Toxicity of cadmium and its competition with mineral nutrients for uptake by plants:A review

Cadmium(Cd)is a toxic heavy metal occurring in the environment naturally and is also generated through various anthropogenic sources and acts as a pollutant.Human health is affected by Cd pollution in farmland soils because food is the main source of Cd intake in the non-smoking population.For crops,Cd toxicity may result from a disturbance in uptake and translocation of mineral nutrients and disturbance in plant metabolism,inhibiting plant growth and development.However,plants have Cd tolerance mechanisms,including restricted Cd uptake,decreased Cd root-to-shoot translocation,enhanced antioxidant enzyme activities,and increased production of phytochelatins.Furthermore,optimal supply of mineral nutrients is one of the strategies to alleviate the damaging effects of Cd on plants and to avoid its entry into the food chain.The emerging molecular knowledge contributes to understanding Cd uptake,translocation,and remobilization in plants.In this review,Cd toxicity and tolerance mechanisms,agricultural practices to minimize Cd accumulation,Cd competition with essential elements(calcium,copper,iron,zinc,and manganese),and genes associated with Cd uptake are discussed in detail,especially regarding how these mineral nutrients and genes play a role in decreasing Cd uptake and accumulation in crop plants.     

Manganese availability and microbial populations in the rhizosphere of wheat genotypes differing in tolerance to Mn deficiency

Plant genotypes differ in their capacity to grow in soils with low manganese (Mn) availability. The physiological mechanisms underlying differential tolerance to Mn deficiency are poorly understood. To study the relationship between Mn content in soil, plant genotypes, and rhizosphere microorganisms in differential Mn efficiency, two wheat (Triticum aestivum L.) cultivars, RAC891 (tolerant to Mn deficiency) and Yanac (sensitive), were grown in a Mn‐deficient soil to which 5, 10, 20 or 40 mg Mn kg^–1 were added. The shoot dry matter of both cultivars increased with increasing Mn addition to the soil. At all soil Mn fertilizer levels, the tolerant RAC891 had a greater shoot dry matter and a higher total shoot Mn uptake than the sensitive Yanac. The concentration of DTPA‐extractable Mn in the rhizosphere soil of RAC891 at Mn20 and Mn40 was slightly lower than in the rhizosphere of Yanac. The population density of culturable microorganisms in the rhizosphere soil was low (log 6.8–6.9 cfu (g soil)^–1) in both cultivars and neither Mn oxidation nor reduction were observed in vitro. To assess the non‐culturable fraction of the soil microbial community, the ribosomal intergenetic spacer region of the bacterial DNA in the rhizosphere soil was amplified (RISA) and separated in agarose gels. The RISA banding patterns of the bacterial rhizosphere communities changed markedly with increasing soil Mn level, but there were no differences between the wheat cultivars. The bacterial community structure in the rhizosphere was significantly correlated with the concentration of DPTA‐extractable Mn in the rhizosphere, fertilizer Mn level, shoot dry matter, and total shoot Mn uptake. The results obtained by RISA indicate that differential tolerance to Mn deficiency in wheat may not be related to changes in the composition of the bacterial community in the rhizosphere.     

Belowground interactions between intercropped wheat and Brassicas in acidic and alkaline soils

Our previous studies showed that, under P-limiting conditions, growth and P uptake were lower in the wheat genotype Janz than in three Brassica genotypes when grown in monoculture. The present study was conducted to answer the question if P mobilised by the Brassicas is available to wheat; leading to improved growth of wheat when intercropped with Brassicas compared to monocropped wheat. To assess if the interactions between the crops depend on soil type, the wheat genotype Janz and three Brassica genotypes (two canolas and one mustard) were grown for 6 weeks in monoculture or wheat intercropped with each Brassica genotype in an acidic and an alkaline soil with low P availability (with two plants per pot). Wheat grew equally well in the two soils, but the Brassicas grew better in the acidic than in the alkaline soil. In the acidic soil, monocropped Brassicas had a 3 to 4 fold greater plant dry weight (dw) and P uptake than wheat; plant dw and P uptake in wheat were decreased or not affected by intercropping and increased in the Brassicas. In the alkaline soil, dw and P uptake of the Brassicas was twice as high as in wheat, with intercropping having no effect on these parameters. The contribution of wheat to the total shoot dw and P uptake per pot was 4-21% and 32-40% in acidic and alkaline soil, respectively. Mycorrhizal colonisation was low in all genotypes in the acidic soil (1-6%). In the alkaline soil, mycorrhizal colonisation of monocropped wheat was 62%, but only 43-47% in intercropped wheat. Intercropping decreased P availability in the rhizosphere of wheat in the acidic soil but had no effect on rhizosphere P availability in the alkaline soil. Intercropping had a variable effect on rhizosphere microbial community composition (assessed by fatty acid methylester analysis (FAME) and ribosomal intergenic spacer amplification (RISA)), ranging from intercropping having no effect on the rhizosphere communities to intercropping resulting in a new and similar rhizosphere community composition in both genotypes. The results of this study show that intercropping with Brassicas does not improve growth and P uptake of wheat; thus there is no indication that P mobilised by the Brassicas is available to wheat.     

