锌肥对不同基因型大麦吸收积累镉的影响

对土壤添加不同Zn、Cd条件下两种基因型(Sahara和Clipper)大麦对Zn、Cd的吸收积累研究表明,在本实验条件下土壤添加Zn、Cd对植物地上部生物量没有显著影响,但土壤添加Zn抑制植物根系生长,在土壤不缺Zn情况下添加Zn<20mg·kg^-1时并没有对大麦体内Cd浓度产生显著影响;当土壤Zn添加量达到40mg·kg^-1时,植物体内Cd浓度明显降低,植物吸收Cd的总量随着土壤添加Zn的增加而显著下降,这主要是由于根系生物量的下降所致,两个基因型大麦品种Zn效率存在显著差异,但这一差异对植物吸收Cd的总量没有影响,Zn高效品种Sahara根部Cd浓度显著低于Clipper。     

The mechanism of boron tolerance for maintenance of root growth in barley (Hordeum vulgare L.)

Cultivar differences in root elongation under B toxic conditions were observed in barley (Hordeum vulgare L.). A significant increase in the length and width of the root meristematic zone (RMZ) was observed in Sahara 3771 (B tolerant) when it was grown under excessive B concentration, compared to when grown at adequate B supply. This coincided with an increase in cell width and cell numbers in the meristematic zone (MZ), whereas a significant decrease in the length and no significant effect on the width of the MZ was observed in Clipper (B intolerant) when it was grown under excessive B supply. This was accompanied by a decrease in cell numbers, but an increase in the length and width of individual cells present along the MZ. Excessive B concentrations led to a significantly lower osmotic potential within the cell sap of the root tip in SloopVic (B tolerant) and Sahara 3771, while the opposite was observed in Clipper. Enhanced sugar levels in the root tips of SloopVic were observed between 48 and 96 h after excess B was applied. This coincided with an increase in the root elongation rate and with a 2.7-fold increase in sucrose level within mature leaf tissue. A significant decrease in reducing sugar levels was observed in the root tips of Clipper under excessive B concentrations. This coincided with significantly lower root elongation rates and lower sucrose levels in leaf tissues. Results indicate a B tolerance mechanism associated with a complex control of sucrose levels between leaf and root tip that assist in maintaining root growth under B toxicity.     

Boron tolerance in barley is mediated by efflux of boron from the roots

Many plants are known to reduce the toxic effects of high soil boron (B) by reducing uptake of B, but no mechanism for limiting uptake has previously been identified. The B-tolerant cultivar of barley (Hordeum vulgare L.), Sahara, was shown to be able to maintain root B concentrations up to 50% lower than in the B-sensitive cultivar, Schooner. This translated into xylem concentrations that were approximately 64% lower and leaf concentrations 73% lower in the tolerant cultivar. In both cultivars, B accumulation was rapid and reached a steady-state concentration in roots within 3 h. In Schooner, this concentration was similar to the external medium, whereas in Sahara, the root concentration was maintained at a lower concentration. For this to occur, B must be actively extruded from the root in Sahara, and this is presumed to be the basis for B tolerance in barley. The extrusion mechanism was inhibited by sodium azide but not by treatment at low temperature. Several anion channel inhibitors were also effective in limiting extrusion, but it was not clear whether they acted directly or via metabolic inhibition. The ability of Sahara to maintain lower root B concentrations was constitutive and occurred across a wide range of B concentrations. This ability was lost at high pH, and both Schooner and Sahara then had similar root B concentrations. A predictive model that is consistent with the empirical results and explains the tolerance mechanism based on the presence of a borate anion efflux transporter in Sahara is presented.     

