Leaf Gas Exchange and Fluorescence of Two Winter Wheat Varieties in Response to Drought Stress and Nitrogen Supply

Water and nitrogen supply are the two primary factors limiting productivity of wheat (Triticum aestivum L.). In our study, two winter wheat varieties, Xinong 979 and large-spike wheat, were evaluated for their physiological responses to different levels of nitrogen and water status during their seedling stage grown in a phytotron. Our results indicated that drought stress greatly reduced the net photosynthetic rate (P_n), transpiration rate (E), and stomatal conductance (G_s), but with a greater increase in instantaneous water use efficiency (WUE). At the meantime, the nitrogen (N) supply improved photosynthetic efficiency under water deficit. Parameters inferred from chlorophyll a measurements, i.e., photochemical quenching coefficient (qP), the maximum photochemical efficiency (F_v/F_m), the quantum yield of photosystemII(Φ_PSII), and the apparent photosynthetic electron transport rate (ETR) decreased under water stress at all nitrogen levels and declined in N-deficient plants. The root–shoot ratio (R/S) increased slightly with water stress at a low N level; the smallest root–shoot ratio was found at a high N level and moderate drought stress treatment. These results suggest that an appropriate nitrogen supply may be necessary to enhance drought resistance in wheat by improving photosynthetic efficiency and relieving photoinhibition under drought stress. However, an excessive N supply had no effect on drought resistance, which even showed an adverse effect on plant growth. Comparing the two cultivars, Xinong 979 has a stronger drought resistance compared with large-spike wheat under N deficiency.     

不同水氮水平下小麦品种对光、水和氮利用效率的权衡

通过再裂区设计田间试验,以3个春小麦品种(和尚头、西旱2号和宁春4号)为材料,设置两个灌溉水平(充分灌水4500 m³·hm^-2和有限灌水3000 m³·hm^-2)和5个施氮水平(0、75、150、225、300 kg N·hm^-2),研究小麦光能利用效率(LUE)、水分利用效率(WUE)、氮素利用效率(NUE)对水氮的响应特性及其相互关系.结果表明: 3个小麦品种间LUE、WUE和NUE差异显著.在一定范围内增加灌水和施氮量则LUE升高,过量施氮则LUE下降.强抗旱和中等抗旱品种(和尚头和西旱2号)WUE受灌水量的影响比不抗旱品种(宁春4号)小.施氮可以调节小麦WUE,中等施氮水平(和尚头和西旱2号在150 kg N·hm^-2时,宁春4号在225 kg N·hm^-2时)有最高的WUE.随施氮量增加,植株氮素累积量先增后减,氮素干物质生产效率(NUE_b)、氮素收获指数(NHI)、氮肥农学利用效率(NAE)和氮肥偏生产力(PFP)均显著降低.灌溉水平对NHI无显著影响;随灌水量增加,小麦氮素积累量显著增加,强抗旱和中等抗旱品种NUE_b和NAE显著降低,不抗旱品种 NUE_b和PFP显著升高,对其他指标无显著影响.3个小麦品种氮素获取能力与氮素利用效率呈极显著负相关,NUE_b与LUE、WUE呈显著负相关,LUE与WUE呈显著正相关,春小麦氮素利用效率与光能利用效率、水分利用效率间存在明显的权衡关系.当灌水量为3000 m³·hm^-2,强抗旱和中等抗旱品种在150 kg N·hm^-2,不抗旱品种在225 kg N·hm^-2时,有较高的资源利用效率.     

冬小麦对有限水分高效利用的生理机制

通过对不同土壤供水条件下的孕穗开花期的冬小麦叶片CO₂/H₂O气体交换参数的系统测定,研究了光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)、细胞间隙CO₂浓度(Ci)、叶温(Tl)与水分利用效率(WUE)间的关系。结果表明,WUE并不随Pn的增长直线增长,而是呈现出二次曲线的变化趋势;只有当蒸腾达到一定程度时,Tr才对WUE产生影响,而Tr过大时WUE则有下降的趋势;WUE与Ci呈负相关,随Ci的增加WUE呈递减趋势;叶温升高对光合和蒸腾都有促进作用,当超过了某种限度则表现为抑制作用,表明在一定温度范围内,Tl升高对水分利用不利;随Gs的增大,WUE增大到一定程度则不再增加,甚至出现一种回落趋势.     

