Improving grasslands: the influence of soil moisture and nitrogen fertilization on the establishment of seedlings

1. In order to investigate the factors influencing the establishment of seedlings in permanent grassland, the influence of soil moisture and nitrogen fertilization on competition between established plants of Lolium perenne and seedlings of Phleum pratense or Trifolium pratense was studied in two experiments under greenhouse conditions using the 'split-box'-technique.
2. There was no difference in the production of plant dry matter of P. pratense or T. pratense between 30% volumetric soil water content (−0·005 MPa) and 22% (−0·04 MPa), but 15% soil moisture (−0·33 MPa) reduced plant growth. L. perenne yields were linearly reduced by reduced soil moisture content.
3. Shoot competition from L. perenne reduced the plant dry matter yield of P. pratense and T. pratense more than did root competition in these experiments. When shoot competition was present, differences between moisture contents were not detected, indicating that light was probably the limiting resource under such conditions. No significant interaction between root competition and soil moisture was observed for plant weight.
4. Root competition was not prevented even though sufficient water and nitrogen were supplied. This indicated either that some other growth factor was limiting or the plants competed for resources at the root hair level even though sufficient resources were supplied at the pot or field scale. Therefore, in the situation of direct drilling of species during grassland renovation, it may be difficult to alleviate competition by adequate provision of water and nitrogen.     

内生真菌感染对干旱胁迫下黑麦草生长的影响

 内生真菌是生活在健康植物的茎叶内,形成不明显感染的一类真菌。以黑麦草（Lolium perenne L.）为实验材料,研究在不同强度的干旱胁迫下内生真菌（Neotyphodium lolii）侵染对其叶片延伸生长、分蘖数和生物量的影响。结果表明,与非感染种群相比,内生真菌感染对黑麦草叶片延伸速率无明显促进作用;内生真菌感染种群具有明显较多的分蘖数;在重度胁迫并经过恢复期后,内生真菌感染种群具有较高的根冠比。因而内生真菌可能通过提高植物的分蘖能力和促进有机物向根系的分配来促进宿主植物的营养生长并提高其抗旱性     

西南喀斯特地区两种草本对干湿交替和N添加的生长响应

喀斯特地区的"岩溶干旱"和频繁的变水环境成为喀斯特地区植被生长和分布的重要选择压力,是该地区植被恢复重建的主要障碍因子。N沉降也会对喀斯特地区的生态系统造成难以预测的影响。为了探究喀斯特地区草本植物对干湿交替和N添加的生长响应,以苍耳(Xanthium sibiricum)和三叶鬼针草(Bidens pilosa)为研究对象,通过盆栽水分受控实验,研究了5种不同水分处理[对照组(CK)、干旱组(D)、1周干湿交替处理组(DW-1)、2周干湿交替处理组(DW-2)和3周干湿交替处理组(DW-3)]与N添加(N+、N-)对两种草本植物生长和生物量的影响。结果表明,干旱胁迫抑制了植物生长和生物量的积累,株高、叶面积、总根长和根体积等生长指标和地上生物量均显著降低,根冠比增大。不同程度的干湿交替对植物的生长和生物量的积累均表现出一定程度的补偿效应,但这种补偿效应的大小随着干旱持续时间的延长而减弱。N添加对植物的生长和生物量积累有显著地促进作用,株高、根表面积、根体积和根生物量较对照组显著增加,但这种促进作用随着干旱历时的增加而减弱,可能与土壤水分状况有关。同时,N添加还影响着植物生物量的分配,在促进两种植物地上和地下生长的同时,还促进了植物根冠比的增加。     

羊草叶片气体交换参数对温度和土壤水分的响应

 采用生长箱控制的方法研究了羊草(Leymus chinensis)幼苗叶片光合参数对5个温度和5个水分梯度的响应和适应。结果表明：轻度、中度土壤干旱并没有限制羊草叶片的生长,对气体交换参数亦无显著影响,反映了羊草幼苗对土壤水分胁迫的较高耐性。叶片生物量以26 ℃时最大,其它依次为23 ℃、20 ℃、29 ℃和32 ℃。温度升高使气孔导度和蒸腾速率增加, 却使光合速率和水分利用效率降低。水分和温度对叶片生物量、光合速率、气孔导度和蒸腾速率存在显著的交互作用,表明高温加强了干旱对叶片生长和气体交换的影响, 降低了羊草对土壤干旱的适应能力。高温和干旱的交互作用将显著减少我国半干旱地区草原的羊草生产力。     