Redistribution of boron in leaves reduces boron toxicity

High soil boron (B) concentrations lead to the accumulation of B in leaves, causing the development of necrotic regions in leaf tips and margins, gradually extending back along the leaf. Plants vary considerably in their tolerance to B toxicity, and it was recently discovered that one of the tolerance mechanisms involved extrusion of B from the root. Expression of a gene encoding a root B efflux transporter was shown to be much higher in tolerant cultivars. In our current research we have shown that the same gene is also upregulated in leaves. However, unlike in the root, the increased activity of the B efflux transporter in the leaves cannot reduce the tissue B concentration. Instead, we have shown that in tolerant cultivars, these transporters redistribute B from the intracellular phase where it is toxic, into the apoplast which is much less sensitive to B. These results provide an explanation of why different cultivars with the same leaf B concentrations can show markedly different toxicity symptoms. We have also shown that rain can remove a large proportion of leaf B, leading to significant improvements of growth of both leaves and roots.Key words: Bor genes, boron tolerance, boron toxicity, efflux pumping, leaf necrosis, membrane transportB-toxic soils are widespread throughout agricultural areas of the world where they cause significant and often substantial reductions in crop quality and yield. The mechanism by which B is toxic to plants is not well understood¹ but toxicity symptoms include reduced root growth which affects uptake of water and nutrients, and the development of necrotic patches on leaves which impairs photosynthesis. Tolerance to B toxicity has been recognized in a number of crops, notably in cereals. In most cases, tolerance is achieved by reduced uptake of B into the root, which then leads to reduced uptake into the shoot. Genetic studies established that in barley, a locus associated with reduced tissue B occurred on chromosome 4 and that this locus could be transferred to other barley cultivars with desirable agronomic traits.²Hayes and Reid³ made a careful study of the characteristics of B uptake in a highly tolerant landrace barley cultivar Sahara, and found that although B was highly permeable, the root B concentration in this cultivar could be maintained at only half that in the external medium, whereas in sensitive cultivars, B was the same in both intracellular and extracellular phases. It was concluded that tolerant cultivars must have a membrane active transporter that exports B from the root. A B exporter, AtBor1 had previously been discovered in Arabidopsis where it was involved in B loading into the xylem⁴ but it was later found to be degraded under high B conditions⁵ and therefore would not be useful in B tolerance.However, other Bor1 homologues were subsequently discovered in Arabidopsis and in rice. Based on homology with rice, Reid⁶ cloned genes from barley and from wheat (HvBor2 and Tabor2 respectively) which were shown to be strongly upregulated in roots of tolerant cultivars, and virtually undetectable in sensitive cultivars. Thus, a simple mechanism to explain tolerance was established; efflux of B from the root reduced the intracellular concentration of B in the root cells, thereby reducing toxicity and improving root growth. At the same time, the lower root content meant that less B was transferred to the shoot, resulting in lower shoot toxicity.Yet there remained several unanswered questions regarding B toxicity. Firstly, it was commonly observed that toxicity symptoms were not reliably correlated with leaf B concentration, and that often after rain, toxicity symptoms became less severe. Nable et al.⁷ had investigated the effect of rain on shoot B concentrations and concluded that although rain did reduce the B concentration in leaves, it did not affect growth and yield. Secondly, field trials with cultivars in which the B tolerance traits were expressed, did not show the improvements in growth and yield that could be observed in glasshouse trials.