Bicarbonate rather than high pH in growth medium induced Fe-deficiency chlorosis in dwarfing rootstock quince A (Cydonia oblonga Mill.) but did not impair Fe nutrition of vigorous rootstock Pyrus betulifolia.

Introduction: Quince A (Cydonia oblonga Mill.), a typical dwarfing rootstock in pear cultivation, is susceptible to iron (Fe) deficiency in calcareous soils. The aim of this study was to compare the strategies in Fe uptake and utilization in dwarfing rootstock quince A (low Fe efficiency) versus a typical vigorous rootstock Pyrus betulifolia (PB) with high Fe efficiency. Methods: Quince A and PB were grown in nutrient solution (pH 6.3) for 4 weeks followed by three pH treatments: pH6.3, pH8.3a (adjusted with hydroxide) and pH8.3b (adjusted with bicarbonate). The Fe uptake and utilization indicators of the rootstocks were assessed at the onset of chlorosis symptoms (after 58 days of treatments). Results and discussion: In contrast to PB, quince A exhibited Fe deficiency chlorosis under bicarbonate (pH8.3b). Bicarbonate stimulated the root proton secretion, inhibited root growth and ferric chelate reductase (FCR) activity in both PB and quince A, whereas high pH without bicarbonate (pH8.3a) stimulated only root proton release. Both species accumulated more Fe in roots under high pH treatments than under pH6.3, resulting in Fe sufficiency in leaves. Both high pH treatments increased the activity of leaf FCR in PB and quince A. However, extractable Fe(II) concentration in leaves was increased by high pH treatments in PB only. This study demonstrated that depressed Fe(III) reduction in leaves caused by bicarbonate rather than high pH explained Fe deficiency in quince A grown in bicarbonate-containing medium.

Soil, leaf and fruit nutrient data for pear orchards located in the Circum-Bohai Bay and Loess Plateau regions.

The data described in this paper were collected from the Circum-Bohai Bay and Loess Plateau regions of northern China. Soil, leaf and fruit nutrients from 225 typical pear orchards in these regions were measured. Soil data included pH, organic matter, total N, alkaline hydrolysable N, available P and available K concentrations of 3 different soil layers, 0-20 cm, 20-40 cm and 40-60 cm, from different orchards. Leaf and fruit data included N, P, K, Ca, Fe, Mn, Cu, Zn and B concentrations of pear trees from different orchards. These data can be used to assess the soil nutrient supply and leaf and fruit nutrient status of pear orchards in two major producing areas, Circum-Bohai Bay and Loess Plateau. Additionally, this dataset provides data to support the development of regionalized and standardized soil nutrient management programs for pear orchards, as well as regionalized layouts of the main varieties in the two producing areas.

Genome-Wide Association Analysis Reveals the Genetic Basis of Iron-Deficiency Stress Tolerance in Maize.

Iron (Fe) is an essential trace element for almost all organisms and is often the major limiting nutrient for normal growth. Fe deficiency is a worldwide agricultural problem, which affects crop productivity and product quality. Understanding the Fe-deficiency response in plants is necessary for improving both plant health and the human diet. In this study, Fe-efficient (Ye478) and Fe-inefficient maize inbred lines (Wu312) were used to identify the genotypic difference in response to low Fe stress during different developmental stages and to further determine the optimal Fe-deficient Fe(II) supply level which leads to the largest phenotypic difference between Ye478 and Wu312. Then, genome-wide association analysis was performed to further identify candidate genes associated with the molecular mechanisms under different Fe nutritional statuses. Three candidate genes involved in Fe homeostasis of strategy II plants (strategy II genes) were identified, including ZmDMAS1, ZmNAAT1, and ZmYSL11. Furthermore, candidate genes ZmNAAT1, ZmDMAS1, and ZmYSL11 were induced in Fe-deficient roots and shoots, and the expression of ZmNAAT1 and ZmDMAS1 responded to Fe deficiency more in shoots than in roots. Beyond that, several genes that may participate in Fe homeostasis of strategy I plants (strategy I genes) were identified, which were either encoding Fe transporters (ZmIRT1 and ZmZIP4), or acting as essential ethylene signal transducers (ZmEBF1). Interestingly, ZmIRT1, ZmZIP4, and ZmEBF1 were significantly upregulated under low Fe stress, suggesting that these genes may be involved in Fe-deficiency tolerance in maize which is considered as strategy II plant. This study demonstrates the use of natural variation in the association population to identify important genes associated with Fe-deficiency tolerance and may further provide insights for understanding the molecular mechanism underlying the tolerance to Fe-deficiency stress in maize.

