Fe Availability Research Articles

Improving the growth, yield and iron concentration of strawberry using sodium hydrosulfide (NaHS) under soilless culture

Required iron (Fe) for the growing plants in soilless culture could be obtained from various Fe sources such as chelates and ferrous sulfate (FeSO4). Despite the low cost of FeSO4, this source is rarely used due to the low Fe solubility at a high pH. Here, the role of sodium hydrosulfide (NaHS) in improving growth and iron availability in strawberry (Fragaria × ananassa cv. Paros) grown in FeSO4 culture solution was reported. The plants were grown hydroponically with two Fe sources, Fe-EDTA and FeSO4 along with three levels (0, 0.2, and 0.5 mM (of H2S donor, NaHS. The results showed that vegetative growth and yield of strawberry under FeSO4 source were lower than Fe-EDTA treated plants. Using NaHS significantly improved vegetative and reproductive growth, yield, chlorophyll SPAD value, maximum efficiency of photosystem II (Fv/Fm), and Fe availability of strawberries under FeSO4 source. Exogenously applied NaHS reduced the hydrogen peroxide (H2O2) and malondialdehyde (MDA) production and electrolyte leakage (EL) in the leaves by enhancing enzymatic antioxidant activity (ascorbate peroxidase, superoxide dismutase, and catalase) in FeSO4 source. Moreover, NaHS affected fruit yield under Fe-EDTA utilization and showed a significant increase in yield amount (by 29.75%) compared to control. In general, there were no significant differences between the two levels of NaHS, 0.2 and 0.5 mM. Our results indicated that H2S affects the maintenance of cell membrane integrity and subsequently iron availability by strengthening the enzymatic antioxidant system and reducing oxidative stress.

Mitigating cadmium accumulation in dicotyledonous vegetables by iron fertilizer through inhibiting Fe transporter IRT1-mediated Cd uptake

The solubility of cadmium (Cd) in soil and its transfer to plants are influenced by soil pH. While increasing soil pH reduces Cd solubility and accumulation in rice plants grown in acidic soils, its effect on Cd accumulation in vegetables remains inconclusive. Here, we investigated the impact of soil pH on Cd accumulation in dicotyledonous vegetables and elucidated the underlying molecular mechanisms. Soils collected from various locations were supplemented with varying quantities of lime to achieve soil pH values of around 5.0, 6.0, 7.0, and 8.0. Raising soil pH from around 5.0 to 8.0 markedly decreased extractable Cd. However, increasing soil pH tended to promote shoot Cd accumulation in dicotyledonous vegetable species including lettuce, pakchoi, and Chinese cabbage, and the model dicotyledonous plant Arabidopsis thaliana. Conversely, soil pH increase resulted in a monotonic decrease in rice Cd accumulation. In our hydroponic experiments, we discovered that iron (Fe) deficiency substantially increased Cd uptake and accumulation in dicotyledonous plants but not in rice. Increasing soil pH reduced soil Fe availability and induced the Fe transporter gene IRT1 expression in dicotyledonous vegetables roots, which led to an increase in IRT1-mediated Cd uptake and subsequently increased Cd accumulation as soil pH increases. A comprehensive model incorporating extractable Cd and root IRT1 expression better explained Cd accumulation in vegetable shoots. The application of 50 mg/kg of Fe fertilizer in neutral or alkaline soils resulted in a significant reduction in Cd accumulation by 34–58% in dicotyledonous vegetables. These findings reveal that increasing soil pH has two opposite effects, decreasing soil Cd availability while promoting Cd uptake through IRT1 upregulation, reconciling the inconsistency in its effect on Cd accumulation in dicotyledonous plants. Our findings provide important insights for understanding the factors affecting Cd uptake in plants and offer a practical solution to mitigate Cd contamination in vegetables.