Linking the patterns in soil moisture to leaf water potential, stomatal conductance, growth, and mortality of dominant shrubs in the Florida scrub ecosystem

Patterns in soil moisture availability affect plant survival, growth and fecundity. Here we link patterns in soil moisture to physiological and demographic consequences in Florida scrub plants. We use data on different temporal scales to (1) determine critical soil moisture content that leads to loss of turgor in leaves during predawn measurements of leaf water status (Ψ _crit), (2) describe the temporal patterns in the distribution of Ψ _crit, (3) analyze the strength of relationship between rainfall and soil moisture content based on 8 years of data, (4) predict soil moisture content for 75 years of rainfall data, and (5) evaluate morphological, physiological and demographic consequences of spring 2006 drought on dominant shrubs in Florida scrub ecosystem in the light of water-uptake depth as determined by stable isotope analysis (δ¹⁸O). Based on 1998–2006 data, the soil moisture content at 50 cm depth explained significant variation in predawn leaf water potential of two dominant shrubs, Quercus chapmanii and Ceratiola ericoides (r ²?=?0.69). During 8 years of data collection, leaves attained Ψ _crit only during the peak drought of 2000 when the soil moisture fell below 1% by volume at 50 and 90 cm depth. Precipitation explained a significant variation in soil moisture content (r ²?=?0.62). The patterns in predicted soil moisture for 75 year period, suggested that the frequency of drought occurrence has not increased in time. In spring 2006, the soil reached critical soil moisture levels, with consequences for plant growth and physiological responses. Overall, 24% of plants showed no drought-induced damage, 51% showed damage up to 50%, 21% had intense leaf shedding and 2% of all plants died. Over the drought and recovery period (May–October 2006), relative height growth was significantly lower in plants with greater die-back. All species showed a significant depression in stomatal conductance, while all but deep-rooted palms Sabal etonia and Serenoa repens showed significantly lower predawn (Ψ _pd) and mid-day (Ψ _md) leaf water potential in dry compared to wet season. Plants experiencing less severe die-back exhibited greater stomatal conductance, suggesting a strong relationship between physiology and morphology. Based on results we suggest that the restoration efforts in Florida scrub should consider the soil moisture requirements of key species.     

Recovery from drought stress in Lolium perenne (Poaceae): are fungal endophytes detrimental?

Perennial ryegrass (Lolium perenne) is a cool-season, perennial species widely used for forage and turf. It is often infected by a clandestine, endophytic fungus (Neotyphodium lolii) that has the potential to affect host growth responses to abiotically stressful conditions. In some species, the grass-endophyte symbiosis is mutualistic, but the relationship is reported to be contingent on environmental conditions and host genotype in L. perenne. The objective of this research was to determine the potential effects of endophyte infection on recovery from severe drought stress in variable genotypes of a perennial ryegrass cultivar. Sixteen infected (+E) and 16 uninfected (-E) ramets were planted in the greenhouse for each of 10 ryegrass genotypes. Eight +E and eight -E plants per genotype were exposed to three sequential droughts where water was withheld for 11-14 d, resulting in <5% soil moisture; the others (control) were watered as needed. Response variables were tiller numbers 1 wk and 4 wk after drought, and leaf area and dry mass of shoots and roots 7 wk after drought. In both control and drought, -E plants had more tillers, and greater leaf area and total mass, than +E plants, suggesting a detrimental effect of endophytic fungi. Fungal hyphae survived the drought and were abundant in post-drought, +E plants. The effects of endophytes were specific for particular host genotypes, as exemplified by significant genotype × endophyte interactions. Root : shoot ratio and percent of mass allocated to tiller bases (a rough measure of resource storage) showed genotype × endophyte × drought interactions. There was plasticity for root : shoot ratio and genetic variation in the ability to restore root growth during recovery from drought. For 7 of 10 genotypes, -E plants showed an equal or greater allocation to tiller bases than +E plants following drought recovery, illustrating a cost to endophyte infection for some genotypes. The symbiotic relationship between L. perenne and its endophyte primarily benefits the fungus, not the host, under many environmental conditions.     

