Aerenchyma Formed Under Phosphorus Deficiency Contributes to the Reduced Root Hydraulic Conductivity in Maize Roots

Root hydraulic conductivity has been shown to decrease under phosphorus （P） deficiency. This study Investigated how the formation of aerenchyma is related to this change. Root anatomy, as well as root hydraulic conductivity was studied In maize （Zea mays L.） roots under different phosphorus nutrition conditions. Plant roots under P stress showed enhanced degradation of cortical cells and the aerenchyma formation was associated with their reduced root hydraulic conductivity, supporting our hypothesis that air spaces that form in the cortex of phosphorusstressed roots Impede the radial transport of water in a root cylinder. Further evidence came from the variation In aerenchyma formation due to genotypic differences. Five maize inbred lines with different porosity in their root cortex showed a significant negative correlation with their root hydraulic conductivity. Shoot relative water content was also found lower In P-deficient maize plants than that in P-sufficient ones when such treatment was prolonged enough, suggesting a limitation of water transport due to lowered root hydraulic conductivity of P-deficient plants.     

Changes in Root Hydraulic Conductivity During Wheat Evolution

A better understanding of the mechanisms of water uptake by plant roots should be vital for improving drought resistance and water use efficiency (WUE). In the present study, we have demonstrated correlations between root system hydraulic conductivity and root characteristics during evolution using six wheat evolution genotypes (solution culture) with different ploidy chromosome sets (Triticum boeoticum Bioss., T. monococcum L.: 2n=2x=14;T. dicoccides Koern., T. dicoccon (Schrank) Schuebl.:2n=4x=28;T. vulgare Vill., T. aestivum L. cv. Xiaoyan No. 6:2n=6x=42). The experimental results showed that significant correlations were found between root system hydraulic conductivity and root characteristics of the materials with the increase in ploidy chromosomes (2x→6x) during wheat evolution. Hydraulic conductivity of the wheat root system at the whole-plant level was increased with chromosome ploidy during evolution, which was positively correlated with hydraulic conductivity of single roots, whole plant biomass,root average diameter, and root growth (length, area), whereas the root/shoot ratio had an inverse correlation with the hydraulic conductivity of root system with increasing chromosome ploidy during wheat evolution. Therefore, it is concluded that that the water uptake ability of wheat roots was strengthened from wild to modern cultivated species during evolution, which will provide scientific evidence for genetic breeding to improve the WUE of wheat by genetic engineering.     

Effects of Local Nitrogen Supply on Water Uptake of Bean Plants in a Split Root System

To study the effects of local nitrogen supply on water and nutrient absorption, French bean （Phaseolus vulgaris L.） plants were grown in a split root system. Five treatments supplied with different nitrogen forms were compared： homogeneous nitrate （NN） and homogenous ammonium （AA） supply, spatially separated supply of nitrate and ammonium （NA）, half of the root system supplied with N-free nutrient solution, the other half with either nitrate （NO） or ammonium （AO）. The results showed that 10 d after onset of treatments, root dry matter （DM） in the nitratesupplied vessels treated with NA was more than two times higher than that in the ammonium-supplied vessels. Water uptake from the nitrate-supplied vessels treated with NA was 281% higher than under ammonium supply. In treatments NO and AO, the local supply of N resulted in clearly higher root DM, and water uptake from the nitratesupplied vessels was 82% higher than in the -N vessels. However, in AO plants, water uptake from the -N nutrient solution was 129% higher than from the ammonium-supplied vessels. This indicates a compensatory effect, which resulted in almost identical rates of total water uptake of treatments AA and AO, which had comparable shoot DM and leaf area. Ammonium supply reduced potassium and magnesium absorption. Water uptake was positively correlated with N, Mg and K uptake.     