Phytoavailability of cadmium in rice amended with organic materials and lime: Effects of rhizosphere chemical changes and cadmium sequestration in iron plaque

Excessive Cd in rice grains produced with acidic paddy soil is receiving increasingly widespread attention because it endangers human health. Applying organic materials (OM) and lime (L) is a common technique used to reduce Cd concentration in grains (CdG). Nevertheless, the mechanism by which their simultaneous application affects the Cd phytoavailability in soilrice systems remains ambiguous. In the current study, we adopted a rhizobag pot culture test to explore the influences of single application of OM [rice straw (RS), milk vetch (MV)], L, and their co-utilization on Cd phytoavailability and the associated mechanisms. The results showed that the application of RS, MV, L, L + RS (LRS), and L + MV (LMV) significantly decreased CdG by 26.9%, 38.2%, 48.6%, 50.0%, and 53.0%, respectively. Fe plaque (IP) formation was not affected by these treatments; however, Cd sequestration in IP (CdIP) was significantly reduced. CdIP was significantly reduced by 18.3%, 23.6%, 43.8%, 33.1%, and 41.4%, after RS, MV, L, LRS, and LMV treatments, respectively. Additionally, available Cd concentrations in rhizospheric soil (RHS) were significantly reduced by 11.5%, 14.8%, 15.1%, and 18.4%, after MV, L, LRS, and LMV treatments, respectively. Cd availability in RHS was significantly influenced by pH, dissolved organic carbon concentration, and Zn, Fe, and Mn availability. The results of the structure equation mode showed that CdG was mainly affected by CdIP, followed by Cd availability and the pH of RHS. In conclusion, the reduction of CdG by OM, L, and their co-utilization was the results of their combined effects of reducing Cd availability in RHS, CdIP, and Cd uptake by the roots. This study emphasizes that the reduction of CdG is a result of the dual effects of reducing Cd availability in RHS and CdIP after amendments application. L application alone or in conjunction with OM is an efficient practice to reduce CdG in acidic Cd-contaminated paddy fields.

Acclimation to medium-level non-lethal iron limitation: Adjustment of electron flow around the PSII and metalloprotein expression in Trichodesmium erythraeum IMS101

The aim of this study was to investigate how acclimation to medium-level, long-term, non-lethal iron limitation changes the electron flux around the Photosystem II of the oceanic diazotroph Trichodesmium erythraeum IMS101. Fe availability of about 5× and 100× lower than a replete level, i.e. conditions common in the natural environment of this cyanobacterium, were applied in chemostats. The response of the cells was studied not only in terms of growth, but also mechanistically, measuring the chlorophyll fluorescence of dark-adapted filaments via imaging fluorescence kinetic microscopy (FKM) with 0.3 ms time resolution. Combining these measurements with those of metal binding to proteins via online coupling of metal-free HPLC (size exclusion chromatography SEC) to sector-field ICP-MS allowed to track the fate of the photosystems, together with other metalloproteins. General increase of fluorescence has been observed, with the consequent decrease in the quantum yields φ of the PSII, while the efficiency ψ of the electron flux between PSII and the PSI remained surprisingly unchanged. This indicates the ability of Trichodesmium to cope with a situation that makes assembling the many iron clusters in Photosystem I a particular challenge, as shown by decreasing ratios of Fe to Mg in these proteins. The negative effect of Fe limitation on PSII may also be due to its fast turnover. A broader view was obtained from metalloproteomics via HPLC-ICP-MS, revealing a differential protein expression pattern under iron limitation with a drastic down-regulation especially of iron-containing proteins and some increase in low MW metal-binding complexes.

The Key Physiological and Biochemical Traits Underlying Common Bean (Phaseolus vulgaris L.) Response to Iron Deficiency, and Related Interrelationships

