Resistance to boron toxicity amongst several barley and wheat cultivars: A preliminary examination of the resistance mechanism

The mechanism of resistance toB toxicity in barley and wheat was studied in a solution culture experiment using several cultivars displaying a large range of sensitivity to excessB supply. Plants were cultured for 35 d atB concentrations ranging from normal to excessive (15 to 5000 M, respectively) then examined for dry matter production and theB distribution between roots and shoots.In both species, increasedB supply was accompanied by increased tissueB concentrations, development ofB toxicity symptoms and depressed growth. At each level ofB supply, however, resistant cultivars accumulated considerably lessB than did sensitive cultivars, in both roots and shoots. Even at the lowestB supply, at which noB toxicity symptoms developed and growth was not affected, resistant cultivars maintained relatively low tissueB concentrations. No cultivar displayed an ability to tolerate high tissueB concentrations.These results indicate that sensitivity toB toxicity in barley and wheat is governed by the ability of cultivars to excludeB. If theB concentrations of tissues is used to indicate resistance toB toxicity, then cultivars have the same ranking whether cultured at a normal or excessB supply.     

The control of boron accumulation by two genotypes of wheat

The mechanism of boron (B) uptake in wheat was studied using two genotypes with known differences in their ability to accumulate B. Influx and efflux of B was measured in the roots of intact 21 d old plants.Roots grown in 15 M B, when transferred to solutions containing 1mM B showed a rapid increase in B content for up to 60 min, after which no further increase was evident up to 4 h. No genotypic difference in B influx was apparent over these time periods. Roots grown in 1mM B for 7 d and then rinsed in B-free solutions quickly lost most of B that they contained within 1 hour; little further efflux was observed over the following three hours. As with the influx, no genotypic difference in B flux was evident.It is suggested that the lack of genotypic difference in the short-term B fluxes could be due to a masking effect of extracellular B bound in the cell walls of the roots.Department of Botany, University of Adelaide     

Early Inhibition of Photosynthesis during Development of Mn Toxicity in Tobacco

Early physiological effects of developing Mn toxicity in young leaves of burley tobacco (Nicotiana tabacum L. cv KY 14) were examined in glass-house/water cultured plants grown at high (summer) and low (winter) photon flux. Following transfer of plants to solutions containing 1 millimolar Mn²⁺, sequential samplings were made at various times for the following 9 days, during which Mn accumulation by leaves increased rapidly from ~70 on day 0 to ~1700 and ~5000 microgram per gram dry matter after 1 and 9 days, respectively. In plants grown at high photon flux, net photosynthesis declined by ~20 and ~60% after 1 and 9 days, respectively, and the onset of this decline preceded appearance (after 3 to 4 days) of visible foliar symptoms of Mn toxicity. Intercellular CO₂ concentrations and rates of transpiration were not significantly affected; moreover, the activity of the Hill and photosystem I and II partial reactions of chloroplasts remained constant despite ultimate development of severe necrosis. Though the activity of latent or activated polyphenol oxidase increased in parallel with Mn accumulation, neither leaf respiration nor the activity of catalase [EC 1.11.1.6] and peroxidase [EC 1.10.1.7] were greatly affected. These effects from Mn toxicity could not be explained by any changes in protein or chlorophyll abundance. Additionally, they were not a consequence of Mn induced Fe deficiency. Therefore, inhibition of net photosynthesis and enhancement of polyphenol oxidase activity are early indicators of excess Mn accumulation in tobacco leaves. These changes, as well as leaf visual symptoms of Mn toxicity, were less severe in plants cultured and treated at low photon flux even though the rates of leaf Mn accumulation at high and low photon flux were essentially equivalent.     

Effects of rainfall on the use of foliar analysis for diagnosing boron toxicity in field-grown wheat

The effect of rain on foliage elemental composition, especially B, was assessed using samples of wheat collected at three harvests from a field trial conducted in soil containing excessive levels of B. Moderate rainfall substantially decreased both the B concentration and content of whole shoots and young leaves. The change in B concentration due to rain suggests that foliar analysis is unreliable for diagnosing B toxicity. For the other elements examined (Ca, Cu, Mg, Mn, Mg, P, S, Zn), rainfall had little effect.     

