphate did not induce chlorosis.     

Effects of moderate and severe iron deficiency chlorosis on fruit yield, appearance and composition in pear (Pyrus communis L.) and peach (Prunus persica (L.) Batsch)

The effects of different levels of Fe-deficiency chlorosis on the fruit yield, appearance and composition of pear and peach trees grown in field orchards have been studied. The major effect of Fe deficiency in both species was a large yield reduction, even when chlorosis was moderate, associated to decreases in fruit tree load. Fruit size increased with moderate chlorosis in both species and decreased with severe chlorosis in peach. In peach, moderate or severe chlorosis affected uniformly all branches, leading to firmer fruits with higher acidity, total phenolics and carboxylates. This indicates a delayed maturity that can be attributed to a low C-availability for fruits. In Fe-deficient pear trees, the majority of fruits (98%) were on non-chlorotic or moderately chlorotic branches, and fruits were less green and firm with an increased sugars/acids ratio. This indicates an advanced fruit maturity that can be attributed to an increased C-availability for fruits. All chlorosis levels increased within-tree variation in fruit appearance.     

Prognosis of iron chlorosis in pear (Pyrus communis L.) and peach (Prunus persica L. Batsch) trees using bud, flower and leaf mineral concentrations

Background and Aims

The possibility of using tree materials in early phenological stages, such as dormant buds and flowers, for the prognosis of Fe deficiency occurring later in the year has been studied in peach and pear trees.

Methods

Thirty-two peach trees and thirty pear trees with different Fe chlorosis degrees were sampled in different commercial orchards. In peach, samples included flower buds, vegetative buds, bud wood, flowers and leaves at 60 and 120?days after full bloom (DAFB). In pear, samples included buds, bud wood, flowers and leaves at 60 and 120?days DAFB. Leaf chlorophyll was assessed (SPAD) at 60 and 120 DAFB. Sampling was repeated for 3–5?years depending on the materials. Mineral nutrients measured were N, P, K, Ca, Mg, Fe, Mn, Zn and Cu.

Results

The relationships between the nutrient concentrations in the different materials and leaf SPAD were assessed using four different statistical approaches: i) comparison of means depending on the chlorosis level, ii) correlation analysis, iii) principal component analysis, and iv) stepwise multiple regression. In all cases, significant associations between nutrients and SPAD were found. The best-fit multiple regression curves obtained for the multi-year data set provided good prediction in individual years.

Conclusions

Results found indicate that it is possible to carry out the prognosis of Fe chlorosis using early materials such as buds and flowers. The relationships obtained were different from those obtained in previous studies using a single orchard. The different methods of analysis used provided complementary data.     

Morpho-physiological parameters affecting iron deficiency chlorosis in soybean (Glycine max L.)

Background and aims

Iron deficiency chlorosis (IDC) leads to severe leaf chlorosis, low photosynthetic rates, and yield reductions of several million metric tonnes each year. In order to devise breeding and genetic transformation programs that aim at generating high-yielding and IDC-tolerant soybean lines, it is necessary to better understand the mechanisms that enable tolerant plants to survive under Fe-limiting conditions.

Methods

An in silico analysis in the USDA soybean collection allowed the identification of a set of novel efficient and inefficient soybean cultivars which can be used in future studies concerning IDC response. Plants were grown in iron deficient and iron sufficient conditions using a bicarbonate system and several IDC-related aspects were studied.

Results

A new set of efficient and inefficient soybean lines were identified in silico, and their tolerance to IDC was confirmed under laboratorial conditions. New plant traits that are highly correlated to IDC scoring were identified: a negative correlation was found between SPAD values and stem weight, weight of the unifoliolates and iron concentration of the first unifoliolates was found; higher SPAD values were correlated with the amount of iron in the first trifoliate leaves. Our data also show that having higher concentrations of iron in the seeds provides increased resistance to IDC. No correlation was found between root iron reductase activity and chlorosis.

