树木抗旱机理研究进展

干旱是树木成活与生长的重要限制因素之一。研究树木的抗旱性,有助于了解树木的干旱适应性机制,探求树木抗旱的适宜调控措施,对充分发挥森林的生态效益具有重要作用。本文从树木的水分关系、净光合速率与蒸腾速率、叶片的δ13C值、木质部导管空穴、光合机制与抗氧化保护机制、渗透调节、脱落酸和C4植物的抗旱机理及树木的抗旱基因与遗传特性(包括水通道蛋白、逆境信号转导、树木抗旱性的遗传基因工程)方面对树木抗旱机理研究进行了综述。     

Effect of heterogeneous distribution of nutrients on root growth, ABA content and drought resistance of wheat plants

Heterogeneous distribution of nutrients influenced morphology of wheat roots; they were shorter and had more laterals at the site of high nutrient content. The roots outside the nutrient-rich patch were longer and penetrated deeper in the soil than those of the plants supplied with the same quantity of nutrients but having them homogeneously distributed. The ABA content of roots sampled from the site of high nutrient content was greater than in those elsewhere. It is suggested that this might be important for inhibition of their extension growth and stimulation of laterals in the nutrient rich patch. Heterogeneously distributed nutrients beneficially influenced accumulation of ¹⁵N and assimilation of CO₂ by the plants, enabling an increased supply of nutrients to the parts of the root system beyond the nutrient rich patch and sustained their rapid extension. Enhanced penetration of roots into the deeper soil layers could contribute to the observed increase in drought resistance of plants grown with localised placement of fertilisers under field conditions.     

Drought timing influences the legacy of tree growth recovery

Whether and how the timing of extreme events affects the direction and magnitude of legacy effects on tree growth is poorly understood. In this study, we use a global database of Ring‐Width Index (RWI) from 2,500 sites to examine the impact and legacy effects (the departure of observed RWI from expected RWI) of extreme drought events during 1948–2008, with a particular focus on the influence of drought timing. We assessed the recovery of stem radial growth in the years following severe drought events with separate groupings designed to characterize the timing of the drought. We found that legacies from extreme droughts during the dry season (DS droughts) lasted longer and had larger impacts in each of the 3 years post drought than those from extreme droughts during the wet season (WS droughts). At the global scale, the average integrated legacy from DS droughts (0.18) was about nine times that from WS droughts (0.02). Site‐level comparisons also suggest stronger negative impacts or weaker positive impacts of DS droughts on tree growth than WS droughts. Our results, therefore, highlight that the timing of drought is a crucial factor determining drought impacts on tree recovery. Further increases in baseline aridity could therefore exacerbate the impact of punctuated droughts on terrestrial ecosystems.     

Quantifying drylands' drought resistance and recovery: the importance of drought intensity,dominant life history and grazing regime

Projected global change will increase the level of land‐use and environmental stressors such as drought and grazing, particularly in drylands. Still, combined effects of drought and grazing on plant production are poorly understood, thus hampering adequate projections and development of mitigation strategies. We used a large, cross‐continental database consisting of 174 long‐term datasets from >30 dryland regions to quantify ecosystem responses to drought and grazing with the ultimate goal to increase functional understanding in these responses. Two key aspects of ecosystem stability, resistance to and recovery after a drought, were evaluated based on standardized and normalized aboveground net primary production (ANPP) data. Drought intensity was quantified using the standardized precipitation index. We tested effects of drought intensity, grazing regime (grazed, ungrazed), biome (grassland, shrubland, savanna) or dominant life history (annual, perennial) of the herbaceous layer to assess the relative importance of these factors for ecosystem stability, and to identify predictable relationships between drought intensity and ecosystem resistance and recovery. We found that both components of ecosystem stability were better explained by dominant herbaceous life history than by biome. Increasing drought intensity (quasi‐) linearly reduced ecosystem resistance. Even though annual and perennial systems showed the same response rate to increasing drought intensity, they differed in their general magnitude of resistance, with annual systems being ca. 27% less resistant. In contrast, systems with an herbaceous layer dominated by annuals had substantially higher postdrought recovery, particularly when grazed. Combined effects of drought and grazing were not merely additive but modulated by dominant life history of the herbaceous layer. To the best of our knowledge, our study established the first predictive, cross‐continental model between drought intensity and drought‐related relative losses in ANPP, and suggests that systems with an herbaceous layer dominated by annuals are more prone to ecosystem degradation under future global change regimes.     