Identification of Zinc Efficiency-Associated Loci (ZEALs) and Candidate Genes for Zn Deficiency Tolerance of Two Recombination Inbred Line Populations in Maize.

Zinc (Zn) deficiency is one of the most common micronutrient disorders in cereal plants, greatly impairing crop productivity and nutritional quality. Identifying the genes associated with Zn deficiency tolerance is the basis for understanding the genetic mechanism conferring tolerance. In this study, the K22×BY815 and DAN340×K22 recombination inbred line (RIL) populations, which were derived from Zn-inefficient and Zn-efficient inbred lines, were utilized to detect the quantitative trait loci (QTLs) associated with Zn deficiency tolerance and to further identify candidate genes within these loci. The BLUP (Best Linear Unbiased Prediction) values under Zn-deficient condition (-Zn) and the ratios of the BLUP values under Zn deficient condition to the BLUP values under Zn-sufficient condition (-Zn/CK) were used to perform linkage mapping. In QTL analysis, 21 QTLs and 33 QTLs controlling the Zn score, plant height, shoot and root dry weight, and root-to-shoot ratio were detected in the K22×BY815 population and the DAN340×K22 population, explaining 5.5-16.6% and 4.2-23.3% of phenotypic variation, respectively. In addition, seventeen candidate genes associated with the mechanisms underlying Zn deficiency tolerance were identified in QTL colocalizations or the single loci, including the genes involved in the uptake, transport, and redistribution of Zn (ZmIRT1, ZmHMAs, ZmNRAMP6, ZmVIT, ZmNAS3, ZmDMAS1, ZmTOM3), and the genes participating in the auxin and ethylene signal pathways (ZmAFBs, ZmIAA17, ZmETR, ZmEIN2, ZmEIN3, ZmCTR3, ZmEBF1). Our findings will broaden the understanding of the genetic structure of the tolerance to Zn deficiency in maize.

Mapping of the Quantitative Trait Loci and Candidate Genes Associated With Iron Efficiency in Maize.

Iron (Fe) is a mineral micronutrient for plants, and Fe deficiency is a major abiotic stress in crop production because of its low solubility under aerobic and alkaline conditions. In this study, 18 maize inbred lines were used to preliminarily illustrate the physiological mechanism underlying Fe deficiency tolerance. Then biparental linkage analysis was performed to identify the quantitative trait loci (QTLs) and candidate genes associated with Fe deficiency tolerance using the recombinant inbred line (RIL) population derived from the most Fe-efficient (Ye478) and Fe-inefficient (Wu312) inbred lines. A total of 24 QTLs was identified under different Fe nutritional status in the Ye478 × Wu312 RIL population, explaining 6.1-26.6% of phenotypic variation, and ten candidate genes were identified. Plants have evolved two distinct mechanisms to solubilize and transport Fe to acclimate to Fe deficiency, including reduction-based strategy (strategy I) and chelation-based strategy (strategy II), and maize uses strategy II. However, not only genes involved in Fe homeostasis verified in strategy II plants (strategy II genes), which included ZmYS1, ZmYS3, and ZmTOM2, but also several genes associated with Fe homeostasis in strategy I plants (strategy I genes) were identified, including ZmFIT, ZmPYE, ZmILR3, ZmBTS, and ZmEIN2. Furthermore, strategy II gene ZmYS1 and strategy I gene ZmBTS were significantly upregulated in the Fe-deficient roots and shoots of maize inbred lines, and responded to Fe deficiency more in shoots than in roots. Under Fe deficiency, greater upregulations of ZmYS1 and ZmBTS were observed in Fe-efficient parent Ye478, not in Fe-inefficient parent Wu312. Beyond that, ZmEIN2 and ZmILR3, were found to be Fe deficiency-inducible in the shoots. These findings indicate that these candidate genes may be associated with Fe deficiency tolerance in maize. This study demonstrates the use of natural variation to identify important Fe deficiency-regulated genes and provides further insights for understanding the response to Fe deficiency stress in maize.

