A first assessment of the impact of the extreme 2018 summer drought on Central European forests

In 2018, Central Europe experienced one of the most severe and long-lasting summer drought and heat wave ever recorded. Before 2018, the 2003 millennial drought was often invoked as the example of a “hotter drought”, and was classified as the most severe event in Europe for the last 500 years. First insights now confirm that the 2018 drought event was climatically more extreme and had a greater impact on forest ecosystems of Austria, Germany and Switzerland than the 2003 drought. Across this region, mean growing season air temperature from April to October was more than 3.3°C above the long-term average, and 1.2°C warmer than in 2003. Here, we present a first impact assessment of the severe 2018 summer drought and heatwave on Central European forests. In response to the 2018 event, most ecologically and economically important tree species in temperate forests of Austria, Germany and Switzerland showed severe signs of drought stress. These symptoms included exceptionally low foliar water potentials crossing the threshold for xylem hydraulic failure in many species and observations of widespread leaf discoloration and premature leaf shedding. As a result of the extreme drought stress, the 2018 event caused unprecedented drought-induced tree mortality in many species throughout the region. Moreover, unexpectedly strong drought-legacy effects were detected in 2019. This implies that the physiological recovery of trees was impaired after the 2018 drought event, leaving them highly vulnerable to secondary drought impacts such as insect or fungal pathogen attacks. As a consequence, mortality of trees triggered by the 2018 events is likely to continue for several years. Our assessment indicates that many common temperate European forest tree species are more vulnerable to extreme summer drought and heat waves than previously thought. As drought and heat events are likely to occur more frequently with the progression of climate change, temperate European forests might approach the point for a substantial ecological and economic transition. Our assessment also highlights the urgent need for a pan-European ground-based monitoring network suited to track individual tree mortality, supported by remote sensing products with high spatial and temporal resolution to track, analyse and forecast these transitions.     

Ecosystem‐scale volatile organic compound fluxes during an extreme drought in a broadleaf temperate forest of the Missouri Ozarks (central USA)

Considerable amounts and varieties of biogenic volatile organic compounds (BVOCs) are exchanged between vegetation and the surrounding air. These BVOCs play key ecological and atmospheric roles that must be adequately represented for accurately modeling the coupled biosphere–atmosphere–climate earth system. One key uncertainty in existing models is the response of BVOC fluxes to an important global change process: drought. We describe the diurnal and seasonal variation in isoprene, monoterpene, and methanol fluxes from a temperate forest ecosystem before, during, and after an extreme 2012 drought event in the Ozark region of the central USA. BVOC fluxes were dominated by isoprene, which attained high emission rates of up to 35.4 mg m^?2 h^?1 at midday. Methanol fluxes were characterized by net deposition in the morning, changing to a net emission flux through the rest of the daylight hours. Net flux of CO₂ reached its seasonal maximum approximately a month earlier than isoprenoid fluxes, which highlights the differential response of photosynthesis and isoprenoid emissions to progressing drought conditions. Nevertheless, both processes were strongly suppressed under extreme drought, although isoprene fluxes remained relatively high compared to reported fluxes from other ecosystems. Methanol exchange was less affected by drought throughout the season, confirming the complex processes driving biogenic methanol fluxes. The fraction of daytime (7–17 h) assimilated carbon released back to the atmosphere combining the three BVOCs measured was 2% of gross primary productivity (GPP) and 4.9% of net ecosystem exchange (NEE) on average for our whole measurement campaign, while exceeding 5% of GPP and 10% of NEE just before the strongest drought phase. The megan v2.1 model correctly predicted diurnal variations in fluxes driven mainly by light and temperature, although further research is needed to address model BVOC fluxes during drought events.     

Contrasting sensitivities of two dominant C4 grasses to heat waves and drought

Heat waves and droughts are predicted to increase in frequency and intensity with climate change. However, we lack a mechanistic understanding of the independent and interactive effects of severe heat and water stress for most ecosystems. In a mesic tallgrass prairie ecosystem, we used a factorial experimental approach to assess ecophysiological and productivity responses of two dominant C₄ grasses, Andropogon gerardii and Sorghastrum nutans, to a season-long drought and a mid-summer heat wave at four intensities. We hypothesized that drought would have greater impacts than heat waves, that combined effects would be greater than either factor alone, and that the dominant grasses would differ in their responses to heat and water heat stress. We detected significant reductions in photosynthesis, leaf water potential, and productivity with drought but few direct responses to the heat waves. Surprisingly, there was no additive effect of heat and water stress on any plant response. However, S. nutans was more sensitive than A. gerardii to drought. In this grassland, water stress will likely dominate photosynthetic and productivity responses caused by discrete drought and heat wave events, rather than direct or additive effects of heat stress, with differential sensitivity in these grasses altering future ecosystem structure and function.     

