Chickpea genotypes differ in their sensitivity to Zn deficiency

Zinc (Zn) deficiency is common in most of the chickpea growing areas of the world and growing Zn-efficient genotypes on Zn-deficient soil is a benign approach of universal interest. Response of 13 chickpea genotypes (10 desi and 3 kabuli) to Zn nutrition was studied in a pot experiment under glasshouse conditions. Plants were grown in a Zn-deficient siliceous sand for 6 weeks and fertilized with 0 (Zn–) and 2.5 mg Zn per kg soil (Zn+). When grown with no added Zn, Zn deficiency symptoms (chlorosis of younger leaves and stipules followed by necrosis of leaf margins) appeared 3–4 weeks after planting and were more apparent in cultivars Tyson, Amethyst and Dooen than Kaniva and T-1587. Zn deficiency reduced shoot growth, but it was less affected in breeding lines T-1587 and CTS 11308 than cultivars Tyson, Dooen, Amethyst and Barwon. Among all genotypes, Tyson produced the lowest root dry weight in Zn– treatment. Zinc efficiency based on shoot dry weight showed marked differences among genotypes; breeding lines CTS-60543, CTS-11308 and T-1587 were 2-fold more Zn-efficient than cultivars Tyson and Dooen. A higher Zn accumulation per plant and higher Zn uptake per g. of root dry weight were recorded in T-1587 and CTS-11308 when compared with Tyson. Root:shoot ratio was increased and proportionally more Zn was transported to the shoot when the soil was deficient. Cultivars that were very sensitive to Zn deficiency tended to have their root:shoot ratio increased by Zn deficiency more than less sensitive cultivars. The insensitive lines T-1587 and CTS-11308 transported more than 70% of the total absorbed Zn to the shoot. It is concluded that chickpea genotypes vary in their sensitivity to Zn deficiency. Advanced breeding lines T-1587 and CTS-11308 are relatively more Zn-efficient compared with Australian chickpea cultivar Tyson. Zn efficiency in chickpea genotypes is probably related to an efficient Zn absorption coupled with a better root to shoot transport.     

Assessment of the Zn status of chickpea by plant analysis

Chickpea (Cicer arietinum L.) is extensively grown in areas where soils are deficient in zinc (Zn). To determine the response of chickpea to Zn nutrition and to diagnose Zn status in plant tissue, two glasshouse experiments were conducted using Zn-deficient siliceous sandy soil. In Experiment 1, two genotypes of desi chickpea (Dooen and Tyson) were grown at five Zn levels (0, 0.04, 0.2, 1.0 and 5.0 mg kg^-1 of soil). After 4 weeks, no difference in growth and no visible symptoms of Zn deficiency were detected. After 6–8 weeks of growth, chlorosis of younger leaves and stipules occured in the Zn₀ treatment, with shoot dry weight being only 24% of that recorded at the highest Zn level. Root growth increased from 0.52 g/plant when no Zn was applied to 1.04 g/plant in the treatment with 0.2 mg Zn kg^-1 of soil; no response to further increase of Zn fertilization occurred. Zinc concentration in the whole shoot increased significantly with increased in Zn application. The critical Zn concentration in the shoot tissue, associated with 90% of maximum growth, was 20 mg kg^-1 for both genotypes at flowering stage.In the second experiment, two genotypes of desi chickpea (Tyson and T-1587) were grown at three Zn levels (0, 0.5 and 2.5 mg kg^-1 of soil) under two moisture regimes (field capacity 12% w/w, and water stress 4% w/w). Shoot growth was influenced by both Zn supply and water stress. The effect of water stress was severe in the 0.5 and 2.5 mg Zn treatments where shoot dry matter was reduced 52 and 46%, respectively. T-1587 was less sensitive to Zn deficiency and produced higher shoot dry weight than Tyson in the Zn₀ treatment. Zinc concentration in shoots increased from 5 mg kg^-1 when no Zn was applied to 40 mg kg^-1 at the highest Zn level. The critical Zn concentration in shoots was 21 mg kg^-1.The results of the two experiments showed that the critical concentration for Zn did not differ amongst the three cultivars used and was not affected by soil moisture. Similar studies should be undertaken with a wider number of genotypes to discover if a critical concentration of 20–21 mg kg^-1 in the shoot can be used to diagose the Zn status of chickpea genotypes.     

