Differential response of rye,triticale, bread and durum wheats to zinc deficiency in calcareous soils

Field and greenhouse experiments were carried out to study the response of rye (Secale cereale L. cv. Aslim), triticale (× Triticosecale Wittmark. cv. Presto), two bread wheats (Triticum aestivum L, cvs. Bezostaja-1 and Atay-85) and two durum wheats (Triticum durum L. cvs. Kunduru-1149 and C-1252) to zinc (Zn) deficiency and Zn fertilization in severely Zn-deficient calcareus soils (DTPA-Zn=0.09 mg kg^-1 soil). The first visible symptom of Zn deficiency was a reduction in shoot elongation followed by the appearance of whitish-brown necrotic patches on the leaf blades. These symptoms were either absent or only slight in rye and triticale, but occurred more rapidly and severely in wheats, particularly in durum wheats. The same was true for the decrease in shoot dry matter production and grain yield. For example, in field experiments at the milk stage, decreases in shoot dry matter production due to Zn deficiency were absent in rye, and were on average 5% in triticale, 34% in bread wheats and 70%, in durum wheats. Zinc fertilization had no effect on grain yield in rye but enhanced grain yield of the other cereals. Zinc efficiency of cereals, expressed as the ratio of yield (shoot dry matter or grain) produced under Zn deficiency compared to Zn fertilization were, on average, 99% for rye, 74% for triticale, 59% for bread wheats and 25% for durum wheats.These distinct differences among and within the cereal species in susceptibility to Zn deficiency were closely related to the total amount (content) of Zn per shoot, but not with the Zn concentrations in shoot dry matter. For example, the most Zn-efficient rye and the Zn-inefficient durum wheat cultivar C-1252 did not differ in shoot Zn concentration under Zn deficiency, but the total amount of Zn per whole shoot was approximately 6-fold higher in rye than the durum wheat. When Zn was applied, rye and triticale accumulated markedly more Zn both per whole shoot and per unit shoot dry matter in comparison to wheats.The results demonstrate an exceptionally high Zn efficiency of rye and show that among the cereals studied Zn efficiency declines in the order rye>triticale>bread wheat>durum wheat. The differences in expression of Zn efficiency are possibly related to a greater capacity of efficient genotypes to acquire Zn from the soil compared to inefficient genotypes.     

Dry matter production and distribution of zinc in bread and durum wheat genotypes differing in zinc efficiency

Six bread wheat (Triticum aestivum cvs. Kiraç-66, Gerek-79, Aroona, ES 91-12, ES-14 and Kirkpinar) and four durum wheat (Triticum durum cvs. BDMM-19, Kunduru-1149, Kiziltan-91 and Durati) genotypes were grown under controlled environmental conditions in nutrient solution for 20 days to study the effect of varied supply of Zn (0 to 1 µM) on Zn deficiency symptoms in shoots, root and shoot dry matter production, and distribution of Zn in roots and shoots.Visual Zn deficiency symptoms, such as whitish-brown lesions on leaves, appeared rapidly and severly in durum wheats, particularly in Kiziltan-91 and Durati. Among the durum wheats, BDMM-19 was less affected by Zn deficiency, and among the bread wheats Kiraç-66, ES 91-12, Aroona and Gerek-79 were less affected than ES-14 and Kirkpinar.Under Zn deficiency, shoot dry matter production was decreased in all genotypes, but more distinctly in durum wheat genotypes. Despite severe decreases in shoot growth, root growth of all genotypes was either not affected or even increased by Zn deficiency. Correspondingly, shoot/root dry weight ratios were lower in Zn-deficient than in Zn-sufficient plants, especially in durum wheat genotypes.The distinct differences among the genotypes in sensitivity to Zn deficiency were closely related with the Zn content (Zn accumulation) per shoot but not with the Zn concentration in the shoot dry matter. On average, genotypes with lesser deficiency symptoms contained about 42% more Zn per shoot than genotypes with severe deficiency symptoms. In contrast to shoots, the Zn content in roots did not differ between genotypes. Shoot/root ratios of total Zn content were therefore greater for genotypes with lesser deficiency symptoms than for genotypes with severe deficiency symptoms (i.e. all durum wheat genotypes).The results suggest that the enhanced capacity of genotypes for Zn uptake and translocation from roots to shoot meristems under deficient Zn supply might be the most important factor contributing to Zn efficiency in wheat genotypes. The results also demonstrate that under severe Zn deficiency, Zn concentration in the shoot dry matter is not a suitable parameter for distinguishing wheat genotypes in their sensitivity to Zn deficiency.     

