马铃薯块茎膨大期不同程度干旱后复水的源库补偿效应

旱后复水的补偿效应在多种作物的不同生育时期都存在,是植物抵抗逆境胁迫和伤害的重要自我调节机制,也是对有限水分高效利用的体现.本研究在马铃薯块茎膨大期进行两轮干旱后复水处理,明确马铃薯补偿效应产生的干旱胁迫阈值,并从源-库角度探索马铃薯旱后复水补偿效应产生的缘由.试验选取‘大西洋’马铃薯脱毒组培苗为材料,设置充分供水(W)、轻度干旱后复水(D₁-W)、中度干旱后复水(D₂-W)和重度干旱后复水(D₃-W)4个水分处理并经过两个循环.结果表明:在经过两轮轻度干旱复水后,马铃薯产量表现出超补偿效应,水分利用效率和产量比充分供水分别提高了17.5%和6.3%;中度水分胁迫表现出近等量补偿效应,产量与充分供水差异不大,而水分利用效率提高了8.4%;而重度水分胁迫没有表现出产量补偿效应.不同程度的干旱胁迫均降低马铃薯叶片叶绿素含量、净光合速率、叶面积等源的大小和活性,而在复水后,轻度和中度胁迫出现了超补偿和补偿效应,增强了源的供应能力.同时,适度干旱后复水显著增强了块茎(库)中蔗糖-淀粉代谢途径关键酶的活性,提高了库活性,进而表现为块茎平均重量的增加.综上,马铃薯块茎膨大期适度的水分亏缺在复水后源-库均存在补偿和超补偿效应,以此来弥补干旱带来的损失,最终在产量上表现为补偿或者超补偿效应,并显著提高了水分利用效率.     

干旱和复水对不同倍性小麦光合生理生态的影响

以不同倍性的小麦为材料,研究干旱和复水对小麦光合生理生态的影响,以期为小麦的抗旱生理和育种提供理论依据.研究发现,在逐步干旱过程中不同倍性小麦的净光合速率都下降,但叶片净光合速率下降时的土壤相对含水量明显不同.复水后,不同倍性小麦净光合速率的恢复能力也有差异,豫麦49的净光合速率恢复最快,并且表现出较强的旱后超补偿效应.当土壤相对含水量减少时,栽培二粒叶片蒸腾速率的降低要明显快于净光合速率的下降,但节节麦蒸腾速率的变化没有变现出类似的适应现象.在正常供水条件下,各组WUE(water use efficiency)的大小为栽培二粒＞豫麦49＞节节麦,变水处理增加了栽培二粒和豫麦49叶片的WUE,但节节麦叶片的WUE在变水条件下却没有增加.     

极端干旱条件下杠柳的落叶相对休眠-复水复苏型生存策略

以3种梯度恒定干旱胁迫(土壤水分分别控制在最大田间持水量的80%、55%和35%)为对照,人工模拟持续自然干旱(干旱15~54 d后复水),研究黄土丘陵区广泛分布的乡土灌木杠柳(Periploca sepium)的形态、耗水、生长、生理在持续干旱、极端干旱以及旱后复水时的变化,探讨长期恒定胁迫及极端干旱下植物的适应性策略。结果表明:(1)在恒定干旱胁迫下,杠柳可调整生长和生理状况使其达到适应有限水分供应的新稳定状态;植株冠层减小,叶片可溶性糖积累以及过氧化物酶活性升高是杠柳适应长期恒定干旱胁迫的重要机制。(2)在长期自然干旱胁迫下,杠柳的形态与生理状况均持续变化。在叶片死亡脱落前,叶片中大量积累的脯氨酸、游离氨基酸和可溶性糖提高了杠柳对干旱胁迫的耐受性;随着干旱胁迫的加剧,杠柳叶片加速衰老死亡、以休眠芽形式进入相对休眠状态。(3)复水后根系和茎复苏长出新芽并补偿生长,新芽中高含量的脯氨酸和游离氨基酸以及高活性状态的SOD和POD表明新芽活跃的生理状态是其补偿生长的重要机制,即杠柳在极端干旱条件下具有落叶相对休眠-复水后复苏型生存策略。     

旱后复水对不同倍性小麦光合及抗氧化特性的影响

利用盆栽控水试验方法,在干旱和复水条件下,于拔节期对不同倍性小麦叶片的光合参数以及抗氧化指标进行测定.结果表明:(1)与干旱处理相比,拔节期恢复供水可使受旱小麦叶片的光合参数增大,叶水势、超氧化物歧化酶(SOD)活性、细胞膜透性以及质膜过氧化程度都有所降低,其中净光合速率(Pn)和蒸腾速率(Tr)的增加超过对照,叶水势和细胞膜透性的降低也超过对照,产生了生理补偿效应;(2)旱后复水条件下,叶水势很快恢复到正常水平,Pn显著提高且高于正常对照水平,其中野生一粒和小偃6号,在干旱条件下,SOD酶活性高,具有较强的活性氧清除能力,复水后,MDA的积累减少,膜透性迅速恢复,抗氧化系统更好地保护和修复了光合机制,表现出较强的光合优势.     