Seed zinc content influences early vegetative growth and zinc uptake in oilseed rape (Brassica napus and Brassica juncea) genotypes on zinc-deficient soil

Low-Zn seed (around 80 ng Zn per seed) and high-Zn seed (around 160 ng Zn per seed) of Zhongyou 821 (a traditional Brassica napus genotype from China found to be Zn-inefficient in our previous experiments), Narendra (Zn-efficient B. napus genotype from Australia) and CSIRO-1 (a Zn-efficient B. juncea genotype from Australia) oilseed rape genotypes were sown in pots containing Zn-deficient siliceous sand fertilized with low Zn supply (0.05 mg Zn kg^–1 soil) or high Zn supply (2.0 mg Zn kg^–1 soil) in a controlled environment. After six weeks, plants derived from the high-Zn seed had better seedling vigour, increased root and shoot growth, more leaf area and chlorophyll concentration in fresh leaf, and higher Zn uptake in shoot compared to those from low-Zn seed at low Zn supply; the impact of high-Zn seed was more marked in Zhongyou 821 compared with CSIRO-1 and Narendra. The influence of high-Zn seed was dissipated at high Zn supply. CSIRO-1 was superior in terms of shoot dry matter production and Zn uptake in shoots at low Zn supply. The results demonstrate that although oilseed rape has very small seeds (about 3 mg per seed weight) compared with wheat (30 mg per seed weight), Zn reserves present in this very small seed still have a strong impact on early vegetative growth as well as on Zn uptake of plants in Zn-deficient soils. The results suggest that sowing high-Zn seed coupled with growing Zn-efficient genotypes may help in sustaining the production of oilseed rape in Zn-deficient soils, and this has implications for improved seed technology.     

Zinc efficiency is correlated with enhanced expression and activity of zinc-requiring enzymes in wheat

Zinc (Zn) is an essential micronutrient for plants. The ability of plants to maintain significant yields under low Zn is termed Zn efficiency (ZE) and its genetic and mechanistic basis is still not well understood. Previously, we showed that root Zn uptake did not play a role in ZE. In the current study, Zn-efficient and -inefficient wheat (Triticum aestivum) genotypes were grown for 13 d in chelate buffer nutrient solutions at low (0.1 pM), sufficient (150 pM), and high (1 microM) Zn(2+) activities and analyzed for root-to-shoot translocation of Zn, subcellular leaf Zn distribution, and activity and expression of the Zn-requiring enzymes in leaves. No correlation between ZE and Zn translocation to the shoot was found. Furthermore, total and water-soluble concentrations of leaf Zn were not associated with ZE, and no differences in subcellular Zn compartmentation were found between Zn-efficient and -inefficient genotypes. However, the expression and activity of the Zn-requiring enzymes copper (Cu)/Zn superoxide dismutase (SOD) and carbonic anhydrase did correlate with differences in ZE. Northern analysis suggested that Cu/ZnSOD gene expression was up-regulated in the Zn-efficient genotype, Kirgiz, but not in inefficient BDME. Under Zn deficiency stress, the very Zn-efficient genotype Kirgiz and moderately Zn-efficient Dagdas exhibited an increased activity of Cu/ZnSOD and carbonic anhydrase when compared with Zn-inefficient BDME. These results suggest that Zn-efficient genotypes may be able to maintain the functioning of Zn-requiring enzymes under low Zn conditions; thus, biochemical Zn utilization may be an important component of ZE in wheat.     