Effect of Osmotic Stress in Early Stages of Ontogenesis on Root Respiration, Growth, Sugar Content, and Cell Injury in Maize Seedlings Differing in Drought Sensitivity

Cultivars of maize （Zea mays L.） with different sensitivity to drought were exposed to 0.3 mol/L sorbitol （-1.4 MPa water potential） for 24 h. Exposure to water deficiency significantly reduced the growth of both shoots （coleoptile and hypocotyl） and roots. Shoot growth was inhibited more than the growth of roots. Osmotic stress enhanced accumulation of soluble sugars. Electrolyte leakage, a cell injury index, was slightly increased after 0.3 mol/L sorbitoh Respiration was measured in the presence and absence of 2,6-dlchloro-phenol indophenoh 2,6-Dichloro-phenol indophenol did not influence respiration rates, because statistically equal results were observed under both conditions. Total respiration （VT） decreased after osmoticum treatment. There were no significant differences in the VT among the cultlvars analysed. The decrease In VT was caused by a decline In the activities and capacities of both cytochrome （Vcyt, Vcyt） and alternative pathway （Valt, Valt） of respiration. A high residual respiration （Vres） was observed, up to 27% of total uninhibited respiration. The result of uncoupler use clearly indicated that coupling was maintained after 24 h of osmotic stress. The recovery of the respiration rate was comparable with that of non-stressed control rates. According to these observations, no possible mltochondrial damage is expected. Water deficiency did not induce a stimulation of the alternative oxidase, so we assume that the stimulation of the alternative pathway is not related to drought stress resistance; rather, the function of the alternative pathway is to balance carbon metabolism and electron transport in a response to a changing environment.     

Root Cluster Formation and Citrate Exudation of White Lupin （Lupinus albus L.） as Related to Phosphorus Availability

A split-root system was used to investigate whether the external or internal P concentration controls root cluster formation and citrate exudation in white lupin (Lupinus albus L.) grown under controlled conditions. In spite of low P concentrations in the shoots and roots of the -P plant, its dry weight was not reduced compared with the P plant. Supplying external P (0.25 mmol/L) to one root half resulted in an increase in P concentration not only in the shoot, but also in the P-deprived root half, indicating P cycling within the plants. Omitting P from both split-root pots stimulated root cluster formation in both root halves,whereas P supply to one root half stimulated root cluster formation at the beginning of the treatment. Neither P supply to just one root half continuously nor resupply of P to one root half after 19 d of P starvation inhibited root cluster formation on the P-deprived side, although the concentration of P in this root half and shoot increased markedly. The results indicate that root cluster formation in L. albus is controlled by both shoot and root P concentrations. The rates of citrate exudation by both root halves with P deficiency were higher than those of the one root half supplied with P only. In the treatment with one root half supplied with P, the rates of citrate exudation by either the P-supplied or -deprived root halves were almost the same,regardless of P concentration in the roots. The results suggest that internal P concentration controls root cluster formation and citrate exudation in white lupin, but these processes may be regulated by different mechanisms.     

Growth,Nitrogen Uptake and Flow in Maize Plants Affected by Root Growth Restriction

The objective of the present study was to investigate the influence of a reduced maize root-system size on root growth and nitrogen （N） uptake and flow within plants. Restriction of shoot-borne root growth caused a strong decrease in the absorption of root ： shoot dry weight ratio and a reduction in shoot growth. On the other hand, compensatory growth and an increased N uptake rate in the remaining roots were observed. Despite the limited long-distance transport pathway in the mesocotyl with restriction of shoot-borne root growth, N cycling within these plants was higher than those in control plants, implying that xylem and phloem flow velocities via the mesocotyl were considerably higher than in plants with an intact root system. The removal of the seminal roots in addition to restricting shoot-borne root development did not affect whole plant growth and N uptake, except for the stronger compensatory growth of the primary roots. Our results suggest that an adequate N supply to maize plant is maintained by compensatory growth of the remaining roots, increased N uptake rate and flow velocities within the xylem and phloem via the mesocotyl, and reduction in the shoot growth rate.     