Iron deficiency is a worldwide nutritional problem affecting crop production. In Tunisia, this mineral disorder hampers the growth and yield of the major crops due to the abundance of calcareous soils that limit iron availability. The common bean is one of these crops suffering from lime-induced iron chlorosis. The exploration of the variability of common bean responses to iron deficiency allows us to screen tolerant cultivars and identify useful traits and indicators for further screening programs. To this end, two common bean cultivars (coco blanc, CB, and coco nain, CN) were cultivated hydroponically in standard nutrient solution (control) or nutrient solution deprived of iron (stressed). Analyses were reported on plant growth, photosynthetic pigments, photosynthesis, iron distribution, H-ATPase, and Fe-chelate reductase (Fe-CR) activities; important indicators were calculated; and convenient correlations were established. Current results demonstrated that iron deficiency stimulated specific symptoms of iron chlorosis on young leaves that were more precocious and severe in CB than CN. Spad index and chlorophyll pigments measurement confirmed these morphological changes and cultivar differences. Net photosynthesis (Pn) showed the same scheme of variation, with a significant decrease in Pn while respecting the previous cultivar’s variability. Plant growth is no exception to this general trend. The biomass decrease was two times higher in CB than CN. Otherwise, this mineral disorder significantly decreased Fe concentration in all plant organs. However, CN accumulated 40% more Fe than CB, resulting from its higher Fe Fe-CR and H-ATPase activities. Our results also demonstrated the close dependence of these metabolic functions on Fe availability in shoots and the strict relationship between Fe-CR and H-ATPase, photosynthesis, and chlorophyll content. Otherwise, CN demonstrated higher efficiency of Fe’s use (FeUE) for the key metabolic functions (photosynthesis, chlorophyll biosynthesis, and plant growth). The relative tolerance of CN as compared to CB was explained by its ability to establish a functional system less vulnerable to iron deficiency that operates effectively under problematic conditions. This system involves metabolic functions in shoots (photosynthesis, chlorophyll biosynthesis, Fe repartition, etc.) and others in roots (H-ATPase, Fe-CR), which are strictly interdependent.

Calcium-dependent protein kinases CPK21 and CPK23 phosphorylate and activate the iron-regulated transporter IRT1 to regulate iron deficiency in Arabidopsis.

Iron (Fe) is an essential micronutrient for all organisms. Fe availability in the soil is usually much lower than that required for plant growth, and Fe deficiencies seriously restrict crop growth and yield. Calcium (Ca2+) is a second messenger in all eukaryotes; however, it remains largely unknown how Ca2+ regulates Fe deficiency. In this study, mutations in CPK21 and CPK23, which are two highly homologous calcium-dependent protein kinases, conferredimpaired growth and rootdevelopment under Fe-deficient conditions, whereas constitutively active CPK21 and CPK23 enhanced plant tolerance to Fe-deficient conditions. Furthermore, we found that CPK21 and CPK23 interacted with and phosphorylated the Fe transporter IRON-REGULATED TRANSPORTER1 (IRT1) at the Ser149 residue. Biochemical analyses and complementation of Fe transport in yeast and plants indicated that IRT1 Ser149 is critical for IRT1 transport activity. Taken together, these findings suggest that the CPK21/23-IRT1 signaling pathway is critical for Fe homeostasis in plants and provides targets for improving Fe-deficient environments and breeding crops resistant to Fe-deficient conditions.

Role of Soil and Foliar-Applied Carbon Dots in Plant Iron Biofortification and Cadmium Mitigation by Triggering Opposite Iron Signaling in Roots.

In China, iron (Fe) availability is low in most soils but cadmium (Cd) generally exceeds regulatory soil pollution limits. Thus, biofortification of Fe along with mitigation of Cd in edible plant parts is important for human nutrition and health. Carbon dots (CDs) are considered as potential nanomaterials for agricultural applications. Here, Salvia miltiorrhiza-derived CDs are an efficient modulator of Fe, manganese (Mn), zinc (Zn), and Cd accumulation in plants. CDs irrigation (1mgmL-1 , performed every week starting at the jointing stage for 12 weeks) increased Fe content by 18% but mitigated Cd accumulation by 20% in wheat grains. This findingwasassociated with the Fe3+ -mobilizing properties of CDs from the soil and root cell wall, as well as endocytosis-dependent internalization in roots. The resulting excess Fe signaling mitigated Cd uptake via inhibiting TaNRAMP5 expression. Foliar spraying of CDs enhanced Fe (44%), Mn (30%), and Zn (19%) content with an unchanged Cd accumulation in wheat grains. This result is attributed to CDs-enhanced light signaling, which triggered shoot-to-root Fe deficiency response. This study not only reveals the molecular mechanism underlying CDs modulation of Fe signaling in plants but also provides useful strategies for concurrent Fe biofortification and Cd mitigation in plant-based foods.

Changes of Hepcidin, Ferritin and Iron Levels in Cycling Purebred Spanish Mares.

