In vitro evidence of aluminum effects on solution movement through root cell walls

Little is known about the primary effects of aluminum (Al) in reducing root growth. However, the sorption of Al by the root cell wall, particularly by calcium (Ca) pectate, has been suggested as being important in the expression of Al toxicity in plants. To overcome problems arising from the close proximity of root cell components that may react with Al, a synthetic Ca pectate membrane was prepared as a model system for Al studies. Solution containing 1 mM Ca (as CaCl2) was passed through the membrane, and the flow rate measured. Solution containing 29 μM Al (as AlCl3) and 1 mM Ca reduced solution flow rate by > 80% from c. 3.5 to c. 0.6 mL/min within 2 min, with a further slight decline over the next 4 min. The Al concentration in solution proximate to the inlet side of the membrane decreased to 15 μM within 10 min, and only 3 μM Al was measured in solution that had passed through the membrane. These results suggest that an important primary effect of toxic Al is a reduction in water movement into the root, with consequent effects on water relations in the plant.     

铝胁迫下植物根尖细胞壁组成物质变化抑制根伸长的生理机制研究

根伸长受抑制是植物受铝毒害的主要症状,铝诱导的细胞壁组成物质的变化是其主要原因。本文主要对铝胁迫下植物根尖细胞壁组成物质如木质素、 胼胝质、 纤维素、 半纤维素、 果胶、 细胞壁多糖蛋白及相关代谢酶类在铝胁迫下的变化对根伸长的影响及生理机理的研究进展进行了综述,明确了铝胁迫诱导的植物根尖细胞壁组成物质含量、 比例及结构的变化导致细胞壁刚性降低,从而抑制细胞伸长,最终抑制根伸长。本文还指出,鉴于缺乏对同一植物甚至同一个种类的植物根尖细胞壁各主要组成物质铝胁迫下变化的系统研究,不能对造成该植物根伸长受抑制的原因做出全面合理的解释,所以今后应侧重于铝胁迫下各细胞壁组分变化在抑制根伸长中的贡献率的研究,尤其要针对主要粮食作物进行系统研究,以有效解决铝胁迫造成的产质量降低。     

Nitrogen Uptake and Allocation in Sweet Viburnum During a Root Growth Flush

Most woody ornamentals exhibit episodic growth flushes and nitrogen (N) uptake has also been demonstrated to be seasonal. However, there is little information on N uptake in relationship to plant growth cycle. In this study, N uptake and allocation of sweet viburnum during periods of low and high root elongation rates were studied. Plants were fertilized with ammonium nitrate (¹⁵NH₄ ¹⁵NO₃) and after 6 d N absorption was determined. Significantly more N was absorbed by plants with low root elongation rate compared with plants with high root elongation rate. About 70% of the N absorbed by plants with low root elongation rate was allocated to the mature leaves compared to 35% on plants with high root elongation rate. It was evident that root growth activity influenced N absorption and allocation. Although only a small amount of the N absorbed by plants with low root elongation rates was allocated to the immature leaves, significantly more N was allocated to the immature leaves by plants with high root elongation rates. It is possible that the N necessary to support immature leaf growth, when root elongation rate is low, is provided by mobilization from other parts, possibly mature leaves.     

高铵胁迫对拟南芥根系向重性的影响及机制初探

向重性是决定植物根系构型的主要因素之一,对根系锚定及水分养分吸收具有重要影响.铵显著影响拟南芥根构型,尽管高铵抑制主根伸长和侧根数,然而铵对主根生长方向的研究却少有报道.本文以拟南芥为材料,利用室内培养基模拟试验,研究了高铵胁迫对根系向重性生长的影响,发现高铵胁迫不仅抑制植物生长,还影响主根的生长方向.0～30h动态观测结果表明:10 mmol/L (NH4)2SO4对根系的向重性反应仅表现为延缓效应,而30 mmol/L(NH4)2SO4则显著减弱主根的向重性弯曲.降低基本培养基养分浓度后,铵对主根向重性反应的抑制效应可发生在较低的NH4+浓度.外源添加Ca2+可部分恢复铵对主根伸长的抑制效应,却不能缓解铵对主根向重性反应的影响.不同部位分开供铵,结果显示铵对拟南芥主根向重性的影响主要是根部铵效应.上述结果表明,铵对主根向重性的影响机制部分独立于其对主根伸长的影响,也不是由于Ca2+缺乏引起的,而有可能与根部高铵诱导的根尖其他生理活动过程改变有关.     

