Effect of zinc deficiency in wheat on the release of zinc and iron mobilizing root exudates

The effect of Zn deficiency in wheat (Triticum aestivum L. cv. Ares) on the release of Zn mobilizing root exudates was studied in nutrient solution. Compared to Zn sufficient plants, Zn deficient plants had higher root and lower shoot dry weights. After visual Zn deficiency symptoms in leaves appeared (15–17 day old plants) there was a severalfold increase in the release of root exudates efficient at mobilizing Zn from either a selective cation exchanger (Zn-chelite) or a calcareous soil. The release of these root exudates by Zn deficient plants followed a distinct diurnal rhythm with a maximum between 2 and 8 h after the onset of light. Re-supply of Zn to deficient plants depressed the release of Zn mobilizing root exudates within 12 h to about 50%-, and after 72 h to the level of the control plants (Zn sufficient plants). The root exudates of Zn deficient wheat plants were equally effective at mobilizing Fe from freshly precipitated Fe^III hydroxide as Zn from Zn-chelite. Furthermore, root exudates from Fe deficient wheat plants mobilized Zn from Zn-chelite, as well as Fe from Fe^III hydroxide. Purification of the root exudates and identification by HPLC indicated that under Zn as well as under Fe deficiency, wheat roots of the cv. Ares released the phytosiderophore 2′-deoxymugineic acid. Additional experiments with barley (Hordeum vulgare L. cv. Europa) showed that in this species another phytosiderophore (epi-3-hydroxymugineic acid) was released under both Zn and Fe deficiencies. These results demonstrate that the enhanced release of phytosiderophores by roots of grasses is not a response mechanism specific for Fe deficiency, but also occurs under Zn deficiency. The ecological relevance of enhanced release of phytosiderophore also under Zn deficiency is discussed.     

Responses of Wheat Genotypes to Zinc Fertilization Under Saline Soil Conditions

ABSTRACT

A greenhouse experiment with four bread wheat [Triticum aestivum L.] genotypes, ‘Rushan,’ ‘Kavir,’ ‘Cross,’ and ‘Falat,’ and a durum wheat [Triticum durum L.] genotype, ‘Dur-3,’ at two zinc (Zn) rates (0 and 15 mg Zn kg^?1 dry soil) and four salinity levels (0, 60, 120, and 180 mM NaCl) was conducted. After 45 d of growth, the shoots were harvested, and Zn, iron (Fe), potassium (K), sodium (Na), and cadmium (Cd) concentrations were determined. In the absence of added Zn, visual Zn deficiency symptoms were observed to be more severe in ‘Dur-3’ and ‘Kavir’ than in other genotypes. The effect of Zn deficiency on shoot dry matter was similar to its effect on visual deficiency symptoms, such that shoot growth was most depressed in ‘Kavir’ and ‘Dur-3.’ At the 180 mM treatment, Zn fertilization had no effect on shoot dry matter of genotypes. Genotypes with high Zn efficiency had greater shoot Zn content than genotypes with low Zn efficiency. In the absence of added Zn, the Dur-3, and ‘Cross’ genotypes had the highest and lowest Cd concentrations, respectively. Application of Zn had a positive effect on salt tolerance of plants.     

Effects of iron nutritional status and time of day on concentrations of phytosiderophores and nico‐tianamine in different root and shoot zones of barley