The role of shoot-localized processes in the mechanism of Zn efficiency in common bean

Zn efficiency (ZE) is the ability of plants to maintain high yield under Zn-deficiency stress in the soil. Two bean (Phaseolus vulgaris L.) genotypes that differed in ZE, Voyager (Zn-efficient) and Avanti (Zn-inefficient), were used for this investigation. Plants were grown under controlled-environment conditions in chelate-buffered nutrient solution where Zn²⁺ activities were controlled at low (0.1 pM) or sufficient (150 pM) levels. To investigate the relative contribution of the root versus the shoot to ZE, observations of Zn-deficiency symptoms in reciprocal grafts of the two genotypes were made. After growth under low-Zn conditions, plants of nongrafted Avanti, self-grafted Avanti and reciprocal grafts that had the Avanti shoot scion exhibited Zn-deficiency symptoms. However nongrafted and self-grafted Voyager, as well as reciprocal grafts with the Voyager shoot scion, were healthy with no visible Zn-deficiency symptoms under the same growth conditions. More detailed investigations into putative shoot-localized ZE mechanisms involved determinations of leaf biomass production and Zn accumulation, measurements of subcellular Zn compartmentation, activities of two Zn-requiring enzymes, carbonic anhydrase and Cu/Zn-dependent superoxide dismutase (Co/ZnSOD), as well as the non-Zn-requiring enzyme nitrate reductase. There were no differences in shoot tissue Zn concentrations between the Zn-inefficient and Zn-efficient genotypes grown under the low-Zn conditions where differences in ZE were exhibited. Shoot Zn compartmentation was investigated using radiotracer (⁶⁵Zn) efflux analysis and suggested that the Zn-efficient genotype maintains higher cytoplasmic Zn concentrations and less Zn in the leaf-cell vacuole, compared to leaves from the Zn-inefficient genotype under Zn deficiency. Analysis of Zn-requiring enzymes in bean leaves revealed that the Zn-efficient genotype maintains significantly higher levels of carbonic anhydrase and Cu/ZnSOD activity under Zn deficiency. While these data are not sufficient to allow us to determine the specific mechanisms underlying ZE, they certainly point to the shoot as a key site where ZE mechanisms are functioning, and could involve processes associated with Zn compartmentation and biochemical Zn utilization.Abbreviations CA Carbonic anhydrase - NR Nitrate reductase - SOD Superoxide dismutase - ZE Zinc efficiency This research was supported by a graduate fellowship awarded to G.H. by The Republic of Turkey     

Wheat genotypes differ in Zn efficiency when grown in chelate-buffered nutrient solution

Ten Triticum aestivum and two Triticum turgidum conv. durum genotypes were grown in chelate-buffered nutrient solution at Zn supplies ranging from deficient to sufficient (free Zn activities from 2 to 200 pM, pZn from 11.7 to 9.7). The critical level of Zn ion activity in solution for healthy growth of wheat plants was around 40 pM. Genotypes differed in the growth response: those classified as Zn-efficient suffered less reduction of shoot growth and did not change the rate of root growth at a Zn supply quite deficient for Zn-inefficient genotypes. Root growth of Zn-inefficient genotypes increased at deficient Zn supply. The shoot/root ratio was the most sensitive parameter of Zn efficiency; Zn-efficient genotypes showed less reduction in the ratio when grown at deficient compared to sufficient Zn supply. Classification of wheat genotypes into Zn-efficient and Zn-inefficient groups after screening in chelate-buffered nutrient solution corresponded well with classification obtained in field experiments on Zn-deficient soil.     

Effect of bicarbonate and root zone temperature on uptake of Zn,Fe, Mn and Cu by different rice cultivars (Oryza sativa L.) grown in calcareous soil

Pot experiments were conducted with a calcareous soil (Inceptisol) to elucidate the effects of bicarbonate (0 and 20 mM) and root zone temperature (15° and 25°C) on the uptake of Zn, Fe, Mn and Cu by "Zn-efficient" and "Zn-inefficient" rice cultivars. Bicarbonate decreased concentrations and total uptake of Zn in shoots of "Zn-inefficient" cultivars, especially of IR 26 at 25°C, but not in Zn-efficient cultivars. Bicarbonate decreased concentrations and uptake of Fe in shoots of Zn inefficient cultivars, particularly in IR 26. Concentrations and total uptake of Mn were lower in bicarbonate treatment in the Zn-inefficient cultivars at 15°C, and in all cultivars at 25°C. However, concentration and uptake of Cu were not affected by bicarbonate in all cultivars. Compared to the 25°C root zone temperature, the concentrations and total uptake of both Zn and Cu in shoots at 15°C were lower in Zn-inefficient than in the Zn-efficient cultivars. The results indicate that Zn-efficiency in rice is causally related to high tolerance of plant to elavated bicarbonate concentrations in soil solution.     

