Nitrogen uptake and utilization efficiency of European maize hybrids developed under conditions of low and high nitrogen input

Maize varieties with improved nitrogen(N)‐use efficiency under low soil N conditions can contribute to sustainable agriculture. Tests were carried to see whether selection of European elite lines at low and high N supply would result in hybrids with differential adaptation to these contrasting N conditions. The objective was to analyze whether genotypic differences in N uptake and N‐utilization efficiency existed in this material and to what extent these factors contributed to adaptation to low N supply. Twenty‐four hybrids developed at low N supply (L × L) were compared with 25 hybrids developed at high N supply (H × H). The N uptake was determined as total above‐ground N in whole plants, and N‐utilization efficiency as the ratio between grain yield and N uptake in yield trials at four locations and at three N levels each. Highly significant variations as a result of hybrids and hybrids × N‐level interaction were observed for grain yield as well as for N uptake and N‐utilization efficiency in both hybrid types. Average yields of the L × L hybrids were higher than those of the H × H hybrids by 11.5% at low N supply and 5.4% at medium N level. There was no significant yield difference between the two hybrid types at high N supply. The L × L hybrids showed significantly higher N uptake at the low (12%) and medium (6%) N levels than the H × H hybrids. In contrast, no differences in N‐utilization efficiency were observed between the hybrid types. These results indicate that adaptation of hybrids from European elite breeding material to conditions with reduced nitrogen input was possible and was mainly the result of an increase in N‐uptake efficiency.     

Variation for nitrogen use efficiency traits in current and historical great plains hard winter wheat

Wheat genotypes that efficiently capture and convert available soil nitrogen into harvested grain protein are key to sustainably meeting the rising global demand for grain protein. The purposes of this study were: to characterize the genetic variation for nitrogen use efficiency (NUE) traits within hard winter wheat adapted to the Great Plains of the United States and evaluate trends in the germplasm with year of release; to explore relationships among traits that may be used for selection within breeding programs; and to identify quantitative trait loci associated with NUE traits in this germplasm. NUE traits were measured in a panel of 299 hard winter wheat genotypes, representing historically important and contemporary germplasm, from across the growing region. Trials were grown in two years at two levels of nitrogen fertility. Genotype and genotype × year interaction effects were highly significant for NUE traits, while genotype × nitrogen rate interactions were non-significant. Strong genetic correlations of plant height and flowering date with NUE traits were observed. Wheat breeders have improved NUE: the subset of 183 genotypes that were released as cultivars after 1960 demonstrated significant trends with year of release for improved grain N yield, grain yield, nitrogen harvest index, nitrogen uptake efficiency, nitrogen utilization efficiency, and post-anthesis nitrogen uptake. In genome-wide association analyses, plant height and flowering date were important covariates in the mixed models, and plant height and flowering date substantially explained the variation in NUE traits in this germplasm. Marker-trait associations were identified that may prove useful in breeding.     

宁夏不同基因型春小麦磷营养的差异

在不同施磷土壤条件下,研究了宁夏春小麦100个基因型的磷营养性状的差异。结果表明,在低磷条件下,小麦籽粒产量和生物学产量在基因型间存在极显著差异（P＜0.01),而在高磷条件下存在显著差异（P＜0.05)。不论施磷与否,茎叶和籽粒中含磷量以及磷吸收累积量、磷利用效率、磷肥利用率和施磷增产率等在基因型间均存在极显著     

Genetic differences for nitrogen uptake and nitrogen utilisation efficiencies in winter wheat

Due to economic and ecological factors, European agricultural practices are likely to go towards extensive systems with lower inputs of nitrogen (N) fertilisers. The objective of this study was to assess varietal differences for N use at two nitrogen levels. A set of 20 winter wheat (Triticum aestivum L.) genotypes was studied over 2 years in northern France on a deep loam soil without (N0) and with 170 kg ha⁻¹ N fertiliser (N+) as ammonium nitrate. Results were consistent on both years as the genotype×year or genotype×year×N level interactions were not significant. The genotype×N level interaction was highly significant except for total N utilisation efficiency (total above-ground dry weight/total above-ground N) and grain N concentration. The genotype×N level interaction for grain yield was mainly due to three contrasting genotypes: Cappelle, a cultivar from the 1940s, had the same yield at N0 and N+; Arche had a high yield at both N levels; and Récital had a high yield with added N and a very low one without N. The number of kernels/ear explained most of the variations of grain yield at N0 (48%) and N+ (80%), and of the interaction (67%). N uptake efficiency (total above-ground N/soil N supply) accounted for 64% of the variation in N use efficiency (grain yield/soil N supply), while at N0 and at N+ it accounted for only 30%. N utilisation efficiency (grain yield/total above-ground N) was then more important at N+ than at N0. Grain N explained most of total plant N variation at both N levels. The interaction for N use efficiency was best explained by the interaction of N uptake (63%). The applications of these results to a breeding programme to create varieties adapted to low-input management systems are discussed.     