Large Scale Spatial Variability of Accumulated Cadmium in the Wheat Farm Grains

Cadmium (Cd) accumulation in edible crops is undesirable due to its hazardous influences on human health. The objectives of this study were: i) to evaluate the spatial variability of grain Cd and its relationships with soil properties in 4000 km² wheat farms; ii) to evaluate the effect of wheat cultivar on the soil properties vs. grain Cd relationships. A number of 255 soil (0–20 cm) and grain samples were taken and Cd concentrations in grain samples and some soil properties were measured. Grain Cd concentrations in 95 percent of the samples exceeded the threshold of 0.2 mg kg^?1. Durum wheat had more potential to accumulate Cd in grain (0.76 mg kg^?1) than bread (0.69 mg kg^?1). There was significant (p < 0.01) correlation between grain Cd and organic carbon (r = 0.66), CEC (r = 0.77) and DTPA-extractable Cd (p < 0.05) (r = 0.57) of the soils. Greater Pearson coefficient values for durum wheat showed that, in the studied calcareous soils, organic carbon, CEC, Cd-DTPA had more effects on durum wheat than bread wheat cultivar. The obtained Kriging map of grain Cd identified three hotspots at the east (durum wheat cultivation), the west (intensive irrigated wheat farms), and south (wheat farms around petrochemical industries) of the region. Agricultural mismanagement due to overusing P-fertilizers increased Cd concentration in the topsoils and grains of wheat farms in the study area.     

Grain zinc, iron and protein concentrations and zinc-efficiency in wild emmer wheat under contrasting irrigation regimes

Micronutrient malnutrition, and particularly deficiency in zinc (Zn) and iron (Fe), afflicts over three billion people worldwide, and nearly half of the world’s cereal-growing area is affected by soil Zn deficiency. Wild emmer wheat [Triticum turgidum ssp. dicoccoides (Körn.) Thell.], the progenitor of domesticated durum wheat and bread wheat, offers a valuable source of economically important genetic diversity including grain mineral concentrations. Twenty two wild emmer wheat accessions, representing a wide range of drought resistance capacity, as well as two durum wheat cultivars were examined under two contrasting irrigation regimes (well-watered control and water-limited), for grain yield, total biomass production and grain Zn, Fe and protein concentrations. The wild emmer accessions exhibited high genetic diversity for yield and grain Zn, Fe and protein concentrations under both irrigation regimes, with a considerable potential for improvement of the cultivated wheat. Grain Zn, Fe and protein concentrations were positively correlated with one another. Although irrigation regime significantly affected ranking of genotypes, a few wild emmer accessions were identified for their advantage over durum wheat, having consistently higher grain Zn (e.g., 125 mg kg^?1), Fe (85 mg kg^?1) and protein (250 g kg^?1) concentrations and high yield capacity. Plants grown from seeds originated from both irrigation regimes were also examined for Zn efficiency (Zn deficiency tolerance) on a Zn-deficient calcareous soil. Zinc efficiency, expressed as the ratio of shoot dry matter production under Zn deficiency to Zn fertilization, showed large genetic variation among the genotypes tested. The source of seeds from maternal plants grown under both irrigation regimes had very little effect on Zn efficiency. Several wild emmer accessions revealed combination of high Zn efficiency and drought stress resistance. The results indicate high genetic potential of wild emmer wheat to improve grain Zn, Fe and protein concentrations, Zn deficiency tolerance and drought resistance in cultivated wheat.     

Root exudation and Fe uptake and transport in wheat genotypes differing in tolerance to Zn deficiency

Tolerance to Zn deficiency in wheat germplasm may be inversely related to uptake and transport of Fe to shoots. The present study examined eight bread (Triticum aestivum) and two durum (T. turgidum L. conv. durum) wheat genotypes for their capacity to take up and transport Fe when grown under either Fe or Zn deficiency. Bread wheat genotypes Aroona, Excalibur and Stilleto showed tolerance to Zn and Fe deficiency, while durum wheat genotypes are clearly less tolerant to either deficiency. Roots of bread wheats tolerant to Zn deficiency exuded more phytosiderophores than sensitive bread and durum genotypes. Greater amounts of phytosideophores were exuded by roots grown under Fe than Zn deficiency. A relatively poor relationship existed between phytosiderophore exudation or the Fe uptake rate and relative shoot growth under Fe deficiency. At advanced stages of Zn deficiency, genotypes tolerant to Zn deficiency (Aroona and Stilleto) had a greater rate of Fe uptake than other genotypes. Zinc deficiency depressed the rate of Fe transport to shoots in all genotypes in early stages, while advanced Zn deficiency had the opposite effect. Compared with Zn-sufficient plants, 17-day-old Zn-deficient plants of genotypes tolerant to Zn deficiency had a lower rate of Fe transport to shoots, while genotypes sensitive to Zn deficiency (Durati, Yallaroi) had the Fe transport rate increased by Zn deficiency. A proportion of total amount of Fe taken up that was transported to shoots increased with duration of either Fe or Zn deficiency. It is concluded that greater tolerance to Zn deficiency among wheat genotypes is associated with the increased exudation of phytosiderophores, an increased Fe uptake rate and decreased transport of Fe to shoots. This revised version was published online in June 2006 with corrections to the Cover Date.     