BVOCs: plant defense against climate warming?

Plants emit a substantial amount of biogenic volatile organic compounds (BVOCs) into the atmosphere. These BVOCs represent a large carbon loss and can be up to approximately 10% of that fixed by photosynthesis under stressful conditions and up to 100gCm(-2) per year in some tropical ecosystems. Among a variety of proven and unproven BVOC functions in plants and roles in atmospheric processes, recent data intriguingly link emission of these compounds to climate. Ongoing research demonstrates that BVOCs could protect plants against high temperatures. BVOC emissions are probably increasing with warming and with other factors associated to global change, including changes in land cover. These increases in BVOC emissions could contribute in a significant way (via negative and positive feedback) to the complex processes associated with global warming.     

陆地生态系统植物挥发性有机化合物的排放及其生态学功能研究进展

综述了国内外生物源挥发性有机化合物 (Biologicalvolatileorganiccompounds, BVOCs) 研究现状及未来的研究方向, 侧重介绍了陆地生态系统中植物排放BVOCs的种类、生物学功能及其对大气化学过程的影响。BVOCs按其化学结构以及在大气中的滞留时间可以分为 4类 :异戊二烯、单萜、其它活性BVOCs和其它次活性BVOCs。不同的植物类群排放不同的BVOCs种类并具有不同的排放特性, 环境条件对植物不同BVOCs的排放影响也不同。BVOCs作为有机物质被排放到体外, 从植物能量代谢的角度来讲要消耗一部分植物光合作用产物从而降低植物的生产力, 因此推测植物排放BVOCs具有一定的生理学或者生态学的功能。其中比较成熟的假说是抗热胁迫假说, 其次是抗氧化假说, 也有一些其它假说例如促氮同化假说等。但这些假说都还缺乏直接的有力证据, 有待更多的研究来支持。BVOCs被排放到大气中对大气化学过程的影响更是科学家关注的问题, BVOCs对大气的影响一方面是在大气对流层中促进臭氧 (O3 ) 的形成, 造成环境污染, 另一方面BVOCs通过对大气中的OH自由基和臭氧等氧化物浓度的调整而影响到大气中甲烷等温室气体的平衡, 对大气温室效应具有间接的贡献。我国在BVOCs的研究上也做了大量的工作, 包括分析鉴定了一些植物排放的BVOCs, 探讨了环境因子对植物BVOCs排放速率的影响, 从不同尺度估测了BVOCs的排放量等等。今后对BVOCs的研究将会集中在以下几个方面 :1) 进一步研究不同植物类群释放的BVOCs种类及其它们在大气中的理化性质 ;2 ) 继续探讨植物排放BVOCs的合成与代谢途径及其生物学功能 ;3) 研究BVOCs对大气化学过程的作用, 以及区域植被变化对BVOCs排放格局进而对区域乃至全球环境变化的影响 ;4 ) 加强对一些研究比较薄弱的生态系统例如在热带地区所进行的BVOCs研究工作 ;5 ) 进一步建立和完善BVOCs排放的理论模型, 以模拟不同陆地生态系统BVOCs排放的时空动态。     

Effects of extreme drought on terrestrial ecosystems: review and prospects

作为地球表层重要的组成部分, 陆地生态系统是人类生存和发展的重要场所。进入21世纪以来, 气候变化导致干旱事件发生的强度、频度和持续时间显著增加, 对陆地生态系统带来深远的影响, 严重制约甚至威胁人类社会的可持续发展。因此, 开展极端干旱对陆地生态系统影响的研究并评估其生态风险效应, 是当前全球变化领域研究的重点问题。该文从植物生理生态过程、生物地球化学循环、生物多样性以及生态系统结构和功能4个方面综述了极端干旱对陆地生态系统的影响, 并对当前的研究热点进行探讨, 深度剖析当前研究中存在的难点问题和未来可能的发展方向, 以期为未来开展干旱对陆地生态系统影响的观测与预测研究提供参考, 为在未来干旱影响下加强陆地生态系统风险评估和管理提供新思路。     