Several studies have demonstrated that in woman the sex hormones such as estrogen (E2) and progesterone (P4) influence iron (Fe) regulation, contributing to variations in Fe parameters along the menstrual cycle. These mechanisms based on the regulation of hepcidin (Hepc) which limits Fe availability during the cycle, remain poorly characterized in healthy mares. The objective of this study was to establish the relationship between Hepc, Fe, ferritin (Ferr), and the primary ovarian hormones E2 and P4 in cycling Purebred Spanish mares. Blood samples were taken from 31 Purebred Spanish mares day -5, on day 0, day +5 and day +16 of the cycle. Fe and Ferr significantly increased and Hepc decreased during pre- and ovulatory periods. The secretion peak of estradiol-17β (E2) was reached on day 0 and progesterone (P4) between days +5 and +16. Fe and Ferr were positively correlated (r = 0.57). Fe and Ferr were negatively correlated with Hepc (r = -0.72 and r = -0.02, respectively). E2 and P4 were negatively and positively correlated with Hepc (r = -0.753 and r = 0.54, respectively). In cycling Purebred Spanish mares there is a measurable relationship between steroid hormones and systemic Fe metabolism. Estrogenic dominance in the pre- and ovulatory period allows for a more effective iron status, mediated by hepcidin inhibition. However, P4 during the luteal phase substantially reduces serum Fe and iron stores, possibly related to Hepc stimulation. Future research is required to clarify the relationship between steroid hormones and iron metabolism at the molecular level in equids.

The influence of light on the availability of photoreactive and photostable iron(III)–ligand complexes to diatoms

AbstractUptake of iron (Fe) by eukaryotic phytoplankton is suggested to proceed by a reductive pathway in which Fe(III) bound to inorganic and organic ligands is reduced prior to Fe internalization. Photochemical reduction of the Fe–ligand complexes can increase Fe bioavailability, but the role of light in regulating physiological processes related to Fe acquisition is not well known. Here, we report how model Fe–ligand complexes differing in photolability affected growth, cellular Fe reduction, and Fe uptake rates of two marine diatoms as a function of irradiance. Growth rates of Thalassiosira oceanica and Phaeodactylum tricornutum in media amended with Fe complexed with ethylenediaminetetraacetic acid (Fe‐EDTA) and desferrioxamine B (FeDFB) increased linearly with light between 50 and 400 μmol photons m−2 s−1 under Fe‐limiting conditions. Steady‐state Fe uptake rates (ρSS) were also light dependent, increasing proportionally with growth irradiance for both Fe–ligand complexes. Photolysis of the Fe‐EDTA complex could explain the increase in ρSS in Fe‐EDTA‐amended medium because it increased [Fe(III)’], but it could not account for the faster rate of Fe uptake from FeDFB at high light because FeDFB is photostable. Short‐term Fe uptake rates measured in the absence of photochemically mediated Fe(III) reduction showed that cells preconditioned to high growth irradiance took up Fe faster than cells grown at low light. Thus, growth at high light induced a physiological response that increased Fe uptake. High growth irradiance also increased the rate of cellular Fe(III)’ reduction in the dark, similar to its stimulatory effect on short‐term Fe uptake. These results identify a previously unrecognized Fe–light interaction that could be important in enhancing Fe availability to diatoms. We suggest that the light effect could be the result of a light‐dependent regulation of cellular Fe(III) reduction.

Potential of animal manure amendments in combating calcareous induced iron deficiency in pearl millet

Calcareous soils are known for their nutritional disorders due to the insoluble form of iron that limits free Fe availability for crops. However, it is well established that organic fertilization improves the physicochemical and biological properties of the soil and can mitigate nutrient deficiency. According to this approach, a greenhouse experiment was conducted on four pearl millet genotypes cultivated on calcareous soil added with an increasing concentration of animal compost as a sustainable approach to mitigate lime-induced Fe deficiency and propose an organic farming system. Fe mobility in the soil-plant system, plant growth, chlorophyll, and photosynthesis were analyzed. All plants grown in calcareous soil developed specific iron deficiency symptoms, with varying intensities and dates of emergence depending on genotype. The addition of compost significantly improved iron availability in the calcareous soil, increased Fe uptake and accumulation in the plant, alleviated symptoms of iron chlorosis, and stimulated photosynthesis and plant growth. Animal compost improved calcareous soil characteristics, followed by Fe availability and uptake. The genotypic differences observed in this study confirmed the specific performance of IP19586 and IP13150, which expressed higher capacities of Fe remobilization in the rhizosphere, Fe uptake, and preferential allocation to shoots.