Uptake of zinc by rye,bread wheat and durum wheat cultivars differing in zinc efficiency

Effect of zinc (Zn) nutritional status on uptake of inorganic ⁶⁵Zn was studied in rye (Secale cereale, cv. Aslim), three bread wheat (Triticum aestivum, cvs. Dagdas, Bezostaja, BDME-10) and durum wheat (Triticum durum, cv. Kunduru-1149) cultivars grown for 13 days in nutrient solution under controlled environmental conditions. The cultivars were selected based on their response to Zn deficiency and to Zn fertilization in calcareous soils under field conditions. When grown in Zn-deficient calcareous soil in the field, the rye cultivar had the highest, and the durum wheat the lowest Zn efficiency. Among the bread wheats, BDME-10 showed higher susceptibility to Zn deficiency and Bezostaja and Dagdas were less affected by Zn deficiency. Similarly to field conditions, in nutrient solution visual Zn deficiency symptoms (i.e. necrotic lesions on leaf blade) appeared to be more severe in Kunduru-1149 and BDME-10 and less severe in rye cultivar Aslim. Under Zn deficiency, shoot concentrations of Zn were similar between all cultivars. Cultivars with adequate Zn supply did not differ in uptake and root-to-shoot translocation rate of ⁶⁵Zn, but under Zn deficiency there were distinct differences; rye showed the highest rate of Zn uptake and the durum wheat the lowest. In the case of bread wheat cultivars, ⁶⁵Zn uptake rate was about the same and not related to their differential Zn efficiency. Under Zn deficiency, rye had the highest rate of root-to-shoot translocation of ⁶⁵Zn, while all bread and durum wheat cultivars were similar in their capacity to translocate ⁶⁵Zn from roots to shoots. When Zn²⁺ activity in uptake solution ranged between 117 p M and 34550 pM, Zn-efficient and Zn-inefficient bread wheat genotypes were again similar in uptake and root-to-shoot translocation rate of ⁶⁵Zn. The results indicate that high Zn efficiency of rye can be attributed to its greater Zn uptake capacity from soils. The inability of the durum wheat cultivar Kunduru-1149 to have a high Zn uptake capacity seems to be an important reason for its Zn inefficiency. Differential Zn efficiency between the bread wheat cultivars used in this study is not related to their capacity to take up inorganic Zn. This revised version was published online in June 2006 with corrections to the Cover Date.     

Phytosiderophore release in bread and durum wheat genotypes differing in zinc efficiency

The effect of the zinc (Zn) nutritional status on the rate of phytosiderophore release was studied in nutrient solution over 20 days in four bread wheat (Triticum aestivum cvs. Kiraç-66, Gerek-79, Aroona and Kirkpinar) and four durum wheat (Triticum durum cvs. BDMM-19, Kunduru-1149, Kiziltan-91 and Durati) genotypes differing in Zn efficiency.Visual Zn deficiency symptoms, such as whitish-brown necrosis on leaves and reduction in plant height appeared first and more severe in Zn-inefficient durum wheat genotypes Kiziltan-91, Durati and Kunduru-1149. Compared to the bread wheat genotypes, all durum wheat genotypes were more sensitive to Zn deficiency. BDMM-19 was the least affected durum wheat genotype. Among the bread wheat genotypes, Kirkpinar was the most sensitive genotype. In all genotypes well supplied with Zn, the rate of phytosiderophore release was very low and did not exceed 1 mol 32 plants^-1 3h^-1, or 0.5 mol g^-1 root dry wt 3h^-1. However, under Zn deficiency, with the onset of visual Zn deficiency symptoms, the release of phytosiderophores was enhanced in bread wheat genotypes up to 7.5 mol 32 plants^-1 3h^-1, or 9 mol g^-1 root dry wt 3h^-1, particularly in Zn-efficient Kiraç-66, Gerek-79 and Aroona. In contrast to bread wheat genotypes, phytosiderophore release in Zn-deficient durum wheat genotypes remained at a very low rate. Among the durum wheat genotypes BDMM-19 had highest rate of phytosiderophore release. HPLC analysis of root exudates showed that 2-deoxymugineic acid (DMA) is the dominating phytosiderophore released from roots of Zn-efficient genotypes. In root extracts concentration of DMA was also much higher in Zn-efficient than in inefficient genotypes. The results demonstrate that enhanced synthesis and release of phytosiderophores at deficient Zn supply is involved in Zn efficiency in wheat genotypes. It is suggested that the expression of Zn efficiency mechanism is causally related to phytosiderophore-mediated enhanced mobilization of Zn from sparingly soluble Zn pools and from adsorption sites, both in the rhizosphere and plants.     

Domesticated emmer wheat [T. turgidum L. subsp. dicoccon (Schrank) Thell.] as a source for improvement of zinc efficiency in durum wheat

Modern durum wheat (AABB) is more sensitive to zinc (Zn) deficiency than bread wheat (AABBDD). One strategy to increase productivity and expansion of durum wheat industry in Zn-deficient soils is to improve its ability to grow and yield in such soils. This ability is termed Zn efficiency. In a growth room experiment using soil culture, we assessed the potential of Triticum turgidum L. subsp. dicoccon (Shrank) Thell. (domesticated emmer wheat, AABB) as a genetic resource for further improvement of Zn efficiency in modern durum wheat. Twenty four accessions of domesticated emmer wheat, four durum landraces/cultivars, and two bread wheat cultivars/ advanced breeders lines of known Zn efficiency were tested under Zn deficiency and Zn sufficiency. Significant variation was observed among genotypes in Zn deficiency symptoms, dry matter production, shoot Zn concentration, shoot Zn content and Zn utilisation efficiency (physiological efficiency). We identified domesticated emmer wheat accessions with greater Zn efficiency than modern durum wheat and even bread wheat genotypes. These accessions could be used in breeding programs to improve Zn efficiency of durum wheat. The results suggest that Zn efficiency of durum or bread wheat is likely to be determined collectively by its progenitors.     