The diurnal pattern in concentrations of phytosiderophores (PS) and its precursor nicotianamine (NA) was studied in different root and shoot zones of iron (Fe)‐sufficient and Fe‐deficient barley (Hordeum vulgare L. cv. Europa) grown in nutrient solution. Roots were separated into apical (0–3 cm) and basal zones (>3 cm) and shoots into young (3 cm basal zones of youngest two leaves) and old (remaining zones of youngest two leaves and oldest leaf) parts. The main PS in barley was identified as epi‐hydroxymugineic acid (epi‐HMA). Regardless of the sampling zone and time of day, epi‐HMA concentrations were several times higher in Fe‐deficient than in Fe‐sufficient plants and several times higher in the roots than in the shoots. In roots and shoots, epi‐HMA concentrations were always higher in the younger compared with the older zones. In both root zones of Fe‐deficient plants, an inverse diurnal rhythm occurred in epi‐HMA concentrations and in its release by the roots. In contrast, such a rhythm was absent in roots of Fe‐sufficient plants and in the shoots regardless of the Fe nutritional status. Nicotianamine concentrations in roots were not affected by the Fe nutritional status in apical zones but slightly enhanced under Fe deficiency in basal zones. In contrast to roots, NA concentrations in both shoot parts were lower in Fe‐deficient than in Fe‐sufficient plants. Regardless of the Fe nutritional status in roots and shoots, NA concentrations were higher in young than in old parts and no consistent diurnal variations were observed. The results suggest that PS are also synthesized in the shoot, although at much lower rates than in roots. As with roots, PS synthesis in the shoot is enhanced under Fe deficiency and is mainly localized in young growing tissue. The distinct diurnal rhythm in PS release in roots is apparently not regulated by variation in the rate of PS synthesis during the day.     

Iron deficiency‐induced zinc uptake by bread wheat

The present study aimed to test the contribution of the iron (Fe) deficiency‐induced uptake system to zinc (Zn) and copper (Cu) uptake by using bread wheat (Triticum aestivum cv. Bezostaja). For this purpose, two different uptake experiments, long‐term and short‐term, were set up in a nutrient solution culture under controlled growth chamber conditions. For the long‐term experiment, wheat cv. plants were grown with different concentrations of Fe or Zn. Results show that there was an uptake system induced under Fe‐limiting conditions which also contributed to Zn and Cu uptake. However, the Zn deficiency‐induced uptake mechanism affected neither Fe nor Cu uptake by wheat. Short‐term uptake experiments indicate that Fe deficiency‐induced Zn²⁺ uptake was more enhanced than the absorption of Zn‐phytosiderophore (PS) complexes. In addition, the Fe‐deficient plants absorbed more Zn in comparison to those plants supplied with sufficient Fe. Similar tendencies in Zn uptake under Fe deficiency in both short‐ and long‐term experiments suggest that there may be a specific Fe uptake system induced under Fe‐limiting conditions for non‐chelated metals in bread wheat. Moreover, this system also contributes to the transport of inorganic forms of some other metals, such as Zn and Cu. Although evidence is still needed involving the use of molecular biological techniques, it is hypothesized that IRT‐like proteins are responsible for this uptake system. Moreover, the release of Fe deficiency‐induced phytosiderophores and uptake of Fe(III)‐phytosiderophore complexes may not be the only mechanisms involved in the adaptation of wheat to Fe‐limiting conditions.     

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

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

Varietal tolerance to Zinc deficiency in wheat and barley grown in chelatorbuffered nutrient solution and its effect on uptake of Cu,Fe, and Mn

Tolerance to zinc (Zn) deficiency was examined for three wheat (Triticum aestivum L.) and three barley (Hordeum vulgare L.) varieties grown in chelator‐buffered nutrient solution. Four indices were chosen to characterize tolerance to Zn deficiency: (1) relative shoot weight at low compared to high Zn supply (“Zn efficiency index”), (2) relative shoot to root ratio at low compared to high Zn supply, (3) total shoot uptake of Zn under deficient conditions, and (4) shoot dry weight under deficient conditions. Barley and wheat exhibited different tolerance to Zn deficiency, with barley being consistently more tolerant than wheat as assessed by all four indices. The tolerance to Zn deficiency in the barley varieties was in the order Thule=Tyra>Kinnan, and that of wheat in the order Bastian=Avle>Vinjett. The less tolerant varieties of both species accumulated more P in the shoots than the more tolerant varieties. For all varieties, the concentrations of Mn, Fe, Cu, and P in shoot tissue were negatively correlated with Zn supply. This antagonism was more pronounced for Mn and P than for Cu and Fe. Accumulation of Cu in barley roots was extremely high under Zn‐deficient conditions, an effect not so clearly indicated in wheat.     