Leaf and root growth dynamics in Eucalyptus globulus seedlings grown in drying soil

Summary Seedlings of Eucalyptus globulus growing in soil columns were subjected to a 24 day soil drying treatment. Water and solute potentials of both young expanding and fully expanded leaves declined under reduced soil water availability, while slightly higher turgor was sustained by the fully expanded leaves. Although leaf area of unwatered seedlings was smaller, the corresponding leaf dry weight was quite similar to that of well-watered seedlings. Soon after rewatering, leaf area of plants experiencing water shortage was comparable to that of well-watered plants. It seems that a difference in wall properties between juvenile and mature leaves allows for an effective pattern of water use by eucalypt plants growing in drying soil. Some stomatal opening is sustained and therefore, presumably, some carbon may be fixed, keeping the carbon balance of the whole plant positive, and allowing a continuous cell division despite the limited water supply. The highest root density of both well-watered and unwatered plants was found in the upper soil layers. However, root growth of unwatered seedlings was gradually increased in the deeper soil layers, where thicker root apices and higher soil water depletion rates per unit root length were recorded. As a consequence, root absorbing surface area was as large in unwatered plants as in well-watered plants.     

H_2O_2胁迫锻炼对小麦幼苗抗旱性的影响

12 d龄的春小麦幼苗在 1 mmol· L- 1及 1 0 mmol· L- 1H2 O2 的胁迫锻炼过程中 ,质膜透性增大 ,O- · 及 H2 O2 含量增多 ,CAT活性升高 ,叶绿素含量降低 ,低浓度 H2 O2 胁迫使SOD活性上升 ,高浓度时却使其活性下降。经过 H2 O2 胁迫锻炼后的小麦遭受干旱胁迫时 ,叶绿素含量、SOD活性、CAT活性均高于对照组 ,而质膜透性、O- · 及 H2 O2 含量却低于对照组。表明 H2 O2 胁迫锻炼 ,提高了小麦幼苗的抗氧化能力 ,增强了其抗旱性     

水分胁迫下AM真菌对柠条锦鸡儿(Caragana korshinskii)生长和抗旱性的影响

利用盆栽实验研究了水分胁迫条件下AM真菌对柠条锦鸡儿(Caragana korshinskii)生长和抗旱性的影响.在土壤相对含水量为80%、60%和40%条件下,分别接种摩西球囊霉(Glomus mosseae)和柠条锦鸡儿根际土著菌,结果表明,水分胁迫对AM真菌的接种效果有显著影响.不同水分条件下,接种AM真菌显著提高了宿主植物根系菌根侵染率.土壤相对含水量为40%～60%时,接种株的株高、茎粗、生物干重和叶片保水力明显高于不接种株;接种AM真菌提高了植株对土壤有效N和有效P的利用率,增加了植株全P、叶片叶绿素和可溶性糖含量以及SOD、POD、CAT等保护酶活性.土壤相对含水量为40%时,叶片MDA含量明显下降.水分胁迫条件下,以接种柠条锦鸡儿根际土著菌的效果最佳.AM真菌增强宿主植物的抗旱性可能源于促进宿主植物根系对土壤水分和矿质元素吸收的直接作用和改善植物体内生理代谢活动、提高保护酶活性的间接作用.     

高温和干旱胁迫对西红柿幼苗生长、养分含量及元素利用效率的影响

全球气候变化将增加未来高温与干旱的发生频率和强度,然而高温与干旱的交互作用对农作物生长、养分含量及其利用效率的影响还不甚清楚。因此,研究高温与干旱交互作用对农作物生理生态的影响将为准确评价农作物对未来极端气候条件的响应提供科学依据。选取全球第四大经济作物——西红柿为研究对象,在人工智能气候箱中模拟高温和干旱环境。共设置两个水分处理(正常浇水;干旱)与两个温度处理(常温-26℃/19℃(白天/夜间);高温-42℃/35℃(白天/夜间)(7d))。主要测定指标包括生物量以及生物量分配、比叶面积、养分含量(全氮、全磷)、光合元素利用效率(光合氮素利用效率、光合磷素利用效率)。研究表明,高温、干旱单独作用以及交互作用均显著降低了根、茎、叶生物量以及总生物量,并且高温干旱交互作用使总生物量降低最多。在生物量分配方面,高温单独作用显著降低了根质量分数以及根冠比,而干旱单独作用增加了根质量分数、茎质量分数以及根冠比,但降低了叶质量分数。在养分含量方面,高温单独作用导致叶片全氮、全磷含量显著降低、茎全磷含量显著增加、根全磷含量显著降低。干旱单独作用导致叶片、茎全磷含量显著降低、根全氮含量显著升高。高温与干旱交互作用对生物量分配及养分含量的影响与干旱胁迫单独作用类似。在光合元素利用效率方面,高温、干旱单独作用均降低了幼苗光合氮素利用效率、光合磷素利用效率,并且高温加剧了干旱对光合磷素利用效率的影响。因此,在未来气候变化情况下,高温与干旱交互作用可能会对农作物产生更大威胁。     