Polymorphism of omega-gliadins in durum wheat as revealed by the two-step APAGE/SDS-PAGE technique

Polymorphism of omega-gliadins was studied in 243 durum wheats from 27 countries using the two-step one-dimensional APAGE/SDS-PAGE technique. A total of 12 bands of different mobility were observed, and four of them were found to be different from those previously detected by Khelifi et al. (1992) in bread wheat. Fifteen alleles, six coded by the Gli-A1 locus and nine coded by the Gli-B1 locus, were identified, accounting for 19 different electrophoretic patterns. Seven new alleles were detected: two at the Gli-A1 locus and five at the Gli-B1 locus. The polymorphism found at the Gli-A1 and Gli-B1 loci was slightly greater than that found in bread wheat. Allelic differences between both species were higher at the Gli-B1 locus. A comparison of the frequencies of alleles in both species was carried out. The null allele, Gli-A1e, was more common in durum wheat than in bread wheat. The Gli-B1b allele, present in 60% of the bread wheats, was found in only 2% of the durum wheats and Gli-B1e, very common in durum wheat (45%), was rare in bread wheat (4%). The Gli-B1IV allele, common in durum wheat (28%), was not detected in bread wheat.     

Uptake and transport of foliar applied zinc (65Zn) in bread and durum wheat cultivars differing in zinc efficiency

Using two bread wheat (Triticum aestivum) and two durum wheat (Triticum durum) cultivars differing in zinc (Zn) efficiency, uptake and translocation of foliar-applied ⁶⁵Zn were studied to characterize the role of Zn nutritional status of plants on the extent of phloem mobility of Zn and to determine the relationship between phloem mobility of Zn and Zn efficiency of the used wheat cultivars. Irrespective of leaf age and Zn nutritional status of plants, all cultivars showed similar Zn uptake rates with application of ⁶⁵ZnSO₄ to leaf strips in a short-term experiment. Also with supply of ⁶⁵ZnSO₄ by immersing the tip (3 cm) of the oldest leaf of intact plants, no differences in Zn uptake were observed among and within both wheat species. Further, Zn nutritional status did not affect total uptake of foliar applied Zn. However, Zn-deficient plants translocated more ⁶⁵Zn from the treated leaf to the roots and remainder parts of shoots. In Zn-deficient plants about 40% of the total absorbed ⁶⁵Zn was translocated from the treated leaf to the roots and remainder parts of shoots within 8 days while in Zn-sufficient plants the proportion of the translocated ⁶⁵Zn of the total absorbed ⁶⁵Zn was about 25%. Although differences in Zn efficiency existed between the cultivars did not affect the translocation and distribution of ⁶⁵Zn between roots and shoots. Bread wheats compared to durum wheats, tended to accumulate more ⁶⁵Zn in shoots and less ⁶⁵Zn in roots, particularly under Zn-deficient conditions. The results indicate that differences in expression of Zn efficiency between and within durum and bread wheats are not related to translocation or distribution of foliar-applied ⁶⁵Zn within plants. Differential compartementation of Zn at the cellular levels is discussed as a possible factor determining genotypic variation in Zn efficiency within wheat.     

Importance of seed Zn content for wheat growth on Zn-deficient soil

Seed nutrient reserves may be important for an early establishment of crops on low-fertility soils. This glasshouse pot study evaluated effects of seed Zn content on vegetative growth of two wheat (Triticum aestivum L.) genotypes differing in Zn efficiency. Low-Zn (around 250 ng Zn per seed) and high-Zn seed (around 700 ng Zn per seed on average) of Excalibur (Zn efficient) and Gatcher (Zn inefficient) wheats were sown in a Zn-deficient siliceous sand fertilised with 0, 0.05, 0.2, 0.8 or 3.2 mg Zn kg^-1 soil. After 3 weeks, plants derived from the high-Zn seed had better root and shoot growth; the cv. Excalibur accumulated more shoot dry matter than the cv. Gatcher. After 6 weeks, greater root and shoot growth of plants grown from the high-Zn seed compared to those from the low-Zn seed was obvious only at nil Zn fertilisation. A fertilisation rate of 0.2 mg Zn kg^-1 soil was required for achieving 90% of the maximum yield for plants grown from the high-Zn seed compared to 0.8 mg Zn kg^-1 soil for plants derived from the low Zn seed. The critical Zn level in youngest expanded leaves for 90% maximum yield was 16 mg Zn kg^-1 dry matter for both genotypes. Zn-efficient Excalibur had greater net Zn uptake rates compared to Zn-inefficient Gatcher after 3 weeks but they were not different at the 6-week harvest. Zinc-deficient plants had greater net uptake rates of Cu, Mn, B, P, and K but a reduced uptake rate of Fe. It is concluded that higher seed Zn content acted similar to a starter-fertiliser effect by improving vegetative growth and dissipating differences in Zn efficiency of wheat genotypes.     