^8,9Our recent work¹⁰ has provided new insights into these phenomena. Sensitive and tolerant cultivars of both wheat and barley were grown in varying levels of B. Then, ignoring the level of B in the growth solution, leaves of the different cultivars that displayed the same degree of leaf necrosis were selected. This revealed that in the tolerant cultivars, necrosis began to appear at leaf B levels that were two-to five-fold higher than in sensitive cultivars. Since no internal tolerance mechanism had been reported, it was hypothesised that in the tolerant cultivars, internal toxicity was reduced by pumping B from the cytoplasm into the cell wall where B is much less toxic. To prove this hypothesis three types of experiment were conducted. Firstly protoplasts were isolated from leaves of tolerant and sensitive cultivars of barley, and it was shown that when incubated in the same concentration of B, the tolerant cultivar was able to reduce the intracellular B concentration to approximately half that of the sensitive cultivar. Secondly, it was reasoned that if more B was accumulated in the apoplast of the tolerant cultivar, then it should be more quickly released by washing of the leaf; this was confirmed. Thirdly, it was shown that the same efflux transporters that were responsible for B export from the root were also highly expressed in leaves of tolerant cultivars of wheat and barley. The combination of these three experiments provided compelling evidence that redistribution of B in the leaf was a significant factor in B tolerance.The elution experiment also highlighted the fact that because B is highly soluble and has high membrane permeability, it can easily be washed from leaves. Obviously in the field B could be removed from leaves by rain, but no positive effect of this on growth had been quantified. In our experiments, we simulated the average rainfall during the early growing season in a high B region of Southern Australia by spraying plants with calibrated amounts of water for 16 d. At high B concentrations, rain reduced leaf B by around 50% while simultaneously improving growth of shoots by up to 90%. Rather surprisingly, the rain treatment, which had no significant effect on root B concentrations, caused a two-fold increase in root growth, presumably by improving the supply of photosynthate from the shoot.This study has enabled an evaluation of the importance of three main factors in determining the severity of B toxicity; two genetically determined processes, efflux pumping of B in roots and leaves, coupled with the variable leaching of B from leaves by rain (Fig. 1). The results also provide an explanation for the poor correlations observed between toxicity and shoot B concentrations in cereals.^7,11Open in a separate windowFigure 1Summary of processes contributing to reduced B toxicity in wheat and barley. The intensity of shading indicates the level of B in different regions of the plant. Boron (B) enters the leaf via the xylem and continues to accumulate as the leaf grows. When plants are grown in high concentrations of B, the older parts of the leaf become necrotic first while the younger basal tissues continue to expand. In tolerant cultivars, B efflux transporters in leaves pump B from the cytoplasm where it is toxic into the cell walls where it can be tolerated at high concentrations. Sensitive cultivars have a very low capacity for B efflux and therefore retain much higher concentrations inside the cell than in tolerant cultivars. rain can remove large amounts of B from leaves, thereby alleviating toxicity. In roots of tolerant cultivars, the same B efflux transporters that occur in leaves are used to pump B from the cells into the external medium. This reduces the toxicity to roots and limits the amount of B entering the xylem and reaching the leaves.     