Importance of P uptake efficiency versus P utilization for wheat yield in acid and calcareous soils in Mexico

There are large agricultural areas in the world where wheat yields are limited by low phosphorus (P) availability. Breeding for P uptake and P utilization efficiency may reduce this problem. This study was conducted to determine the contribution of P uptake and utilization efficiency to grain yield of selected spring wheat genotypes in different environments. Thirty-eight semidwarf spring bread wheat (Triticum aestivum) genotypes were grown in two experiments in Mexico, each on an acid Andisol under rainfed conditions and on a calcareous Aridisol with irrigation, without (−P) and with 35 kg P per ha fertilized (+P). Without P fertilization, grain yield ranged from 0.8 to 4.6 t ha⁻¹ in the acid soil and from 2.4 to 5.2 t ha⁻¹ in the calcareous soil. With P fertilization, this range was even larger. Under conditions of P deficiency, i.e. in the acid soil at −P and +P (high P adsorption) and calcareous soil at −P (P-depleted soil), P uptake explained 71–100% of the variation in grain yield, and was highly correlated with grain yield (r=0.79–0.95). In contrast, at +P in the calcareous soil, P utilization efficiency explained 60–63% of the variation in grain yield. Here, low grain P concentration was related to high grain yield (r=−0.40 to −0.59). In the calcareous soil, the harvest index was correlated with grain yield, irrespective of the P level. In the acid soil, post-anthesis P accumulation was important. It was positively correlated with grain yield, whereas in the calcareous soil, no post-anthesis-P accumulation occurred. Here, grain P accumulation at maturity was completely determined by translocation of pre-anthesis shoot P. We conclude that the combination of improved P uptake and P utilization efficiency in the same genotypes requires selection under both high and low-P conditions.     

茄子氮效率基因型差异的研究

为了解茄子的氮素吸收利用特性和筛选氮高效基因型,采用田间小区试验和测试分析方法,通过测定茄子产量、氮利用效率、氮响应度和氮素吸收总量等指标,对10个茄子基因型进行了氮效率方面的研究。结果表明,不同基因型茄子的产量、氮利用效率、氮响应度和氮素吸收总量均存在显著差异。根据供试材料在施氮和不施氮处理下氮效率的不同将供试10个基因型划分为5种类型:高氮高效-低氮高效型(HH-LH)包括06-991;高氮高效-低氮中效型(HH-LM)包括06-917;高氮高效-低氮低效型(HH-LL)包括06-961和06-867;高氮中效-低氮低效型(HM-LL)包括06-972,06-947,06-854,06-830和06-909;高氮低效-低氮低效型(HL-LL)包括06-910。对氮效率构成因素通径分析表明,氮素吸收总量对氮效率的直接作用大于氮利用效率的直接作用,氮素吸收总量是决定氮效率的主要因素。     

Efficiency in the use of phosphorus,nitrogen and potassium in topless faba beans (Vicia faba L.) – variability and inheritance*

In a field experiment with topless faba beans, parental inbred lines and FI hybrids of a 7 × 7-diallel were evaluated for traits related to the P, N and K efficiency. In addition, the parental inbreds were cultivated in a pot experiment at two P levels (100 vs. 700 mg P per pot). Significant heterosis for grain yield, uptake and utilization efficiency of nutrients in the field experiment highlighted the enhancement of nutrient efficiency in the hybrids. Amongst both parental inbreds and hybrids, significant genotypic variation was found for nearly all recorded traits. In the diallel analysis, the GCA effects generally proved to be highly significant and in most cases considerably higher than the SCA effects. In the pot experiment, grain yield ranking of the parental lines at the high P level (= P700) was very similar to that in the field experiment, whereas at the low P level (= P100, – P deficiency) the ranking of the lines changed considerably. Correspondingly, the P level-line interaction was highly significant. High tolerance towards P deficiency was found for those two parental inbreds derived from cultivars bred under Syrian conditions. At the high P supply, P efficiency was more favoured by a high P uptake, but at P deficiency by a high internal P utilization.     