Seed priming with iron and zinc in bread wheat: effects in germination,mitosis and grain yield

Currently, the biofortification of crops like wheat with micronutrients such as iron (Fe) and zinc (Zn) is extremely important due to the deficiencies of these micronutrients in the human diet and in soils. Agronomic biofortification with Fe and Zn can be done through different exogenous strategies such as soil application, foliar spraying, and seed priming. However, the excess of these micronutrients can be detrimental to the plants. Therefore, in the last decade, a high number of studies focused on the evaluation of their phytotoxic effects to define the best strategies for biofortification of bread wheat. In this study, we investigated the effects of seed priming with different dosages (1 mg L^?1 to 8 mg L^?1) of Fe and/or Zn in germination, mitosis and yield of bread wheat cv. ‘Jordão’ when compared with control. Overall, our results showed that: micronutrient dosages higher than 4 mg L^?1 negatively affect the germination; Fe and/or Zn concentrations higher than 2 mg L^?1 significantly decrease the mitotic index and increase the percentage of dividing cells with anomalies; treatments performed with 8 mg L^?1 of Fe and/or 8 mg L^?1 Zn caused negative effects in germination, mitosis and grain yield. Moreover, seed priming with 2 mg L^?1 Fe?+?2 mg L^?1 Zn has been shown to be non-cytotoxic, ensuring a high rate of germination (80%) and normal dividing cells (90%) as well as improving tillering and grain yield. This work revealed that seed priming with Fe and Zn micronutrients constitutes a useful and alternative approach for the agronomic biofortification of bread wheat.     

Expression of high zinc efficiency of Aegilops tauschii and Triticum monococcum in synthetic hexaploid wheats

The effect of varied zinc (Zn) supply on shoot and root dry matter production, severity of Zn deficiency symptoms and Zn tissue concentrations was studied in two Triticum turgidum (BBAA) genotypes and three synthetic hexaploid wheat genotypes by growing plants in a Zn-deficient calcareous soil under greenhouse conditions with (+Zn=5 mg kg^-1 soil) and without (−Zn) Zn supply. Two synthetic wheats (BBAADD) were derived from two different Aegilops tauschii (DD) accessions using same Triticum turgidum (BBAA), while one synthetic wheat (BBAAAA) was derived from Triticum turgidum (BBAA) and Triticum monococcum (AA). Visible symptoms of Zn deficiency, such as occurrence of necrotic patches on leaves and reduction in shoot elongation developed more rapidly and severely in tetraploid wheats than in synthetic hexaploid wheats. Correspondingly, decreases in shoot and root dry matter production due to Zn deficiency were higher in tetraploid wheats than in synthetic hexaploid wheats. Transfer of the DD genome from Aegilops tauschii or the AA genome from Triticum monococcum to tetraploid wheat greatly improved root and particularly shoot growth under Zn-deficient, but not under Zn-sufficient conditions. Better growth and lesser Zn deficiency symptoms in synthetic hexaploid wheats than in tetraploid wheats were not accompanied by increases in Zn concentration per unit dry weight, but related more to the total amount of Zn per shoot, especially in the case of synthetic wheats derived from Aegilops tauschii. This result indicates higher Zn uptake capacity of synthetic wheats. The results demonstrated that the genes for high Zn efficiency from Aegilops tauschii (DD) and Triticum monococcum (AA) are expressed in the synthetic hexaploid wheats. These wheat relatives can be used as valuable sources of genes for improvement of Zn efficiency in wheat. This revised version was published online in June 2006 with corrections to the Cover Date.     

Genotypic variation in zinc efficiency and resistance to crown rot disease (Fusarium graminearum Schw. Group 1) in wheat

A crown rot disease in wheat caused by the fungusFusarium graminearum Schw. Group 1 is a widespread problem in chronically Zn-deficient Australian soils. A link between crown rot and Zn deficiency was established by Sparrow and Graham (1988). This paper reports a test of a further hypothesis, that wheat genotypes more efficient at extracting zinc from low-zinc soils are more resistant to infection by this pathogen. Three wheat cultivars (Excalibur, Songlen and Durati) of differential Zn efficiency were tested at three zinc levels (0.05, 0.5 and 2.0 mg Zn kg⁻¹ of soil) and three levels ofF. graminearum S. Group 1 inoculum (0.1 g and 0.3 g kg⁻¹ live chaff-inoculum and control having 0.1 g kg⁻¹ dead chaff inoculum). Six weeks after sowing dry matter production of shoots and roots was decreased byFusarium inoculation at 0.05 mg and 0.5 mg kg⁻¹ applied Zn.Fusarium inoculum at 0.1 g was as effective as 0.3 g kg⁻¹ for infection and decreasing dry matter. The infection at the basal part of culm decreased significantly by increasing the rate of Zn application. Excalibur, a Zn-efficient cultivar (tolerant to Zn deficiency) produced significantly more shoot and root dry matter, and showed less disease infection compared with Zn-inefficient cultivars (Durati and Songlen) at low (0.05 mg Zn kg⁻¹ soil) and medium (0.5 mg Zn kg⁻¹ soil) Zn fertilization rates. Higher rate of Zn fertilization (2.0 mg Zn kg⁻¹ soil) reduced the disease level in Durati to the level of Excalibur but the disease level of Songlen was still high, indicating its high Zn requirement and or sensitivity to crown rot. The data on Zn uptake show that Excalibur, being Zn-efficient, was able to scavenge enough Zn from Zn-deficient soil, we suggest that besides sustaining growth Excalibur was able to build and maintain resistance to the pathogen; inefficient cultivars needed extra Zn fertilization to achieve performance comparable to that of Excalibur. The present study indicates that growing Zn-efficient cultivars of wheat along with judicious use of Zn fertilizer in Zn-deficient areas where crown rot is a problem may sustain wheat production by reducing the severity of the disease as well as by increasing the plant vigour through improved Zn nutrition. ei]Section editor: R Rodriques-Kalana     