DROUGHT EFFECTS ON CARBON EXCHANGE IN A SUBTROPICAL CONIFEROUS PLANTATION IN CHINA

 干旱对陆地生态系统的影响已成为全球变化研究的焦点问题之一。该研究基于生态系统过程模型——CEVSA2, 结合涡度相关通量观测, 分析了不同程度干旱对亚热带人工针叶林碳交换的影响及其关键控制因素。结果表明: 1)干旱使生态系统碳交换显著下降, 2003和2004年的干旱使得年净生态系统生产力(Net ecosystem production, NEP)相比无干旱影响情景的模拟结果分别减少了63%和47%; 2)光合和呼吸对干旱具有不同的响应, 干旱时光合的下降比呼吸更为显著, 这导致了NEP的显著下降; 3)当饱和水气压差(Vapor pressure deficit, VPD)达到1.5 kPa以上时, 生态系统的光合、呼吸和净碳吸收均开始下降, 当VPD大于2.5 kPa、土壤相对含水量(土壤含水量/土壤饱和含水量)(Relative soil water content, RSW)低于40%时, 生态系统的碳收支由碳汇转为碳源; 4)土壤干旱是造成碳交换下降的主要驱动因素, 对年NEP下降的平均贡献率为46%, 而大气干旱的贡献率仅为4%。     

Improvement of heat and drought photosynthetic tolerance in wheat by overaccumulation of glycinebetaine

Within their natural habitat, crops are often subjected to drought and heat stress, which suppress crop growth and decrease crop production. Causing overaccumulation of glycinebetaine (GB) has been used to enhance the crop yield under stress. Here, we investigated the response of wheat (Triticum aestivum L.) photosynthesis to drought, heat stress and their combination with a transgenic wheat line (T6) overaccumulating GB and its wild-type (WT) Shi4185. Drought stress (DS) was imposed by controlling irrigation until the relative water content (RWC) of the flag leaves decreased to between 78 and 82%. Heat stress (HS) was applied by exposing wheat plants to 40°C for 4 h. A combination of drought and heat stress was applied by subjecting the drought-stressed plants to a heat stress as above. The results indicated that all stresses decreased photosynthesis, but the combination of drought and heat stress exacerbated the negative effects on photosynthesis more than exposure to drought or heat stress alone. Drought stress decreased the transpiration rate (Tr), stomatal conductance (Gs) and intercellular CO₂ concentration (Ci), while heat stress increased all of these; the deprivation of water was greater under drought stress than heat stress, but heat stress decreased the antioxidant enzyme activity to a greater extent. Overaccumulated GB could alleviate the decrease of photosynthesis caused by all stresses tested. These suggest that GB induces an increase of osmotic adjustments for drought tolerance, while its improvement of the antioxidative defense system including antioxidative enzymes and antioxidants may be more important for heat tolerance.     

A climatic threshold triggers the die‐off of peat mosses during an extreme heat wave

Heat waves, which are projected to be more frequent and intense in a warmer climate, could become a serious threat to plants that rely on water surplus availability, such as bryophytes. Here, I take the advantage of the European summer 2003 climate anomaly to assess the impact of an extreme heat wave on peat mosses of the genus Sphagnum, a group of bryophytes forming the bulk of living and dead biomass in peatlands. With this aim, 20 selected bogs in the Italian Alps were checked for Sphagnum survival in the years following the heat wave. Over the study area, the period May–September 2003 was characterized by higher mean monthly air temperature (13.5 °C) and lower mean monthly precipitation (87 mm) compared with normal climatic conditions (11.5 °C and 117 mm, respectively) so that the heat wave coincided with a drought spell. As a consequence of the unusual water stress, I documented an increased mortality of peat mosses forming high hummocks. In particular, at habitat scale, the distribution of desiccated peat mosses was restricted to the hummock face receiving the greatest amount of solar irradiation. However, at regional scale, the present study identified a climatic threshold, simply defined by the ratio of precipitation to temperature (P : T), which triggered an irreversible desiccation of peat mosses when mean monthly P : T dropped below 6.5 (mm : °C) during May–September 2003. The absence of any sign of recovery after 4 years since the drought must be seen as a harbinger of the deleterious effects of extreme heat waves on organisms not adapted to cope with abrupt climate anomaly.     

