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.     

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.     

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.     

Comparison of Winter Wheat Genotypes Differing in Zinc Efficiency and Origin: The Zinc Uptake and Enzyme Activity

Zinc (Zn)–efficient wheat genotypes yield well on Zn-deficient soil. In this study, two Chinese wheat genotypes, Kenong9204 and Han6172, and two reference genotypes, Bezostaja (Zn efficient) and BDME10 (Zn inefficient), from Turkey were conducted to measure their physiological responses to Zn deficiency in the greenhouse. Results showed obvious genetic variation among the genotypes with Zn efficiency from 76% to 105%. Bezostaja and Kenong9204 had greater shoot dry weight and accumulated more shoot Zn content than BDME10 and Han6172 without Zn application. In one aspect of enzyme activities, Bezostaja and Kenong9204 presented significantly greater activities of superoxide dismutase while maintaining similar activities of catalase, ascorbate peroxidase, and glutathione reductase compared with inefficient genotypes BDME10 and Han6172 under Zn-deficient condition. Zinc-efficient genotypes are recommended to satisfy the sustainable grain yield in China and other areas, where Zn deficiency in soil is spread and multiple stresses may happen at times.     

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.     

Effect of different zinc application methods on grain yield and zinc concentration in wheat cultivars grown on zinc‐deficient calcareous soils

The effect of six different zinc (Zn) application methods on grain yield and concentrations of Zn in whole shoots and grain was studied in wheat cultivars (Triticum aestivum, L. cvs. Gerek‐79, Dagdas‐94 and Bezostaja‐1 and Triticum durum, Desf. cv. Kunduru‐1149) grown on severely Zn‐deficient calcareous soils (DTPA‐extractable Zn: 0.12 mg‐kg^‐1 soil) of Central Anatolia which is the major wheat growing area of Turkey. Zinc application methods tested were: a) control (no Zn application), b) soil, c) seed, d) leaf, e) soil+leaf, and f) seed+leaf applications. Irrespective of the method, application of Zn significantly increased grain yield in all cultivars. Compared to the control, increases in grain yield were about 260% with soil, soil+leaf, and seed+leaf, 204% with seed and 124% with leaf application of Zn. In a similar manner, biomass production (dry weight of above‐ground parts) was increased by Zn treatments. The highest increase (109%) was obtained with the soil application and the lowest increase (40%) with the leaf application. Significant effects of Zn application methods were also found on the yield components, i.e., spike number.m^‐2, grain number‐spike^‐1, and thousand kernel weight. Spike number.m^‐2 was affected most by Zn applications, particularly by soil and soil+leaf applications. Concentrations of Zn in whole shoots and grain were greatly affected by different Zn treatments. In plants without added Zn, concentrations of Zn were about 10 mg‐kg^‐1 both in shoots and grain and increased to 18 mg‐kg^‐1 dry weight (DW) by soil application of Zn, but not affected by seed application of Zn. Soil+leaf application of Zn had the highest increase in concentration of Zn in shoot (82 mg‐kg^‐1 DW) and grain (38 mg‐kg^‐1 DW). Soil application of Zn was economical and had long‐term effects for enhancing grain yield of wheat grown on Zn deficient soils. When high grain yield and high Zn concentration in grains are desired, soil+leaf application of Zn was most effective method of Zn application.     

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.     

Release of phytosiderophores from roots of wheat and triticale under nickel‐deficient conditions

Despite numerous studies on phytosiderophores (PS) there is still an open question whether nickel (Ni) deficiency induces release of PS from graminaceous plant roots. Seedlings of two wheat cultivars (Triticum aestivum L. cvs. Rushan and Kavir) and a triticale cultivar (X. triticosecale) were grown in Ni‐free nutrient solution (Ni‐deficient, Ni–) and with 10 µM NiSO₄ (Ni‐sufficient, Ni+, control). Root exudates were collected weekly for 4 weeks and the amount of PS in the root exudates was measured. The response to Ni deficiency on the release of PS differed between species. Roots of Rushan and triticale exuded higher PS in response to Ni‐deficient conditions. Nickel deficiency significantly enhanced shoot Fe and Zn concentrations in wheat, while it decreased shoot Fe and Zn concentrations in triticale. In Kavir, PS exudation was decreased by Ni deficiency at weeks 3 and 4 and the reduced release of PS from roots of Kavir was accompanied by lower concentrations of Fe and Zn in plant roots but higher Fe and Zn concentrations in shoot tissue. The PS release by Kavir was triggered by a Ni‐induced Zn deficiency particularly in the shoots. According to the results, it is suggested that in the studies concerning the phytosiderophore release under Ni deficiency, special attention should be given to different responses among and within cereals and to the plant Zn or Fe nutritional status.     