Accumulation of cadmium and selected elements in flax seed grown on a calcareous soil

Seed of flax (Linum usitatissimum L.) grown on calcareous and neutral soils sometimes accumulates relatively high concentrations of Cd. The influence of a post-flowering application of NH₄NO₃ (115 mg N kg^-1), CdSO₄ (1 mg Cd kg^-1), FeEDDHA (2 mg Fe kg^-1), NaH₂PO₄ (120 mg P kg^-1) and ZnSO₄ (8 mg Zn kg^-1) on seed accumulation of Cd, Fe, N, Mn, P and Zn by flax grown on a Calciaquoll was studied in two experiments under greenhouse conditions. Seed yields were increased by the N and Zn treatments, and the N×Zn interaction was positive. Zinc deficiency delayed flowering and boll formation by up to 20 days and reduced seed size. In the absence of added Cd, seed accumulated up to 0.33 mg Cd kg^-1. This Cd accumulation was reduced by approximately 50 and 17% by added Zn and Fe, respectively, but was little affected by P fertilizer and post-flowering N stress. In the presence of added Cd, seed Cd exceeded 3.3 mg Cd kg^-1, and the antagonistic effects of Fe and Zn on seed Cd were absent. Seed N, P, Fe and Zn concentrations were increased on average by 10, 45, 31 and 97% by the N, P, Fe and Zn fertilizer treatments, respectively. FeEDDHA reduced seed Mn concentration by approximately 58%. However, seed Mn concentration was much less than that found in vegetative tissue at flowering. Soil-applied Zn may reduce seed Cd concentration in flax under field conditions, and may increase marketability of flax for food use.     

Differences in Zinc Efficiency among and within Diploid, Tetraploid and Hexaploid Wheats

Greenhouse experiments were carried out with six diploid, ninetetraploid and seven hexaploid wheats, including wild and primitivegenotypes, to study the influence of varied zinc (Zn) supplyon the severity of Zn deficiency symptoms, shoot dry matterproduction and shoot Zn concentrations. In addition to wildand primitive genotypes, one modern tetraploid cultivar withhigh sensitivity to Zn deficiency and two modern hexaploid cultivars,one highly sensitive to and one resistant to Zn deficiency,were included for comparison. Plants were grown for 44 d ina severely Zn-deficient calcareous soil, with (+Zn; 5 mg Znkg^-1soil) and without (-Zn) Zn fertilization. Visible Zn deficiencysymptoms, including whitish-brown necrotic patches on leaf blades,appeared very rapidly and severely in all tetraploid wheat genotypes.Compared with tetraploid wheats, diploid and hexaploid wheatswere less sensitive to Zn deficiency. With additional Zn, shootdry matter production was higher in tetraploid than diploidand hexaploid wheats. However, under Zn-deficient conditionstetraploid wheats had the lowest shoot dry matter production,indicating the very high sensitivity of tetraploid wheats toZn deficiency. Consequently, Zn efficiency expressed as theratio of shoot dry matter produced under Zn deficiency to Znfertilization, was much lower in tetraploid wheats than in diploidand hexaploid wheats. On average, Zn efficiency ratios were36% for tetraploid, 60% for diploid and 64% for hexaploid wheats.Differences in Zn efficiency among and within diploid, tetraploidand hexaploid wheats were positively related to the amount ofZn per shoot of the genotypes, but not to the amount of Zn perunit dry weight of shoots or seeds used in the experiments.The seeds of the accessions of tetraploid wild wheats containedup to 120 mg Zn kg^-1, but the resulting plants showed very highsensitivity to Zn deficiency. By contrast, hexaploid wheatsand primitive diploid wheats with much lower Zn concentrationsin seeds had higher Zn efficiencies. It is suggested that notonly enhanced Zn uptake capacity but also enhanced internalZn utilization capacity of genotypes play important roles indifferential expression of Zn efficiency. The results of thisstudy also suggest the importance of the A and D genomes asthe possible source of genes determining Zn efficiency in wheat.Copyright 1999 Annals of Botany Company Seeds, Triticum aestivum, Triticum monococcum, Triticum turgidum, zinc concentrations, zinc deficiency, zinc efficiency.     