Iron Nutrition in Plants: Towards a New Paradigm?

Iron (Fe) is an essential micronutrient for plant growth and development. Fe availability affects crops' productivity and the quality of their derived products and thus human nutrition. Fe is poorly available for plant use since it is mostly present in soils in the form of insoluble oxides/hydroxides, especially at neutral to alkaline pH. How plants cope with low-Fe conditions and acquire Fe from soil has been investigated for decades. Pioneering work highlighted that plants have evolved two different strategies to mine Fe from soils, the so-called Strategy I (Fe reduction strategy) and Strategy II (Fe chelation strategy). Strategy I is employed by non-grass species whereas graminaceous plants utilize Strategy II. Recently, it has emerged that these two strategies are not fully exclusive and that the mechanism used by plants for Fe uptake is directly shaped by the characteristics of the soil on which they grow (e.g., pH, oxygen concentration). In this review, recent findings on plant Fe uptake and the regulation of this process will be summarized and their impact on our understanding of plant Fe nutrition will be discussed.

Photomorphogenic tomato mutants high-pigment 1 and aurea responses to iron deficiency

• There is a close interaction between Fe nutrition and light perception in plants. • The tomato response mechanisms to Fe deficiency is highly light-dependent. • hp-1 tomato mutant is more tolerant to Fe deficiency. • au tomato mutant tend to accumulate less nutrients. Iron (Fe) is a micronutrient for plant development, as constituent of several photosynthesis- and respiration-related proteins and enzymes. Consequently, Fe deficiency leads to chlorosis in leaves and plant growth impairment. It has become increasingly evident that light signals coordinate iron homeostasis in plants. To further address new insights into how light is a fundamental part of Fe deficiency responses, we employed Micro-Tom (wild type, WT) tomato ( Solanum lycopersicum L.) plants and high-pigment 1 ( hp1 ) and aurea ( au ) photomorphogenic mutants, which exhibit an excessive light response and low light perception, respectively. Plant growth, pigment contents, oxidative status, and nutrient profile were analyzed. The results revealed the influence of the different genotypes on Fe deficiency responses. WT and au exhibited plant growth reduction under Fe deficiency. WT, hp1 and au demonstrated that Fe availability and light perception play fundamental roles in chlorophyll and anthocyanin biosynthesis. Lipid peroxidation was not increased for any genotype under Fe deficiency, indicating that mutations in light perception and signaling differentially modulate H 2 O 2 production and scavenging under this condition. Additionally, macronutrients and micronutrients were taken up and distributed differently among the different plant genotypes, tissues and Fe conditions analyzed. In general, the au plants accumulated lower amounts of nutrients (Ca, S, P, Mg, B and Zn) than the WT and hp1 genotypes regardless of the Fe concentrations. Our data clearly indicates that light perception and signaling influence Fe-dependent morphophysiological responses in plants, suggesting possibilities for biotechnological improvement of crops grown under Fe shortage.

Techniques to Study Common Root Responses to Beneficial Microbes and Iron Deficiency.

Iron (Fe) plays a central role in the vital processes of a plant. The Fe status of a plant influences growth and immunity, but it also dictates interactions of roots with soil microbiota through the production of Fe mobilizing, antimicrobial fluorescent phenolic compounds called coumarins. To adapt to low Fe availability in the soil, plants deploy an efficient Fe deficiency response. Interestingly, this Fe deficiency response is hijacked by root-colonizing microbes in the root microbiome to establish a mutually beneficial relationship. In this chapter, we describe how we cultivate plants and microbes to study the interaction between plants, beneficial rhizobacteria, and the plant's Fe deficiency response. We describe (a) how we study activity and localization of these responses by assessing gene-specific promoter activities using GUS assays, (b) how we visualize root-secreted coumarins in response to Fe deficiency and colonization by beneficial rhizobacteria, and (c) how we prepare our samples for metabolite extraction and reverse-transcriptase quantitative PCR to analyze the expression of marker genes.

The interactions of iron nutrition, salinity and ultraviolet-B radiation on the physiological responses of wheat (Triticum aestivum L.)