渗透胁迫下不同抗旱性小麦幼苗氨同化差异

在渗透胁迫下,测定了不同抗旱性小麦（抗旱性强的品种洛旱6号和抗旱性弱的品种周麦18）幼苗氨同化酶及相关参数的变化.结果表明:小麦生物量在渗透胁迫下明显降低,且抗旱性弱的周麦18降幅较大.铵态氮含量随胁迫程度的增加而增加,且周麦18增加较明显;谷氨酰胺合成酶(GS)活性在不同抗旱性品种间表现不同,抗旱性强的洛旱6号在低渗透胁迫下显著增加,在高渗透胁迫下明显降低,而周麦18随胁迫程度的增加逐渐降低;依赖还原型辅酶Ⅰ的谷氨酸脱氢酶(NADH-GDH)活性随胁迫程度的增加逐渐加大,低渗透胁迫下周麦18增加较明显,高渗透胁迫下洛旱6号增幅较大;依赖氧化型辅酶Ⅰ的谷氨酸脱氢酶(NAD^+-GDH)和依赖氧化型辅酶Ⅱ的异柠檬酸脱氢酶(NADP-ICDH)活性均随胁迫程度的增加而增加,周麦18的NAD^+-GDH活性、洛旱6号的NADP-ICDH活性增幅较大.表明小麦抗旱性的提高与铵态氮同化的增强有关,低渗透和高渗透胁迫下分别依赖GS和NADH-GDH活性的增加.     

干旱胁迫下4种常用植物幼苗的光合和荧光特性综合评价

老鸭嘴,山毛豆,假连翘和葛藤是常见的矿山植被恢复植物。采用盆栽控制土壤水分的方法,测定了这 4 种幼苗叶片的光合和荧光指标,以期为矿山植被的恢复植物筛选提供依据。结果表明（1）干旱胁迫处理期间,4 种幼苗叶片的净光合速率（P_n）、气孔导度（G_s）、蒸腾速率（T_r）和胞间CO₂ 浓度（C_i）持续下降,复水8 d后P_n均显著增加,老鸭嘴和山毛豆的G_s和T_r恢复到对照水平,山毛豆和葛藤的C_i恢复到对照水平;假连翘幼苗叶片的气孔限制值（L_s）随干旱胁迫时间的延长而逐渐增加,其他3 种幼苗略降后增加,复水8 d后均恢复到对照水平。（2）干旱胁迫处理期间,4 种幼苗叶片的photosystemⅡ（PSⅡ）光能捕获效率（F_v’/F_m’）、实际光化学量子效率（Yield）和表观电子传递速率（ETR）持续降低,8 d时显著小于对照。复水8 d后,F_v’/F_m’小幅波动,Yield和ETR有所恢复,其中葛藤的Yield和山毛豆的ETR恢复到对照水平;4种幼苗叶片的非化学淬灭（NPQ）持续上升,在第8天显著大于对照,复水8 d后,均持续下降,其中葛藤恢复到对照水平。（3）对光合和荧光指标进行主成分分析表明,干旱胁迫下4 种幼苗的抗旱性能大小顺序为山毛豆 > 老鸭嘴 > 假连翘 > 葛藤。     