Estimation of quantitative genetic parameters for grain yield and quality in winter wheat under high and low nitrogen fertilization

The main question connected with developing wheat cultivars adapted to low nitrogen (N) is whether separate breeding programs for low and high input conditions are necessary. Nineteen wheat cultivars were grown over three years in Croatia in a total of eight environments at high N and low N in order to determine the effect of two N levels on means, variances and heritability of grain yield and bread-making quality and to assess the relative efficiency of indirect selection under high N in improving trait means under low N. Means of grain yield and grain protein content decreased under low N 10 and 13 %, respectively compared to the high N, whereas higher reductions of means due to lower fertilization were observed for grain N yield (21 %), wet gluten content (20 %), Zeleny sedimentation value (27 %) and for most rheological parameters, whose mean values were reduced from 20 % to as much as 57 %. Heritabilities for grain yield and grain N yield at high N were 0.82 and 0.76, respectively, and 0.77 and 0.43 at low N. Heritability for dough development time, stability and resistance also tended to be higher at high N than at low N due to a decrease in genetic variance and an increase in error variance at low N. The genetic correlation coefficients between high and low N were higher than 0.90 and the efficiency of indirect selection under high N for performance under low N was near 1.0 for grain yield and for most bread-making quality traits.     

Intraspecific Variation in Nitrogen Uptake and Nitrogen Utilization Efficiency in Wheat (Triticum aestivum L.)

Twenty wheat ( Triticum aestivum L.) varieties differing in plant height were grown in soil culture and evaluated for differences in nitrogen uptake and nitrogen utilization efficiency (NUE) at limited (40 kg N ha⁻¹) and normal (120 kg N ha⁻¹) nitrogen supply. Nitrogen uptake showed 1.4- and 1.5-fold varietal variation at harvest for limited and normal N supply, respectively. NUE for dry matter production (NE1) exhibited 1.28- and 1.38-fold genotypic variation while NUE for grain production (NE2) varied by 1.25- and 1.21-fold at limited and normal N supply, respectively. Tall varieties were found to have higher N uptake and NUE for dry matter production, while dwarf cultivars had greater NUE for grain production. Nitrogen uptake was found to be strongly positively associated with dry matter production (r=0.85 and r =0.77 at limited and normal N supply, respectively), indicating an important effect of growth rate on N uptake. NUE for biomass production, as well as for grain production, was reduced as the supply of nitrogen was increased.     

不同基因型玉米硫素吸收利用效率的研究 Ⅱ.硫素吸收利用的基因型差异

在两个硫素供应水平下,研究34个玉米不同基因型的硫素吸收利用情况,通过相关分析和聚类分析,对玉米硫素利用类型进行鉴定。研究结果表明,不同基因型玉米的籽粒产量、硫素吸收和利用效率指标都存在很大差异,根据这些指标,通过聚类分析将参试基因型划分为4类。自交系和甜玉米的有关指标均低,与普通玉米杂交种有很大区别     

Remobilization of Nitrogen and Carbohydrate from Stems of Bread Wheat in Response to Heat Stress during Grain Filling

When wheat (Triticum aestivum L.) is grown under heat-stress conditions during grain filling, preanthesis stored total non-structural carbohydrates (TNC) and nitrogen (N) could serve as alternative source of assimilates. This study was performed to evaluate wheat genotypes for their ability to accumulate and remobilize TNC and N stored in their stem to support grain filling under heat stress. Eighteen genotypes were used for N remobilization study while nine of them were used for TNC remobilization study. They were grown in pots and placed in a vinyl house with the maximum temperature kept below 30 °C. Five days after anthesis (5DAA), half of the pots were taken to phytotrons where temperature was gradually increased and the maximum was set at 38 °C. Grain yield and grain weight decreased by about 35 % under heat stress. Significant differences were found among genotypes in percentage reduction in grain yield, grain weight, grain filling duration and harvest index because of heat stress. The N and TNC concentrations of the stem at 5DAA were significantly different among genotypes. Heat stress significantly reduced the N remobilization efficiency of most of genotypes. However, heat stress significantly increased TNC remobilization efficiency and significant variation were observed among genotypes. N remobilization efficiency across treatments significantly correlated with grain yield, grain weight, harvest index and grain filling duration. TNC at 5DAA negatively correlated with N at 5DAA and harvest index, but the TNC remobilization efficiency under heat stress positively correlated with mainstem grain yield, grain weight and harvest index. The rate of chlorophyll loss from flag leaf positively correlated with N and TNC remobilization efficiencies under heat stress suggesting a link between leaf senescence and remobilization efficiency. The results indicate that heat stress negatively affected grain yield, its components and N remobilization while it increased TNC remobilization because of the increasing demand for resources.     