Increasing human population, land degradation and global change is intensifying pressure on agricultural production in marginal environments. Many marginal environments experience multiple stressors, however, few studies have investigated the interaction of more than two stressors. This is the first study examining the interaction of Fe nutrition, salinity and ultraviolet-B (UV-B) radiation in plants. Wheat (Triticum aestivum L. cv. 1862) was grown in a three-way factorial design: iron (75 µM Fe and 3.75 µM Fe as FeCl3), salinity (0 mM and 75 mM NaCl) and UV-B radiation (1 kJ m−2 d−1, and 15 kJ m−2 d−1 biologically effective UV-B) in chelator-buffered solutions. Examination of the stress interactions showed that phytosiderophore release and root Fe accumulation were limited by UV-B and NaCl stress under Fe deficiency. The findings demonstrated Fe dependency of plant responses to salinity and to UV-B, e.g. sufficient Fe is required for UV-B-induced stomatal opening. Other interactions included that elevated salinity alleviated Fe deficiency-induced decreases in water use efficiency. In addition, Fe deficiency increased the accumulation of UV-absorbing compounds under elevated UV-B radiation. UV-B radiation affected belowground morphology and physiology, including a 47 % reduction in phytosiderophore release. This is the first study to demonstrate a role for wavelengths other than photosynthetically active radiation in phytosiderophore release. Our findings demonstrate that aboveground environmental factors such as UV-B radiation can affect important aspects of belowground plant function, and that Fe availability needs to be considered when growing plants in saline or high UV-B radiation environments.

Impacts of Long-Term Micronutrient Fertilizer Application on Soil Properties and Micronutrient Availability

Deficiencies of micronutrients in calcareous soils have been reported in different areas of China's Loess Plateau. The objective of this research was to study the influence of the continuous application of micronutrient fertilizers on soil properties and micronutrient availability in this region. The micronutrient fertilizer field plot experiment began in 1984. It included Zn, Mn and Cu fertilizer treatments and the control treatment. The crop system was continuously cropped winter wheat. The soil properties and available Zn, Mn, Cu and Fe were measured. Their relationships were analyzed through correlation and path analysis. After 31 years, the soil pH, CaCO3 and available P levels decreased; in contrast, the organic matter, fulvic acid, reducing substances and soil moisture levels in the surface soil increased in the micronutrient fertilized treatments compared to the control treatment. Cu and Zn fertilizers promoted the available Cu and Zn levels in the surface and deep soil, but available Mn was not significantly affected by Mn fertilizer. It can be seen from the interaction between the micronutrient availability and micronutrient fertilizers that Zn, Cu and Mn fertilizers can increase the available Fe level; Mn fertilizer can increase the available Cu level, and Cu and Mn fertilizers can increase the available Zn level. This means that Fe, Cu and Zn availability were easy to implement, whereas the soil-available Mn was difficult to improve in calcareous soils on the Loess Plateau. Fulvic acid and organic matter showed a significant and direct effect on the available Zn; the available Mn and Fe were mainly affected by the soil CaCO3 and moisture; the available Cu was mainly affected by the soil organic matter, available P and reducing substances. These results indicate the importance of organic matter in calcareous soils; it can not only directly affect the availability of micronutrients but also indirectly affect their availability through the indirect interaction with fulvic acid, reducing substances, available P and CaCO3. The above conclusions suggest that the long-term micronutrient fertilizers changed some important soil properties and increased the micronutrient availability in the loess-derived soil.

The low ferric chelate reductase activity and high apoplastic pH in leaves cause iron deficiency chlorosis in ‘Huangguan’ pears grafted onto quince A grown in calcareous soil