Response of shoot growth and transpiration to soil drying in sugarcane

The relationship between plant-available water (PAW) and shoot extension and transpiration is required to model crop response to water stress, and has not been previously defined for sugarcane (Saccharum spp. (L.)). We subjected sugarcane plants at the 5–6 leaf stage to a continuous drying cycle in large (42 L) pots to determine the threshold fraction of plant available water (PAW_t) at which plants slowed shoot extension and transpiration relative to plants watered daily. Transpiration rate was measured as the daily mass loss from the pots and shoot extension as the height increase from ground level to the tip of the youngest actively expanding leaf. Three experiments were conducted with cultivar Q115 covering a range of soil types (and hence PAW) and rates of soil drying. To compare the response with sugarcane, sorghum (Sorghum bicolor (L.) Moench s.lat.), a species that has been well characterized for the relationship between PAW and transpiration and shoot extension, was grown in two additional experiments. For the same species, response curves and PAW_t for either shoot extension or transpiration were very similar for the different experiments. This similarity occurred despite there being different soils, different environmental conditions, different PAW, different times taken for the pots to dry down, and hence different rates of stress development. In sugarcane, there was almost no threshold in PAW_t (0.92) for shoot extension and a very small threshold in PAW_t for transpiration (0.85), while in sorghum PAW_t for sorghum shoot extension (0.54) and plant transpiration (0.47) were consistent with those published previously. The present data extend previous reports that sugarcane stalk extension is very sensitive to water stress, and we discuss several factors that could provide the physiological basis for the sensitivity. This revised version was published online in June 2006 with corrections to the Cover Date.     

Further evidence that zinc is required throughout the root zone for optimal plant growth and development

The effect of variable Zn supply with depth in a soil profile was examined in two wheat genotypes differing in their Zn efficiency. Gatcher (Zn-inefficient) and Excalibur (Zn-efficient) were grown in a low Zn soil in pots with two treatment zones. The upper zone (10 cm) was supplied with Zn while Zn was either supplied or withheld from the lower zone (25 cm). In both genotypes, withholding Zn from the lower zone had no effect on root growth in either the upper or lower zones; neither did it affect plant appearance prior to booting. However, withholding Zn from the lower zone delayed head emergence in Gatcher by some 10 d and depressed grain yield by 20%. In Excalibur, Zn treatment had no effect on head emergence or grain yield. In Gatcher, withholding Zn from the lower zone depressed water usage by 12% during a 60 d period preceding maturity. No effect of Zn treatment on water usage was seen in Excalibur. Tissue Zn concentrations closely reflected the lower zone Zn treatments in both genotypes. Irrespective of the Zn treatment, Excalibur had higher Zn concentrations in flag leaves but lower concentrations in grain than Gatcher. In whole shoots, genotypic differences in Zn concentration only occurred when Zn was added to the lower zone; Excalibur having almost twice that of Gatcher. Clearly, if Zn is not supplied to the entire root system there is the potential for impaired root function and plant development, and for reduced grain yield. Approaches to managing this problem are discussed.     

Induction of sesquiterpenoid biosynthesis in tobacco cell suspension cultures by fungal elicitor

Large amounts of the sesquiterpenoid capsidiol accumulated in the media of tobacco (Nicotiana tabacum L. cv KY14) cell suspension cultures upon addition of fungal elicitor. Capsidiol accumulation was proportional to the amount of elicitor added. The accumulation of capsidiol was preceded by a transient increase in the capsidiol de novo synthesis rate as measured by the incorporation of exogenous [¹⁴C]acetate. Changes in 3-hydroxy-3-methylglutaryl-CoA reductase activity (HMGR; EC 1.1.1.34), an enzyme of general isoprenoid metabolism, paralleled the changes in [¹⁴C]acetate incorporation into capsidiol. Incubation of the cell cultures with mevinolin, a potent in vitro inhibitor of the tobacco HMGR enzyme activity, inhibited the elicitor-induced capsidiol accumulation in a concentration dependent manner. [¹⁴C]Acetate incorporation into capsidiol was likewise inhibited by mevinolin treatment. Unexpectedly, [³H] mevalonate incorporation into capsidiol was also partially inhibited by mevinolin, suggesting that mevinolin may effect secondary sites of sesquiterpenoid biosynthesis in vivo beyond HMGR. The data indicated the importance of the induced HMGR activity for capsidiol production in elicitor-treated tobacco cell suspension cultures.     