Iron deprivation results in a rapid but not sustained increase of the expression of genes involved in iron metabolism and sulfate uptake in tomato （Solanum lycopersicum L.） seedlings

Characterization of the relationship between sulfur and iron in both Strategy I and Strategy II plants, has proven that low sulfur availability often limits plant capability to cope with iron shortage. Here it was investigated whether the adaptation to iron deficiency in tomato （Solanum lycopersicum L.） plants was associated with an increased root sulfate uptake and translocation capacity, and modified dynamics of total sulfur and thiols accumulation between roots and shoots. Most of the tomato sulfate transporter genes belonging to Groups 1, 2, and 4 were significantly upregulated in iron-deficient roots, as it commonly occurs under S-deficient conditions. The upregulation of the two high affinity sulfate transporter genes, SlST1.1 and SlST1.2, by iron deprivation clearly suggests an increased root capability to take up sulfate. Furthermore, the upregulation of the two low affinity sulfate transporter genes SlST2.1 and SlST4.1 in iron-deficient roots, accompanied by a substantial accumulation of total sulfur and thiols in shoots of＆amp;nbsp;iron-starved plants, likely supports an increased root-to-shoot translocation of sulfate. Results suggest that tomato plants exposed to iron-deficiency are able to change sulfur metabolic balance mimicking sulfur starvation responses to meet the increased demand for methionine and its derivatives, al owing them to cope with this stress.     

Cultivar variation in the effect of chlorsulfuron in depressing the uptake of copper in wheat

The application of herbicides has induced symptoms of nutrient deficiencies under some circumstances. This glasshouse study examined the effect of chlorsulfuron on the uptake and utilization of copper (Cu) in four cultivars of wheat plants (Triticum aestivum L. cvs. Kulin, Cranbrook, Gamenya and Bodallin) on a Cu-responsive soil. Application of chlorsulfuron depressed the concentration of Cu in wheat plants receiving either inadequate or adequate Cu. In plants with inadequate Cu supply, chlorsulfuron increased the severity of Cu deficiency. Shoot weight was markedly decreased by chlorsulfuron at all levels of Cu, through decreasing the number of tillers and the elongation of leaves. This decreased growth of shoots occurred prior to the effect on Cu concentration in tissues. The retranslocation of Cu in old tissues over time was unaffected by chlorsulfuron. In all wheat cultivars, the decreased growth of shoots were correlated with the concentration of Cu in the youngest fully emerged leaf blade with critical levels of 1.6−1.7 at day 25 and 0.9−1.0 μg g⁻¹ d. wt. at day 60. The application of chlorsulfuron tended to increase the critical level at day 25 but not at day 60. In addition, Kulin seems to be most, and Cranbrook least, sensitive to chlorsulfuron. This sensitivity was associated with the sensitivity of the cultivars to Cu deficiency. It is suggested that chlorsulfuron application induces Cu deficiency in wheat plants mainly due to effects on the uptake of Cu. This revised version was published online in June 2006 with corrections to the Cover Date.     

Genetic Transformation of Tobacco with the Trehalose Synthase Gene from Grifola frondosa Fr. Enhances the Resistance to Drought and Salt in Tobacco

Trehalose is a non-reducing disaccharide of glucose that functions as a protectant in the stabilization of biological structures and enhances the tolerance of organisms to abiotic stress. In the present study, we report on the expression of the Grifolafrondosa Fr. trehalose synthase (TSase) gene for manipulating abiotic stress tolerance in tobacco (Nicotiana tabaccum L.). The expression of the transgene was under the control of two tandem copies of the CaMV35S promoter and was transferred into tobacco by Agrobacterium tumefaciens EHA105. Compared with non-transgenic plants, transgenic plants were able to accumulate high levels of products of trehalose, which were increased up to 2.126-2.556 mg/g FW, although levels were undetectable in non-transgenic plants. This level of trehalose in transgenic plants was 400-fold higher than that of transgenic tobacco plants cotransformed with Escherichia coli TPS and TPP on independent expression cassettes, twofold higher than that of transgenic rice plants transformed with a bifunctional fusion gene (TPSP) of the trehalose-6-phosphate (T-6-P) synthase (TPS) and T-6-P phosphatase(TPP) of E. coil, and 12-fold higher than that of transgenic tobacco plants transformed the yeast TPS1 gene.It has been reported that transgenic plants with E. coil TPS and/or TPP were severely stunted and had morphological alterations of their roots. Interestingly, our transgenic plants have obvious morphological changes, including thick and deep-coloured leaves, but show no growth inhibition; moreover, these morphological changes can restore to normal type in T2 progenies. Trehalose accumulation in 35S-35S:TSase plants resulted in increased tolerance to drought and salt, as shown by the results of tests on drought, salt tolerance, and drought physiological indices, such as water content in excised leaves, malondialdehyde content, chlorophyll a and b contents, and the activity of superoxide dismutase and peroxidase in excised leaves. These results suggest that transgenic plants transformed with the TSase gene can accumulate high levels of trehalose and have enhanced tolerance to drought and salt.     