Response of Contrasting Rice Genotypes to Zinc Sources under Saline Conditions

Abiotic stresses are among the major limiting factors for plant growth and crop productivity. Among these, salinity is one of the major risk factors for plant growth and development in arid to semi-arid regions. Cultivation of salt tolerant crop genotypes is one of the imperative approaches to meet the food demand for increasing population. The current experiment was carried out to access the performance of different rice genotypes under salinity stress and Zinc (Zn) sources. Four rice genotypes were grown in a pot experiment and were exposed to salinity stress (7 dS m⁻¹), and Zn (15 mg kg⁻¹ soil) was applied from two sources, ZnSO₄ and Zn-EDTA. A control of both salinity and Zn was kept for comparison. Results showed that based on the biomass accumulation and K⁺/Na⁺ ratio, KSK-133 and BAS-198 emerged as salt tolerant and salt sensitive, respectively. Similarly, based on the Zn concentration, BAS-2000 was reported as Zn-in-efficient while IR-6 was a Zn-efficient genotype. Our results also revealed that plant growth, relative water content (RWC), physiological attributes including chlorophyll contents, ionic concentrations in straw and grains of all rice genotypes were decreased under salinity stress. However, salt tolerant and Zn-in-efficient rice genotypes showed significantly higher shoot K⁺ and Zn concentrations under saline conditions. Zinc application significantly alleviates the harmful effects of salinity by improving morpho-physiological attributes and enhancing antioxidant enzyme activities, and the uptake of K and Zn. The beneficial effect of Zn was more pronounced in salt-tolerant and Zn in-efficient rice genotypes as compared with salt-sensitive and Zn-efficient genotypes. In sum, our results confirmed that Zn application increased overall plant’s performance under saline conditions, particularly in Zn in-efficient and tolerant genotypes as compared with salt-sensitive and Zn efficient rice genotypes.     

内生真菌感染对宿主植物高羊茅锌耐受性的影响

以感染内生真菌(Neotyphodium coenophialum)和未感染内生真菌的高羊茅(Festuca arundinacea Schreb.)为实验材料,在营养液中加入ZnSO4进行锌胁迫实验,分析内生真菌对宿主植物锌耐受性的影响。与未感染内生真菌的植株相比,内生真菌感染对高羊茅的总生物量没有显著增益作用,但增加了分蘖数和叶片延伸生长累积值。内生真菌感染降低了高羊茅中Zn2+的总含量,改变了锌在高羊茅中的分配,增加叶鞘中锌的含量,减少叶片中锌的含量。在高锌浓度下,内生真菌感染对净光合速率的变化没有影响,但是显著提高了其宿主的PSⅡ光化学效率(Fv/Fm)。总体来看,内生真菌感染改善宿主高羊茅的锌耐受性。     