荒漠河岸林植物木质部导水与栓塞特征及其对干旱胁迫的响应

 以同处于干旱区的塔里木河下游(铁干里克)和黑河下游(乌兰图格)断面为研究区, 比较了荒漠河岸林主要建群种胡杨(Populus euphratica)、柽柳(Tamarix spp.)、疏叶骆驼刺(Alhagi sparsifolia)和花花柴(Karelinia caspia)在长期遭受不同干旱胁迫下的根、枝条木质部导水力和栓塞化程度的变化特征, 并分析了木质部导水对干旱胁迫的响应及适应策略。结果表明: 1) 黑河下游荒漠河岸林植物的导水能力显著高于塔里木河下游, 其中柽柳、胡杨、疏叶骆驼刺和花花柴根木质部的初始比导率(K_s0)分别高11.97、6.74、7.10和3.73倍, 枝条的K_s0分别高9.48、3.65、2.07和1.88倍, 地下水埋深导致的干旱胁迫程度不同是诱发荒漠植物导水能力差异的根本原因; 2)柽柳耐干旱能力最强, 适应范围较宽, 而花花柴、疏叶骆驼刺的耐旱性相对较弱, 适生范围较窄, 这可能与植物的根系分布有关; 3)干旱胁迫较轻时, 枝条木质部是荒漠河岸林植物水分传输的主要阻力部位, 干旱胁迫严重时, 根木质部是限制植株水流的最大阻碍部位; 4)荒漠河岸林植物主要通过调节枝条木质部的水流阻力来适应干旱胁迫, 且其适应策略与干旱胁迫程度有关, 干旱胁迫轻时, 植物通过限制枝条木质部水流来协调整株植物的均匀生长; 干旱胁迫严重时, 植物通过牺牲劣势枝条、增强优势枝条水流来提高植株整体生存的机会。     

An experimental device for studying seed responses to naturally fluctuating temperature of surface soil under a constant water table

1. A simple experimental device was designed, in which seeds can be exposed to natural fluctuation of surface soil temperature under a constant soil moisture condition maintained by an automatic water supply system based on the principle of a Mariotte siphon.
2. Except for the period during summer drought, surface soil temperature at a depth of 5 cm and its fluctuation within the device were largely similar to the temperature of the surface soil subjected to natural fluctuation of moisture.
3. The seeds of Persicaria lapathifolia placed at the depth of 0·5 cm in the soil within the device germinated during the natural germination season of the species, while the seeds placed at the depth of 5 cm or beneath the 10 cm-thick layer of litter failed to germinate.
4. The ungerminated seeds retrieved in August did not germinate at the favourable alternating temperature in the laboratory test unless exposed to previous moist chilling, suggesting the induction of secondary dormancy. Therefore, higher summer temperatures but not summer drought or moisture fluctuation seem to have been responsible for the dormancy induction, because the dormancy was induced in the fully hydrated seeds.     

呼吸活性的下降可解释水分亏缺对银合欢根瘤固氮酶活性的抑制(英)

在温室条件下研究了水分亏缺对银合欢根瘤的水分关系、固氮酶活性（乙炔还原活性）、呼吸活性以及蔗糖代谢有关酶活性的影响。随着土壤含水量的下降,根瘤水势也相继下降。土壤干旱不但显著地抑制了根瘤乙炔还原的活性,而且对根瘤的呼吸活性、ATP的产生以及催化蔗糖降解的碱性转化酶和蔗糖合成酶的活性也具强烈的抑制作用。然而,根瘤可溶性总糖含量则不受土壤干旱的影响。用呼吸抑制剂DNP处理根瘤后,其固氮酶活性、呼吸活性及ATPI含量都受到极显著的抑制。这都表明,水分胁迫对根瘤呼吸活性及ATP产生的抑制可解释干旱条件下固氮酶活性的下降。     

Drought-induced changes in xylem pH, ionic composition, and ABA concentration act as early signals in field-grown maize (Zea mays L.).