Effects of the supply levels and ratios of nitrogen and phosphorus on seed traits of Chenopodium glaucum北大核心CSCD

Aims Global nitrogen (N) deposition not only alters soil N and phosphorus (P) availability, but also changes their ratio. The levels and ratios of N and P supply and their interaction may simultaneously influence plant seed traits. However, so far there has been no experiments to distinguish these complex impacts on plant seed traits in the field. Methods A pot experiment with a factorial design of three levels and ratios of N and P supply was conducted in the Nei Mongol grassland to explore the effects of levels and ratios of N and P supply and their interaction on seed traits of Chenopodium glaucum. Important findings We found that the relative contribution (15%–24%) of N and P supply levels in affecting the N concentrations, P concentrations and germination rates of seeds was larger than that (3%–7%) of N:P supply ratios, whereas seed size was only significantly influenced by N:P. Simultaneously, seed N and P concentrations were impacted by the interaction of N and P supply levels and ratios. At the same N:P, decrease in nutrient supply levels increased seed N concentrations, P concentrations and germination rates. N:P supply ratios only had a significant effect on seed size and germination rates under low nutrient levels. Overall, these results indicate that different seed traits of C. glaucum show different sensitivities to N or P limitations, leading to adaptive and passive responses under different nutrient limitations. This study presents the the first field experiment to distinguish the effects of nutrient supply levels, ratios and their interactions on plant seed traits, which provides a new case study on the influences of global N deposition on future dynamics of plant population and community. © Chinese Journal of Plant Ecology.     

Effects of Phosphorus Nutrient on the Hydraulic Conductivity of Sorghum (Sorghum vulgare Pers.) Seedling Roots Under Water Deficiency

Hydroponic experiments were conducted in a growth chamber and changes in the hydraulic conductivity of sorghum (Sorghum vulgare Pers.) roots (Lpr) at the three-leaf stage were measured using the pressure chamber method. Water deficiency was imposed with polyethylene glycol (PEG) 6000 and the phosphorus (P) levels were controlled by complete Hoagland solution with and without P nutrient. The objective of this study was to investigate the effect of P nutrition on root Lpr under water deficiency. The results showed that the Lpr in P deficiency treatments decreased markedly, but the Lpr recovered to the same value as that of control when sufficient P was supplied for 4-24 h. Water deficiency decreased Lpr, but the hydraulic conductivity of the roots with sufficient P supply was still higher than that of plants without P supply. When resuming water supply, the Lpr of the water-deficient plants under P supply recovered faster than that of plants without P supply, which indicates that plants with sufficient P nutrient are more drought tolerant and have a greater ability to recover after drought. The treatment of HgCl2 indicated that P nutrient could regulate the Lpr by affecting the activity and the expression levels of aquaporins.     

Water uptake by roots of Hordeum marinum: formation of a barrier to radial O2 loss does not affect root hydraulic conductivity

The adventitious roots of Hordeum marinum grown in stagnant deoxygenated solution contain a barrier to radial O2 loss (ROL) in basal zones, whereas roots of plants grown in aerated solution do not. The present experiments assessed whether induction of the barrier to ROL influences root hydraulic conductivity (Lpr). Wheat (Triticum aestivum) was also studied since, like H. marinum, this species forms aerenchyma in stagnant conditions, but does not form a barrier to ROL. Plants were grown in either aerated or stagnant, deoxygenated nutrient solution for 21-28 d. Root-sleeving O2 electrodes were used to assess patterns of ROL along adventitious roots, and a root-pressure probe and a pressure chamber to measure Lpr for individual adventitious roots and whole root systems, respectively. Lpr, measured under a hydrostatic pressure gradient, was 1.8-fold higher for individual roots, and 5.6-fold higher for whole roots systems, in T. aestivum than H. marinum. However, there was no difference in Lpr between the two species when measured under an osmotic driving force, when water moved from cell to cell rather than apoplastically. Root-zone O2 treatments during growth had no effect on Lpr for either species (measured in aerobic solution). It is concluded that induction of the barrier to ROL in H. marinum did not significantly affect the hydraulic conductivity of either individual adventitious roots or of the whole root system.     