高产冬小麦水分利用效率及其组分特征分析

依据8年田间和盆栽试验,分析了冬小麦基因型、密度和供水对水分利用效率(WUE)的影响。结果表明:当供水量变化范围较大时,总耗水(ET)和WUE呈显著线性负相关,ET对WUE的负效应明显大于产量(GY)对WUE的正效应。GY和ET有正相关趋势。当供水量变化范围较小时,GY和WUE呈显著线性正相关。蒸散效率(ETE)和收获指数(HI )都和WUE有正     

Diversity of crop development traits and nitrogen use efficiency among potato cultivars grown under contrasting nitrogen regimes

Potato (Solanum tuberosum L.) requires abundant nitrogen (N) to perform well and has low nitrogen use efficiency (NUE). We assessed phenotypic variation among 189 potato cultivars for NUE and the association between NUE and ecophysiological variables describing canopy development (CDv), under high and low N input. In 2009 and 2010, 189 cultivars were grown with N supply (soil N + fertiliser N) of 75 or 180 kg N/ha at Bant, the Netherlands. CDv was assessed weekly as the percentage of soil covered by green potato leaves (%SC). Data were analysed using a model that described CDv as a function of thermal time, based on the Beta function and estimates of cardinal temperatures. Nitrogen significantly affected model-derived, biologically relevant, curve-fit parameters for each cultivar. The t 1 (i.e., thermal time required to reach maximum soil cover (Vx)) was higher at low than at high N. Other parameters were higher at high than at low N, especially Vx and the period over which it was maintained. Nitrogen also affected tuber dry matter yield, tuber size and weight distributions, N content and N uptake but not tuber dry matter percentage. The total area under the %SC curve was highly correlated with yield in both years. Cultivars performing well under high N also performed well under low N. There was large variation in NUE component traits among cultivars; maturity type partially explained this variation. Variables of the CDv model captured this variation, N effects on light interception and its correlation with yield.     

水氮管理模式对不同氮效率水稻氮素利用特性及产量的影响

以高产氮高效品种(德香4103)和中产氮低效品种(宜香3724)为材料,通过“淹水灌溉+氮肥优化运筹(W1N1)”、“控制性交替灌溉+氮肥优化运筹(W2N1)”、“旱种+氮肥优化运筹(W3N2)” 3种水氮管理模式处理,研究其对氮素利用及产量的影响及其生理特性,并探讨氮素利用及产量与生理响应间的关系。结果表明,氮效率品种间的差异与水氮管理模式对水稻氮素利用特征、灌溉水生产效率、生理特性及产量均存在显著影响;不同氮效率品种间在氮肥利用效率方面的差异明显高于水氮管理模式的调控效应;而水氮管理模式对灌溉水生产效率、总吸氮量、氮素干物质生产效率及稻谷生产效率的调控作用显著。W2N1相对于W1N1及W3N2水氮管理模式能促进不同氮效率水稻拔节至抽穗期、抽穗至成熟期氮素的累积,提高功能叶谷氨酰胺合成酶(GS)活性、光合速率(P_n)及根系活力,进而提高稻谷产量及氮肥利用率,且对中产氮低效品种的调控效应显著高于对高产氮高效品种,为本试验最佳的水氮管理模式。高产氮高效品种的平均总颖花数、拔节至抽穗期稻株氮累积量、功能叶GS活性、P_n及根系活力均显著高于氮低效品种,尤其结实期高产氮高效品种更有利于维持叶片及根系的代谢同化能力,利于氮素转运、再分配到籽粒中提高稻谷生产效率及氮肥利用效率,是氮高效品种相对于氮低效品种高产、氮高效利用的重要原因。相关分析表明,水氮管理模式下不同氮效率水稻主要生育时期功能叶GS活性、P_n及根系活力与氮素利用及稻谷产量均存在显著或极显著的正相关;尤其以水稻抽穗期剑叶GS活性及根系活力与氮素利用及稻谷产量的正相关性最高。     