Zinc nutrition effect on the tolerance of wheat genotypes to Fusarium root-rot disease in a solution culture experiment

Abstract

Zinc (Zn) nutrition and plant genotype are two factors that may affect the tolerance of wheat to root-rot diseases. The aim of the present study was to determine the effect of Zn on shoot yield, root permeability and infection by Fusarium solani in six wheat genotypes with different Zn efficiency. A greenhouse (solution culture) experiment was carried out in which five bread wheat genotypes (Triticum aestivum L. cvs Rushan, Kavir, Cross, Pishtaz and Falat) and one durum wheat genotype (Triticum durum L. cv. Yav79), which are common in Zn-deficient soils of Iran and were exposed to two levels of Zn (0 and 1?μmol?L^–1?Zn?kg^?1, as ZnSO₄.7H₂O) and two F. solani infection levels (0 and 10⁶?spore?mL^?1). Zinc deficiency significantly decreased shoot dry matter in five of the genotypes (Yav79, Kavir, Rushan, Cross and Falat), but had no effect on shoot growth in Pishtaz. Infection with F. solani significantly decreased the shoot dry matter in Yav79, but did not affect the shoot dry weight of the other wheat genotypes. Root membrane permeability was lower in the Zn treatments than in the Zn-free treatments. Zinc deficiency caused a decrease in root reactive sulfhydryl (SH) groups, particularly in the Cross genotype. Root sulfhydryl groups decreased with Fusarium infection. Zinc application sharply increased the Zn content and decreased the Mn content of the shoots. Application of Zn had a positive effect on the tolerance of wheat to F. solani root rot. The relationship between Zn nutrition and disease tolerance suggests that Zn deficiency should be treated before evaluating the cost-effectiveness of fungicides. No correlation was found between the Zn efficiency of the wheat genotypes and Fusarium root-rot disease severity in this solution culture experiment.     

Phytosiderophore release from manganese‐induced iron deficiency in barley

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

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

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

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

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

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

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

Effects of zinc fertilization and irrigation on grain yield and zinc concentration of various cereals grown in zinc‐deficient calcareous soils

Effects of varied irrigation and zinc (Zn) fertilization (0, 7, 14, 21 kg Zn ha^‐1 as ZnSO₄7.H₂O) on grain yield and concentration and content of Zn were studied in two bread wheat (Triticum aestivum), two durum wheat (Triticum durum), two barley (Hordeum vulgare), two triticale (xTriticosecale Wittmark), one rye (Secale cereale), and one oat (Avena sativa) cultivars grown in a Zn‐deficient soil (DTPA‐extractable Zn: 0.09 mg kg^‐1) under rainfed and irrigated field conditions. Only minor or no yield reduction occurred in rye as a result of Zn deficiency. The highest reduction in plant growth and grain yield due to Zn deficiency was observed in durum wheats, followed by oat, barley, bread wheat and triticale. These decreases in yield due to Zn deficiency became more pronounced under rainfed conditions. Although highly significant differences in grain yield were found between treatments with and without Zn, no significant difference was obtained between the Zn doses applied (7–21 kg ha^‐1), indicating that 7 kg Zn ha^‐1 would be sufficient to overcome Zn deficiency. Increasing doses of Zn application resulted in significant increases in concentration and content of Zn in shoot and grain. The sensitivity of various cereals to Zn deficiency was different and closely related to Zn content in the shoot but not to Zn amount per unit dry weight. Irrigation was effective in increasing both shoot Zn content and Zn efficiency of cultivars. The results demonstrate the existence of a large genotypic variation in Zn efficiency among and within cereals and suggest that plants become more sensitive to Zn deficiency under rainfed than irrigated conditions.     

Phytosiderophore release in relation to multiple micronutrient metal deficiency in wheat

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

Alien cytoplasm effects on phytosiderophore release in two spring wheats (Triticum aestivum L.)