Non-Methane Biogenic Volatile Organic Compound Emissions from a Subarctic Peatland Under Enhanced UV-B Radiation

Boreal and subarctic peatlands have been extensively studied for their major role in the global carbon balance. However, study efforts have so far neglected the contribution of these ecosystems to the non-methane biogenic volatile organic compound (BVOC) emissions, which are important in the atmospheric chemistry and feedbacks on climate change. We aimed at estimating the BVOC emissions from a subarctic peatland in northern Finland. Furthermore, our aim was to assess how these emissions are affected by enhanced UV-B radiation, the amount of which has increased especially at high latitudes as a result of stratospheric ozone depletion. The contribution of BVOC emissions to the total net carbon exchange and correlations between the emission of different BVOCs and net ecosystem CO₂ exchange, CH₄ emission, total green leaf area, and abiotic factors were also studied. The UV-B exposure, simulating a 20% depletion of stratospheric ozone, was started in 2003, and measurements were performed during the growing seasons of 2006 and 2008. The subarctic peatland proved to be a small source of BVOCs and the dominant moss, Warnstorfia exannulata, emitted a diverse compound spectrum. The water table level exerted a major influence on the BVOC emissions surpassing the effect of enhanced UV-B. In fact, no overall UV-B effect was established on the BVOC emissions, apart from toluene and 1-octene, emissions of which were doubled and tripled by enhanced UV-B in 2008, respectively. The contribution of BVOCs to the total net carbon exchange was below 1%.     

Midsummer heat wave effects on lacustrine plankton: Variation of assemblage structure and fatty acid composition

The effects of the 2003 European heat wave on a freshwater plankton assemblage and its fatty acid (FA) composition were investigated. Composition and FA profiles of four size categories of planktonic organisms collected in 2003 were compared to those of the colder year 2002.     

外来植物入侵对土壤生物多样性和生态系统过程的影响

随着科学家对生态系统地下部分的重视,评价外来植物入侵对土壤生态系统的影响成为当前入侵生态学领域的研究热点之一。本文综述了外来植物入侵对土壤微生物、土壤动物以及土壤碳、氮循环动态影响的研究,并探讨了其影响机制。已有的研究表明,植物入侵对土壤生物多样性及相关生态系统过程的影响均存在不一致的格局,影响机制也是复杂多样的。外来植物与土著植物凋落物的质与量、根系特征、物候等多种生理生态特性的差异可能是形成格局多样性和影响机制复杂性的最主要原因。今后,加强多尺度和多生态系统的比较研究、机制性研究、生物多样性和生态系统过程的整合性研究及土壤生态系统对植物入侵的反馈研究是评价外来植物入侵对土壤生态系统影响的发展趋势。     

供氮水平对落叶松幼苗光合作用的影响

为探讨落叶松光合作用对外界供氮环境变化的响应规律,采用砂培方法在温室内设置了4种供氮浓度(1、4、8mmol/L和16mmol/L),对生长在不同供氮水平下落叶松(Larixgmelinii)1年生幼苗的气体交换参数、叶绿素荧光参数以及一些生化指标进行了测定。结果显示,随供氮水平的提高,落叶松幼苗针叶氮含量、叶绿素含量、类胡罗卜素含量、可溶性蛋白(TSP)含量和光饱和净光合速率(Pmax)均显著增加,同时伴随着叶磷含量和胞间与外界CO2浓度之比(Ci/Ca)的降低。然而,当供氮水平超过8mmol/L增至16mmol/L时,TSP含量及Pmax不再增加,反而略有下降。供氮不足显著降低了幼苗针叶光系统最大光能转换效率(Fv/Fm)、光系统量子效率(ΦPS)和光化学猝灭系数(qP),却增加了非光化学猝灭系数(NPQ)。而增加供氮可使Fv/Fm和ΦPS回升,同时qP升高,NPQ下降,但当供氮水平超过8mmol/L后,各叶绿素荧光参数变化幅度较小。结果表明,增加供氮可显著提高落叶松幼苗的光合能力,增加光系统天线色素捕获的光能用于光化学电子传递的份额,减缓光抑制。然而,16mmol/L已经超过落叶松幼苗最适的供氮水平,过量供氮引起的负面效应可能主要与过低的叶磷含量导致的营养失衡有关。     

Increasing climate sensitivity of beech and pine is not mediated by adaptation and soil characteristics along a precipitation gradient in northeastern Germany