不同基因型旱稻早期锌吸收与利用差异研究

为了研究不同基因型旱稻早期对锌的吸收、利用差异,以‘巴西陆稻’（籽粒低锌）与‘秦爱-3’（籽粒富锌）为材料,在低、中、高(0μmol/L,0.5μmol/L,100μmol/L)不同供锌水平下,对基因型间最大分蘖期植株锌吸收、分布以及锌的利用效率的差异进行了比较分析。结果表明,两基因型根部和地上部干重随着供锌水平增加均呈先增加后降低趋势;植株锌吸收量随着供锌水平升高而增加,而锌利用效率却呈先增加后降低趋势,在低、中供锌水平下,‘秦爱-3’锌吸收量和锌利用效率均高于‘巴西陆稻’,其中锌利用效率差异达到显著水平(P≤0.05);无论中锌还是高锌水平下,‘秦爱-3’锌相对吸收量均高于‘巴西陆稻’,虽然差异未达到显著。因此与籽粒低锌基因型‘巴西陆稻’相比,籽粒富锌基因型‘秦爱-3’在低、中锌水平下具有较强的锌吸收能力以及较高的锌利用效率。     

高氮条件下不同黄瓜品种氮素吸收利用的差异

为了研究高氮条件下黄瓜对氮素吸收利用的特性,以5个黄瓜品种为材料,在高氮地块中研究施氮和不施氮水平下黄瓜品种氮素吸收利用的差异。结果表明：施氮处理中黄瓜品种产量、干物质量及总吸氮量无显著差异,在氮素利用效率上有显著差异;在不施氮处理中黄瓜品种产量、干物质量、总吸氮量及氮素利用效率均有显著差异。相关分析及通径分析表明,土壤高氮情况下不论施肥与否,氮素利用效率对黄瓜产量起主导作用。     

Influence of genotype on phosphate uptake and utilization efficiencies in spring barley

To investigate the genetic variation of phosphate (P) uptake and P utilization efficiency, 24 high-yielding spring barley cultivars were grown in two pot experiments on a loess loam-sand mixture. In the first experiment, the plants grew until maturity under P stress (50% of the maximum yield), and in the second experiment, the plants grew until the stage of tillering (DC 25) at a low or at a high P supply.

At maturity, the range between cultivars with the highest and the lowest values were 30% for total dm yield (grain and straw), 28% for grain yield, 24% for P uptake efficiency (P in grain and straw), 26% for P concentration in grains and 24% for P utilization efficiency quotient PEQ (g dm grain per mg P in shoots) (mean of all cultivars = 100%). Grain yield was correlated with P uptake per plant, r = 0.71^***, and with PEQ, r = 0.60^**. Between P uptake and PEQ, there was only a weak relationship (r = −0.14). Therefore, a combination of high uptake efficiency and high PEQ in a cultivar may be possible.

At growth stage DC 25, the cultivars showed a significant variability in shoot biomass, P concentration, P removal, P influx, acitvity of acid phophatases (P_ase) and root length. The ranking of the cultivars, however, was very different at the two P levels, but the root-length and the P_ase activity were more influenced by genotype than by the P supply.

As the relationships between grain yield, P removal, PEQ and the characters of the young plants cultivated under P stress were very weak (r < 0.43), selection for P efficiency at the stage of tillering cannot be recommended.     