Distribution and remobilization of Zn and Mn during grain development in wheat

Wheat (Triticum aestivum cv. Aroona) was grown in siliceoussand with essential nutrients for unlimited growth except forthe following treatments: controls (sufficient Zn and Mn), lowMn (sufficient Zn) and low Zn (sufficient Mn) until anthesis.Replicate plants were harvested at anthesis; the remaining plantswere transferred to a chelate-buffered nutrient solution containingall essential nutrients except Zn and Mn to allow monitoringof the remobilization of existing Zn and Mn reserves withinthe plant. These plants were harvested 14 d post-anthesis andat grain maturity. At each harvest plants were separated intoindividual components. There were no growth differences between any of the treatmentsat the three harvests. Large amounts of Zn and Mn found in theroots and stems at anthesis were rapidly depleted during graindevelopment. The Zn content of the leaves increased from anthesisto 14 d post-anthesis, but then declined. The Mn content ofthe leaves increased throughout grain development in the controlswhilst remaining constant in the plants pre-grown at low Mn.The Zn and Mn content of the glumes, palea and lemma rose inthe controls from anthesis to 14 d post-anthesis; thereafterZn content declined but Mn content continued to increase. TheZn and Mn content of the grain rose sharply toward grain maturity.We conclude that Mn was not remobil-ized from the leaves ofwheat during grain development. Zinc was remobilized from theleaves, especially the flag leaf and from the leaves of thelow Zn plants. The post-anthesis accumulation of Zn and Mn withinthe glumes will be discussed in relation to the transport pathwaythat Zn and Mn use to enter the developing seed. Key words: Zinc, manganese, wheat, distribution, remobilization     

微量元素硼和锌在作物营养平衡中的作用

许多土壤中B、Zn由于含量或有效性低而供给不足,成为作物营养平衡与大面积产量提高的重要限制因素。采取各种方法直接施用B、Zn,或者配制在尿素中施用,可以调节作物营养平衡,改善作物-环境生态关系,提高产量和品质。在水稻旱育苗地区还可把Zn配在育苗土壤调制剂中施用。在缺Zn褐土上,施入的Zn迅速转化为各种形态,其中碳酸盐结合态是有效Zn的主要仓库。配合施用P肥对Zn的有效性无不良影响,Zn与N、P配合施用有益于作物营养平衡。     

Discrimination Against 13CO2 in Leaves,Pod Walls,and Seeds of Water-stressed Chickpea

The rate of photosynthesis (P _N) in leaves and pods as well as carbon isotope content in leaves, pod walls, and seeds was measured in well-watered (WW) and water-stressed (WS) chickpea plants. The P _N, on an area basis, was negligible in pods compared to leaves and was reduced by water stress (by 26%) only in leaves. WS pod walls and seeds discriminated less against ¹³CO₂ than did the controls. This response was not observed for leaves as is usually the case. Pod walls and seeds discriminated less against ¹³CO₂ than did leaves in both WW and WS plants. Measurement of carbon isotope composition in pods may be a more sensitive tool for assessing the impact of water stress on long-term assimilation than is the instantaneous measurement of gas exchange rates.     

Sulfhydryl groups in root-cell plasma membranes of wheat genotypes differing in Zn efficiency

Sulfhydryl groups were quantified in root-cell plasma membranes of two genotypes of wheat ( Triticum aestivum cv. Warigal and T. turgidum conv. durum cv. Durati) differing in Zn efficiency. Smaller amounts of 5,5'-dithio-bis(2-nitrobenzoic acid)-reactive sulfhydryl groups were found in Zn-deficient than in Zn-sufficient roots; and also in Zn-inefficient genotype Durati compared to Zn-efficient Warigal, regardless of Zn supply. Upon transfer of 15-day-old Zn-deficient plants into solutions containing various Zn²⁺ activities, a Zn-dependent increase in the amount of reactive sulfhydryl groups was evident in roots of both genotypes, but occurred only in Warigal when 20-day-old plants were used, indicating irreversible physiological damage in Durati plants due to prolonged Zn deficiency. Upon transfer into solutions of increasing Zn²⁺ activities, the increase in total Zn concentration in roots was about an order of magnitude smaller than the increase in amounts of reactive sulfhydryl groups in the roots of both genotypes, suggesting that, in wheat roots, a relatively small amount of Zn is required for preventing oxidation of sulfhydryl groups into disulfides. The amount of reactive sulfhydryl groups in the roots is positively related to Zn efficiency of wheat genotypes and may be one of the mechanisms that, under conditions of Zn deficiency, allow better growth and productivity of Zn-efficient genotypes in comparison to Zninefficient ones.     