Early signals potentially regulating leaf growth and stomatal aperture in field-grown maize (Zea mays L.) subjected to drought were investigated. Plants grown in a field lysimeter on two soil types were subjected to progressive drought during vegetative growth. Leaf ABA content, water status, extension rate, conductance, photosynthesis, nitrogen content, and xylem sap composition were measured daily. Maize responded similarly to progressive drought on both soil types. Effects on loam were less pronounced than on sand. Relative to fully-watered controls, xylem pH increased by about 0.2 units one day after withholding irrigation (DAWI) and conductivity decreased by about 0.25 mS cm(-1) 1-3 DAWI. Xylem nitrate, ammonium, and phosphate concentrations decreased by about 50% at 1-5 DAWI and potassium concentration decreased by about 50% at 7-8 DAWI. Xylem ABA concentration consistently increased by 45-70 pmol ml(-1) at 7 DAWI. Leaf extension rate decreased 5 DAWI, after the changes in xylem chemical composition had occurred. Leaf nitrogen significantly decreased 8-16 DAWI in droughted plants. Midday leaf water potential and photosynthesis were significantly decreased in droughted plants late in the drying period. Xylem nitrate concentration was the only ionic xylem sap component significantly correlated to increasing soil moisture deficit and decreasing leaf nitrogen concentration. Predawn leaf ABA content in droughted plants increased by 100-200 ng g(-1) dry weight at 7 DAWI coinciding with a decrease in stomatal conductance before any significant decrease in midday leaf water potential was observed. Based on the observed sequence, a chain of signal events is suggested eventually leading to stomatal closure and leaf surface reduction through interactive effects of reduced nitrogen supply and plant growth regulators under drought.     

Effects of soil moisture regimes on photosynthesis and growth in cattail (Typha latifolia)

Both waterlogging and water deficiency are major environmental factors affecting plant growth and functioning in many wetland and floodplain ecosystems across North America. Wetland plants possess various characteristics that enable them to survive and function in the intermittently flooded wetland environments, while their sensitivity to drought has received less attention. The present study quantified the photosynthetic and growth responses of cattail (Typha latifolia), an important species of freshwater wetlands, to a wide range of soil moisture regimes. In addition, changes in the efficiency of photosynthetic apparatus following initiation of the treatments were investigated. Under greenhouse conditions, seedlings were subjected to four soil moisture regimes: (1) drained (control), (2) continuous flooding, (3) periodic flooding, and (4) periodic drought. Results indicated that dark fluorescence yield was increased in response to periodic drought, while it showed decreases under continuous flooding. Net photosynthesis and stomatal conductance were enhanced by continuous flooding and periodic flooding. In contrast, these parameters exhibited reduction under periodic drought. In addition, leaf chlorophyll content was adversely affected by periodic drought. Recovery of net photosynthesis was noted, along with enhanced height growth, in both continuously and periodically flooded plants. Meanwhile, continuous flooding enhanced biomass production while periodic drought led to biomass reduction. Periodic drought also contributed to substantial reduction in root growth compared with shoot growth. Therefore, the combined photosynthetic performance and growth responses of cattail are likely to contribute to the ability of this species to thrive in flooded condition but be susceptive to periodic drought.     

Seedling growth response to added nutrients depends on seed size in three woody genera

1 We tested whether seedlings of small‐seeded species were more reliant on soil nutrients than large‐seeded ones by growing 21 species from three woody genera ( Eucalyptus, Hakea and Banksia ) along a gradient of nutrient availability.
2 At very low nutrient availability, larger seeds produced larger seedlings. This was seen especially among the eucalypts, but the difference was eliminated at optimal soil nutrient levels. Hakea species with large seed mass, and all Banksia species, appeared unable to exploit additional soil nutrients for growth, whatever the level supplied.
3 Larger seeds tended to have proportionately higher contents of N, P and K and, under nutrient‐poor conditions, supplied more of these to their seedlings, although at a diminishing rate.
4 We suggest that large‐seededness could be an adaptation to the high‐light, nutrient‐impoverished habitats in which these species occur by providing the seedling with the mineral nutrients, rather than carbon‐based metabolites, needed for maximizing initial root growth. Reaching reliable moisture before summer (drought avoidance) is an alternative strategy to physiological tolerance of drought.     

苜蓿与沙打旺苗期生长和水分利用对土壤水分变化的反应

通过室内生长箱内盆栽实验,比较了苜蓿和沙打旺苗期的根冠生长和水分利用对种土壤水分环境变化的响应和差异.结果表明,充分供水下苜蓿和沙打旺苗期生物量和蒸腾效率均最高,苜蓿均显著高于沙打旺.土壤水分减少后苜蓿苗期生物量和蒸腾效率下降幅度均大于沙打旺.从低水到阶段低水处理后土壤水分逐渐降低和降低后再复水到低水处理,苜蓿和沙打旺的生物量分别较持续低水处理显著减少47.8%和27.9%.旱后复水后苜蓿根冠比和单位根量耗水量较显著增加,蒸腾效率显著下降; 沙打旺根冠比显著下降,单位根量耗水量和蒸腾效率变化不显著.     