根系对酸胁迫的应激反应及硼调控耐酸机制的研究进展

过量施用氮肥在提高作物产量的同时加速了土壤酸化进程，近20年间我国农田土壤pH下降了0.42个单位。过多的H+可破坏根系细胞壁结构稳定性，诱导细胞死亡，进而抑制根系伸长，降低其对养分和水分的吸收，限制农产品产量和品质的提高。因此，探究植物对酸胁迫的应激反应机制，对制定缓解酸胁迫措施十分必要。本文综述植株内部调控和耐受酸胁迫的响应，以及硼提高植株耐酸性的机制。通常，植物通过Ca2+和K+通道进行的信号传导可感应介质pH的改变，并同时在基因和蛋白水平进行调控，从而快速响应酸胁迫；近期多项研究表明，硼可调控植物根系有机酸分泌，通过改变细胞壁特性来维持细胞壁结构稳定，并通过刺激乙烯和Ⅲ类过氧化物酶 (CIII Prxs) 代谢等共同作用缓解植物酸胁迫。目前，对植物酸胁迫的研究逐步深入，但酸胁迫下植物根系代谢产物变化及诱导机制尚不清楚，响应低pH的特定基因尚不明确，仍需进一步研究。     

Time course study of aluminum-induced callose formation in barley roots as observed by digital microscopy and low-vacuum scanning electron microscopy

To clarify the mechanism(s) involved in the short-term inhibition of root elongation by AI, we monitored the morphological changes of barley roots by digital microscopy. Within 30 min after exposure to 37 µM AI, the surface of the root epidermis in the region of a distance of 1.5 mm from the root tip became rough and began to show signs of damage. After 38 min, callose was rapidly excreted from the junction between the root cap and the root epidermis, and formed a spherical lump approximately 60 µm in diameter. The fine structure of the callose deposits on the root surface was analyzed by low-vacuum scanning electron microscopy. After 50 min, there was a significant increase in the callose contents in the distal 0.6 mm part. At the same time, root elongation stopped completely. Fluorescence staining indicated that callose was localized on the surface of the cell elongation area (the elongation zone of primary roots and root hairs), but not on the surface of the meristem. The root growth reduction associated with AI treatment may be due to the use of sugar substrates for callose formation instead of cellulose formation.     

Growth and functioning of roots and of root systems subjected to soil compaction. Towards a system with multiple signalling?

The effect of soil mechanical impedance on root growth is discussed on several levels from the apex to the root system. At the individual root level, the balance of pressures on the root apex cannot account for observed reductions in root elongation rate. Furthermore, soil mechanical impedance affects the elongation rate of non-impeded organs, such as leaves or non-impeded roots. A chemical message originating in roots could account for such an effect, probably via changes in cell wall rheological properties in all growing zones of the plant. Changes in carbon allocation could also have a role. At a whole-plant level, indirect effects linked to changes in the plant structure contribute in a major way to the effect of mechanical impedance on root growth. Although only a small proportion of roots of fieldgrown plants are in contact with compact soil, geometrical characteristics of root systems are considerably affected. In particular, root deepening is delayed and roots tend to have a clumped spatial arrangement. Experimental evidence and modelling suggest that this change in root system architecture could cause water stress, even in relatively wet soil, because of an increase in resistance to the soil-root water flux. As a consequence, root water status and water flux decrease, and stomatal conductance is reduced as a consequence of a chemical message originating in the roots. This secondary message is superimposed onto the direct message linked to mechanical impedance. Under some climatic conditions, whole-plant growth rate, carbon allocation and phenologic development can then be significantly affected by a compaction in the ploughed layer, while only small changes can be expected under more favourable conditions.     