‘On-farm’ seed priming with zinc in chickpea and wheat in Pakistan

A series of on-station trials was implemented between 2002 and 2006 to assess the response of wheat (Triticum aestivum L.) and chickpea (Cicer arietinum) to zinc (Zn) added by soaking seeds (priming) in solutions of ZnSO₄ before sowing. Wheat seed was primed for 10 h in 0.3% Zn and chickpea for 6 h in 0.05% Zn. Seed treatments increased the seed concentration in wheat from 27 to 470 mg/kg and in chickpea from 49 to 780 mg/kg. Priming wheat seeds with 0.3% Zn significantly increased the mean shoot dry mass, Zn concentration and Zn uptake of 15-day-old seedlings relative to non-primed controls and seeds primed with water alone. Using 0.4% Zn further increased shoot Zn concentration but depressed shoot dry mass to the level of the non-primed control. In seven trials, mean grain yield of wheat was significantly increased from 2.28 to 2.42 t/ha (6%) by priming with water alone and to 2.61 t/ha (14%) by priming with 0.3% Zn. Mean grain yield of chickpea in seven trials was increased significantly from 1.39 to 1.65 t/ha (19%) by priming seeds with 0.05% Zn. The effect of priming chickpea seeds with water was intermediate (1.49 t/ha) and not statistically separable from the non-primed and zinc-primed treatments. Increased grain yield due to priming in both crops was associated with increases in total biomass but there was no significant effect of priming on harvest index. In addition to increasing yield, priming seeds with Zn also significantly increased grain zinc concentration, by 12% in wheat (mean of three trials) and by 29% in chickpea (one trial) and the total amount of Zn taken up by the grain (by 27% in wheat and by 130% in chickpea). Using ZnSO₄ to prime seeds was very cost-effective, with net benefit-to-cost ratios of 75 for wheat and 780 for chickpea. An erratum to this article can be found at     

Improved nitrogen nutrition enhances root uptake, root-to-shoot translocation and remobilization of zinc ((65) Zn) in wheat

This study focussed on the effect of increasing nitrogen (N) supply on root uptake and root-to-shoot translocation of zinc (Zn) as well as retranslocation of foliar-applied Zn in durum wheat (Triticum durum). Nutrient solution experiments were conducted to examine the root uptake and root-to-shoot translocation of (65) Zn in seedlings precultured with different N supplies. In additional experiments, the effect of varied N nutrition on retranslocation of foliar-applied (65) Zn was tested at both the vegetative and generative stages. When N supply was increased, the (65) Zn uptake by roots was enhanced by up to threefold and the (65) Zn translocation from roots to shoots increased by up to eightfold, while plant growth was affected to a much smaller degree. Retranslocation of (65) Zn from old into young leaves and from flag leaves to grains also showed marked positive responses to increasing N supply. The results demonstrate that the N-nutritional status of wheat affects major steps in the route of Zn from the growth medium to the grain, including its uptake, xylem transport and remobilization via phloem. Thus, N is a critical player in the uptake and accumulation of Zn in plants, which deserves special attention in biofortification of food crops with Zn.     

Uptake of zinc from chelate-buffered nutrient solutions by wheat genotypes differing in zinc efficiency

Zinc-efficient Triticum aestivum (cv. Warigal) and Zn-inefficientTriticum turgidum conv. durum (cv. Durati) were grown in chelate-buffered,complete nutrient solutions providing either deficient or sufficientZn supply. When transferred to fresh chelatebuffered nutrientsolutions containing a wide range of Zn supplies (0–1.28µmol m^–3 Zn²⁺ activity) for 24–48 h, bothgenotypes increased net Zn uptake linearly with an increasein solution Zn²⁺ activities. Zincefficient Warigal accumulatedZn at a greater rate than Zn-inefficient Durati. The greaterrate of net Zn uptake was observed by plants of both genotypeswhen pretreated at deficient Zn supply. Net loss of Zn to thesolution was higher in plants pretreated with sufficient Znand was inversely related to Zn²⁺ activity in the external solution.When continuously supplied with 40 nmol m^–3 Zn²⁺, netZn uptake by Zn-efficient Warigal was significantly greaterthan that of Zn-inefficient Durati, but the difference diminishedwith plant age. Shoot concentrations of Fe, Mn and Cu were higherwhen plants were grown at deficient than at sufficient Zn supply.The Zn-efficient genotype transported less Zn and Fe to shootsand had higher Fe concentrations in roots than the Zn-inefficientgenotype, supporting the hypothesis that Zn efficiency may beconnected with inefficient transport of Fe from roots to shootsand thus initiation of the Fe-deficiency response resultingin increased release of Zn- and Fe-binding phytosiderophores.It is concluded that differential Zn efficiency of wheat genotypesis at least partly due to a greater ability of efficient genotypesto accumulate Zn. Key words: Chelate-buffering, genotypes, micronutrients, Triticum spp., uptake, zinc efficiency     