Conclusions

Soybean differential chlorosis susceptibility between different accessions is linked to specific morpho-physiological parameters such as unifoliolate leaf size, stem weigh, concentration of iron in the seeds, and tissue iron partitioning.     

Tertiary structural changes associated with iron binding and release in hen serum transferrin: a crystallographic and spectroscopic study

The iron binding and release of serum transferrin are pH-dependent and accompanied by a conformational change between the iron-bound (holo-) and iron-free (apo-) forms. We have determined the crystal structure of apo-hen serum transferrin (hAST) at 3.5A resolution, which is the first reported structure to date of any full molecule of an apo-serum transferrin and studied its pH-dependent iron release by UV-vis absorption and near UV-CD spectroscopy. The crystal structure of hAST shows that both the lobes adopt an open conformation and the relative orientations of the domains are different from those of apo-human serum transferrin and human apolactoferrin but similar to that of hen apo-ovotransferrin. Spectroscopic analysis reveals that in hen serum transferrin, release of the first iron starts at a pH approximately 6.5 and continues over a broad pH range (6.5-5.2). The complete release of the iron, however, occurs at pH approximately 4.0. The near UV-CD spectra show alterations in the microenvironment of the aromatic residues surrounding the iron-binding sites.     

Leaf xeromorphy as related to physiological and structural influences

The Botanical Review -     

缺铁和矫治缺铁对梨树叶片结构的影响

通过对不同缺铁黄化程度的酥梨叶片进行解剖结构分析和矫治缺铁后叶片解剖结构的恢复情况的研究,结果表明,缺铁黄化使叶片厚度,叶肉厚度,栅栏组织厚度,栅栏组织／海绵组织的比值均明显下降;栅栏组织细胞排列疏松,细胞变短甚至断裂成块状,叶绿体数量明显减少甚至无,并出现解体细胞;栅栏组织与海绵组织界限模糊,海绵组织细胞排列变密;维管束的导管口径变小,维管束鞘和韧皮部的细胞番红染色加重。进行缺铁黄化矫治后,叶片结构恢复正常。这些结构的变化,可作为梨树缺铁诊断和矫治的指标。     

Physiological and biochemical responses of the iron chlorosis tolerant grapevine rootstock 140 Ruggeri to iron deficiency and bicarbonate

Background and aims

Iron (Fe) deficiency chlorosis associated with high levels of soil bicarbonate is one of the main nutritional disorders observed in sensitive grapevine genotypes. The aim of the experiment was to assess both the independent and combined effects of Fe and bicarbonate nutrition in grapevine.

Methods

Plants of the Fe chlorosis tolerant 140 Ruggeri rootstock were grown with and without Fe(III)-EDTA and bicarbonate in the nutrient solution. SPAD index, plant growth, root enzyme (PEPC, MDH, CS, NADP⁺ ?IDH) activities, kinetic properties of root PEPC, organic acid concentrations in roots and xylem sap and xylem sap pH were determined. A factorial statistical design with two factors (Fe and BIC) and two levels of each factor was adopted: +Fe and ?Fe, and +BIC and ?BIC.

Results

This rootstock strongly reacted to Fe deficiency by activating several response mechanisms at different physiological levels. The presence of bicarbonate in the nutrient solution changed the activity of PEPC and TCA related enzymes (CS, NADP⁺-IDH) and the accumulation/translocation of organic acids in roots of Fe-deprived plants. Moreover, this genotype increased root biomass and root malic acid concentration in response to high bicarbonate levels in the substrate. Bicarbonate also enhanced leaf chlorophyll content.

Conclusions

Along with a clear independent effect on Fe nutrition, our data support a modulating role of bicarbonate on Fe deficiency response mechanisms at root level.     