镉诱导拟南芥根尖过氧化氢积累导致植物根生长抑制

以模式植物拟南芥为材料研究了植物主根对不同浓度镉胁迫的响应。结果表明,随镉浓度的升高,植物主根生长受到明显抑制,胎盘兰染色表明高剂量的镉造成主根根尖细胞死亡。进一步二氨基联苯胺(DAB)染色发现镉胁迫诱导植物根尖大量积累过氧化氢,而在胁迫培养基中加入维生素C可显著改善植物根的生长、降低过氧化氢积累,并减少镉诱导的根尖细胞死亡。上述结果表明,镉胁迫诱导的拟南芥主根生长抑制很可能是由于根尖细胞过氧化物积累所致。     

铝胁迫对不同耐铝小麦品种根伸长生长影响的研究

为探讨铝胁迫抑制根生长的机理,以耐铝型小麦品种ET8和铝敏感型ES8为试验材料,研究了铝胁迫对小麦根相对伸长率,根尖细胞显微结构的影响以及细胞壁木质素含量及苯丙氨酸解氨酶(PAL)、肉桂醇脱氢酶(CAD)、过氧化物酶(POD)活性的变化。结果表明,ET8和ES8经50μmol/L铝胁迫6、122、4 h后,根相对伸长率随铝胁迫时间延长而变小。利用植物显微技术发现,ET8和ES8经50μmol/L胁迫24 h后,根尖伸长区皮层细胞变扁平,细胞间隙变小,细胞壁褶皱,并呈齿轮状交合;ES8细胞受伤害程度较ET8显著。经50μmol/L铝胁迫6、12、24 h后,ET8和ES8根尖细胞长度受铝胁迫的程度随时间延长而加强,根尖细胞相对长度与根相对伸长率呈显著正相关的关系(r=0.9911**)。50μmol/L铝胁迫24 h后,ET8和ES8根尖苯丙氨酸解氨酶(PAL)、肉桂醇脱氢酶(CAD)、过氧化物酶(POD)活性及细胞壁木质素合成显著增加。上述结果表明,铝胁迫通过增加苯丙氨酸解氨酶、肉桂醇脱氢酶、过氧化物酶活性,促进根尖细胞壁木质素合成,加快细胞的木质化,细胞壁延展性变小,从而抑制细胞的伸长,减小根的生长。由于铝胁迫下ES8中木质素合成显著高于ET8,且根尖细胞结构受破坏较ET8显著,造成铝胁迫下ES8根生长受抑制比ET8显著,是ET8较ES8耐铝胁迫的主要原因。     

Boron-Calcium Synergically Alleviates Aluminum Toxicity in Wheat Plants (Triticum aestivum L.)

The effects of B and Ca treatments on root growth, nutrient localization and cell wall properties in wheat ( Triticum aestivum L.) plants with and without Al stress were investigated. Seedlings were grown hydroponically in a complete nutrient solution for 7 d and then treated with B (0, 40 μM), Ca (0, 2,500 μM), and Al (0, 100 μM) in a 500 μM CaCl₂ solution for 8 d. The cell wall materials (CWM) were extracted with a phenol: acetic acid: water (2:1:1 w/v/v) solution and used for subsequent pectin extraction with trans -1,2-diami-nocyclohexane- N,N,N,N -tetraacetic acid (CDTA) and Na₂CO₃ solutions. Boron, Ca, and B + Ca treatments enhanced root growth by 19.5, 15.2, and 27.2%, respectively, compared to the control (pH 4.5). Calcium and B+Ca treatments enhanced root growth with Al stress by 43 and 54%, respectively, while B did not exert any effect. The amounts of CWM and pectin per unit of root fresh weight increased by Al treatment, whereas the Ca and B+Ca treatments slightly reduced the contents of these components. Seventy-four percent of total B, 69% of total Ca, and 85% of total Al were located in the cell wall in the B, Ca, and Al treatments, respectively and 32% of total B, 33% of total Ca, and 33% of total Al were located in the CDTA-soluble and Na₂CO₃-soluble pectin fractions. A more conspicuous localization of B was observed in the presence of Al. Aluminum treatment markedly decreased the Ca content in the cell wall as well as pectin fractions, mainly in the case of the CDTA-soluble pectin fraction. Boron + Ca treatment decreased the Al content in the cell wall and pectin fractions compared to the Ca treatment alone in the presence of Al. It is concluded that the B+Ca treatment enhanced root growth and, B and Ca uptake, and helped to maintain a normal B and Ca metabolism in the cell walls even in the presence of Al.     