Rising temperature and altered precipitation regimes will lead to severe droughts and concomitant extreme events in the future. Forest ecosystems have shown to be especially prone to climate change. In assessing climate change impacts, many studies focus on high altitude or ecological edge populations where a climate signal is supposedly most pronounced. While these studies represent only a fraction of the forest ecosystems throughout Europe, findings on climate sensitivity of lowland core populations remain comparatively underrepresented.By using tree-ring widths of a large region-wide network of European beech and Scots pine populations along a precipitation gradient in northeastern Germany, we identify main climatic drivers and spatio-temporal patterns in climate sensitivity. Further, we analyze the resistance of tree growth towards drought. Detailed data on soil characteristics was used to interpret climate-growth relationships.Beech was found to be most sensitive to summer drought during early summer at dry sites, whereas pine displayed highest sensitivity for winter temperature at wet sites. The resistance to extreme drought was lower for beech. By splitting the observation period (1964–2017) into an early and late period, we found non-stationary climate-growth relationships for both study species with beech showing an increase in drought sensitivity and pine in winter temperature sensitivity.Overall, beech populations seem to be especially endangered by prospective climate changes, whereas climate-growth relationships of pine seem more ambiguous with a possible trade-off between enhanced photosynthetic activity caused by early photosynthesis in late winter and reduced activity due to summer drought.     

Performance of High Arctic tundra plants improved during but deteriorated after exposure to a simulated extreme temperature event

Arctic ecosystems are known to be extremely vulnerable to climate change. As the Intergovernmental Panel on Climate Change scenarios project extreme climate events to increase in frequency and severity, we exposed High Arctic tundra plots during 8 days in summer to a temperature rise of approximately 9°C, induced by infrared irradiation, followed by a recovery period. Increased plant growth rates during the heat wave, increased green cover at the end of the heat wave and higher chlorophyll concentrations of all four predominating species (Salix arctica Pall., Arctagrostis latifolia Griseb., Carex bigelowii Torr. ex Schwein and Polygonum viviparum L.) after the recovery period, indicated stimulation of vegetative growth. Improved plant performance during the heat wave was confirmed at plant level by higher leaf photochemical efficiency (F_v/F_m) and at ecosystem level by increased gross canopy photosynthesis. However, in the aftermath of the temperature extreme, the heated plants were more stressed than the unheated plants, probably because they acclimated to warmer conditions and experienced the return to (low) ambient as stressful. We also calculated the impact of the heat wave on the carbon balance of this tundra ecosystem. Below‐ and aboveground respiration were stimulated by the instantaneous warmer soil and canopy, respectively, outweighing the increased gross photosynthesis. As a result, during the heat wave, the heated plots were a smaller sink compared with their unheated counterparts, whereas afterwards the balance was not affected. If other High Arctic tundra ecosystems react similarly, more frequent extreme temperature events in a future climate may shift this biome towards a source. It is uncertain, however, whether these short‐term effects will hold when C exchange rates acclimate to higher average temperatures.     

植物干旱胁迫下水分代谢、碳饥饿与死亡机理

植物在生长发育过程中受众多环境因子共同作用。随着全球气候变化,气温升高、降水量下降等问题频繁出现。目前气象学家一致预测未来环境变暖会使干旱更加频繁剧烈,这一环境改变使植物死亡更加严重。植物在水分胁迫、特别是干旱胁迫条件下,体内水分代谢与碳代谢会发生失衡现象:光合速率降低、蒸腾速率降低,带来生长降低;为维持植物新陈代谢,植物呼吸作用必然下调。在长期干旱胁迫条件下植物体内碳水化合物储存发生失衡现象,这种失衡使植物陷入碳饥饿现象。另外,由于水分失衡而出现的木质部栓塞和空穴会进一步加剧水分运输障碍,而修复空穴则需要大量非结构性碳水化合物(NSC),这使植物陷入两难选择。总结了植物干旱胁迫下,碳饥饿与水分代谢、植物死亡关系的相关研究,对未来的研究方向和重点提出建议,以期对未来的植物死亡研究提供帮助。     