八种针阔叶幼树清晨叶水势与土壤含水量的关系及其抗旱性研究

一、前言土壤是植物根系的生活环境,是供给植物赖以生存的水分和养分的源泉。干旱区的植物尤其是树木常常由于土壤供水不足而限制其自然分布和生长。根据土壤-植物-大气连续体系     

Responses of Cyperus brevifolius (Rottb.) Hassk. and Cyperus kyllingia Endl. to varying soil water availability

Different responses of Cyperus brevifolius and Cyperus kyllingia to varying soil water regimes were examined to explain their successful existence in a diverse range of habitats throughout the year in Indonesia. Thirty 43-day sprouts of each species were grown in three soil conditions, namely drought, field capacity, and flooding under greenhouse conditions. Plant height, leaf length, tiller number, and flower number were measured twice a week, from 45 to 98 days after sowing (DAS), while the other 12 traits were recorded at the end of the observation time. Ten out of the twelve traits were substantially influenced by the soil water content. Both species exhibited their best growth, production, and reproduction under field capacity conditions, and these traits were greatly subdued under drought conditions. Under drought conditions, both species manifested reduced growth and leaf expansion; however, stomatal aperture and frequency did not exhibit strong response to the soil water content. C. brevifolius showed a significantly greater biomass production and reproductive traits in field capacity and flooded conditions, but under drought conditions, growth was greatly hampered and only vegetative propagation occurred. On the other hand, C. kyllingia showed a higher tolerance to drought conditions, indicated by both a higher biomass and a higher number of flowers. The results obtained suggest that survival ability when faced with drought conditions was more apparent in C. kyllingia than in C. brevifolius, but where there was sufficient soil water, C. brevifolius was more prolific. This could be the explanation for the dominance of C. brevifolius in flooded areas and during the rainy season, and the occurrence of C. kyllingia in a wider range of habitats throughout the year.     

Differences in osmotic adjustment,foliar abscisic acid dynamics,and stomatal regulation between an isohydric and anisohydric woody angiosperm during drought

Species are often classified along a continuum from isohydric to anisohydric, with isohydric species exhibiting tighter regulation of leaf water potential through stomatal closure in response to drought. We investigated plasticity in stomatal regulation in an isohydric (Eucalyptus camaldulensis) and an anisohydric (Acacia aptaneura) angiosperm species subject to repeated drying cycles. We also assessed foliar abscisic acid (ABA) content dynamics, aboveground/belowground biomass allocation and nonstructural carbohydrates. The anisohydric species exhibited large plasticity in the turgor loss point (Ψ_TLP), with plants subject to repeated drying exhibiting lower Ψ_TLP and correspondingly larger stomatal conductance at low water potential, compared to plants not previously exposed to drought. The anisohydric species exhibited a switch from ABA to water potential‐driven stomatal closure during drought, a response previously only reported for anisohydric gymnosperms. The isohydric species showed little osmotic adjustment, with no evidence of switching to water potential‐driven stomatal closure, but did exhibit increased root:shoot ratios. There were no differences in carbohydrate depletion between species. We conclude that a large range in Ψ_TLP and biphasic ABA dynamics are indicative of anisohydric species, and these traits are associated with exposure to low minimum foliar water potential, dense sapwood and large resistance to xylem embolism.     

Changes in the concentration of ABA in xylem sap as a function of changing soil water status can account for changes in leaf conductance and growth

Abstract. Maize seedlings ( Zea mays L. John Innes F1 hybrid) were grown in a greenhouse in l-m-long tubes of soil. When the plants were well established, water was withheld from half of the tubes. Control plants were watered every day during the 20-d experimental period. The soil drying treatment resulted in a substantial restriction of stomatal conductance and a limitation in shoot growth, even though there was no detectable difference in the water relations of watered and unwatered plants. From day 7 of the soil drying treatment, xylem ABA concentrations (measured using the sap exuded from detopped plants) were substantially increased in unwatered plants compared to values recorded with sap from plants watered every day. Measurements of water potential through the profile of unwatered soil suggest that xylem ABA concentrations reflects the extent of soil drying. Leaf ABA content was a much less sensitive indicator of the effect of soil drying and during the whole of experimental period there was no significant difference between ABA concentration in leaves of well watered and unwatered plants. In a second set of experiments, ABA was fed to part of the roots of potted maize plants to manipulate xylem ABA concentration. These manipulations suggested that the increases in ABA concentration in xylem sap, which resulted from soil drying, were adequate to explain the observed variation in stomatal conductance and might also explain the restriction in leaf growth rate. These results are discussed in the light of recent work which suggests that stomatal responses to soil drying are partly attributable to an as-yet unidentified inhibitor of stomatal opening.     