CO2浓度、氮和水分对春小麦光合、蒸散及水分利用效率的影响

研究了不同土壤氮和土壤水分条件下,大气CO2浓度升高对春小麦光合作用、气孔导度、蒸散和水分利用效率的影响。结果表明,CO2浓度升高,干旱处理的春小麦（Triticum aestivum L.）叶片光合作用速率幅度增加大于湿润处理,随着氮肥用量增加光合速率相应增加,而不施氮脂增加有限;干旱处理气孔导度幅度减少大于湿润处理,不施氮肥的大于氮肥充足的CO2浓度升高,干旱处理的蒸散量减少比湿润处理多,不施氮肥的蒸散量减少较为明显;但干旱处理单叶WUE增加大于湿润处理;随着氮肥用量增加,冠层WUE提高,而不施氮肥的冠层WUE最低。因而CO2浓度升高、光合速率增加和蒸散量减少会减缓干旱的不利影响,增强作物对干旱胁迫的抵御能力。     

Impact of drought on productivity and water use efficiency in 29 genotypes of Populus deltoides x Populus nigra

We examined the relationships among productivity, water use efficiency (WUE) and drought tolerance in 29 genotypes of Populus x euramericana (Populus deltoides x Populus nigra), and investigated whether some leaf traits could be used as predictors for productivity, WUE and drought tolerance. At Orléans, France, drought was induced on one field plot by withholding water, while a second plot remained irrigated and was used as a control. Recorded variables included stem traits (e.g. biomass) and leaf structural (e.g. leaf area) and functional traits [e.g. intrinsic water use efficiency (Wi) and carbon isotope discrimination (Delta)]. Productivity and Delta displayed large genotypic variability and were not correlated. Delta scaled negatively with Wi and positively with stomatal conductance under moderate drought, suggesting that the diversity for Delta was mainly driven by stomatal conductance. Most of the productive genotypes displayed a low level of drought tolerance (i.e. a large reduction of biomass), while the less productive genotypes presented a large range of drought tolerance. The ability to increase WUE in response to water deficit was necessary but not sufficient to explain the genotypic diversity of drought tolerance.     

Genotypic and nutrition-dependent variation in water use efficiency and photosynthetic activity of leaves in winter wheat (Triticum aestivum L.)

Effects of high and reduced NPK nutrition on the genetic variation of components of water use efficiency at the leaf and whole-plant levels were examined in pot-grown old and modern cultivars of winter wheat (Triticum aestivum L.). At the subsequent growth stages, the photosynthetic rate (A), transpiration rate (E), leaf area (LA) and gas exchange efficiency (A/E) were measured on fully developed 4th, 5th, penultimate and flag leaves. At the plant canopy level, the total amount of water transpired was recorded during the whole life cycle to determine the efficiency of water use in the vegetative (WUEveg) and grain (WUEgen) matter formation. Considerable genotypic differences were found for the characters studied. The limited NPK supply caused a decrease in LA, A and A/E, but contributed to an increase in WUEgen. Examined cultivars (C) did not interact with nutrition levels (N) for these characteristics. However, the position of leaves (L) and the C x L and N x L interactions significantly affected the variance in leaf photosynthetic characteristics. A and A/E were negatively correlated with LA, and the flag leaves were photosynthetically less active and less efficient per unit area than the lower leaves. The whole-plant components of WUE were found to be more genetically stable than the photosynthetic leaf characteristics. Some modern cultivars tended to form leaves of higher A and A/E than the older ones, and this corresponds with a more efficient use of water in grain formation (WUEgen) of the former. Stay-green duration of flag leaves and harvest index showed positive correlations with WUEgen. However, no close associations were noticed between WUE components and stress tolerance, and the modern cultivars were usually less tolerant to NPK shortage.     