Interaction Between Root and Leaf Disease Development in Barley Cultivars after Inoculation with Different Isolates of Bipolaris sorokiniana

The relationship between root and leaf infection in 11 cultivars of barley ( Hordeum vulgare ) by different isolates of Bipolaris sorokiniana was investigated in young plants. Roots of 10-day-old seedlings, grown in filterpaper rolls, and the third leaf of 17-day-old seedlings were inoculated with the different isolates and a Disease Development Index (DDI) was calculated.
The rate of lesion development in leaves was higher than in roots, indicated by generally higher DDI after leaf inoculation than after root inoculation. Significant differences in resistance were found among the barley cultivars. Inoculation with different isolates of B. sorokiniana caused significant differences in DDI for both roots and leaves. In the leaves, but not in the roots, a significant cultivar–isolate interaction was found. No significant correlations, neither in isolate aggressiveness nor in cultivar reaction between root and leaf, were observed.     

Phosphorus efficiencies and responses of barley (<Emphasis Type="Italic">Hordeum vulgare</Emphasis> L.) to arbuscular mycorrhizal fungi grown in highly calcareous soil

Two experiments were carried out to investigate phosphorus efficiencies and mycorrhizal responsiveness in an improved cultivar (Clipper) and a landrace (Sahara) of barley (Hordeum vulgare L.). In experiment 1, two pot sizes were used to evaluate the effect of soil volume on P uptake and mycorrhizal responsiveness. In experiment 2, a compartmented ("cross-pot") system was used to monitor (32)P delivery by external hyphae of arbuscular mycorrhizal fungi (AMF) to the host plant. Results showed that, irrespective of growth conditions, Sahara had much larger root biomass than Clipper and consequently substantially more P was allocated to roots in Sahara than in Clipper. Specific root length in Clipper was much longer than in Sahara. Increase in soil volume enhanced percentage root length colonised by AMF, plant growth and P uptake, and Sahara was more sensitive to changes in soil volume than Clipper. Pot size (soil volume) used to assess responsiveness to AMF by different plant species or genotypes with different root/shoot ratios might be a confounding factor. Clipper was more responsive to AMF than Sahara in terms of tissue P concentrations, which is partly related to their differences in root/shoot ratios. However, increases in SPU [specific P uptake, mg P (g root biomass)(-1)] caused by AMF were bigger in Clipper, suggesting that AMF played a larger role in P uptake. In accordance with the larger increase in SPU, Clipper took up more (32)P via AMF hyphae than Sahara. The compartmented system using radioactive P might be an alternative approach to directly investigate mycorrhizal responsiveness of different plant species or varieties than conventional pot experiments, provided that the same AM fungus is used.     

Apoplastic peroxidases and ascorbate are involved in manganese toxicity and tolerance of Vigna unguiculata

Excessive manganese (Mn) supply induced the formation of brown spots on leaves as typical Mn toxicity symptoms in cowpea ( Vigna unguiculata L. Walp.) grown in hydroponics. Differences in Mn resistance between cv. TVu 91 (Mn-sensitive) and cv. TVu 1987 (Mn-tolerant) expressed in the density of brown spots in older leaves were due to higher Mn tissue tolerance. Apoplastic water-soluble peroxidase (POD) in the apoplastic washing fluid (AWF) was enhanced by increasing Mn leaf content and generally significantly higher in leaves of cv. TVu 91 than in cv. TVu 1987. Electrophoresis of AWF revealed the presence of several water-soluble POD isoenzymes. At toxic Mn supply, the activities of these and additional POD isoenzymes increased more in the Mn-sensitive cultivar. Levels of ascorbic acid in the apoplast and cytoplasm of the Mn-sensitive cv. TVu 91 decreased with increasing leaf Mn contents, whereas Mn-tolerant cv. TVu 1987 was not affected. Mn treatment lead to a stimulation of the enzymes of the ascorbic acid regeneration system (monodehydroascorbic acid reductase and glutathione reductase) in both cultivars, but the activation of glutathione reductase was clearly more enhanced in the Mn-tolerant cultivar TVu 1987. The results provide circumstantial evidence that apoplastic ascorbate and peroxidases are involved in the expression of Mn toxicity and genotypic Mn tolerance.     

Boron accumulation and partitioning in wheat cultivars with contrasting tolerance to boron deficiency

Two pot experiments at the Plant Environment Laboratory (PEL), Reading, UK investigated sterility, boron (B) accumulation and B partitioning of wheat cultivars grown with limited B in the growing medium. The first experiment evaluated nine cultivars of spring wheat with diverse field responses to low available soil B, supplied with or without 20 μM B. A second experiment examined the response of a susceptible (SW-41) and a tolerant (Fang-60) cultivar to B-deficiency. These cultivars were supplied with either 20 μM B from sowing to flag leaf emergence and no added B thereafter, or 20 μM B from sowing to maturity. When B was not supplied in the nutrient solution, the number of grains ranged from 4 per ear (cv. BL-1135) to 32 per ear (cv. BL-1249) and sterility of competent florets ranged from 39% to 93%. Boron concentration in the flag leaf at anthesis did not differ greatly when the growing medium contained limited B, but differences between cultivars were evident when B was unlimited. Tolerance of B-deficiency was not related to the B concentration in the flag leaf. Some cultivars produced viable pollen and set grains while others failed to do so at similar B concentrations in the flag leaf. The two contrasting cultivars did not differ much in their pattern of B partitioning when B supply was restricted from flag leaf emergence onwards. Similarly, little evidence was found that the tolerant cultivars translocated B from their leaves, roots or stems when the supply in the growing medium was restricted. The proportion of total B partitioned in different organs was the same irrespective of B supply and cultivar. On average, leaves contained 68% of the total B content in the whole plant compared to 16% in the roots, 10% in the ears and only 6% in the stems. Tolerant or susceptible cultivars of wheat could not be distinguished based on the B concentration and B content of the flag leaf. This revised version was published online in June 2006 with corrections to the Cover Date.     