玉米根系水流导度差异及其与解剖结构的关系

在人工气候室水培条件下,从单根水平研究了不同水分条件下玉米根系水流导度的基因型差异及解剖结构之间的关系.结果表明,抗旱性的杂交种户单四号具有水流导度上的杂种优势现象,不抗旱的父本803根系水流导度最低,3个品种根系水流导度大小为F₁代户单四号＞母本天四＞父本803;水分胁迫普遍降低了根系直径、导管直径和皮层厚度.同时,玉米品种根系的解剖结构和根系水流导度有关,正常水分条件下,根系导管直径与3个玉米品种的根系水流导度呈正相关,胁迫条件下则呈负相关.无论是在胁迫还是正常水分条件下,根系皮层厚度占根系直径的比例与根系水流导度都呈负相关,说明根系皮层是根系吸收水分的主要阻力部位.     

Phosphorus deficiency-induced reduction in root hydraulic conductivity in Medicago falcata is associated with ethylene production

Plants grown in phosphorus-deficient solutions often exhibit disruption of water transport due to reduction in root hydraulic conductivity (Lpr) and enhanced ethylene production. To uncover the relationship between the reduction in Lpr and increase in ethylene production, we investigated effect of phosphorus (P) deficiency on ethylene production and Lpr in legume plants of Medicago falcata L. There was an increase in ethylene production and a reduction of Lpr of M. falcata roots when M. falcata seedlings grown in P sufficient solutions (0.5 mM H₂PO₄^2?) were transferred to P-deficient solutions (5 μM H₂PO₄^2?). Antagonists of ethylene biosynthesis, CoCl₂ and aminoethoxyvinylglycine (AVG), abolished the P deficiency-induced ethylene production. Root hydraulic conductivity of M. falcata seedlings grown in P-sufficient solutions was insensitive to CoCl₂ and AVG, while the two chemicals enhanced Lpr for those grown in P-deficient solutions, suggesting that P deficiency-induced decrease in Lpr can be reversed by inhibiting ethylene production. Ethylene precursor 1-amino cyclopropane-1-carboxylic acid (ACC) and ethylene donor ethephon had greater inhibitory effect on Lpr of P-sufficient seedlings than that of P-deficient seedlings. Root hydraulic conductivity of P-sufficient seedlings was more sensitive to HgCl₂ than that of P-deficient seedlings. Taken together, these findings suggest that ethylene induced by P deficiency may play an important role in modulation of root hydraulic conductivity by affecting aquaporins in plants.     

向日葵根系水通道蛋白活性与苗龄关系的研究

利用压力室结合水通道蛋白抑制剂氯化汞(HgCl2)检测了不同苗龄(15d、25d和35d)向日葵根系水通道的活性,结果显示此生长期间根系导水率保持相对恒定,但0．1mmol／L氯化汞使所有苗龄根系的水流速率和根系导水率迅速降低,而降幅随根龄的增大而增大,表明向日葵根存在调节水分进入根系的水通道蛋白,其活性随根龄的增大而提高,质外体水流随根龄的增大而减小。结论是：在根系生长过程中,细胞到细胞途径水通道蛋白活性的提高可以补偿由于质外体途径导水度降低所致根系导水率的降低,从而维持根系导水率的相对稳定。     

Plant morphology and root hydraulics are altered by nutrient deficiency in Pistacia lentiscus (L.)