Identification of Quantitative Trait Loci Associated With Iron Deficiency Tolerance in Maize.

Iron (Fe) is a limiting factor in crop growth and nutritional quality because of its low solubility. However, the current understanding of how major crops respond to Fe deficiency and the genetic basis remains limited. In the present study, Fe-efficient inbred line Ye478 and Fe-inefficient inbred line Wu312 and their recombinant inbred line (RIL) population were utilized to reveal the physiological and genetic responses of maize to low Fe stress. Compared with the Fe-sufficient conditions (+Fe: 200 µM), Fe-deficient supply (-Fe: 30 µM) significantly reduced shoot and root dry weights, leaf SPAD of Fe-efficient inbred line Ye478 by 31.4, 31.8, and 46.0%, respectively; decreased Fe-inefficient inbred line Wu312 by 72.0, 45.1, and 84.1%, respectively. Under Fe deficiency, compared with the supply of calcium nitrate (N1), supplying ammonium nitrate (N2) significantly increased the shoot and root dry weights of Wu312 by 37.5 and 51.6%, respectively; and enhanced Ye478 by 23.9 and 45.1%, respectively. Compared with N1, N2 resulted in a 70.0% decrease of the root Fe concentration for Wu312 in the -Fe treatment, N2 treatment reduced the root Fe concentration of Ye478 by 55.8% in the -Fe treatment. These findings indicated that, compared with only supplying nitrate nitrogen, combined supply of ammonium nitrogen and nitrate nitrogen not only contributed to better growth in maize but also significantly reduced Fe concentration in roots. In linkage analysis, ten quantitative trait loci (QTLs) associated with Fe deficiency tolerance were detected, explaining 6.2-12.0% of phenotypic variation. Candidate genes considered to be associated with the mechanisms underlying Fe deficiency tolerance were identified within a single locus or QTL co-localization, including ZmYS3, ZmPYE, ZmEIL3, ZmMYB153, ZmILR3 and ZmNAS4, which may form a sophisticated network to regulate the uptake, transport and redistribution of Fe. Furthermore, ZmYS3 was highly induced by Fe deficiency in the roots; ZmPYE and ZmEIL3, which may be involved in Fe homeostasis in strategy I plants, were significantly upregulated in the shoots and roots under low Fe stress; ZmMYB153 was Fe-deficiency inducible in the shoots. Our findings will provide a comprehensive insight into the physiological and genetic basis of Fe deficiency tolerance.

Identification and Analysis of Zinc Efficiency-Associated Loci in Maize.

Zinc (Zn) deficiency, a globally predominant micronutrient disorder in crops and humans, reduces crop yields and adversely impacts human health. Despite numerous studies on the physiological mechanisms underlying Zn deficiency tolerance, its genetic basis of molecular mechanism is still poorly understood. Thus, the Zn efficiency of 20 maize inbred lines was evaluated, and a quantitative trait locus (QTL) analysis was performed in the recombination inbred line population derived from the most Zn-efficient (Ye478) and Zn-inefficient inbred line (Wu312) to identify the candidate genes associated with Zn deficiency tolerance. On this basis, we analyzed the expression of ZmZIP1-ZmZIP8. Thirteen QTLs for the traits associated with Zn deficiency tolerance were detected, explaining 7.6-63.5% of the phenotypic variation. The genes responsible for Zn uptake and transport across membranes (ZmZIP3, ZmHMA3, ZmHMA4) were identified, which probably form a sophisticated network to regulate the uptake, translocation, and redistribution of Zn. Additionally, we identified the genes involved in the indole-3-acetic acid (IAA) biosynthesis (ZmIGPS) and auxin-dependent gene regulation (ZmIAA). Notably, a high upregulation of ZmZIP3 was found in the Zn-deficient root of Ye478, but not in that of Wu312. Additionally, ZmZIP4, ZmZIP5, and ZmZIP7 were up-regulated in the Zn-deficient roots of Ye478 and Wu312. Our findings provide a new insight into the genetic basis of Zn deficiency tolerance.