Ca—GA合剂浸种对水稻萌发及幼苗期抗旱性的影响

研究在模拟干旱胁迫下,钙赤合剂（Ca-GA）浸种后对水稻种子萌发及幼苗抗旱性的影响。结果表明：经Ca-GA处理后的水稻种子萌发时α-淀粉酶活性显著升高,发芽势、发芽率均高于对照。幼苗的根冠比增大,可溶性糖、Pro含量升高。叶片中SOD、CAT保持较高活性,膜透性较低,MDA、O2^-积累较少,叶片保持较高的PSⅡ光化学活性和光合速率。表明Ca-GA合剂浸种能促进水稻种子的萌发和提高幼苗的抗旱性。     

水分胁迫下AM真菌对沙打旺生长和抗旱性的影响

利用盆栽试验研究了水分胁迫条件下接种AM真菌对优良牧草和固沙植物沙打旺(Astragalus adsurgens Pall.)生长和抗旱性的影响。在土壤相对含水量为70%、50%和30%条件下,分别接种摩西球囊霉(Glomus mosseae)和沙打旺根际土著菌,不接种处理作为对照。结果表明,水分胁迫显著降低了沙打旺植株(无论接种AM真菌与否)的株高、分枝数、地上部干重和地下部干重,并显著提高了土著AM真菌的侵染率,对摩西球囊霉的侵染率无显著影响。接种AM真菌可以促进沙打旺生长和提高植株抗旱性,但促进效应因土壤含水量和菌种不同而存在差异。不同水分条件下,接种AM真菌显著提高了植株菌根侵染率、根系活力、地下部全N含量和叶片CAT活性。土壤相对含水量为30%和50%时,接种株地上部全N、叶片叶绿素、可溶性蛋白、脯氨酸含量和POD活性显著高于未接种株;接种AM真菌显著降低了叶片MDA含量;接种土著AM真菌的植株株高、分枝数、地上部和地下部干重显著高于未接种株。土壤相对含水量为30%时,接种AM真菌显著增加了地上部全P含量和叶片相对含水量;接种摩西球囊霉的植株株高、分枝数、地上部和地下部干重显著高于未接种株。水分胁迫40d,接种AM真菌显著提高了叶片可溶性糖含量。水分胁迫80d,接种株叶片SOD活性显著增加。菌根依赖性随水分胁迫程度增加而提高。沙打旺根际土著菌接种效果优于摩西球囊霉。水分胁迫和AM真菌的交互作用对分枝数、菌根侵染率、叶片SOD、CAT和POD活性、叶绿素、脯氨酸、可溶性蛋白、地上部全N和全P、地下部全N和根系活力有极显著影响,对叶片丙二醛和地下部全P有显著影响。AM真菌促进根系对土壤水分和矿质营养的吸收,改善植物生理代谢活动,从而提高沙打旺抗旱性,促进其生长。试验结果为筛选优良抗旱菌种,充分利用AM真菌资源促进荒漠植物生长和植被恢复提供了依据。     

干旱胁迫对小麦幼苗根系生长和叶片光合作用的影响

采用水培试验方法,以2个耐旱性不同的小麦品种(敏感型望水白和耐旱型洛旱7号)为材料,研究了干旱胁迫对小麦幼苗根系形态、生理特性以及叶片光合作用的影响,以期揭示小麦幼苗对干旱胁迫的适应机制.结果表明： 干旱胁迫下,2个小麦品种幼苗的根系活力显著增大,而根数和根系表面积受到抑制;干旱胁迫降低了望水白的叶片相对含水量,提高了束缚水/自由水,而对洛旱7号无显著影响;干旱胁迫降低了2个小麦品种叶片的叶绿素含量、净光合速率、蒸腾速率、气孔导度和胞间CO2浓度,但随胁迫时间的延长,洛旱7号的叶绿素含量和净光合速率与对照差异不显著;干旱胁迫降低了2个小麦品种幼苗的单株叶面积,以及望水白的根系、地上部和植株生物量,而对洛旱7号无显著影响.水分胁迫下,耐旱型品种可以通过提高根系活力、保持较高的根系生长量来补偿根系吸收面积的下降,保持较高的根系吸水能力,进而维持较高的光合面积和光合速率,缓解干旱对生长的抑制.     