Field screening of barley cultivars to soil salinity using a sprinkler and a drip irrigation system

The establishment of proper agronomical practices and plant breeding programs for saline environments is limited by the lack of adequate field screening methods. We assessed the relationships between leaf ion concentration and grain yield in a set of barley cultivars and compared their ranking for salinity tolerance established with a triple-line-source (TLS) sprinkler system, where the absorption of salts is through the leaves and the roots, with that obtained with a drip-irrigation (DI) system, where the absorption of salts is only through the roots. The saline solution in both systems was made up of sodium and hydrated calcium chloride (1:1 w/w). Except for the highest saline treatments, direct leaf absorption of toxic Na⁺ and Cl^- was minor or negligible, but it was substantial for Ca²+. Irrespective of barley cultivar and leaf age, the accumulation of Cl^- in the TLS was 1.5–2.5 times greater than Na⁺. There was no significant correlation between grain yield and leaf sap ion concentration among eighteen barley cultivars. Thus, leaf ion concentrations should not be used as screening tools in breeding programs for increasing salinity tolerance in barley. The highest-yielding cultivars under non-saline conditions were also most productive under moderately saline conditions, though not under high-saline conditions. Although grain yields of the eighteen barley cultivars in the TLS were substantially lower than in the DI, the salinity tolerances estimated in both systems were significantly correlated (P < 0.05), indicating that the simple and inexpensive TLS irrigation system could be successfully used in screening for salinity tolerance in barley.     

Phenylalanine Ammonia-lyase Activity in Barley After Infection with Bipolaris sorokiniana or Treatment with its Purified Xylanase

Phenylalanine ammonia-lyase (PAL) activity was determined from leaves and roots of two barley (Hordeum vulgare L.) cultivars after infection with a necrotrophic pathogen, Bipolaris sorokiniana (Sacc.) Shoem., and treatment with its purified xylanase. PAL activity increased in leaves of both cultivars 16 h after fungal inoculation but two phases, with activity peaks at 24–32 h and 40 h, were recorded only for the more resistant cultivar, Agneta. Attempts to use a PAL inhibitor, χ-amin, ooxyacetic acid, to increase susceptibility to B. sorokiniana in barley leaves were unsuccessful. Treatments of leaves with purified xylanase resulted in more rapid (4–12 h after injection), although reduced, induction of PAL compared with fungal injection. The higher the concentration of xylanase applied the earlier the activity peaks were detected. Fungal inoculation only slightly increased PAL activity in barley roots while xylanase treatment had no effect. The basal level of PAL was however much higher in roots than in leaves. In wheat, Triticum aestivum L. resistant to B. sorokiniana, the time-course of PAL induction after fungal infection and xylanase treatment resembled that for cv. Agneta, while in oats, Avena sativa L. (non-host) PAL activity did not change after the treatments. The results suggest that the second phase of PAL induction, associated only with responses of barley cv. Agneta and wheat, is linked with their resistance to B. sorokiniana infection. The possible role of xylanase as an elicitor of PAL is discussed.     