氮肥后移对抽穗后水分胁迫下冬小麦光合特性及产量的影响

采用子母桶栽土培法模拟冬小麦抽穗后不同的水分胁迫状态,研究了氮肥后移对冬小麦光合特性及产量的影响.设置3个氮肥处理,分别为N₁(基肥∶拔节肥∶开花肥=10∶0∶0)、N₂(6∶4∶0)和N₃(4∶3∶3),模拟冬小麦抽穗后2种水分胁迫(渍水胁迫、干旱胁迫),设正常供水为对照.结果表明：相同供水条件下,N₂和N₃处理较N₁处理显著提高冬小麦灌浆期旗叶的SPAD和光合速率,确保了收获时较高的穗数、穗粒数和地上部分生物量;氮肥后移处理显著提高了冬小麦的耗水量,但其籽粒产量和水分利用效率也显著提高.相同氮肥条件下,干旱胁迫和渍水胁迫处理较正常供水显著降低了冬小麦开花期和灌浆期旗叶的光合速率、千粒重、穗粒数和产量.与正常供水相比,各氮肥条件下干旱胁迫和渍水胁迫处理花后旗叶光合速率及籽粒产量的减小幅度均表现为N₁＞N₂＞N₃.表明氮肥后移通过提高旗叶SPAD、减缓花后旗叶光合速率的下降幅度、增加地上部分干物质积累量,调控产量及其构成要素,以减轻逆境灾害(干旱和渍水胁迫)对产量的影响.     

氮对苹果幼树水分利用效率的影响

以２年生盆栽新红星／平邑甜茶苹果树为试材,初步探讨了土壤不同水分状况下氮肥对植株水分利用效率（ＷＵＥ）及有关参数的影响,结果表明,充足供水时,随施Ｎ量的增加,植株ＷＵＥ降低,施Ｎ导致气孔导率（Ｇｓ）增大,对蒸腾（Ｔｒ）的提高幅度大于光合（Ｐｎ）;供水不足时,施Ｎ植株的ＷＵＥ明显高于对照,不同施Ｎ水平的ＷＵＥ表现为：高Ｎ〉中Ｎ〉低Ｎ,ＷＵＥ改善是由于叶肉羧化能力提高,导致光合增强。     

Responses of photosynthesis and carbohydrate-partitioning to limitations in nitrogen and water availability in field-grown sunflower*

Abstract. Sunflower plants (Helianthus annuus L., cv. CGL 208) were field-grown in adjacent plots of varying resource availability. Control plants received irrigation (on a 4–5 d interval) and high levels of fertilizer nitrogen. Nutrient-stress (N-stress) plants received control levels of irrigation but no nutrient amendments and were determined to be nitrogen-limited. Water-stress (H₂O-stress) plants received control levels of fertilizer nitrogen, but no irrigation after approximately 6 weeks of plant growth. Both stress treatments reduced maximum and diurnal net photosynthesis (A) but resulted in different physiological or biochemical adjustments that tended to maintain or increase A per unit of resource (nitrogen or water) in shortest supply while decreasing the ratio of A per unit of abundant resource. Nutrient-stress reduced total foliar nitrogen, foliar chlorophyll, and initial and total RuBPCase activities, thereby enhancing or preserving photosynthetic nitrogen-use efficiency (NUE), defined as the maximum A observed per unit of leaf nitrogen, relative to the control and H₂O-stress treatments. In addition, N-stress reduced photosynthetic water-use efficiency (WUE), defined as the ratio of A to stomatal conductance to water vapour (g). The slope of A versus g increased with H₂O-stress. In addition, sunflower plants responded to H₂O-stress by accumulating foliar glucose and sucrose and by exhibiting diurnal leaf wilting, which presumably provided additional improvements in photosynthetic WUE through osmoregulation and reduction of midday radiation interception respectively. Photosynthetic NUE was decreased by H₂O-stress in that control levels of total nitrogen, foliar chlorophyll, and RuBPCase activities were maintained even after mean diurnal levels of A had fallen to less than 50% of the control level. We conclude that field-grown sunflower manages a trade-off between photosynthetic WUE and NUE, increasing use efficiency of the scarce resource while decreasing use efficiency of the abundant resource.     