Growth,P uptake and rhizosphere properties of wheat and canola genotypes in an alkaline soil with low P availability

The aim of the present study was to assess the role of soil type on growth, P uptake and rhizosphere properties of wheat and canola genotypes in an alkaline soil with low P availability. Two wheat (Goldmark and Janz) and two canola genotypes (Drum and Outback) were grown in a calcareous soil (pH 8.5) at two P levels [no P addition (0P) or addition of 200 mg kg⁻¹ P as Ca₃(PO₄)₂ (200P)] and harvested at flowering or maturity. Shoot and root dry weight, root length and shoot P content were greater in the two canola genotypes than in wheat. There were no consistent differences in available P, microbial P and phosphatase activity in the rhizosphere of the different genotypes. Shoot P content was significantly positively correlated with root length, pH and phosphatase activity in the rhizosphere. The microbial community composition, assessed by fatty acid methylester analysis, of the canola genotypes differed strongly from that of the wheat genotypes. The weight percentage bacterial fatty acids, the bacteria/fungi (b/f) ratio and the diversity of fatty acids were greater in the rhizosphere of the canolas than in the rhizosphere of the wheat genotypes. In contrast to the earlier studies in an acidic soil, only small differences in growth and P uptake between the genotypes of one crop were detected in the alkaline soil used here. The results confirmed the importance of root length for P uptake in soils with low P availability and suggest that the rhizosphere microbial community composition may play a role in the better growth of the canola compared to the wheat genotypes.     

Cultivation effects on temporal changes of organic carbon and aggregate stability in desert soils of Hexi Corridor region in China

Sustainable agricultural use of cultivated desert soils has become a concern in Hexi Corridor in Gansu Province of China, because loss of topsoil in dust storms has been recently intensified. We chose four desert sites to investigate the effects of cultivation (cropping) on (i) soil organic C and its size fractions and (ii) soil aggregate stability (as a measure of soil erodibility). These parameters are of vital importance for evaluating the sustainability of agricultural practices.

Total organic C as well as organic C fractions in soil (coarse organic C, 0.1–2 mm; young organic C, 0.05–0.1 mm; stable organic C, <0.05 mm) generally increased with the duration of the cultivation period from 0 (virgin soil, non-cultivated) to more than 30 years (p < 0.05). Compared to total organic C in virgin soils (2.3–3.5 g kg⁻¹ soil), significantly greater values were found after 10 to >20 years of cultivation (6.2–7.1 g kg⁻¹ soil). The increase in organic C in desert soils following prolonged cultivation was mainly the consequence of an increase in the coarse organic C. The increase in total organic C in soil was also dependent on clay content [total organic C = 0.96 + 0.249 clay content (%) + 0.05 cultivation year, R² = 0.48, n = 27, p < 0.001]. This indicates that clay protected soil organic C from mineralization, and also contributed to the increase in soil organic C as time of cultivation increased.

There was a significant positive correlation between aggregate stability and total organic C across all field sites. The water stability of aggregates was low (with water-stable aggregate percentage 4% of dry-sieved aggregates of size 1–5 mm). There was no consistent pattern of increase in the soil aggregate stability with time of cultivation at different locations, suggesting that desert soils might remain prone to wind erosion even after 50 years of cultivation. Alternative management options, such as retaining harvested crop residues on soil surface and excluding or minimizing tillage, may permit sustainable agricultural use of desert soils.     

Ca2+ Effects on K+ Fluxes in Arabidopsis Seedlings Exposed to Al3+

Potassium transport was investigated in the root elongation zone of Arabidopsis seedlings during the first minutes of Al³⁺ exposure, using the non-invasive MIFE microelectrode technique. To prevent pH changes during Al³⁺ application, and to separate aluminium from acidic stress, plants were pre-treated with 5 mM homoPIPES before addition of AlCl₃ (pH 4.2). The 30-min treatment with 50 or 500 μM AlCl₃ led to a significant increase in K⁺ efflux in solutions containing 100 μM CaCl₂. This efflux was suppressed by high concentrations of Ca²⁺ (10 mM) in the bathing solution. Our results suggest that elevated external Ca²⁺ activities can sustain K⁺ influx in the root elongation zone during Al³⁺ exposure either by maintaining [Ca²⁺]_cyt or by affecting Al³⁺ uptake across the plasma membrane.     