Additive and non-additive responses of seedlings to simulated herbivory and drought

Drought is a global threat, increasing in severity and frequency throughout tropical ecosystems. Although plants often face drought in conjunction with biotic stressors, such as herbivory or disease, experimental studies infrequently test the simultaneous effects of drought and biotic stress. Because multiple simultaneous stressors may have non-additive and complex effects on plant performance, it is difficult to predict plant responses to multiple threats from research examining one stress at a time. Using an experimental approach in the greenhouse, we investigated potential non-additivity in seedling growth and survival to simulated drought and herbivory across a phylogenetically diverse pool of ten Hawaiian plant species. Overall, seedlings showed limited tolerance, defined as similar growth and survival in stressed compared with control (non-stressed) plants, to simulated herbivory and drought, with the combined effects of both stressors to be generally additive and negative across species. Significant variation in stress tolerance was detected among species, and species variation was explained, at least in part, by functional traits such that species with larger root/shoot ratios and smaller seeds, tended to demonstrate greater herbivory and drought tolerance. Future research incorporating additional trait analysis and different stressors could shed light on mechanisms underlying seedling stress tolerance and clarify whether additivity, as detected in this study, extends across other combinations of stressors. Such work will provide needed insights into the regeneration of seedlings in tropical forests under threats of herbivory and climate change.     

土壤干旱对沙棘苗木生长及水分利用的影响

研究了土壤干旱条件下盆栽沙棘苗木生长、光合和水分关系的变化。结果表明,同样环境条件下,蒸腾速率随干旱胁迫程度加重而降低,而蒸腾日进程差异不大,均为单峰曲线,峰值出现在11：00;轻、中度干旱胁迫下沙棘叶含水量、水势、渗透势、膨压下降幅度较小,光合速率、侧枝生长速率保持在较高水平,能维持基本的生长量,表现出耐旱植物的生理特征;重度干旱胁迫使上述指标下降较显著,且长时间胁迫后,33．3％沙棘幼苗死亡。     

Effects of increasing root carbon investment on the mortality and resprouting of Haloxylon ammodendron seedlings under drought

• Tree mortality induced by drought is one of the most complex processes in ecology. Although two mechanisms associated with water and carbon balance are proposed to explain tree mortality, outstanding problems still exist.
• Here, in order to test how the root system benefits survival and resprouting of Haloxylon ammodendron seedlings, we examined the various water‐ and carbon‐related physiological indicators (shoot water potential, photosynthesis, dark respiration, hydraulic conductance and non‐structural carbohydrates [NSC]) of H. ammodendron seedlings, which were grown in drought and control conditions throughout a grow season in greenhouse.
• The survival time of the seedling root system (died 70 days after drought) doubled the survival time of the shoot (died at 35 days). Difference in survival time between shoot and root resulted from sustained root respiration supported by increased NSC in roots under drought. Furthermore, investment into the root contributed to resprouting following drought. Based on these results, a death criterion is proposed for this species. The time sequence of major events indicated that drought shifted carbon allocation between shoot and root and altered the flux among different sinks (growth, respiration or storage). The interaction of water and carbon processes determined death or survival of droughted H. ammodendron seedlings.
• These findings revealed that the ‘root protection’ strategy is critical in determining survival and resprouting of this species, and provided insights into the effects of carbon and water dynamics on tree mortality.

     

种子大小和干旱胁迫对辽东栎幼苗生长和生理特性的影响

在温室内遮阴条件下,设置80%、60%、40%和20%田间持水量(对照、轻度、中度和重度干旱)4个处理,研究种子大小和干旱胁迫对盆栽辽东栎幼苗生长和生理特性的影响。结果表明:大种子(3.05±0.38 g)幼苗的单株叶面积、总干质量和根冠比在所有处理均显著大于小种子(1.46±0.27 g)幼苗,前者的株高、基径、叶片数、比叶面积、相对生长率和净同化率等生长参数在轻度、中度和重度干旱处理均不同程度大于后者。大种子幼苗叶片过氧化物酶(POD)、过氧化氢酶(CAT)和超氧化物歧化酶(SOD)活性均高于小种子幼苗,前者叶片丙二醛(MDA)、可溶性蛋白、游离脯氨酸含量和叶绿素总量在部分干旱处理显著大于后者。除根冠比外,其他生长参数均随干旱胁迫增强逐渐减小,重度干旱处理大、小种子幼苗总干质量分别比对照降低19.4%和20.0%。POD、CAT和SOD活性均随干旱胁迫增强先升后降,在中度干旱处理,大、小种子幼苗POD活性分别显著高于对照126.7%和142.1%,CAT活性分别显著高于对照170.0%和151.9%。在重度干旱处理,大、小种子幼苗MDA含量分别显著高于对照86.5%和68.9%。可...