Interactions between elevated CO2 concentration, nitrogen and water: effects on growth and water use of six perennial plant species

Two experiments are described in which plants of six species were grown for one full season in greenhouse compartments with 350 or 560 μ mol mol^–1 CO₂. In the first experiment two levels of nitrogen supply were applied to study the interaction between CO₂ and nitrogen. In the second experiment two levels of water supply were added to the experimental set-up to investigate the three-way interaction between CO₂, nitrogen and water. Biomass and biomass distribution were determined at harvests, while water use and soil moisture were monitored throughout the experiments. In both experiments a positive effect of CO₂ on growth was found at high nitrogen concentrations but not at low nitrogen concentrations. However, plants used much less water in the presence of low nitrogen concentrations. Drought stress increased the relative effect of elevated CO₂ on growth. Available soil moisture was used more slowly at high CO₂ during drought or at high nitrogen concentrations, while at low nitrogen concentrations decreased water use resulted in an increase in soil moisture. The response to the treatments was similar in all the species used. Although potentially faster growing species appeared to respond better to high CO₂ when supplied with a high level of nitrogen, inherently slow-growing species were more successful at low nitrogen concentrations.     

贵州特有植物美丽红山茶水分胁迫的生理响应

采用水分抗逆性指标主成分分析法,选择4年实生苗,测定和评价了连续30 d水涝和干旱胁迫的生理响应.结果表明：(1)水涝处理的叶片生长旺盛,光泽度和观赏性无明显变化,叶片相对含水量、MDA含量、SOD活性、POD活性、CAT活性的变化差异在5％以内,土壤含水量增加3．47％,细胞膜透性减少1．59％,短期水涝对美丽红山茶伤害不明显;(2)随着干旱时间延长,新叶卷缩、老叶脱落,土壤含水量、叶片相对含水量随时间呈负相关缓慢减少,细胞膜透性呈正相关缓慢增长,但复水10 d 后可恢复正常生长,无明显生理变化响应;(3)SOD活性和MDA含量分别增加了0．66％和5．31％,POD活性随干旱时间上下波动而增加6．47％, CAT活性随干旱时间延长而增加,20 d增加33．33％达最大值20．8 U??g-1??min-1.在抗性生理中,维持膜稳定性的叶片含水量、POD和CAT的变化对抗旱能力大小起主要作用.     

Influence of Nitrogen Supply and Water Stress on Growth and Nitrogen, Phosphorus, Potassium and Calcium Contents in Pearl Millet

Influence of supra-optimal concentrations of N on growth and accumulation of N, K, P and Ca in the shoots and roots in Pennisetum glaucum (L.) R.Br. under water stress was assessed in a pot experiment under glasshouse conditions. Thirty four-day-old plants of two lines, ICMV94133 and WCA-78, were subjected to 224, 336, or 448 mg(N) kg^–1(soil) and soil moisture 100 or 30 % of field capacity for 30 d. Increasing soil N supply decreased growth of both lines under water deficit. Nitrogen content in the shoots of both lines was not affected by supra-optimal levels of N or different watering regimes, but in contrast, the root N content was increased consistently in WCA-78 with increase in soil N content. Shoot P content increased considerably in WCA-78 at the two higher N contents, but it was significantly lower at drought stress than at well-watered treatment. In contrast, shoot or root P content in ICMV94133 did not differ under both watering regimes. Potassium content in the shoots of WCA-78 was considerably increased at the two higher N contents under drought conditions. Root K content was increased in WCA-78 at the highest N content under well-watered conditions, whereas the reverse was true in ICMV94133. Calcium content in the shoots of ICMV94133 was higher under drought stress compared with that at well-watered conditions, but such pattern was not observed in WCA-78. However, root Ca content increased in both lines with increase in N supply.