Metabolic changes associated with selenium deficiency in mice

Selenium (Se), which is a central component for the biosynthesis and functionality of selenoproteins, plays an important role in the anti-oxidative response, reproduction, thyroid hormone metabolism and the protection from infection and inflammation. However, dietary Se effects have not well been established to date and the available studies often present contradictory results. To obtain a better understanding of Se intake and its influence on the metabolism of living systems, we have utilized a metabolomics approach to gain insight into the specific metabolic alterations caused by Se deficiency in mice. Serum samples were collected from two groups of C57BL/6 mice: an experimental group which was fed a Se-deficient diet and controls consuming normal chow. The samples were analyzed by ¹H nuclear magnetic resonance spectroscopy and gas chromatography-mass spectrometry. The resulting metabolite data were examined separately for both analytical methods and in a combined manner. By applying multivariate statistical analysis we were able to distinguish the two groups and detect a metabolite pattern associated with Se deficiency. We found that the concentrations of 15 metabolites significantly changed in serum samples collected from Se-deficient mice when compared to the controls. Many of the perturbed biological pathways pointed towards compensatory mechanisms during Se deficiency and were associated with amino acid metabolism. Our findings show that a metabolomics approach may be applied to identify the metabolic impact of Se and reveal the most impaired biological pathways as well as induced regulatory mechanisms during Se deficiency.     

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。     

Rhizobitoxine-induced chlorosis occurs in coincidence with methionine deficiency in soybeans

BACKGROUND AND AIMS: Rhizobitoxine, produced by the legume symbiont Bradyrhizobium elkanii, inhibits cystathionine-beta-lyase (EC 4.4.1.8) in methionine biosynthesis and 1-aminocyclopropane-1-carboxylate synthase (ACC) in ethylene biosynthesis. Rhizobitoxine production by B. elkanii enhances nodulation of host legumes via the inhibition of ethylene synthesis, but causes foliar chlorosis in susceptible soybeans, though how it does so remains to be investigated. The aim of this study was to examine the physiological basis of rhizobitoxine-induced chlorosis in soybeans. METHODS: Wild-type B. elkanii and a rhizobitoxine-deficient mutant were inoculated in Glycine max 'Lee'. Thirty days after inoculation, the upper parts of soybean shoots were analysed for amino acid contents. Chlorotic soybeans inoculated with wild-type B. elkanii were treated with methionine and ACC to assess the effects of the chemicals on the chlorosis. KEY RESULTS: Chlorotic upper shoots of soybeans inoculated with wild-type B. elkanii had a lower methionine content and higher accumulation of the methionine precursors than those with the rhizobitoxine-deficient mutant. In addition, the foliar chlorosis was alleviated by the application of methionine. CONCLUSIONS: Rhizobitoxine-induced chlorosis occurs in coincidence with methionine deficiency as a result of cystathione-beta-lyase inhibition during methionine biosynthesis.     

Iron deficiency-induced changes in the photosynthetic pigment composition of field-grown pear (Pyrus communis L) leaves

In this work we characterize the changes induced by iron deficiency in the pigment composition of pear (Pyrus communis L.) leaves grown under high light intensities in field conditions in Spain. Iron deficiency induced decreases in neoxanthin and β-carotene concomitantly with decreases in chlorophyll a, whereas lutein and carotenoids within the xanthophyll cycle were less affected. Iron deficiency caused major increases in the lutein/chlorophyll a and xanthophyll cycle pigments/chlorophyll a molar ratios. The chlorophyll a/chlorophyll b ratio increased in response to iron deficiency. The carotenoids within the xanthophyll cycle in iron-deficient and in iron-sufficient (control) leaves underwent epoxidations and de-epoxidations in response to ambient light conditions. In control leaves dark-adapted for several hours, most of the xanthophyll cycle pigment pool was in the epoxidated form vio-laxanthin, whereas iron-deficient leaves had significant amounts of zeaxanthin. Iron-deficient leaves also exhibited an increased non-photochemical quenching, supporting the possibility of a role for pigments within the xanthophyll cycle in photoprotection.