A mechanistic model for the inhibiting effects of aluminium on the uptake of cations

Reported laboratory observations mention inhibiting effects of Al on uptake of cations. However, the diverse conditions of these experiments do not allow conclusions to be drawn about the extent of this inhibiting effect. An uptake model for Ca and Mg was therefore developed to study the interactions of Al and these cations. The model is based on surface complexation of the root surface with both specific adsorption reactions and exchange reactions included. Application of this model to experiments using a nutrient flux growth technique showed that the observed effects are well simulated. The behaviour of the model agreed with reported inhibiting effects of Al. The model showed that Al effects might be reversed by increased nutrient solution concentrations of cations. However, there exists a strong competitive interaction between Ca and Mg. Therefore, high Ca concentrations have a negative effect on the uptake of Mg. Plants with a high growth rate and a high shoot to root ratio appear to be more sensitive to Al effects than more slowly growing plants.     

硝态氮供应下植物侧根生长发育的响应机制

旱地土壤上硝态氮是作物吸收和利用的主要无机氮形态。硝态氮不仅是植物营养的主要氮源,而且还可以作为信号物质调节植物根系生长发育。为适应土壤中硝态氮非均衡供应,植物侧根发育往往呈现出可塑性反应。本文综述了植物侧根生长发育对硝态氮供应的响应机制。在拟南芥、玉米、大麦等植物上研究表明,硝态氮对植物侧根发育具有双向调节途径,即:1)局部供应硝态氮,硝态氮自身作为信号物质通过信号传导通路发生作用,对侧根具有伸长的刺激效应,硝态氮转运蛋白AtNRT1.1作用于转录因子ANR1的上游,ANR1的转录调节侧根发育;2)植物组织中高浓度的硝态氮对侧根分裂组织活动具有抑制效应,植物激素如生长素和脱落酸可能参与其中的信号传导过程。近些年来研究发现小RNA也参与调控硝态氮供应下植物侧根发育。     

Root to shoot translocation of macronutrients in relation to shoot demand in maize (Zea mays L.) grown at different root zone temperatures

Root growth and nutrient uptake rates of maize (Zea mays L.) are decreased at low root zone temperatures (RZT) and thus, shoot growth may be limited by nutrient deficiency. The objectives of this research were to characterize the shoot demand for nutrients per unit root at suboptimal RZT and to relate net translocation rates of N, P, K, and Ca from the roots to the shoot to shoot demand. Maize plants were grown for 11 days in soil or 8 days in nutrient solution at uniform shoot (24°/20°C, day/night) but different RZT (12°, 18°, and 24°C). The shoot base of the plants (apical shoot meristem and zone of leaf extension) was either kept within or above the cooled root zone. Shoot and root growth were significantly reduced at suboptimal RZT (12°, 18°). Lifting the shoot base above the cooling zone increased shoot growth markedly, whereas root growth was not significantly influenced. Thus, the shoot fresh weight increment day^?1 g^?1 root fresh weight (i.e. the shoot demand per unit root) was increased by a factor of up to 9 for plants with their shoot base above as compared to within the cooling zone. At suboptimal RZT, translocation rates of N, K, and Ca to the shoot remained low in plants with the shoot base in the cooling zone but were higher than in 24°C-grown plants, when the shoot base was above the cooling zone. In both nutrient solution- and soil-grown plants translocation rates of N, K, and Ca were closely correlated with the shoot demand per unit root but less to RZT. In contrast, the translocation rate of P was mainly affected by RZT but insensitive to shoot demand and, therefore, was always higher at a RZT of 24° than of 12°C. From these results it is suggested, that at low RZT the root-to-shoot translocation rates of N, K, and Ca are mainly determined by the shoot demand, whereas the translocation rate of P, regardless of the shoot demand, is reduced by a direct effect of low temperature on the roots.     

柑橘果实发育中果胶酸钙、草酸钙和果胶动态的研究

以单性结实的龟井蜜柑和自花结实的鄂柑1号橘为试材,对整个果实发育期的子房(幼果)、果皮和果肉的果胶酸钙、草酸钙和果胶含量变化进行了测定。结果表明,1)两品种子房(幼果)果胶酸钙含量呈类似的下降趋势;草酸钙则相反,龟井花后趋下降,而鄂柑1号却明显上升;而且鄂柑1号子房(幼果)果胶酸钙、草酸钙和果胶含量均相对较高。2)在果实增大期内,两品种果皮和果肉的果胶酸钙含量均出现显著上升,对应果皮草酸钙含量虽有波动但居相对较高水平,而果肉草酸钙则趋明显下降。3)两品种果皮和果肉水溶性果胶含量均在增大期内呈显著上升,对应原果胶含量均相对较高,进入增大后期均明显下降。     