干旱胁迫对不同施氮水平麻疯树幼苗光合特性及生长的影响

采用盆栽控水的方法,研究了干旱胁迫（连续干旱0 d,5 d,10 d,…,45 d）对不同施氮水平(对照 0 kg N·hm^-2、低氮 96 kg N·hm^-2、中氮 288 kg N·hm^-2、高氮 480 kg N·hm^-2)麻疯树幼苗光合特性及其生长的影响.结果表明: 随干旱胁迫强度的增加,各施氮水平麻疯树幼苗叶片相对含水量、苗高生长量、地径生长量、叶面积、净光合速率、蒸腾速率和气孔导度均降低,且各水分处理间差异极显著(P<0.01);随干旱时间的延长,叶绿素含量和水分利用效率表现出先升高后降低的趋势,而胞间CO₂浓度呈先降低后升高的趋势.正常供水时,施氮处理均不同程度提高了麻疯树幼苗的光合能力,促进了麻疯树幼苗的生长,且施氮量越高效果越好;干旱条件下,氮素营养对植株光合能力和生长的影响与干旱程度和施氮水平有关.轻度干旱时,提高施氮水平对植株光合能力和生长具有明显的促进作用;中度干旱时,中氮的促进作用明显高于其他施氮水平;严重干旱时,低氮的促进效果最好,高氮的促进作用减弱并逐渐转向抑制.     

EFFECTS OF ROOT PRUNING AT THE REGREENING STAGE ON DROUGHT TOLERANCE AND WATER USE EFFICIENCY OF WINTER WHEAT IN LATE GROWING STAGE

 通过盆栽试验研究了返青期根修剪对冬小麦(Triticum aestivum)后期耐旱性及水分利用效率的影响。在返青期设置了两个根修剪处理: 1)小剪根, 在植株一侧切去部分侧生根; 2)大剪根, 在主茎四周切去部分侧生根。不剪根者设为对照(CK)。研究结果显示, 两个根修剪处理均显著减少了小麦的根系, 但对根冠比没有显著影响。在花期, 两个根修剪处理的小麦旗叶的叶绿素荧光参数最大光化学效率(The maximum photochemical efficiency of PSⅡ, F_v/F_m)、 PSⅡ潜在活性 (PSⅡ potential activity, F_v/F_o)、实际光化学量子产量(Effective PSⅡ quantum yield, Φ_PSⅡ)、表观光合电子传递速率(Apparent rate of photosynthetic electron transport, ETR)、光化学淬灭系数( Coefficient of photochemical quenching, q_P)和非光化学淬灭系数(Coefficient of non-photochemical quenching, NPQ)值, 在停止供水7 d后, 均显著高于对照, 这表明根修剪小麦的耐旱性强于对照, 因此在干旱胁迫下有较高的光化学活性。小剪根处理在高水条件下对小麦产量无显著影响, 而在中度干旱条件下显著提高了小麦的产量, 因此, 小剪根处理显著提高了小麦的抗旱系数; 小剪根处理在高水分处理(土壤水分含量为田间持水量的85%)和中度干旱胁迫处理(土壤水分含量为田间持水量的55%)条件下, 均显著提高了小麦的水分利用效率。但大剪根处理由于严重影响了群体数量和产量, 水分利用效率和抗旱系数均没有提高。可见, 适当地减少根系有助于小麦的耐旱性和水分利用效率的提高。     

中国东部南北样带森林优势植物叶片的水分利用效率和氮素利用效率

通过测定中国东部南北样带主要森林生态系统中10种优势植物(兴安落叶松、蒙古栎、水曲柳、紫椴、色木槭、红松、杉木、木荷、马尾松、锥栗)叶片的碳氮含量(Cmass、Nmass)、同位素丰度(δ13C、δ15N)以及光合响应曲线,分析了不同优势植物叶片的水分利用效率和氮素利用效率之间的差异及其相互关系.结果表明： 不同生活型植物叶片的Nmass和δ15N差异显著,表现为阔叶植物>针叶植物,落叶植物>常绿植物;最大光合速率(Pn max)表现为针叶植物>阔叶植物,落叶植物>常绿植物;植物叶片的瞬时水分利用效率(WUEi)和长期水分利用效率(WUE)均表现为阔叶植物>针叶植物,常绿植物>落叶植物;植物叶片的瞬时氮素利用效率(NUEi)和长期氮素利用效率(NUE)则表现出相反的规律,且常绿植物和落叶植物叶片的NUE差异显著;WUEi和WUE之间相关性不显著,而NUEi和NUE之间呈显著正相关.植物叶片的水分利用效率与氮素利用效率显著负相关.两种资源利用效率均受植物生活型的影响,并且存在一定的制约关系.     