Soil solution dynamics and plant uptake of cadmium and zinc by durum wheat following phosphate fertilization

A growth chamber study was conducted to evaluate the effect of application of phosphate fertilizer on soil solution dynamics of cadmium (Cd) and Cd accumulation in durum wheat (Triticum turgidum L. var. durum). Treatments consisted of three phosphate fertilizer sources containing 3.4, 75.2, and 232 mg Cd kg^?1 applied at three rates (20, 40 and 80 mg P kg^?1) plus a no fertilization control. An unplanted treatment at 40 mg P kg^?1 was included to separate the effects on soil solution Cd dynamics of the crop from that of the fertilizer. Soil solution samples were obtained using soil moisture samplers every 10 days after germination. The experimental results indicated that plant biomass significantly increased with P application rates and decreased with increased Cd concentration in the phosphate fertilizers. Total cadmium concentration in soil solution was not consistently affected by phosphate fertilization rate and fertilizer sources, and therefore Cd concentration in the fertilizer. Application of phosphate fertilizer, however, increased the concentration and accumulation of Cd and shoot Cd/Zn ratio, and decreased shoot Zn concentration in durum wheat. Phosphate sources had a marginally significant effect (P?=?0.05) on shoot Cd concentration and did not affect Cd accumulation in durum wheat. Concentration of Cd in soil solution was unrelated to Cd concentration in durum wheat. These results suggest that the immediate increase in Cd concentration and Cd accumulation in durum wheat with phosphate application is due more to competition between Zn and Cd for absorption into plants, enhanced root to shoot translocation and enhanced root development, than to a direct addition effect from Cd contained in phosphate fertilizer. In the short term, application of phosphate fertilizers can increase Cd concentration in the crops, regardless of the Cd concentration of the fertilizer. An optimal P fertilization, possibly in combination with Zn application, may offer an important strategy for decreasing Cd concentration and accumulation in crops.     

Characterization of Zinc Uptake, Binding, and Translocation in Intact Seedlings of Bread and Durum Wheat Cultivars

Durum wheat (Triticum turgidum L. var durum) cultivars exhibit lower Zn efficiency than comparable bread wheat (Triticum aestivum L.) cultivars. To understand the physiological mechanism(s) that confers Zn efficiency, this study used ⁶⁵Zn to investigate ionic Zn²⁺ root uptake, binding, and translocation to shoots in seedlings of bread and durum wheat cultivars. Time-dependent Zn²⁺ accumulation during 90 min was greater in roots of the bread wheat cultivar. Zn²⁺ cell wall binding was not different in the two cultivars. In each cultivar, concentration-dependent Zn²⁺ influx was characterized by a smooth, saturating curve, suggesting a carrier-mediated uptake system. At very low solution Zn²⁺ activities, Zn²⁺ uptake rates were higher in the bread wheat cultivar. As a result, the Michaelis constant for Zn²⁺ uptake was lower in the bread wheat cultivar (2.3 μm) than in the durum wheat cultivar (3.9 μm). Low temperature decreased the rate of Zn²⁺ influx, suggesting that metabolism plays a role in Zn²⁺ uptake. Ca inhibited Zn²⁺ uptake equally in both cultivars. Translocation of Zn to shoots was greater in the bread wheat cultivar, reflecting the higher root uptake rates. The study suggests that lower root Zn²⁺ uptake rates may contribute to reduced Zn efficiency in durum wheat varieties under Zn-limiting conditions.Soils that contain insufficient levels of the essential plant micronutrient Zn are common throughout the world. As a result, Zn deficiency is a widespread problem in crop plants, especially cereals (Graham et al., 1992). The importance of plant foods as sources of Zn, particularly in the marginal diets of developing countries, is well established (Welch, 1993). The development of crop plants that are efficient Zn accumulators is therefore a potentially important endeavor. In addition to its effects on nutrition, Zn deficiency in crops is relevant to other areas of human health. Another consequence of Zn-deficient soils is the tendency for plants grown in such soils to accumulate heavy metals. For example, in the Great Plains region of North America, where soil Zn levels are low and naturally occurring Cd is present, durum wheat (Triticum turgidum L. var durum) grains accumulate Cd to relatively high concentrations (Wolnik et al., 1983). The presence of Cd in food represents a potential human health hazard and, in response, international trade standards have been proposed to limit the levels of Cd in exported grain (Codex Alimentarius Commission, 1993). Thus, there is a need to understand the physiological processes that control acquisition of Zn from soil solution by roots and mobilization of Zn within plants.It has been demonstrated in recent years that crop plants vary in their ability to take up Zn, particularly when its availability to roots is limited. Zn efficiency, defined as the ability of a plant to grow and yield well in Zn-deficient soils, varies among wheat cultivars (Graham and Rengel, 1993). In field trials, durum wheat cultivars have been shown to be consistently less Zn efficient than bread wheat (Triticum aestivum L.) cultivars (Graham et al., 1992). Similarly, durum wheat varieties were reported to be less Zn efficient than bread wheat varieties when grown in chelate-buffered hydroponic nutrient culture (Rengel and Graham, 1995a).The physiological mechanism(s) that confers Zn efficiency has not been identified. Processes that could influence the ability of a plant to tolerate limited amounts of available Zn include higher root uptake, more efficient utilization of Zn, and enhanced Zn translocation within the plant. Cakmak et al. (1994) showed that a Zn-inefficient durum wheat cultivar exhibited Zn-deficiency symptoms earlier and more intensely than a Zn-efficient bread wheat cultivar even though the Zn tissue concentrations were similar in both lines, suggesting differential utilization of Zn in the two cultivars. Rates of Zn translocation to shoots were shown to vary among sorghum cultivars, although correlations with Zn efficiency were not established (Ramani and Kannan, 1985). Root uptake kinetics have been reported to vary between rice cultivars having different Zn requirements, with high-Zn-requiring cultivars exhibiting consistently higher root uptake rates (Bowen, 1986). In contrast, a correlation between Zn efficiency and rates of root Zn uptake in bread and durum wheat cultivars could not be demonstrated (Rengel and Graham, 1995b).In grasses Zn influx into the root symplasm has been hypothesized to occur as the free Zn²⁺ ion (Halvorson and Lindsay, 1977), as well as in the form of Zn complexes with nonprotein amino acids known as phytosiderophores (Tagaki et al., 1984) or phytometallophores (Welch, 1993). Concentration-dependent uptake of free Zn²⁺ ions has been shown to be saturable in several species, including maize (Mullins and Sommers, 1986), barley (Veltrup, 1978), and wheat (Chaudhry and Loneragan, 1972), suggesting that ionic uptake in grasses occurs via a carrier-mediated system. However, several of these studies have been criticized on the basis that excessively high (and physiologically unrealistic) Zn²⁺ concentrations were used (Kochian, 1993).This study was undertaken to examine unidirectional Zn²⁺ influx and translocation to shoots in Zn-efficient bread wheat lines and Zn-inefficient durum wheat lines. Experiments were performed in the absence of added phytometallophores and results are presumed to represent influx of ionic Zn²⁺. Zn activities in the nanomolar range were used to more closely mimic free Zn²⁺ levels occurring naturally in soil solution. The results presented here indicate that a Zn-efficient bread wheat cultivar maintained higher rates of Zn uptake than a Zn-inefficient durum wheat cultivar, particularly at low (and physiologically relevant) solution Zn²⁺ activities.     