不同氮素处理对中麦175和京冬17产量相关性状和氮素利用效率的影响

本研究旨在了解我国黄淮和北部冬麦区不同施氮量和施氮模式对氮高效吸收和利用的影响,以及中麦175和京冬17产量对不同施氮处理的响应。2013-2014和2014-2015连续两年在河北吴桥和北京顺义两地种植两品种,观测不同施氮量和基追比处理下,冬小麦的群体特性、产量相关性状,以及氮素吸收效率(NUpE)和氮素利用效率(NUtE)。在吴桥点设0、60+0、120+0、120+60、120+120、120+180 kg hm?2 (基肥+拔节肥) 6个处理,在顺义点仅设前5个处理。在总施氮量0~240 kg hm?2 (吴桥)和0~180 kg hm?2 (顺义)范围内,随施氮量增加,归一化植被指数(NDVI)和气冠温差(CTD)提高,群体总粒数和成熟期生物量增加,进而产量提高;但继续增加施氮量会导致粒重、开花前干物质向籽粒转运量、转运率、对籽粒贡献率、收获指数、氮肥偏生产力、氮素吸收和利用效率降低。在不同施氮水平下,中麦175的产量和稳定性均优于京冬17,表现出穗数多、穗粒重稳定性好、群体活力持久、生物量和收获指数高、花前干物质积累量高和花后干物质转运能力强、氮素吸收效率高,这可能是其高产高效的重要基础。考虑到产量回报和经济效益,推荐中麦175和京冬17在黄淮麦区(北片)施氮量为180~240 kg hm?2,在北部冬麦区施氮量为120~180 kg hm?2。灌浆中后期,NDVI和CTD与穗数、产量和生物量相关性高,可作为快速评价品种氮肥敏感性的指标。     

典型玉米自交系氮素吸收利用特点的研究

在大田条件下,对8个代表性的玉米自交系在不同施氮水平下的氮素吸收与利用特点及与氮效率相关的一些性状进行了研究。结果表明：玉米自交系在产量、生物量和氮累积量上存在显著的差异。通过玉米自交系各性状与氮效率的关系的分析表明,低氮胁迫下,应选择抽丝期穗位叶叶绿素含量高、穗位叶叶面积大、籽粒含氮量高和氮收获指数大的自交系,高氮处理条件下,应选择抽丝期生物量大和成熟期生物量大的自交系。通径分析表明,氮吸收和氮利用效率对氮效率的作用均是直接作用大于间接作用。3个施氮水平条件下,对氮效率的直接作用是氮吸收效率大于氮利用效率。在低氮条件下,氮吸收效率的直接作用更强,而氮利用效率对氮效率的直接作用是随着施氮量的增加逐渐增强。在氮2、氮3处理中,吸收效率和利用效率的共同影响导致了产量差异,并且是吸收效率的影响较大;高氮处理则主要是吸收效率的影响。     

不同水肥条件下二倍体、四倍体和六倍体小麦养分吸收、利用效率的研究

试验选用了3个二倍体（Triticum boeoticum，AA；Aegilops speltoides，BB和Ae. tauschii，DD）、2个四倍体（T. dicoccoides，AABB和T. dicoccum，AABB）和1个六倍体（T. aestivum，AABBDD）的小麦进化材料，在不同水肥条件下研究小麦进化中养分效率的差异及水肥条件对小麦养分利用的影响。试验表明，随小麦的进化，养分（N、P和K）吸收量和吸收效率显著增加，但是籽粒养分含量却减少。水分胁迫和低肥处理减少小麦籽粒养分含量、（整株）养分吸收量和吸收效率。小麦的生物量养分利用效率和产量养分利用效率都存在显著的种间差异，而且后者的差异更大。随小麦的进化，生物量养分利用效率和产量养分利用效率均显著增加。6个小麦进化材料的生物量氮、磷利用效率（NUTEb和PUTEb）的大小顺序相同，均为：Ae. speltoides > T. dicoccum、T. dicoccoides、Ae. tauschii、T. boeoticum > T. aestivum，而生物量钾利用效率（KUTEb）是T. aestivum、Ae. tauschii、Ae. speltoides > T. dicoccoides > T. dicoccum > T. boeoticum。Ae. tauschii的产量养分利用效率显著高于其他两个二倍体小麦（AA和BB），表明D组染色体上存在有控制高效利用养分的基因。产量氮、磷、钾利用效率（NUTEg、PUTEg、KUTEg）和收获指数（HI）的大小排序均为：T. aestivum > T. dicoccum、T. dicoccoides、Ae. tauschii > T. boeoticum、Ae. speltoides。水分与养分对生物量养分利用效率有显著影响，而对产量养分利用效率的作用却不显著，说明小麦产量养分利用效率是较为稳定的特性，主要由基因型决定。水分胁迫有利于提高小麦生物量养分利用效率，但是增加施肥量却对其有负作用。