不同肥料组合对冬小麦水分供需状况的研究

采用Ｄ－饱和最优设计对半湿润易旱区冬小麦在各生育阶段土壤水分平衡状况以及不同肥料组合对冬小麦产量和水分利用效率的影响进行了分析和研究。结果表明,根据水量平衡法计算不同施肥条件下土壤供水量明显不同,以Ｎ、Ｐ、有机肥组合土壤供水占的比例最大为２７％,单施氮肥利用降水补给占的比例最高为１００％。整个生育期,单施氮肥冬小麦水分亏缺最严重,而磷肥与有机肥组合水分亏缺最小,Ｎ、Ｐ、有机肥组合能显著提高冬小麦产     

冬小麦光合特征及叶绿素含量对保水剂和氮肥的响应

以不施保水剂和氮（N）肥为对照,测定了保水剂（60 kg·hm^-2）与不同N肥水平（0、225、450 kg·hm^-2）及其配施条件下大田小麦的光合特征、叶绿素含量和水分利用效率等指标,研究了冬小麦拔节期和灌浆期光合生理特征、叶绿素含量及水分利用对保水剂和N肥的响应.结果表明：灌浆期各处理的光合速率、气孔导度、胞间CO₂浓度、叶片水分利用效率及叶绿素含量均大于拔节期.在拔节期,单施N肥条件下,随施N量的增加,单叶水分利用效率提高,光合速率、气孔导度、胞间CO₂浓度及蒸腾速率均先增后减;225 kg·hm^-2 N肥处理的叶绿素含量最高.施用保水剂后,随施N量的增加,胞间CO₂浓度降低,而光合速率等均提高;单施保水剂及其与N肥配施提高了叶绿素含量,而过多N肥效果不显著在灌浆期,单施N肥显著提高了小麦的光合速率及水分利用效率,降低了气孔导度、胞间CO₂浓度及蒸腾速率;叶绿素含量随N肥用量的增加而增加.施用保水剂后,随N肥用量的增加,光合速率和叶片水分利用效率均先增后减,而胞间CO₂浓度和蒸腾速率先减后增,但均低于对照,气孔导度随施N量的增加而提高.单施保水剂的叶绿素含量显著提高,但其与N肥配施叶绿素含量有所降低.保水剂与N肥配合施用显著提高了小麦的千粒重、产量及水分生产效率.其中,保水剂与225 kg·hm^-2N肥配施处理的产量及水分生产效率均最高.     

不同类型保水剂对冬小麦水分利用效率和根系形态的影响

采用田间小区试验,研究5种不同类型保水剂及2个施用水平对冬小麦产量、水分利用效率、根形态的影响.结果表明: 冬小麦茎蘖数、旗叶面积、产量和水分利用效率在不同类型保水剂及施用量处理之间存在显著差异.与对照相比,5种保水剂处理冬小麦产量增加1.3%～7.9%,水分利用效率由对照的17.1 kg·hm^-2·mm^-1提高到18.0～20.7 kg·hm^-2·mm^-1.保水剂对冬小麦根系的平均直径、总根长和总表面积的影响均达到显著水平,在0～20和20～40 cm土层,总根长分别增加3.7%～19.1%和6.3%～27.3%,总表面积分别增加6.5%～21.7%和2.9%～18.5%.冬小麦根系形态特征与冬小麦产量和水分利用效率均呈显著正相关.丙烯酰胺/无机矿物复合型保水剂对提高冬小麦水分利用效率和促进根系生长效果最为明显.     