Quince (Cydonia oblonga Mill.) is a dwarfing rootstock often used in pear cultivation. However, the pear cultivar ‘Huangguan’ (Pyrus bretschneideri Rehd. cv) grafted onto quince A (HG-QA) suffered iron (Fe) deficiency chlorosis in the calcareous soils of North China, while ‘Huangguan’ grafted onto Pyrus betulifolia (HG-PB) did not. We hypothesized that the higher leaf apoplastic pH of HG-QA than HG-PB causes Fe deficiency chlorosis by restricting Fe transportation within leaves. Three-year-old rootstocks quince A and Pyrus betulifolia and the combinations HG-QA and HG-PB were grown in the same orchard in a calcareous soil. Newly expanded leaves were analysed for Fe related properties. Micro X-ray fluorescence analysis was used for Fe mapping within the individual leaves. The extractable Fe(II) concentrations in quince A and HG-QA leaves were much lower than those in Pyrus betulifolia and HG-PB, even though the total leaf Fe concentrations were similar. More Fe was located near the leaf midrib in quince A and HG-QA than in Pyrus betulifolia and HG-PB. These were correlated with the lower ferric chelate reductase (FCR) activity in the chlorotic leaves of the quince A rootstock than Pyrus betulifolia and the lower FCR activity and higher apoplastic pH in the chlorotic leaves of the HG-QA rootstock-scion combination than HG-PB. In conclusion, the reduced Fe availability within the leaves caused by the lower FCR activity and higher apoplastic pH compared with those of HG-PB led to HG-QA leaf chlorosis in calcareous soils. This result implies that rootstocks are involved in the regulation of leaf Fe utilization by scions.

Physical and biogeochemical responses in the Southern Ocean to a simple parameterization of Langmuir circulation

Langmuir circulation (LC) plays an important role in deepening the mixed layer, especially when the ocean is weakly stratified. LC parameterization has been known to improve the accuracy of the mixed-layer depth, sea surface temperature, and ocean ventilation in the Southern Ocean. However, changes in ocean dynamics and biogeochemical processes owing to LC mixing have rarely been investigated. In this study, we implemented LC parameterization to a physical–biogeochemical coupled model and examined the changes in water properties and circulation and their effects on biogeochemical processes in the Southern Ocean. The LC effect enhances the turbulent mixing length scale and turbulent kinetic energy, especially in the subantarctic region, resulting in the deepening of the mixed layer. Increased vertical mixing causes deeper warm and saline water to reach the surface layer and weakens the surface meridional velocity by transferring momentum deeper. The weakening of the meridional velocity decreases equatorward freshwater transport, causing the surface water to become saltier and denser north of 50°S adjacent to the formation site of Subantarctic Mode Water, which enhances ocean ventilation eventually. Additionally, the weak equatorward velocity drives the retreat of sea-ice south of 60°S, leading to cooler and fresher water in the southern region of 60°S. Changes in the sea-ice distribution, mixed-layer depth, and dissolved inorganic carbon distribution alter the air-sea CO2 exchange, suggesting that the LC parameterization in the model can ameliorate 10% of the air-sea CO2 exchange. The LC effects on biogeochemical tracers, such as iron (Fe) and dissolved inorganic carbon, are mainly determined by changes in ocean circulation, and the modest improvement in primary productivity is mainly attributed to the changes in the Fe distribution in the high-nutrient, low-chlorophyll region, where Fe availability limits primary production. Although the direct LC effects occur near the surface, the altered meridional circulation spreads the effects to a deeper layer and improves the overall representation of physical, biogeochemical tracers and air-sea CO2 exchange in the Southern Ocean, suggesting that a simple LC parameterization can improve the ocean model.

The bHLH Transcription Factor OsbHLH057 Regulates Iron Homeostasis in Rice

Many basic Helix-Loop-Helix (bHLH) transcription factors precisely regulate the expression of Fe uptake and translocation genes to control iron (Fe) homeostasis, as both Fe deficiency and toxicity impair plant growth and development. In rice, three clade IVc bHLH transcription factors have been characterised as positively regulating Fe-deficiency response genes. However, the function of OsbHLH057, another clade IVc bHLH transcription factor, in regulating Fe homeostasis is unknown. Here, we report that OsbHLH057 is involved in regulating Fe homeostasis in rice. OsbHLH057 was highly expressed in the leaf blades and lowly expressed in the roots; it was mainly expressed in the stele and highly expressed in the lateral roots. In addition, OsbHLH057 was slightly induced by Fe deficiency in the shoots on the first day but was not affected by Fe availability in the roots. OsbHLH057 localised in the nucleus exhibited transcriptional activation activity. Under Fe-sufficient conditions, OsbHLH057 knockout or overexpression lines increased or decreased the shoot Fe concentration and the expression of several Fe homeostasis-related genes, respectively. Under Fe-deficient conditions, plants with an OsbHLH057 mutation showed susceptibility to Fe deficiency and accumulated lower Fe concentrations in the shoot compared with the wild type. Unexpectedly, the OsbHLH057-overexpressing lines had reduced tolerance to Fe deficiency. These results indicate that OsbHLH057 plays a positive role in regulating Fe homeostasis, at least under Fe-sufficient conditions.