Mineral bioavailability in grains of Pakistani bread wheat declines from old to current cultivars

Estimating variation in grain mineral concentration and bioavailability in relation to grain yield and the year of cultivar release is important for breeding wheat with increased content of bioavailable minerals. The grain yield and yield components, grain phytate concentration, and concentration and bioavailability of minerals (zinc Zn, iron Fe and calcium Ca) in wheat grains were estimated in 40 wheat cultivars released in Punjab (Pakistan) during the last five decades. Mean grain Zn and Ca concentrations in current-cultivars were significantly lower (≥14%) than in obsolete cultivars released during the Green Revolution (1965–1976). Much of this variation was related to increased grain weight in current-cultivars. There was a positive correlation among minerals (r = 0.39 or higher, n = 40) and minerals with phytate in wheat grains (r = 0.38 or higher, n = 40). The tested cultivars varied widely in grain yield and grain phytate-to-mineral molar ratios (phytate:mineral). Compared to obsolete cultivars, the current-cultivars had a higher phytate:mineral ratio in grains, indicating poor bioavailability of minerals to humans. The study revealed a non-significant relationship between grain yield and phytate:mineral ratios in grains. Therefore, breeding for lower phytate:mineral ratios in wheat grains can ensure increased mineral bioavailability without significant reduction in the yield potential. Future breeding should be focused on developing new genotypes suitable for mineral biofortification and with increased mineral bioavailability in grains.     

Genetic Variation in Cadmium Accumulation and Tolerance among Wheat Cultivars at the Seedling Stage

Exploitation of genetic differences to select wheat cultivars’ pollution-safe from cadmium (Cd) contamination requires better understanding of Cd uptake and translocation patterns. For this purpose, 15 wheat cultivars were grown in nutrient solution and exposed to four levels of Cd, i.e., 0, 15, 30, and 45 µM. The plants were harvested after 2 weeks of Cd exposure. Root and shoot relative dry matter (DM) was decreased in most of the cultivars, but some cultivars did not exhibit any toxic symptoms. The lowest Cd concentration in shoots was recorded for Lasani-2008 and Iqbal-2000 while the highest for Sehar-2006 and Inqlab-91. Both root absorption and translocation accounted for regulating Cd concentration in shoots. There was no relationship between relative DM and Cd concentrations in roots and shoots. The results suggest that wheat cultivars with low shoot Cd concentration but higher tolerance, i.e. Lasani-2008 and Iqbal-2000, could be used in breading programs for low Cd wheat.     

Crop species differ in root plasticity response to localised P supply

The effect of localised phosphorus (P) fertiliser placement and in particular, deep P fertiliser placement, on the comparative root growth and P uptake of fibrous vs tap‐rooted crops is not known. In this study, we examined the root growth and P uptake of wheat (Triticum aestivum L.), canola (Brassica napus L.), and narrow‐leaf lupin (Lupinus angustifolius L.) in a split‐root system and in columns with deep (19 cm) or shallow (5 cm) P fertiliser sources in glasshouse conditions. In the split‐root system, plants of all three species grown under heterogeneous soil P conditions absorbed more P and produced greater root and shoot biomass than those under homogeneous P supply. Root plasticity differed between species under heterogeneous soil P supply: canola and wheat allocated relatively more root biomass and root length to the high P zone than narrow‐leaf lupin. In the column experiment, there was no difference in the amount of P accumulated in shoots of any crops grown in the deep vs shallow P fertiliser treatments. Root proliferation occurred within the shallow and deep‐P fertiliser bands in all three species; however, root distribution above or below the bands did not differ between deep or shallow P fertiliser treatments in any species. Whilst root plasticity responses to heterogeneous soil P supply differed among species, root architecture (fibrous vs taproot) did not confer any advantage or disadvantage to the acquisition of P from deep vs shallow P fertiliser bands. Moreover, whilst roots proliferate in the vicinity of P fertiliser bands, root distribution outside of the bands appears to remain unaltered in both fibrous and tap‐rooted crops during early growth.     

Growth and nutrient uptake of temperate perennial pastures are influenced by grass species and fertilisation with a microbial consortium inoculant

Background: The low fertility of sandy soils in South‐Western Australia is challenging for the establishment of temperate perennial pastures. Aims: To assess whether microbial consortium inoculant may improve plant growth by increasing nutrient supply, root biomass and nutrient uptake capacity. Methods: Five temperate perennial pasture grasses–cocksfoot (Dactylis glomerata L. cv. Howlong), phalaris (Phalaris aquatica L. cv. Atlas PG), tall fescue (Festuca arundinacea L. cv. Prosper), tall wheatgrass (Thinopyrum ponticum L. cv. Dundas), and veldt grass (Ehrharta calycina Sm. cv. Mission) were tested in a controlled environment on the growth and nutrition with the microbial consortium inoculant and rock mineral fertiliser. Results: Veldt grass produced the highest shoot and root growth, while tall fescue yielded the lowest. Rock mineral fertiliser with or without microbial consortium inoculant significantly increased root and shoot biomass production across the grass species. The benefit of microbial consortium inoculation applied in conjunction with rock mineral fertiliser was significant regarding shoot N content in tall wheatgrass, cocksfoot and tall fescue. Shoot P and K concentrations also increased in the five grass species by microbial consortium inoculation combined with rock mineral fertiliser in comparison with the control treatment. Arbuscular mycorrhizal (AM) colonisation decreased with rock mineral fertilisation with or without microbial consortium inoculant except in cocksfoot. Conclusions: The response to microbial consortium inoculation, either alone or in combination with rock mineral fertiliser, was plant species‐dependent, indicating its potential use in pasture production.