Physiological and genetic control of the tolerance of wheat to high concentrations of boron and implications for plant breeding

Physiological and genetic studies have been undertaken to further the understanding of genetic variation in response to high concentrations of B in the soil and so facilitate the breeding of tolerant varieties for cultivation in high B regions. Genetic variation in response to high concentrations of B has been identified for a number of crop and pasture species of southern Australia, including wheat, barley, oats, field peas and annual pasture medics. The wheat variety Halberd, which was the most widely grown variety in Australia during the 1970s and early 1980s, is the most tolerant of the current Australian wheat varieties. The mechanism of tolerance for all species studied is reduced accumulation of B by tolerant genotypes in both roots and shoots. Results from experiments of uptake kinetics indicate that control of B uptake is a non-metabolic process. The response of wheat to high B supply is under the control of several major additive genes, one of which has been located to chromosome 4A.     

Boron toxicity

Whilst of lesser prevalence than B deficient soils, B-rich soils are important, causing B toxicity in the field and decreased crop yields in different regions of the world. The highest naturally occurring concentrations of soil B are in soils derived from marine evaporites and marine argillaceous sediment. In addition, various anthropogenic sources of excess B may increase soil B to levels toxic for plants. The most important source is irrigation water, but others include wastes from surface mining, fly ash, and industrial chemicals. Ameliorating high-B soils is extremely difficult. A commonly used method of reclaiming high B soils is to extensively leach with low B water. Though used successfully, leaching may not be a permanent solution and causes difficulties with the disposal of the leachates. Other amelioration methods include the use of soil amendments (e.g. lime, gypsum) and the planting of plant genotypes that are tolerant of high external B concentrations. Although there are various methods available to determine the levels of B in soils, soil analysis can provide little more than a general risk assessment for B toxicity. Similarly, diagnosing B toxicity in plants, either by visible symptoms or tissue analysis has limited applicability. Thus at present, neither soil nor plant analysis can be recommended to precisely predict the growth of plants on high soil B. Recent physiological and genetic studies have provided some understanding of genetic variation in the response of plants to high concentrations of B. Moreover, these studies have facilitated the breeding of tolerant genotypes for cultivation on high B soils. Considerable genetic variation in response to high B has been identified in a wide range of plant species, most of which share a similar tolerance mechanism – reduced uptake of B in both shoots and roots. The tolerance mechanism appears to be under the control of several major additive genes, and specific chromosomal locations have been identified for the genes in some species. Considerable success has been achieved in breeding for tolerance to B toxicity, a process that is greatly aided by the ease with which genotypic variation for this characteristic can be assessed and the range of methods available to screen breeding populations.     

Problems associated with the use of foliar analysis for diagnosing boron toxicity in barley

The effects of evapotranspiration conditions on the distribution of B in leaves and critical values for B toxicity measured in shoots were examined in solution culture experiments with barley.Increased water use resulted in increased B accumulation by plants and B was concentrated in the leaf tips. The relationship between shoot DM production and shoot B concentrations was markedly affected by evapotranspiration conditions, but the effect could be removed by not analysing leaf tips. Excluding the leaf tips also decreased the shoot B concentration at which shoot DM production was depressed.Regularly spraying plants with water removed considerable B from leaves without affecting DM production.The present results indicate a number of problems in the establishment of critical values and the use of foliar analysis for diagnosing B toxicity. These problems may account for conflicting reports of critical values and discrepencies between results from glasshouse- and field-cultured plants.