Biomass and Accumulation of Potassium,Calcium, and Magnesium in Gladiolus as Affected by Heat Units and Corm Size

In México, gladiolus is an important crop cultivated in the central part of the country. In ornamental geophytes, the size of bulbs is a critical factor, impacting plant growth and quality. The objective of the present study was to model the accumulation of potassium (K), calcium (Ca) and Magnesium (Mg) in gladiolus plants as affected by corm size (3.5, 3.0 and 2.5 cm). Regardless of corm size, total dry mass of plants accumulated in three different stages. The initial stage occurred during the corm sprouting phase and continued through the vegetative phase, ending either shortly before (plants from 3.5 cm corms), at (plants from 3.0 cm corms) or shortly after (plants from 2.5 cm corms) the heading phase. The second stage, on which the plants exhibited a rapid dry mass accumulation, occurred shortly before or shortly after the heading phase, finishing with the elongation of the flowering stem. The third stage showed the highest dry mass accumulation rate, coinciding with the elongation of the flowering stem and ending at the blooming of the spike. To complete all the phenological phases, gladiolus required 1818 heat units regardless of corm size. Not considering the nutrients provided by the corm, gladiolus required a total of 6.33, 5.59 and 5.01 mmol plant^?1 of K, 2.22, 1.69 and 1.38 mmol plant^?1 of Ca and 2.49, 2.00 and 1.68 mmol plant^?1 of Mg when grown from 3.5, 3.0, and 2.5 cm corms. At the beginning of the vegetative phase, plants tended to increase Ca content at the expense of K, whereas the proportion of Mg remained unaffected, however, between the vegetative and heading phases, the proportion of Ca declined while that of K and Mg increased. At the end of the study, the proportion of K was much higher than that of Ca and Mg, whereas the proportion of Ca and Mg was similar.     

Aluminium Induced Magnesium Deficiency in Oats

It was the objective to study the effect of Al on Mg uptake by plants, precluding as far as possible the effect of Al on root growth. Oat plants were grown in a complete standard nutrient solution without any differential treatment, in order to obtain a set of plants which did not differ in the size, the morphology and the physiology of the root system. After the first harvest at the beginning of the stem elongation stage 4 different treatments were introduced: pH 5.5-6.0, pH 5.5-6.0 without Mg, pH 3.8-4.1, pH 3.8-4.1 + 0.3 mmole Al/l. Apart from these variations the composition of the nutrient solution remained unaltered. After another 10 days 2 vessels of each treatment were harvested. The final harvest was 14 days after the beginning of the differential treatments. The growth (in terms of dry matter yield) of neither the shoots nor the roots was adversely affected by the differential treatments, although the plants in the Al and Mg₀ treatments showed distinct symptoms of nutritional disorder. The plants in the low and the high pH treatments differed neither in Mg uptakte nor in Mg concentration in the plants. However, the addition of Al to the nutrient solution reduced Mg uptake in the shoots to about 30% of that in the Al₀ treatments, while there was a net loss of Mg in the roots in spite of the fact that dry matter increased. This means that net uptake of Mg was less than was translocated to the shoot during the period of differential treatments. With no Al in the nutrient solution the Mg concentration in the shoots declined by 3–8% between the first and the final harvest, whereas it increased by 22–35% in the roots. If, however, Al was added to the nutrient solution the Mg concentration dropped by 46% in the shoots and 70% in the roots. With the exception of Ca in the roots, the differential treatments had no effect on the uptake and concentration of Ca, K and P in the plants. In terms of dry matter the differential treatments did not influence root growth and it was concluded that Al had a direct effect on Mg uptake by either inactivating or competing for uptake sites or carriers.     

NAA对大白菜吸收和积累~(45)Ca的效应

用ＮＡＡ处理大白菜根部和叶部,对其吸收和转运＾４５Ｃａ的变化进行了研究。结果表明,不论根部施用ＮＡＡ或叶面喷洒ＮＡＡ均使大白菜根对＾４５Ｃａ的吸收增加,提高茎叶＾４５Ｃａ的积累,尤其心叶中＾４５Ｃａ的积累显提高,同是改变心叶＾４５Ｃａ分布,使心叶叶缘的＾４５Ｃａ积累明显增加,且根部施用效果比叶面喷洒效果更明显。     

Effects of Zinc Application on Growth,Absorption and Distribution of Mineral Nutrients Under Salinity Stress in Soybean (Glycine Max L.)