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.     

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.     

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.     

Phytosiderophore release as a criterion for genotypic evaluation of iron efficiency in oat

Iron (Fe) deficiency in small grains grown on calcareous soils results in reduced yields, is difficult and expensive to treat with fertilizer, and is complicated to overcome by genetic field screening due to heterogeneous soil and environmental conditions. Recently, phytosiderophore release has been linked to ability of species and genotypes to resist Fe‐deficiency chlorosis. We propose a laboratory technique to measure phytosiderophore release by Fe‐deficient oat (Avena sativa L.) genotypes as a selection method for Fe‐deficiency chlorosis resistance in oat. Plants were grown in Fe‐limiting nutrient solution and phytosiderophore release was measured on 11 days. Summations of daily phytosiderophore release by 17 oat genotypes correlate well with Fe‐deficiency chlorosis scores in the field (r = ‐0.70, p = 0.01). The proposed method consistently identified the genotypes most susceptible to Fe deficiency but did not clearly separate the moderately susceptible genotypes. In these latter genotypes, other factors such as active uptake sites, root growth rate, utilization of acquired Fe, or soil interactions may be modifying factors to phytosiderophore in Fe efficiency. Quantification of phytosiderophore provides a useful selection criterion for oat by eliminating the most inefficient types and with refinement, may become a powerful tool for identifying Fe efficiency in grass crops.     

Iron deficiency stress response of various c‐3 and c‐4 grain crop genotypes: Strategy II mechanism evaluated

The relative amount of phytosiderophore produced by various Strategy II plants has been categorized as follows: barley (Hordeum vulgare L.) > wheat (Triticum aestivum L.) > oat (Avena byzantina C. Koch.) > rye (Secale cereale L.) >> corn (Zea mays L.) >> sorghum (Sorghum bicolor (L.) Moench) > rice (Oryza sativa L.). With the exception of rice, these plants developed under oxidized soil conditions, and the C‐3 species produce more phytosiderophore than C‐4 species under Fe‐deficiency stress. Iron‐efficient Coker 227 oat produced phytosiderophore in response to Fe‐deficiency stress, while Fe‐inefficient TAM 0–312 oat did not. Although Fe‐efficient WF9 corn and Fe‐inefficient ys₁ corn differed in their ability to obtain Fe, neither produced sufficient quantities of phytosiderophore to explain these differences. The objectives of this research were to determine: (a) if phytosiderophore production of Fe‐deficiency stressed C‐4 species millet (Panicum miliaceum L.) and corn is low or absent compared to identically stressed C‐3 species oat and barley, and (b) if native, inbred and hybrid corn cultivars differ in ability to produce and utilize phytosiderophores.

Although release of phytosiderophore for Fe‐stressed corn and millet was generally lower than oat, quantity of release was not always related to obtaining Fe and maintaining green plants. Barley maintained high leaf Fe and low chlorosis with a minor release of phytosiderophore. Oat with increased release acted similarly to barley, whereas a relatively high release of phytosiderophore from White maize did not effect Fe uptake or greening. Likewise, small amounts of phytosiderophore were produced by all corn types, but corn was generally unable to obtain adequate Fe from the growth medium. Two of the native corns, Coneso and Tepecintle, maintained relatively low chlorosis, but they differed in phytosiderophore release. Thus, it appears that the C‐4 plants studied herein generally release a lower amount of phytosiderophore than do C‐3 species, but overcoming Fe‐deficiency chlorosis is not guaranteed by such release. The Strategy II mechanism of mere release of phytosiderophore and consequential Fe acquisition appears simplistic. There is a need for understanding what other factors are involved.     

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.     

Diurnal rhythm of release of phytosiderophores and uptake rate of zinc in iron-deficient wheat

Abstract

The diurnal rhythm of release of phytosiderophores and uptake rate of zinc (Zn) was studied in iron (Fe) deficient wheat (Triticum aestivum L. cv. Ares) plants grown in nutrient solution under controlled environmental conditions. Different forms of Zn (e.g. ZnSO₄, ZnEDTA) were used to obtain different degrees of loading of the root apoplasmic pool with Zn.