Traffic and industrial activities around Riyadh cause the accumulation of heavy metals in legumes: A case study

The objective of this study was to analyse the effect of the continuously increasing anthropogenic activities around Riyadh, Saudi Arabia on the accumulation of heavy metals in leguminous crops. This study determined whether four legume crops, Pisum sativum L., Vicia faba L., Glycine max and Vigna sinensis, could accumulate the heavy metals Cu, Mn, Pb and Zn in their leaves, pods and grains during the summer when grown under conditions with ambient air pollution from heavy traffic and industrial activities in Riyadh, Saudi Arabia. The effect of the air pollution was examined by quantifying the protein and trace element Cu, Mn, Pb and Zn concentrations in the leaves, pods and grains of the four plant species. Analysis of the results indicated that air pollution significantly increased the heavy metal concentrations in the leaves, pods and grains. Toxic concentrations of the heavy metals were found in the plants grown at L3, L4 and L5. In conclusion, the air pollution increases as the traffic, industrial activities and population density increase.     

Alfalfa genotypes differ in their ability to tolerate zinc deficiency

Response of 13 alfalfa (Medicago sativa L.) genotypes to varied Zn supply (+Zn: 2 mg kg⁻¹ soil, −Zn: no added Zn) was studied in a pot experiment under controlled environmental conditions. Plants were grown for four weeks in a Zn-deficient siliceous sandy soil. Plants grown at no added Zn showed typical Zn deficiency symptoms i.e. interveinal chlorosis of leaves, yellowish-white necrotic lesions on leaf blades, necrosis of leaf margins, smaller leaves and a marked reduction in growth. There was solute leakage from the leaves of Zn-deficient plants, while no solute leakage from Zn-sufficient plants. The ratios of P:Zn, Fe:Zn, Cu:Zn and Mn:Zn in Zn-deficient plants were extremely high compared with Zn-sufficient plants indicating disturbance of P:Zn, Fe:Zn, Cu:Zn and Mn:Zn balance within plant system by Zn deficiency. Genotypes differed markedly in Zn efficiency based on shoot dry matter production. Alfalfa genotypes also differed markedly in P:Zn ratio, Cu:Zn ratio and Fe:Zn ratio under —Zn treatment. The shoot dry weight, shoot:root ratio, chlorophyll content of fresh leaf tissue, solute leakage from the leaves, Zn uptake and distribution of Zn in shoots and roots were the most sensitive parameters of Zn efficiency. Zn-efficient genotypes had less solute leakage but higher shoot:root ratio and higher Zn uptake compared with Zn-inefficient genotypes. Under —Zn treatment, Zn-inefficient genotypes had less Zn partitioning to shoots (33–37%) and more Zn retained in roots (63–67%), while Zn-efficient genotypes had about equal proportions of Zn in roots (50%) and shoots (50%). This revised version was published online in June 2006 with corrections to the Cover Date.     