Effects of Multi-Stage Continuous Drought on Photosynthetic Characteristics,Yield and Water Use Efficiency of Winter Wheat

A drought event can cause entire crops to fail or yield loss. In order to study the effects of continuous drought on photosynthetic characteristics, yield, and water use efficiency (WUE) of winter wheat (Triticum aestivum L.), the winter wheat variety “Aikang 58” was selected as test material with controlling the water of the pot-planted winter wheat under a mobile rainout shelter. Based on foot planting and safe wintering, winter wheat was evaluated under different drought conditions, including light, moderate and severe drought at the jointing (B), heading (C), and filling (G) stages. The soil water content was controlled in a range of 60% to 70%, 50% to 60%, and 40% to 50% of the field capacity, respectively. In the experiment, there were 9 single-stage droughts, 3 three-stage droughts, and 1 test control (totaling 13 trials). The results are as follows: Under a single-stage drought, the change of net photosynthetic rate (Pn) and stomatal conductance (Gs) have similar trends, and they both decrease significantly with the severity of the drought. Under three-stage continuous droughts, the change curve of Gs shows a constant downward trend; the change curve of Pn showed a “valley shape,” and the minimum value of Pn appeared at the heading stage. All droughts will reduce the yield of winter wheat. Under the three-stage continuous drought conditions, except for light drought, moderate drought and severe drought will cause significant yield reduction, mainly due to lack of water at the jointing and heading stages. Continuous drought will reduce the WUE, and the difference will reach a significant level under moderate and severe drought. The present results suggested that when water resources are scarce, it is a better irrigation model to save water and achieve high grain yield by applying appropriate water stress (60%–70% FC) during the critical growth period of winter wheat.     

The combined effect of constant water deficit and nitrogen supply on WUE, NUE and Δ13C in durum wheat potted plants

Water scarcity and nitrogen shortage are the main constraints on durum wheat productivity. This paper examines the combined effects of a constant water deficit and nitrogen supply (NS) on growth, photosynthesis, stomatal conductance (g_s) and transpiration, instantaneous and time‐integrated water use efficiency (WUE) and nitrogen use efficiency (NUE) and carbon isotope discrimination (Δ¹³C) in durum wheat genotypes grown in pots under greenhouse conditions. Three water levels (40%, 70% and 100% container capacity), two nitrogen doses (high and low N) and four genotypes were assayed in a total of 24 experimental treatments. Water and nitrogen treatments were imposed 2 weeks after plant emergence. The growth, nitrogen content and Δ¹³C of the shoot and the gas exchange in the flag leaf were determined about 2 weeks after anthesis. As expected, both water and NS had a strong positive effect on growth. However, a reduction in water supply had low effect decreasing photosynthesis and transpiration, Δ¹³C and NUE and increasing WUE. On the contrary, increasing the level of nitrogen supplied had a significant negative effect on g_s, which decreased significantly the ratio of intercellular to ambient CO₂ concentrations and Δ¹³C, and increased both instantaneous and time‐integrated WUE. In addition, a higher N level also negatively affected the instantaneous and time‐integrated NUE. The Δ¹³C of shoots correlated significantly and negatively with either instantaneous or time‐integrated measurements of WUE. Moreover, within each NS, Δ¹³C also correlated negatively with the integrated NUE. We concluded that under our experimental conditions, Δ¹³C gives information about the efficiency with which not just water but also nitrogen are used by the plant. In addition, this study illustrates that a steady water limitation may strongly affect biomass without consistent changes in WUE. The lack of effect of the different water regimes on gas exchange, WUE and Δ¹³C illustrate the importance of how stress is imposed during growth.     

Carbon isotope discrimination and bundle sheath leakiness in three C(4) subtypes grown under variable nitrogen,water and atmospheric CO(2) supply