In the Fe-deficient plants the release of phytosiderophores from the roots followed a distinct diurnal rhythm with a steep peak about 4 h after the onset of the light period. These plants also showed a similar pattern in the rates of Zn uptake over the 24 h day-night cycle. During the light period there was a steep transient peak (factor 3.8) in Zn uptake rate in the Fe-deficient plants supplied with ZnSO₄. This transient peak was much less distinct in plants supplied with ZnEDTA (factor 1.8) and absent in plants supplied with ZnEDTA plus free chelator (+ NaEDTA) in excess. The peak in Zn uptake coincided with the maximum rate of phytosiderophore release in the Fe-deficient plants. In the Fe-sufficient plants the release of phytosiderophores was very low and no such peak in Zn uptake rates could be observed.

These results demonstrate that phytosiderophores mobilize Zn not only in the rhizosphere, but also from the root apoplast. Thus, the apoplasmic pool of micronutrient cations has to be taken into account as potential source for both uptake and diurnal variation in uptake rates of Micronutrient cations.     

Folate in wheat genotypes in the HEALTHGRAIN Diversity Screen

As part of the diversity screen of the HEALTHGRAIN project, the total folate contents of bread wheat (130 winter and 20 spring wheat genotypes), durum wheat (10 genotypes), earlier cultivated diploid einkorn and tetraploid emmer wheat (5 genotypes of each), and spelt (5 genotypes), grown in the same location in a controlled manner, were determined by a microbiological assay. The total folate contents ranged from 364 to 774 ng/g of dm in winter wheat and from 323 to 741 ng/g of dm in spring wheat, thus showing a marked variation. The highest mean for total folate content was measured in the durum wheat genotypes, whereas the earlier cultivated diploid and tetraploid wheat genotypes and spelt were shown to possess comparable or even higher folate contents than bread wheat. HPLC analysis of selected genotypes showed that 5-formyltetrahydrofolate was the major vitamer. The data provide a basis for breeding wheat genotypes with improved folate content.     

Interactions of Genotype and Glutenin Subunit Composition on Breadmaking Quality of Durum 1AS•1AL‐1DL Translocation Lines

Dual‐purpose durum (Triticum turgidum L. subsp. durum) wheat, having both good pasta and breadmaking quality, would be an advantage in the market. In this study, we evaluated the effects of genotype and varying HMW and LMW glutenin subunit composition on durum breadmaking quality. Genotypes included five near‐isogenic backgrounds that also differed by variability at the Glu‐D1d (HMW subunits 1Dx5+1Dy10), Glu‐B1 (presence or absence of subunit 1By8), and Glu‐B3 (LMWI or LMWII pattern) loci. Quality tests were conducted on genotypes grown at five North Dakota locations. Genotype had a stronger influence on free asparagine content than glutenin subunit composition. Genotypes carrying Glu‐D1d had higher glutenin content than lines that did not carry Glu‐D1d. Among Rugby translocation genotypes, lines carrying LMWI had higher gliadin content and better loaf volume than genotypes carrying LMWII. Absence of 1By8 produced major reductions in loaf volume in nontranslocation lines regardless of whether LMWI or LMWII was present. In contrast, the presence of Glu‐D1d compensated well for the absence of 1By8 regardless of which LMW pattern was present. The durum genotypes did not have loaf volumes equal to bread wheat cultivars, and results suggest that improved extensibility is needed to improve durum breadmaking quality.     