Iron (Fe) deficiency is a difficult nutrient problem particularly in crop plants grown on calcareous soils. Recently, phytosiderophore (PS) release has been linked to the ability of graminaceous species and genotypes to withstand Fe-deficiency chlorosis. So enhancing PS release is a critical step to improve iron efficiency of plants grown on iron stressed soils. The effects of alien cytoplasm on PS release in spring wheat were studied by analyzing PS release from twenty wheat genotypes, including two spring wheat 881 and 352-35, and their 18 alloplasmic lines with the participation of cytoplasms from the Aegilops and Triticum species. Different genotypes were grown in iron sufficient and deficient nutrient solution under controlled environmental conditions. PS release rates were determined at two or three days intervals after onset of iron deficiency symptoms by the measurement of iron mobilizing capacity of root exudates from freshly precipitated Fe^III hydroxide. High amounts of phytosiderophores were released from roots of all wheat genotypes without iron supplied, and the amount progressively increased with the development of iron deficiency chlorosis. The results revealed that (1) the release rate of phytosiderophores from roots of common wheat could be considerably influenced by alien cytoplasms. Some alien cytoplasms exerted positive effects, some ones did negative effects, and the other ones had no significant effects. (2) the same alien cytoplasm could affect similarly or oppositely the phytosiderophores release from different wheat. (3) some alien cytoplasms, such as Chinese Spring, Ae. speltoides Tausch and Ae. cylindrica Host showed promising and potential in improving the rate of phytosiderophore release in common wheat. These cytoplasms which showed the desired effect should be given priority in interspecific and intergeneric hybridization to develop and reconstruct the needed wheat cultivars.     

Root exudation of sugars,amino acids,and organic acids by maize as affected by nitrogen,phosphorus, potassium,and iron deficiency

Root exudates play a major role in the mobilization of sparingly soluble nutrients in the rhizosphere. Since the amount and composition of major metabolites in root exudates from one plant species have not yet been systematically compared under different nutrient deficiencies, relations between exudation patterns and the type of nutrient being deficient remain poorly understood. Comparing root exudates from axenically grown maize plants exposed to N, K, P, or Fe deficiency showed a higher release of glutamate, glucose, ribitol, and citrate from Fe‐deficient plants, while P deficiency stimulated the release of γ‐aminobutyric acid and carbohydrates. Potassium‐starved plants released less sugars, in particular glycerol, ribitol, fructose, and maltose, while under N deficiency lower amounts of amino acids were found in root exudates. Principal‐component analysis revealed a clear separation in the variation of the root‐exudate composition between Fe or P deficiency versus N or K deficiency in the first principal component, which explained 46% of the variation in the data. In addition, a negative correlation was found between the amounts of sugars, organic and amino acids released under deficiency of a certain nutrient and the diffusion coefficient of the respective nutrient in soils. We thus hypothesize that the release of dominant root exudates such as sugars, amino acids, and organic acids by roots may reflect an ancient strategy to cope with limiting nutrient supply.     

The role of root plasma membrane ATPase and rhizosphere acidification in zinc uptake by two different Zn-deficiency-tolerant wheat cultivars in response to zinc and histidine availability

ABSTRACT

We investigated the effect of histidine (His) and Zn deficiency on H⁺-ATPase activity and H⁺ release from wheat roots. Two bread wheat (Triticum aestivum L. cvs. Kavir and Back Cross Roshan) were grown in a nutrient solution for four weeks before being transferred to treatment solutions consisting of two concentrations of His (0 and 50 µM) and two concentrations of Zn (0 and 10 μM). The Zn-only and the Zn+His treatments were observed to release more H⁺ in the root media than did the control ones, with the highest achieved under the Zn+His treatment which was roughly 2.1 times higher than that under the control conditions. The H⁺ release from wheat roots increased slightly but significantly in the presence of only His when compared with the control solutions. The hydrolytic and transport activities of H⁺-ATPase were affected by both Zn deficiency and His supply. In both cultivars, application of Zn and His resulted in a higher hydrolytic activity of H⁺-ATPase when compared with the control solutions. The highest hydrolytic activity of H⁺-ATPase in the root plasma membrane vesicles was achieved with the Zn+His treatment. The ‘Back Cross Roshan’ exhibited a higher (PM) H⁺-ATPase activity and H⁺ pumping than did ‘Kavir’.     