Bacterial inoculation speeds zinc release from ground tire rubber used as Zn fertilizer for corn and sunflower in a calcareous soil

Background and Aims

We tested the utility of some biological treatments to hasten degradation of waste tire rubber in soil and thus the release of zinc and sulfur for plant uptake.

Methods

Three rates of ground tire rubber (0, 150, and 300?mg?kg^?1) were incorporated into a Zn-deficient calcareous soil. Before addition to the soil, ground rubber was given four microbial treatments including no inoculation, inoculation with Rhodococcus erythropolis, inoculation with R. erythropolis+Escherichia coli, and inoculation with R. erythropolis+E. coli+Acinobacter calcoaceticus. In the pot experiment, corn (Zea mays L. Hybrid Single Cross 500) and sunflower (Helianthus annuus L. cv. Record) plants were exposed to three rates of ground rubber (0, 150, and 300?mg?kg^?1) or 3?mg zinc kg^?1 as ZnSO₄. Before addition to the soil, ground rubber and ZnSO₄ were inoculated or non-inoculated with R. erythropolis+E. coli+A. calcoaceticus.

Results

Ground rubber and microbial inoculation treatments reduced soil pH and the magnitude of this reduction increased over time. Ground rubber in combination with microbial inoculation increased DTPA-extractable soil Zn and Fe. The amount of DTPA-extractable Zn and Fe of rubber-amended soils increased over time so that the highest concentration of available Zn and Fe was found at week 10. Application of microbial inoculated ground tire rubber significantly increased shoot Zn concentration of each plant species.

Conclusions

Bacterial inoculation of ground rubber was effective in hastening increase in DTPA-extractable Zn in the studied calcareous soil and in enhancing Zn uptake by plants.     

Energy-dispersive X-ray fluorescence spectrometry as a tool for zinc, iron and selenium analysis in whole grain wheat

Background and aims

Crop biofortification programs require fast, accurate and inexpensive methods of identifying nutrient dense genotypes. This study investigated energy-dispersive X-ray fluorescence spectrometry (EDXRF) for the measurement of zinc (Zn), iron (Fe) and selenium (Se) concentrations in whole grain wheat.

Methods

Grain samples were obtained from existing biofortification programs. Reference Zn, Fe and Se concentrations were obtained using inductively coupled plasma optical emission spectrometry (ICP-OES) and/or inductively coupled plasma mass spectrometry (ICP-MS). One set of 25 samples was used to calibrate for Zn (19–60?mg?kg^–1) and Fe (26–41?mg?kg^–1), with 25 further samples used to calibrate for Se (2–31?mg?kg^–1 ). Calibrations were validated using an additional 40–50 wheat samples.

Results

EDXRF limits of quantification (LOQ) were estimated as 7, 3 and 2?mg?kg^–1 for Zn, Fe, and Se, respectively. EDXRF results were highly correlated with ICP-OES or -MS values. Standard errors of EDXRF predictions were ±2.2?mg Zn kg^–1, ±2.6?mg Fe kg^–1, and ±1.5?mg Se kg^–1.

Conclusion

EDXRF offers a fast and economical method for the assessment of Zn, Fe and Se concentration in wheat biofortification programs.