Study of Viscoelastic Properties of Wheat Kernels Using Compression Load Method

The method to measure hardness and other viscoelastic properties of intact wheat kernels is presented. Wheat with 9.3% moisture showed high elastic behavior compared with wheat tempered at 22.5% moisture that showed a plastic behavior. Load‐deformation curves showed that bread wheat behaves as a more plastic material than durum wheat, which is a more elastic material. Yield point of all the wheat samples was ≈18.5 N, independent of wheat type and moisture content. The height of the wheat kernel increased linearly, and the compression area increased exponentially, with increasing moisture content. The modulus of elasticity of wheat ranged from 99.2 MPa for 22.5% moisture content to 394.8 MPa for 9.3% moisture content. Young's modulus range for soft wheat such as Salamanca, Saturno, and Cortazar cultivars was 232.2–308.5 MPa, as compared with Rayón bread wheat at 321.5 MPa and the Altar, Sofía, and Rafi cultivars of durum wheat that had elastic moduli of 438.7–485.8 MPa. The compression force and final stress decreased from 69.9 N and 40.1 MPa in soft wheat to 90 N and 78.9 MPa in durum, respectively. Total work range was 14.7 MPa/sec in soft wheat to 19.7 MPa/sec for durum wheat and, as expected, was higher in the durum and bread wheat than in soft wheat. The plastic part ranged from 2.4 MPa/sec in soft wheat to 0.6 MPa/sec in durum wheat.     

根分泌物与禾本科植物对缺铁胁迫的适应机理

本文系统地总结了自然基金重点项目根分泌物在根际微生态系统中的营养机理的研究进展和部分主要成果。研究表明,养分专一性根分泌物是植物营养遗传特性控制基因的标记物,它受某一养分缺乏的诱导,是在植物体内合成并可通过主动分泌作用进入根际的代谢产物。它的合成和分泌只受该养分胁迫的专一诱导和控制,只要改善这一营养状况就能抑制或阻止其合成和分泌。当植物缺乏这一养分时,植物体可通过自身的调节能力,合成专一性物质并自根分泌到根际,促进该养分的活化并提高植物对其吸收利用效率,从而达到克服或缓解该营养胁迫的目的。用单基因突变材料进行的研究表明,植物铁载体的生物合成和吸收利用是受单基因控制的过程。这一发现不仅使人类有可能运用生物学研究技术来解决营养缺乏问题,而且也为有效地利用自然资源、降低生产成本、减少环境污染提供了可能性。麦根酸类(mugineic.acids)植物铁载体(phytosiderophores)在缺铁禾本科植物体内的生物合成,从植物根内向根际的分泌、在根际环境中对铁的活化及植物对F63+植物铁载体螯合体的吸收四个过程组成了禾本科植物对缺铁胁迫的适应机理。植物铁载体只在早晨日出后2～6h内大量分泌的节律性增加了它们在根际土壤中的相对浓度,减少了它们与土壤颗粒的接触和被吸附;分泌部位集中在微生物尚未侵染的根尖避开了微生物的破坏和分解,同时也增加了它们在土壤微区中的相对浓度。分泌作用和螯合作用不受介质pH值和Ca2+离子浓度影响的特性使该机理在经常出现缺铁现象的石灰性土壤上具有特殊意义。研究结果还表明,缺铁可以诱导激活根细胞原生质膜上可能存在的Fe3+植物铁载体复合体的专一性吸收和运载蛋白,高pH值和CaCO3对这一蛋白的载体功能只有很小的抑制作用,即使在pH值和CaCO3含量都较高的石灰性土壤上,也能有效地发挥作用。这一结论揭示了小麦等禾本科植物适应铁胁迫的实质及其生态学意义。     

Inheritance of manganese efficiency in durum wheat

Manganese (Mn) deficiency is a widespread problem on the alkaline soils, particularly for durum wheat (Triticum turgidum L. var. durum), which is more sensitive than either bread wheat or barley. The existence of considerable genetic variation in current germplasm of durum wheat (a relative yield of 58% in Stojocri 2 compared to 15% in check cv Yallaroi) and the development of a consistent selection criterion (Mn content of 35‐day‐old seedlings) has made breeding for Mn efficiency feasible. The development of Mn‐efficient durum wheat would be facilitated if the mode of inheritance was well understood. F1 hybrid, F2, and F2‐derived families from a cross between Stojocri 2 (moderately efficient) and Hazar (inefficient) were studied under controlled‐environment conditions. F1 hybrid was intermediate to the parents, indicating incomplete dominance and dependence on external Mn concentration. Analysis of 110 F2 and 220 F3 single plants (including 20 F2‐derived F3 families) showed that the observed variance was in agreement with the expected variance of a population segregating for two genes. Chi‐square analysis of the segregation ratios of F3 families also supported the digenic segregation hypothesis. Currently Stojocri 2 is used in a breeding program for the transfer of Mn efficiency to commercial varieties, by backcrossing (two backcrosses retain about 88% of recurrent parent genotype).