Effects of Salinity on the Extensibility and Ca Availability in the Expanding Region of Growing Barley Leaves

The growth of barley (Hordeum vulgare L.) leaves is reduced by salinity. We used the Instron extensometric technique to measure the reversible and irreversible compliance of the expanding regions of growing barley leaves from plants exposed to 1, 40, 80 and 120 mM NaCl in nutrient solution. Two barley cultivars differing in salinity resistance (cv ‘Arivat’ and cv ‘Briggs’) were compared over 5d of leaf growth. During the period of most active leaf expansion, salinity reduced reversible compliance and increased compliance in the leaf segments, although responses to salinity were complex and changed over the course of leaf expansion. Salinity increased irreversible compliance more in the salt-sensitive cultivar Arivat than in the more salt-tolerant cultivar Briggs. Elemental analysis of the basal leaf segments used for extensometry revealed an accumulation of Na and a depletion of Ca in segments from salinized plants, resulting in very high Na: Ca ratios in salinized expanding tissue. The concentrations of K and Mg in basal leaf tissue were elevated by salinity. Our data do support the hypothesis that the inhibition of leaf expansion by salinity stress is mediated by a decline in irreversible extensibility. We suggest that reduced Ca availability in expanding leaf tissue may contribute to growth reduction in salt-stressed barley seedlings.     

Genetic and molecular analyses of resistance to a variant of Puccinia striiformis in barley

Seedlings of 62 Australian barley cultivars and two exotic barley genotypes were assessed for resistance to a variant of Puccinia striiformis, referred to as “Barley Grass Stripe Rust” (BGYR), first detected in Australia in 1998, which is capable of infecting wild Hordeum species and some genotypes of cultivated barley. Fifty-three out of 62 cultivated barley cultivars tested were resistant to the pathogen. Genetic analyses of seedling resistance to BGYR in six Australian barley cultivars and one Algerian barley landrace indicated that they carried either one or two major resistance genes to the pathogen. A single recessive seedling resistance gene, rpsSa3771, identified in Sahara 3771, was located on the long arm of chromosome 1 (7 H), flanked by the restriction fragment length polymorphism (RFLP) markers Xwg420 and Xcdo347 at genetic distances of 12.8 and 21.9 cM, respectively. Mapping resistance to BGYR at adult plant growth stages using the doubled haploid (DH) population Clipper × Sahara 3771 identified two major quantitative trait loci (QTL), one on the long arm of chromosome 3 (3 H) and the second on the long arm of chromosome 1 (7 H), accounting for 26 % and 18 % of the total phenotypic variation, respectively. The QTL located on chromosome 7HL corresponded to seedling resistance gene rpsSa3771 and the second QTL was concluded to correspond to a single APR gene, designated rpsCl, contributed by cultivar Clipper.     

Silicification in sorghum (Sorghum bicolor) cultivars with different drought tolerance

Sorghum belongs to a group of economically important, silicon accumulating plants. X-ray microanalysis coupled with environmental scanning electron microscopy (ESEM) of fresh root endodermal and leaf epidermal samples confirms histological and cultivar specificity of silicification. In sorghum roots, silicon is accumulated mostly in endodermal cells. Specialized silica aggregates are formed predominantly in a single row in the form of wall outgrowths on the inner tangential endodermal walls. The density of silica aggregates per square mm of inner tangential endodermal cell wall is around 2700 and there is no significant difference in the cultivars with different content of silicon in roots. In the leaf epidermis, silicon deposits were present in the outer walls of all cells, with the highest concentration in specialized idioblasts termed 'silica cells'. These cells are dumb-bell shaped in sorghum. In both the root endodermis and leaf epidermis, silicification was higher in a drought tolerant cultivar Gadambalia compared with drought sensitive cultivar Tabat. Silicon content per dry mass was higher in leaves than in roots in both cultivars. The values for cv. Gadambalia in roots and leaves are 3.5 and 4.1% Si, respectively, and for cv. Tabat 2.2 and 3.3%. However, based on X-ray microanalysis the amount of Si deposited in endodermal cell walls in drought tolerant cultivar (unlike the drought susceptible cultivar) is higher than that deposited in the leaf epidermis. The high root endodermal silicification might be related to a higher drought resistance.     