The plants in arid and semiarid areas are often limited by water and nutrients. Morpho-functional adjustments to improve nutrient capture may have important implications on plant water balance, and on plant capacity to withstand drought. Several studies have shown that N and P deficiencies may decrease plant hydraulic conductance. Surprisingly, studies on the implications of nutrient limitations on water use in xerophytes are scarce. We have evaluated the effects of strong reductions in nitrogen and phosphorus availability on morphological traits and hydraulic conductance in seedlings of a common Mediterranean shrub, Pistacia lentiscus L.. Nitrogen deficiency resulted in a decrease in aboveground biomass accumulation, but it did not affect belowground biomass accumulation or root morphology. Phosphorus-deficient plants showed a decrease in leaf area, but no changes in aboveground biomass. Root length, root surface area, and specific root length were higher in phosphorus-deficient plants than in control plants. Nitrogen and phosphorus deficiency reduced both root hydraulic conductance and root hydraulic conductance scaled by total root surface area. On the other hand, nutrient limitations did not significantly affect root conductance per unit of foliar surface area. Thus, adaptation to low nutrient availability did not affect seedling capacity for maintaining water supply to leaves. The implications for drought resistance and survival during seedling establishment in semi-arid environments are discussed.     

Colonisation with the arbuscular mycorrhizal fungus Glomus mosseae (Nicol. & Gerd.) enhanced phosphorus uptake from dry soil in Sorghum bicolor (L.)

The aim of the present study was to quantify the contribution of AMF to phosphorus (P) nutrition of the host plant when the P availability in the soil was limited by drought. To investigate the potential of AMF hyphae in taking up P from dry soil, mycorrhizal [+M] and nonmycorrhizal [?M]Sorghum bicolor L. plants were grown in a vertical split root system that consisted of two compartments placed upon one the other. The upper compartment was filled with well fertilised soil and the plant roots were allowed to grow into the lower compartment through a perforated bottom. The lower compartment was filled with an expanded clay substrate and nutrient solution, to supply the plants with water and all nutrients except P. The soil in the upper compartments was either dried [?W] or kept moist [+W] during a period of four weeks before harvest. The total plant P content did not differ significantly between the [?M] and the [+M] plants within the [+W] treatment. In contrast, the P content of the [+M] plants was almost twice as high as the [?M] plants when the soil in the upper compartment was dried. The concentrations of all elements except P in plant shoot tissue were sufficient for adequate plant growth. Phosphorus concentrations in the shoots of [?M/?W] plants indicated P deficiency, and these plants also had significantly lower dry matter and transpiration compared to the plants in all other treatments. From the results of the present experiment it can be concluded that mycorrhizal colonisation seems to be particularly benefical to P uptake from dry soil     

Water Transport in Onion (Allium cepa L.) Roots (Changes of Axial and Radial Hydraulic Conductivities during Root Development)

The hydraulic architecture of developing onion (Allium cepa L. cv Calypso) roots grown hydroponically was determined by measuring axial and radial hydraulic conductivities (equal to inverse of specific hydraulic resistances). In the roots, Casparian bands and suberin lamellae develop in the endodermis and exodermis (equal to hypodermis). Using the root pressure probe, changes of hydraulic conductivities along the developing roots were analyzed with high resolution. Axial hydraulic conductivity (Lx) was also calculated from stained cross-sections according to Poiseuille's law. Near the base and the tip of the roots, measured and calculated Lx values were similar. However, at distances between 200 and 300 mm from the apex, measured values of Lx were smaller by more than 1 order of magnitude than those calculated, probably because of remaining cross walls between xylem vessel members. During development of root xylem, Lx increased by 3 orders of magnitude. In the apical 30 mm (tip region), axial resistance limited water transport, whereas in basal parts radial resistances (low radial hydraulic conductivity, Lpr) controlled the uptake. Because of the high axial hydraulic resistance in the tip region, this zone appeared to be "hydraulically isolated" from the rest of the root. Changes of the Lpr of the roots were determined by measuring the hydraulic conductance of roots of different length and referring these data to unit surface area. At distances between 30 and 150 mm from the root tip, Lpr was fairly constant (1.4 x 10-7 m s-1 MPa-1). In more basal root zones, Lpr was considerably smaller and varied between roots. The low contribution of basal zones to the overall water uptake indicated an influence of the exodermal Casparian bands and/or suberin lamellae in the endodermis or exodermis, which develop at distances larger than 50 to 60 mm from the root tip.