The effect of experimental ecosystem warming on CO2 fluxes in a montane meadow

Climatic change is predicted to alter rates of soil respiration and assimilation of carbon by plants. Net loss of carbon from ecosystems would form a positive feedback enhancing anthropogenic global warming. We tested the effect of increased heat input, one of the most certain impacts of global warming, on net ecosystem carbon exchange in a Rocky Mountain montane meadow. Overhead heaters were used to increase the radiative heat flux into plots spanning a moisture and vegetation gradient. We measured net whole-ecosystem CO₂ fluxes using a closed-path chamber system, relatively nondisturbing bases, and a simple model to compensate for both slow chamber leaks and the CO₂ concentration-dependence of photosynthetic uptake, in 1993 and 1994. In 1994, we also measured soil respiration separately. The heating treatment altered the timing and magnitude of net carbon fluxes into the dry zone of the plots in 1993 (reducing uptake by ≈100 g carbon m^–2), but had an undetectable effect on carbon fluxes into the moist zone. During a strong drought year (1994), heating altered the timing, but did not significantly alter the cumulative magnitude, of net carbon uptake in the dry zone. Soil respiration measurements showed that when differences were detected in dry zone carbon fluxes, they were caused by changes in carbon input from photosynthesis, not by temperature-driven changes in carbon output from soil respiration. When differences were detected in dry-zone carbon fluxes, they were caused by changes in carbon input from photosynthesis, not by a temperature-driven changes in carbon output from soil respiration. Regression analysis suggested that the reduction in carbon inputs from plants was due to a combination of two soil moisture effects: a direct physiological response to decreased soil moisture, and a shift in plant community composition from high-productivity species to low-productivity species that are more drought tolerant. These results partially support predictions that warming may cause net carbon losses from some terrestrial ecosystems. They also suggest, however, that changes in soil moisture caused by global warming may be as important in driving ecosystem response as the direct effects of increased soil temperature.     

湿地氮素传输过程研究进展

综述了湿地氮素传输过程的研究进展。湿地氮素传输过程包括物理过程、化学过程和生物过程 ,与土壤、植物的发生、发育紧密联系在一起 ,并形成了空气 -水 -土 -生命系统中物质循环和能量流动的复杂网络。湿地硝态氮的淋失直接威胁着湿地地下水水质安全 ,N2 O源汇转变受土壤和水体等环境因子的制约 ,氨挥发则与水体 p H值密切相关排放。湿地氮素的化学转化过程是矿质养分供给和 N2 O产生的主要机制 ,受环境因子和人类活动干扰的影响 ;动力学模型可用于描述氮素的化学转化过程。湿地植物的吸收和累积以及微生物的分解过程是湿地氮素循环的重要环节。最后分析了当前国内外研究中存在的不足 ,并对未来研究的重点领域进行了展望     

Nitrogen Modulates the Effects of Heat,Drought, and Combined Stresses on Photosynthesis,Antioxidant Capacity,Cell Osmoregulation,and Grain Yield in Winter Wheat

Longer and more intense heat and drought stresses will occur in terrestrial ecosystems in the future. Although the effects of individual heat or drought stress on wheat plants have been largely explored, the regulatory effect of nitrogen (N) on winter wheat under heat, drought, and combined stresses and whether N alleviates damage to wheat plants caused by these stresses remain unclear. Therefore, the objective of the present study was to investigate the growth, photosynthesis, antioxidant enzyme and N metabolism-related enzyme activity, cell membrane system, osmoregulatory substance, and yield responses to heat, drought, and combined stresses in wheat plants and to clarify the regulatory effects of N on the growth, physiological and biochemical characteristics, and yield of wheat plants under stress conditions. The results showed that wheat plant exposure to individual heat or drought stress reduced photosynthesis and N metabolism-related enzyme activities and increased antioxidant enzyme activities, electrolyte leakage (EL), and the contents of MDA (malondialdehyde) and O₂^? (superoxide anion). The above parameters showed typical superposition effects under combined stress. Under individual heat or drought stress, wheat plants treated with a medium (N₂) or high (N₃) N supply maintained higher photosynthesis and N metabolism-related enzyme activities than did those treated with a low N supply (N₁). Enhanced heat and drought tolerance in wheat plants under an appropriate N supply may be attributed to improved antioxidant capacity, as exemplified by increased activities of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), glutathione reductase (GR) and ascorbate peroxidase (APX), and to enhanced osmoregulation capacity, as signified by increased contents of soluble sugar (SS), soluble protein (SP), and proline (Pro). Variable importance in projection (VIP) analysis indicated that efficient SOD, POD, CAT, and GR activities and an increased Pro content had superior potential to alleviate heat, drought, and combined stress stresses in wheat plants, and the improvements in growth and grain yield in wheat plants further confirmed the oxidative stress alleviation and stress tolerance enhancement. However, positive effects of N on wheat growth and grain yield under combined stress were usually observed under a low N supply. These results may facilitate future research on the effects of N fertilizer on the stress resistance of winter wheat.

Graphical Abstract