Gas exchange by pods and subtending leaves and internal recycling of CO(2) by pods of chickpea (Cicer arietinum L.) subjected to water deficits

Terminal drought markedly reduces leaf photosynthesis of chickpea (Cicer arietinum L.) during seed filling. A study was initiated to determine whether photosynthesis and internal recycling of CO(2) by the pods can compensate for the low rate of photosynthesis in leaves under water deficits. The influence of water deficits on the rates of photosynthesis and transpiration of pods and subtending leaves in chickpea (cv. Sona) was investigated in two naturally-lit, temperature-controlled glasshouses. At values of photosynthetically active radiation (PAR) of 900 micromol m(-2) s(-1) and higher, the rate of net photosynthesis of subtending leaves of 10-d-old pods was 24 and 6 micromol m(-2) s(-1) in the well-watered (WW) and water-stressed (WS) plants when the covered-leaf water potential (Psi) was -0.6 and -1.4 MPa, respectively. Leaf photosynthesis further decreased to 4.5 and 0.5 micromol m(-2) s(-1) as Psi decreased to -2.3 and -3.3 MPa, respectively. At 900--1500 micromol m(-2) s(-1) PAR, the net photosynthetic rate of 10-d-old pods was 0.9-1.0 micromol m(-2) s(-1) in the WW plants and was -0.1 to -0.8 micromol m(-2) s(-1) in the WS plants. The photosynthetic rates of both pods and subtending leaves decreased with age, but the rate of transpiration of the pods increased with age. The rates of respiration and net photosynthesis inside the pods were estimated by measuring the changes in the internal concentration of CO(2) of covered and uncovered pods during the day. Both the WW and WS pods had similar values of internal net photosynthesis, but the WS pods showed significantly higher rates of respiration suggesting that the WS pods had higher gross photosynthetic rates than the WW pods, particularly in the late afternoon. When (13)CO(2) was injected into the gas space inside the pod, nearly 80% of the labelled carbon 24 h after injection was observed in the pod wall in both the WW and WS plants. After 144 h the proportion of (13)C in the seed had increased from 19% to 32% in both treatments. The results suggest that internal recycling of CO(2) inside the pod may assist in maintaining seed filling in water-stressed chickpea.     

Mobilization of Cu and Zn by root exudates of dicotyledonous plants in resin-buffered solutions and in soil

It has been frequently suggested that root exudates play a role in trace metal mobilization and uptake by plants, but there is little in vivo evidence. We studied root exudation of dicotyledonous plants in relation to mobilization and uptake of Cu and Zn in nutrient solutions and in a calcareous soil at varying Cu and Zn supply. Spinach (Spinacia oleracea L.) and tomato (Lycopersicon esculentum L.) were grown on resin-buffered nutrient solutions at varying free ion activities of Cu (pCu 13.0–10.4) and Zn (pZn 10.1–6.6). The Cu and Zn concentrations in the nutrient solution increased with time, except in plant-free controls, indicating that the plant roots released organic ligands that mobilized Cu and Zn from the resin. At same pCu, soluble Cu increased more at low Zn supply, as long as Zn deficiency effects on growth were small. Zinc deficiency was observed in most treatment solutions with pZn ≥ 9.3, but not in nutrient solutions of a smaller volume/plant ratio in which higher Zn concentrations were observed at same pZn. Root exudates of Zn-deficient plants showed higher specific UV absorbance (SUVA, an indicator of aromaticity and metal affinity) than those of non-deficient plants. Measurement of the metal diffusion flux with the DGT technique showed that the Cu and Zn complexes in the nutrient solutions were highly labile. Diffusive transport (through the unstirred layer surrounding the roots) of the free ion only could not explain the observed plant uptake of Cu and of Zn at low Zn²⁺ activity. The Cu and Zn uptake by the plants was well explained if it was assumed that the complexes with root exudates contributed 0.4% (Cu) or 20% (Zn) relative to the free ion. In the soil experiment, metal concentrations and organic C concentrations were larger in the solution of planted soils than in unplanted controls. The SUVA of the soil solution after plant growth was higher for unamended soils, on which the plants were Zn-deficient, than for Zn-amended soils. In conclusion, root exudates of dicotyledonous plants are able to mobilize Cu and Zn, and plants appear to respond to Zn deficiency by exuding root exudates with higher metal affinity.