Maize Genotypes With Different Zinc Efficiency in Response to Low Zinc Stress and Heterogeneous Zinc Supply.

All over the world, a common problem in the soil is the low content of available zinc (Zn), which is unevenly distributed and difficult to move. However, information on the foraging strategies of roots in response to heterogeneous Zn supply is still very limited. Few studies have analyzed the adaptability of maize inbred lines with different Zn efficiencies to different low Zn stress time lengths in maize. This study analyzed the effects of different time lengths of low Zn stress on various related traits in different inbred lines. In addition, morphological plasticity of roots and the response of Zn-related important gene iron-regulated transporter-like proteins (ZIPs) were studied via simulating the heterogeneity of Zn nutrition in the soil. In this report, when Zn deficiency stress duration was extended (from 14 to 21 days), under Zn-deficient supply (0.5 µM), Zn efficiency (ZE) based on shoot dry weight of Wu312 displayed no significant difference, and ZE for Ye478 was increased by 92.9%. Under longer-term Zn deficiency, shoot, and root dry weights of Ye478 were 6.5 and 2.1-fold higher than those of Wu312, respectively. Uneven Zn supply strongly inhibited the development of some root traits in the -Zn region. Difference in shoot dry weights between Wu312 and Ye478 was larger in T1 (1.97 times) than in T2 (1.53 times). Under heterogeneous condition of Zn supply, both the -Zn region and the +Zn region upregulated the expressions of ZmZIP3, ZmZIP4, ZmZIP5, ZmZIP7, and ZmZIP8 in the roots of two inbred lines. These results indicate that extended time length of low-Zn stress will enlarge the difference of multiple physiological traits, especially biomass, between Zn-sensitive and Zn-tolerant inbred lines. There were significant genotypic differences of root morphology in response to heterogeneous Zn supply. Compared with split-supply with +Zn/+Zn, the difference of above-ground biomass between Zn-sensitive and Zn-tolerant inbred lines under split-supply with -Zn/+Zn was higher. Under the condition of heterogeneous Zn supply, several ZmZIP genes may play important roles in tolerance to low Zn stress, which can provide a basis for further functional characterization.

GENERAL CONTROL NONREPRESSED PROTEIN5-Mediated Histone Acetylation of FERRIC REDUCTASE DEFECTIVE3 Contributes to Iron Homeostasis in Arabidopsis.

Iron homeostasis is essential for plant growth and development. Here, we report that a mutation in GENERAL CONTROL NONREPRESSED PROTEIN5 (GCN5) impaired iron translocation from the root to the shoot in Arabidopsis (Arabidopsis thaliana). Illumina high-throughput sequencing revealed 879 GCN5-regulated candidate genes potentially involved in iron homeostasis. Chromatin immunoprecipitation assays indicated that five genes (At3G08040, At2G01530, At2G39380, At2G47160, and At4G05200) are direct targets of GCN5 in iron homeostasis regulation. Notably, GCN5-mediated acetylation of histone 3 lysine 9 and histone 3 lysine 14 of FERRIC REDUCTASE DEFECTIVE3 (FRD3) determined the dynamic expression of FRD3. Consistent with the function of FRD3 as a citrate efflux protein, the iron retention defect in gcn5 was rescued and fertility was partly restored by overexpressing FRD3. Moreover, iron retention in gcn5 roots was significantly reduced by the exogenous application of citrate. Collectively, these data suggest that GCN5 plays a critical role in FRD3-mediated iron homeostasis. Our results provide novel insight into the chromatin-based regulation of iron homeostasis in Arabidopsis.