Antioxidant defense system and cadmium uptake in barley genotypes differing in cadmium tolerance

Using two barley (Hordeum vulgare) cultivars (cvs. Tokak and Hamidiye) nutrient solution experiments were conducted in order to study the genotypic variation in tolerance to Cd toxicity based on (i) development of leaf symptoms, (ii) decreases in dry matter production, (iii) Cd concentration and (iv) changes in antioxidative defense system in leaves (i.e., superoxide dismutase, ascorbate peroxidase, glutathione reductase, catalase, ascorbic acid and non-protein SH-groups). Plants were grown in nutrient solution under controlled environmental conditions, and subjected to increasing concentrations of Cd (0, 15, 30, 60 and 120 micromol/L Cd) for different time periods. Of the barley cultivars Hamidiye was particularly sensitive to Cd as judged by the severity and earlier development of Cd toxicity symptoms on leaves. Within 48 h of Cd application Hamidiye rapidly developed severe leaf Cd toxicity symptoms whereas in Tokak the leaf symptoms appeared only slightly. Hamidiye also tended to show more decrease in growth caused by Cd supply when compared to Tokak. The differences in sensitivity to Cd between Tokak and Hamidiye were not related to Cd concentrations in roots and shoots or Cd accumulation per plant. With the exception of catalase, activities of the enzymes involved in detoxification of reactive oxygen species (ROS) were markedly enhanced in Hamidiye by increasing Cd supply. By contrast, in Tokak there was either only a slight increase or no change in the activities of the antioxidative enzymes. Similarly, levels of ascorbic acid and especially non-protein SH-groups were increased in Hamidiye by Cd supply, but not affected in Tokak. The results indicate the existence of a large genotypic variation between barley cultivars for Cd tolerance. The differential Cd tolerance found in the barley cultivars was not related to uptake or accumulation of Cd in plants, indicating importance of internal mechanisms in expression of differential Cd tolerance in barley. As a response to increasing Cd supply particular increases in antioxidative mechanisms in the Cd-sensitive barley cultivar Hamidiye suggest that the high Cd sensitivity of Hamidiye is related to enhanced production and oxidative damage of ROS.     

Antioxidant defense mechanisms in response to cadmium treatments in two safflower cultivars

By using two safflower (Carthamus tinctorius L.) cultivars, Arak2811 and Goldasht, the experiments were conducted in order to study (i) the genotypic variation in cadmium (Cd) tolerance, (ii) Cd concentrations in plants, and (iii) changes in the antioxidant defense systems in leaves, including antioxidant enzymes and nonenzymatic antioxidants. Plants were grown under controlled environmental conditions and subjected to Cd treatments (0, 25, 50, 75, and 100 μM Cd) for different time periods. Cd concentrations and cultivar-dependent response to Cd were assessed. Of the two cultivars, Goldasht showed a greater sensitivity to Cd toxicity as judged from the severity of Cd toxicity symptoms on leaves, much stronger enhancement in the MDA level, and decreases in dry matter production. Increasing Cd supply markedly reduced the shoot and root dry weights in both cultivars, but at the higher Cd concentrations and longer exposure durations, this decrease was more marked in cv. Goldasht. Plants accumulated substantial amount of Cd, especially in the roots, the highest being in the roots of cv. Arak2811 at 100 μM Cd after 4 days. Cd-induced oxidative stress as was indicated by the increase in lipid peroxidation with the increase in metal concentration and exposure duration. Under different Cd stress levels, activities of antioxidant enzymes differed in the two cultivars. The results indicated that Cd tolerance of cv. Arak2811 was related to the retention of Cd in the roots and avoiding the toxic effect by activation of the antioxidant system.     