The purpose of the present work was to evaluate effects of zinc application on growth and uptake and distribution of mineral nutrients under salinity stress [0, 33, 66, and 99 mM sodium chloride (NaCl)] in soybean plants. Results showed that, salinity levels caused a significant decrease in shoot dry and fresh weight in non-zinc application plants. Whereas, zinc application on plants exposed to salinity stress improved the shoot dry and fresh weight. Potassium (K) concentration, K/sodium (Na) and calcium (Ca)/Na ratios significantly decreased, while sodium (Na) concentration increased in root, shoot, and seed as soil salinity increased. Phosphorus (P) concentration significantly decreased in shoot under salinity stress. Moreover, calcium (Ca) significantly decreased in root, but increased in seed with increased salinization. Iron (Fe) concentration significantly decreased in all organs of plant (root, shoot, and seed) in response to salinity levels. Zinc (Zn) concentration of plant was not significantly affected by salinity stress. Copper (Cu) concentration significantly decreased by salinity in root. Nonetheless, manganese (Mn) concentration of root, shoot, and seed was not affected by experimental treatments. Zinc application increased Ca/Na (shoot and seed) ratio and K (shoot and seed), P (shoot), Ca (root and seed), Zn (root, shoot, and seed) and Fe (root and shoot) concentration in soybean plants under salinity stress. Zinc application decreased Na concentration in shoot tissue.     

The maize primary cell wall microfibril: a new model derived from direct visualization

Understanding the molecular architecture of the plant cell wall is critical to reducing the biomass recalcitrance problem, which currently impedes economic bioconversion processing. The parenchyma cell walls from field senesced, maize stem pith have been directly visualized without extraction processes using high-resolution atomic force microscopy (AFM). By imaging the cell wall inner surfaces from different cells and different faces of the same cell, we were able to map the native primary cell wall ultrastructures. Depending on the thickness of non-cellulosic deposition, the parallel-microfibrils appear in various morphologies ranging from clearly defined to completely embedded in the wall matrixes forming cell wall lamella. Macrofibrils were found to exist only on the uppermost layer of the native primary cell wall and appeared to be bundles of elementary fibrils. This novel observation led us to a new hypothesis for the cell wall fibrillar network and biosynthesis processes. Put concisely, a number of elementary fibrils are synthesized at one locus, that of the cellulose synthase complex (CelS), and coalesce into much larger macrofibrils. These macrofibrils eventually split at the ends to form parallel microfibrils with deposition of other cell wall components (i.e. hemicelluloses, pectin, etc.) also evident. On the basis of these AFM surface measurements and current supportive evidence from cell wall biophysics, biosynthesis, and genomics, we propose a new molecular model consisting of a 36-glucan-chain elementary fibril, in which the 36-glucan chains form both crystalline and subcrystalline structures. We also propose a modified model of CelS based on recently reported experimental evidence from plant cell wall biosynthesis.     

Response of wheat cultivars to different soil nitrogen and moisture regimes: I. Dry matter partitioning and root growth 1

The balance between root and shoot growth is one of the mechanisms used by plants to adapt to a particular environment. This balance is affected by nutrient supply and water availability. The objective of this study was to understand how nitrogen (N) affects root and shoot growth of three cultivars of wheat (Triticum aestivum L.) grown under two regimes of soil moisture ('non water stressed’ and ‘stressed’ during stem elongation') in the greenhouse and growth chamber. Data showed that before stem elongation, shoot growth was less sensitive to high soil N levels than root growth. In fact, root growth was inhibited by excessive soil N concentration. The cultivar ‘Nesma’ produced more dry matter, absorbed N more rapidly and avoided the negative effect of high soil N concentrations on root growth. ‘Merchouch 8’ produced less dry matter and tolerated more water stress, and appeared to require less water. Severe water stress masked the effect of N on root and shoot growth, and the negative effect of water stress on growth was more important under high N. Plants which were water stressed during stem elongation could not fully recover when they were rewatered from boot stage to anthesis. Nitrogen application improved shoot but not root growth under this soil moisture situation. This study suggested that shoot and root growth were affected differently by N availability and that high soil N levels enhanced the negative effect of water stress on growth.