Early diagenetic processes in an iron-dominated marine depositional system

The early diagenetic interplay between reactive iron, sulfur, and organic matter in the bathymetrically isolated Santa Monica Basin (SMB) sediments are investigated in this study. We explore solid-phase and porewater profiles from the basin, supplemented with a transect from 71 to 907 m water depth that includes oxygenated (>60 μM O2) bottom waters near the coast and oxygen-deficient waters (∼4 µM O2) in the basin. The geochemical data of the basin sediments are further scrutinized by means of reactive transport modeling.The results show that the basin sediments do not follow the traditional geochemical signatures of oxygen-deficient settings. A lack of dissolved sulfide accumulation and sulfurized iron persists despite the sediments being deposited under reducing conditions (without bioturbation/bioirrigation), strong organic carbon input (TOC up to 5.0 wt%), and active dissimilatory sulfate reduction. Not only did we find an exceptional enrichment in highly reactive Fe in the surface sediments (∼45 % of total Fe), but the enrichment of reactive Fe, including ferrihydrite, persists downcore and coexists with high levels of dissolved Fe. The enhanced preservation of Fe oxides and lack of iron-sulfide precipitation is in part explained by detection via Mössbauer spectra of iron oxides bounded to organic matter (Fe[III]-OM coprecipitates).The modeled Fe budget shows that most of the Fe oxides in the surface sediments are internally recycled by upward diffusion and subsequent oxidation of Fe2+. Sulfide oxidation coupled to Fe reduction effectively precludes sulfide accumulation while enhancing build-up of dissolved Fe, fueling the Fe cycle within the first 5 cm depth. Continuous reoxidation of Fe2+ enhances the formation of Fe(III)-OM coprecipitates, limiting the amount of reactive organic matter. In the unavailability of labile organic matter, other than within the uppermost layers, the organic-rich sediment profiles are dominated by Fe cycling that limits the production and preservation of sulfides and enhances the preservation of Fe oxides and organic carbon. This study highlights key local controls on Fe availability in marginal basins and describes an intricate biogeochemical C-Fe-S cycling in modern and possibly ancient marine systems with important implications for Fe availability in the marine realm.

Nitric oxide acts as an inducer of Strategy-I responses to increase Fe availability and mobilization in Fe-starved broccoli (Brassica oleracea var. oleracea)

Iron (Fe) deficiency causes reduced growth and yield in broccoli. This study elucidates how sodium nitroprusside (SNP), known as nitric oxide (NO) donor, mitigates the retardation caused by Fe deficiency in broccoli. The SNP caused substantial nitric oxide accumulation in the roots of Fe-deficient plants, which resulted in a significant improvement in chlorophyll levels, photosynthetic efficiency, and morphological growth parameters, showing that it has a favorable influence on recovering broccoli health. Ferric reductase activity and the expression of BoFRO1 (ferric chelate reductase) gene in roots were consistently increased by SNP under Fe deficiency, which likely resulted in increased Fe mobilization. Furthermore, proton (H+) extrusion and BoHA2 (H+-ATPase 2) expression were significantly increased, suggesting that they may be involved in lowering rhizospheric pH to restore Fe mobilization in roots of bicarbonate-treated broccoli plants. The levels of Fe in root and shoot tissues and the expression of BoIRT1 (Fe-regulated transporter) both increased dramatically after SNP supplementation under Fe deprivation. Furthermore, SNP-induced increase in citrate and malate concentrations suggested a role of NO in improved Fe chelation in Fe-deficient broccoli. A NO scavenger (cPTIO) ceased the elevated FCR activity and IAA (indole-3-acetic acid) concentration in Fe-starved plants treated with SNP. These findings suggest that SNP may play a role in initiating Fe availability by elevated IAA concentration and BoEIR1 (auxin efflux carrier) expression in the roots of broccoli during Fe shortage. Therefore, SNP may improve Fe availability and mobilization by increasing Strategy-I Fe uptake pathways, which may help broccoli tolerate Fe deficiency.