Correlation of gibberellin-induced growth, polyamine levels and amine oxidases in epicotyl, root and leaf blade of barley seedlings

The potential role of diamine oxidase (DAO) and polyamine oxidase (PAO) in relation to polyamines was investigated in epicotyls, roots and leaf blades at 3 and 6 days after gibberellic acid (GA) application in barley (Hordeum vulgare L.) seedlings of cvs. Maythorpe (non-mutant parent) and Golden Promise (semi-dwarf mutant). There was a significant increase in epicotyl and leaf-blade elongation rates in GA-treated seedlings of cv. Maythorpe as compared to cv. Golden Promise. DAO and PAO were detectable in all segments of the leaf blade, but the highest activities were present in basal segments. These enzymes, which are thought to have a role in the elimination of cellular polyamines, increased in activity following GA application compared to controls. Application of 10⁻⁶ M GA to the first leaf, significantly increased endogenous bound putrescine (Put) levels in both the epicotyl and leaf blade of cv. Maythorpe. In contrast, there was only a slight increase in cv. Golden Promise. Levels of soluble Put increased in roots and leaf blades of both cultivars following GA treatment but the effect was greatest in leaves of cv. Maythorpe. It is suggested that polyamines may play a role in GA-induced epicotyl and leaf-blade elongation in barley.     

Cold-induced accumulation of protein in the leaves of spring and winter barley cultivars

Electrophoretic pattern and quantitative changes in soluble proteins were determined in the leaves of spring and winter cultivars of barley (Hordeum vulgare L., cv. Makouei and cv. Reyhan, respectively) exposed to 4 degrees C for 14 d. Seedlings were grown in a controlled growth chamber for 2 weeks at a constant air temperature of 20 degrees C and then transferred to constant 4 degrees C for 14 d followed by returning to 20 degrees C (cold treatment), or they were maintained throughout at 20 degrees C during the experimental period of 40 d (control treatment). Plants were sampled every 48 h for leaf fresh weight measurements. Total leaf soluble proteins were extracted and their concentration was either determined by a colorimetric method, or size-fractionated on SDS-PAGE. Low temperature-induced increases in protein amount occurred over the second week of exposure to cold treatment irrespective of cultivar: the winter cultivar was 2 d prior in this response. The protein patterns and their density showed differences between-cultivars and between-temperature treatments. A new cold-induced polypeptide was recognized in the leaves of winter barley cultivar on day 22 (8 d at 4 degrees C) compared to the control. This polypeptide was produced earlier over the first 48 h of low temperature in the winter cultivar compared with the spring one, recognizing in the leaves of cold-treated seedling until day 26. This more rapid response to a low temperature by the winter barley cultivar indicates a more sensitive response compared with the spring barley, probably cold-shock protein is a component of this cold-induced response.     

Root responses to cereal cyst nematode (Heterodera avenae) in hosts with different resistance genes

The development of cereal cyst nematode (CCN; Heterodera avenae ) induced syncytia in the host roots of infected resistant bread wheat ( Triticum aestivum cv. AUS10894), diploid wheat ( Aegilops tauschii ), barley ( Hordeum vulgare cv. Chebec and cv. Galleon) and in the susceptible wheat cv. Meering and barley cv. Clipper were studied over a period of 13 d. The resistance to CCN in these cereal plants is conferred by the resistance genes Cre1 in the wheat cv. AUS10894, Cre3 in A. tauschii , Ha2 in barley cv. Chebec and Ha4 in barley cv. Galleon. Anatomical observations were made on the development of the syncytia in CCN-infected wheat and barley roots, which carry each of these four sources of resistance genes. Accelerated development of the syncytia in resistant plants, especially in the barley cultivars, was observed. The sites of syncytia development in susceptible wheat and barley were also closely associated with the vascular tissues in the stele, but less so in the resistant plants. The syncytia in the infected susceptible wheat and barley were also metabolically active at day 13. By contrast, the syncytia of resistant wheat plants carrying the Cre1 or Cre3 genes remained extensively vacuolated and less metabolically active. In barley plants with the Ha2 or Ha4 genes, the syncytia appeared non-functional and in early stages of degeneration by day 13 after inoculation.