Differences in shoot growth and zinc concentration of 164 bread wheat genotypes in a zinc‐deficient calcareous soil

Abstract

A greenhouse experiment was carried out to study severity of the zinc (Zn) deficiency symptoms on leaves, shoot dry weight and shoot content and concentration of Zn in 164 winter type bread wheat genotypes (Triticunt aestivum L.) grown in a Zn‐deficient calcareous soil with (+Zn=10 mg Zn kg^?1 soil) and without (‐Zn) Zn supply for 45 days. Tolerance of the genotypes to Zn deficiency was ranked based on the relative shoot growth (Zn efficiency ratio), calculated as the ratio of the shoot dry weight produced under Zn deficiency to that produced under adequate Zn supply. There was a substantial difference in genotypic tolerance to Zn deficiency. Among the 164 genotypes, 108 genotypes had severe visible symptoms of Zn deficiency (whitish‐brown necrotic patches) on leaves, while in 25 genotypes Zn deficiency symptoms were slight or absent, and the remaining genotypes (e.g., 31 genotypes) showed mild deficiency symptoms. Generally, the genotypes with higher tolerance to Zn deficiency originated from Balkan countries and Turkey, while genotypes originating from the breeding programs in the Great Plains of the United States were mostly sensitive to Zn deficiency. Among the 164 wheat genotypes, Zn efficiency ratio varied from 0.33 to 0.77. The differences in tolerance to Zn deficiency were totally independent of shoot Zn concentrations, but showed a close relationship to the total amount (content) of Zn per shoot. The absolute shoot growth of the genotypes under Zn deficiency corresponded very well with the differences in tolerance to Zn deficiency. Under adequate Zn supply, the 10 most Zn‐ inefficient genotypes and the 10 most Zn‐efficient genotypes were very similar in their shoot dry weight. However, under Zn deficiency, shoot dry weight of the Zn‐efficient genotypes was, on average, 1.6‐fold higher compared to the Zn‐inefficient genotypes. The results of this study show large, exploitable genotypic variation for tolerance to Zn deficiency in bread wheat. Based on this data, total amount of Zn per shoot, absolute shoot growth under Zn deficiency, and relative shoot growth can be used as reliable plant parameters for assessing genotypic variation in tolerance to Zn deficiency in bread wheat.     

Influence of Gyttja on Shoot Growth and Shoot Concentrations of Zinc and Boron of Wheat Cultivars Grown on Zinc‐Deficient and Boron‐Toxic Soil

Abstract

Greenhouse experiments were carried out to study the influence of gyttja, a sedimentary peat, on the shoot dry weight and shoot concentrations of zinc (Zn) and boron (B) in one bread wheat (Triticum aestivum L., cv. Bezostaja) and one durum wheat (Triticum durum L., cv. Kiziltan) cultivar. Plants were grown in a Zn‐deficient (DTPA‐Zn: 0.09 mg kg^?1 soil) and B‐toxic soil (CaCl₂/mannitol‐extractable B: 10.5 mg kg^?1 soil) with (+Zn = 5 mg Zn kg^?1 soil) and without (?Zn = 0) Zn supply for 55 days. Gyttja containing 545 g kg^?1 organic matter was applied to the soil at the rates of 0, 1, 2.5, 5, and 10% (w/w). When Zn and gyttja were not added, plants showed leaf symptoms of Zn deficiency and B toxicity, and had a reduced growth. With increased rates of gyttja application, shoot growth of both cultivars was significantly enhanced under Zn deficiency, but not at sufficient supply of Zn. The adverse effects of Zn deficiency and B toxicity on shoot dry matter production became very minimal at the highest rate of gyttja application. Increases in gyttja application significantly enhanced shoot concentrations of Zn in plants grown without addition of inorganic Zn. In Zn‐sufficient plants, the gyttja application up to 5% (w/w) did not affect Zn concentration in shoots, but at the highest rate of gyttja application there was a clear decrease in shoot Zn concentration. Irrespective of Zn supply, the gyttja application strongly decreased shoot concentration of B in plants, particularly in durum wheat. For example, in Zn‐deficient Kiziltan shoot concentration of B was reduced from 385 mg kg^?1 to 214 mg kg^?1 with an increased gyttja application. The results obtained indicate that gyttja is a useful organic material improving Zn nutrition of plants in Zn‐deficient soils and alleviating adverse effects of B toxicity on plant growth. The beneficial effects of gyttja on plant growth in the Zn‐deficient and B‐toxic soil were discussed in terms of increases in plant available concentration of Zn in soil and reduction of B uptake due to formation of tightly bound complexes of B with gyttja.     