The changes in composition and productivity of semi-arid C(4) grassland, anticipated with rising atmospheric CO(2), will depend on soil water and nutrient availability. The interactive effects of soil resource limitation and elevated CO(2 )on these grasses, furthermore, may vary among C(4) biochemical subtypes (NADP-ME, NAD-ME, PCK) that differ in bundle sheath leakiness (Phi) responses to drought and nitrogen supply. To address C(4) subtype responses to soil resource gradients, the carbon isotope discrimination (Delta), bundle sheath leakiness (Phi), leaf gas exchange (A, g(s), c(i)/c(a)) and above-ground biomass accumulation were measured on three dominant grasses of semi-desert grassland in south-eastern Arizona. Bouteloua curtipendula (PCK), Aristida glabrata (NADP-ME) and the non-native Eragrostis lehmanniana (NAD-ME) were grown in controlled-environment chambers from seed under a complete, multi-factorial combination of present ambient (370 ppm) and elevated (690 ppm) CO(2) concentration and under high and low water and nitrogen supply. E. lehmanniana (NAD-ME) had the highest photosynthetic rate (A) and lowest Phi compared to the other two grasses when grown under low nitrogen and water availability. However, favourable water and nitrogen supply and elevated atmospheric CO(2) enhanced photosynthetic performance and above-ground biomass production of B. curtipendula (PCK) to a greater extent than in A. glabrata and E. lehmanniana. Contrary to pre vious studies, Phi and Delta in the NADP-ME subtype (A. glabrata) were most affected by changing environmental conditions compared to the other subtypes; deviations from the classic NADP-ME anatomy in Aristida could have accounted for this result. Overall, response of semi-arid grasslands to rising atmospheric CO(2) may depend more on species-specific responses to drought and nitrogen limitation than on general C(4) subtype responses.     

Effect of Pre-Anthesis Drought Hardening on Post-Anthesis Physiological Characteristics,Yield and WUE in Winter Wheat

A drought event can cause yield loss or entire crops to fail. In order to study the effects of continuous drought on physiological characteristics, yield, and water use efficiency (WUE) of winter wheat (Triticum aestivum L.), the variety “Zhoumai 22” was grown in controlled water conditions of the pot-planted winter wheat under a mobile rainout shelter. Foot planting and safe wintering were used to evaluate, winter wheat under different drought conditions, including light, moderate and severe drought at the jointing, heading, and filling stages. The soil water content was controlled at 60–70%, 50–60%, or 40–50% of field capacity. Experimental trials included 3 pre-anthesis drought hardening, 3 three-stage continuous drought, and 1 test control conditions. Under drought stress conditions, winter wheat leaf water potential, soil plant analysis development (SPAD), photosynthesis parameters, and yield declined due to pre-anthesis drought hardening. And the degree of decline: continuous drought > pre-anthesis drought hardening. Changes in the post-anthesis photosynthetic capacity of winter wheat were mainly related to the pre-anthesis drought level, rather than post-anthesis rehydration. The threshold of non-stomata limiting factors caused by photosynthesis at the filling stage is 40–50%FC, while comprehensive yield and WUE affected, the yield in severe drought treatments was the most significant, B3C3 and B3C3G3 decreased by 55.68% and 55.88%, respectively. Pre-anthesis drought was the main reason for the decreased crop yield. Thus, severe drought should be avoided during planting, while pre-anthesis light drought is a suitable choice for water-saving and crop production, as proper pre-anthesis drought hardening (60–70% FC) is feasible and effective.     

Effects of N and water supply on water use-efficiency of a semiarid grassland in Inner Mongolia

Productivity of semiarid grasslands is primarily limited by seasonal rainfall amount and becomes increasingly limited by nutrient availability under wet conditions. Interactive effects of water and N availability on grassland productivity and parameters related to water use were studied on a grassland site in Inner Mongolia, China, in a 2-factorial experiment with two levels of water (rainfed: 158 mm; irrigated: 839 (N0) and 972 (N1) mm) and N supply (0 or 180 kg N ha^?1). RUE was calculated from ANPP and cumulative water supply. Bare soil evaporation (E) was calculated from climatic data and leaf area dynamics, and percolation (D) and transpiration (T) were estimated with HYDRUS-1D. Water-use efficiency (WUE, ANPP / (T + D)) and transpiration efficiency (TE, ANPP / T) were calculated. Resource availability had pronounced effects on the water-use efficiency of semiarid grassland. RUE, WUE, and TE all decreased under irrigated compared to rainfed conditions and were significantly increased with N fertilizer application at both levels of water supply. While the irrigation effect on parameters of water-use efficiency were accordingly reflected in stable carbon isotope signatures, N supply resulted in significantly less negative δ¹³C-values under rainfed but not irrigated conditions. It is concluded, that spatial or temporal gradients in resource availability have pronounced effects on the water-use efficiency of semiarid grassland. The decrease of water use-efficiency under high water supply was related to differences in TE and not to a relative increase of unproductive water loss. Carbon isotope discrimination was highly correlated with WUE and TE, but can be a poor predictor of RUE.