Metal concentrations,growth, and yield of potato produced from In Vitro plantlets or microtubers and grown in municipal solid‐waste‐amended substrates

In vitro plantlets or microtubers (in vitro produced tubers) of ‘Spunta’ potato (Solanum tuberosum L.) were planted in a 3 soil: 2 peat moss: 1 sand substrate (by volume) amended with municipal solid waste (MS W) compost at 0, 10, 20, or 30 g 4^‐1 L pot. Three months later, plant growth and tuber yield were evaluated and concentrations of shoot and tuber tin (Sn), arsenic (As), copper (Cu), zinc (Zn), nickel (Ni), lead (Pb), manganese (Mn), cadmium (Cd), and iron (Fe) were determined. Amending with MSW resulted in significant increases in concentrations of all tested metals in the substrate. Number of proliferated shoots of plants started from rooted plantlets was greatest at 10 g pot^‐1 MSW, whereas shoot weight of plants started from microtubers was greatest at 10 and 20 g pot^‐1 MSW. Tuber yield of plants started from rooted plantlets or microtubers was greatest at 10 or 30 g pot^‐1 MSW, respectively. In all instances, amending with MSW at 30 g pot^‐1 resulted in significant increases in concentrations of all tested metals in shoots and tubers. Concentrations of shoot Ni and tuber Zn and Fe for plants started from rooted plantlets and concentrations of shoot Fe and tuber As, Cu and Pb for plants started from microtubers increased consistently with increasing MSW percentage of the substrate. Plants started from rooted plantlets produced shoots with sufficient Zn, Mn, and Ni concentrations regardless of the substrate but with toxic Cu content at 30 g pot^‐1 MSW. Plants started from microtubers produced shoots with sufficient Mn and Ni concentrations regardless of the substrate but with low Zn and deficient Cu in unamended substrates. All plants had shoot Fe content higher than the sufficiency range. Although there were significant differences in concentrations of some nutrients among MSW treatments, no symptoms of nutrient toxicity or deficiency were observed. In all instances, tested elements did not accumulate in tubers to levels hazardous to human health. Concentrations of Cd, the most hazardous element, in potato tubers was not high enough to pose a threat to human. Our results indicate that there is a potential use of MSW in satisfying the needs of potato growth with negligible increases in heavy metal concentrations in tubers.     

Zinc‐biofortified seeds improved seedling growth under zinc deficiency and drought stress in durum wheat

High zinc (Zn) concentration of seeds has beneficial effects both on seed vigor and human nutrition. This study investigated the effect of Zn biofortification on growth of young durum wheat (Triticum durum cv. Yelken) seedlings under varied Zn and water supply. The seeds differing in Zn concentrations were obtained by spraying ZnSO₄ to durum wheat plants at different rates under field conditions. Three groups of seeds were obtained with the following Zn concentrations: 9, 20, and 50 mg Zn kg^?1. The seeds differing in Zn were tested for germination rate, seedling height, shoot dry matter production, and shoot Zn concentration under limited and well irrigated conditions in a Zn‐deficient soil with and without Zn application. In an additional experiment carried out in solution culture, root and shoot growth and superoxide dismutase activity (SOD) of seedlings were studied under low and adequate Zn supply. Low seed Zn concentration resulted in significant decreases in seedling height both in Zn‐deficient and sufficient soil, but more clearly under water‐limited soil condition. Decrease in seed germination due to low seed Zn was also more evident under limited water supply. Increasing seed Zn concentration significantly restored impairments in seedling development. Drought‐induced decrease in seedling growth at a given seed Zn concentration was much higher when soil was Zn‐deficient. Increasing seed Zn concentration also significantly improved SOD activity in seedlings grown under low Zn supply, but not under adequate Zn supply. The results suggest that using Zn‐biofortified seeds assures better seed vigor and seedling growth, particularly when Zn and water are limited in the growth medium. The role of a higher antioxidative potential (i.e., higher SOD activity) is discussed as a possible major factor in better germination and development of seedlings resulting from Zn‐biofortified seeds.