极端干旱条件下杠柳的落叶相对休眠-复水复苏型生存策略（英文）

以3种梯度恒定干旱胁迫(土壤水分分别控制在最大田间持水量的80%、55%和35%)为对照,人工模拟持续自然干旱(干旱15~54d后复水),研究黄土丘陵区广泛分布的乡土灌木杠柳(Periploca sepium)的形态、耗水、生长、生理在持续干旱、极端干旱以及旱后复水时的变化,探讨长期恒定胁迫及极端干旱下植物的适应性策略。结果表明:(1)在恒定干旱胁迫下,杠柳可调整生长和生理状况使其达到适应有限水分供应的新稳定状态;植株冠层减小,叶片可溶性糖积累以及过氧化物酶活性升高是杠柳适应长期恒定干旱胁迫的重要机制。(2)在长期自然干旱胁迫下,杠柳的形态与生理状况均持续变化。在叶片死亡脱落前,叶片中大量积累的脯氨酸、游离氨基酸和可溶性糖提高了杠柳对干旱胁迫的耐受性;随着干旱胁迫的加剧,杠柳叶片加速衰老死亡、以休眠芽形式进入相对休眠状态。(3)复水后根系和茎复苏长出新芽并补偿生长,新芽中高含量的脯氨酸和游离氨基酸以及高活性状态的SOD和POD表明新芽活跃的生理状态是其补偿生长的重要机制,即杠柳在极端干旱条件下具有落叶相对休眠-复水后复苏型生存策略。     

土壤大气湿度组合对玉米生长和WUE效应研究

采用人工气候生长箱盆栽试验,模拟土壤和大气湿度不同组合处理,发现土壤干旱和大气干旱显著抑制玉米生长。大气湿度提高,在一定程度能改善作物的水分状况,降低蒸腾,解除土壤干旱危害,产生生长和生理补偿效应,明显提高水分利用效率。     

盐分和干旱对沙枣幼苗生理特性的影响

以沙枣(Elaeagnus angustifolia L.)幼苗为实验材料,分别对其进行轻度干旱(土壤含水量7%—9%)、重度干旱(土壤含水量3%—5%)、100 mmol/L NaCl以及100 mmol/L NaCl处理下不同程度的盐旱共胁迫处理,处理2周后测其生理指标,包括生长指标、光合指标、渗透调节指标以及复水后生长指标,研究盐旱共胁迫对沙枣幼苗生理特性的影响。结果表明:和对照相比,轻度干旱对沙枣幼苗的生物量没有显著影响,重度干旱处理明显降低了沙枣的生物量,无论是轻度干旱还是重度干旱,都显著降低了沙枣幼苗的净光合速率、K~+含量,显著增加了Na~+含量、脯氨酸含量、可溶性糖含量、有机酸含量、总酚和类黄酮含量;和对照相比,100 mmol/L NaCl处理显著降低了沙枣幼苗的生物量、净光合速率和K~+含量,显著增加了Na~+含量、脯氨酸含量、可溶性糖含量、有机酸含量、总酚和类黄酮含量;和盐处理相比,轻度干旱和盐分共胁迫对沙枣幼苗的各项指标没有显著差异,而重度干旱和盐分共胁迫明显降低了沙枣幼苗的生物量、净光合速率;复水一周后,只有轻度干旱可以回复到对照水平。以上结果表明,盐分和干旱处理明显抑制了沙枣幼苗的生长,轻度干旱和盐分共胁迫条件下,沙枣幼苗表现出一定的交叉适应现象,而重度干旱却加重了盐害。     

谷子全生育期抗旱性鉴定及抗旱指标筛选

作物全生育期抗旱性对发掘和利用抗旱品种和抗旱基因至关重要,但谷子全生育期抗旱鉴定至今未有报道,抗旱鉴定缺乏鉴定指标。本试验在干旱池模拟干旱条件下,对苗期抗旱性表现不同的谷子品种进行了全生育期抗旱性研究,调查分析了根干重等形态和生理性状在干旱胁迫条件下的变化;采用相关和灰色关联度等方法,分析这些指标与抗旱性(DRI)的关系。结果表明,相对根冠比、相对单穗粒重和灌浆期光合速率、蒸腾速率同抗旱性表现极显著相关,可以作为谷子全生育期抗旱性鉴定的指标;而相对根干重、相对单穗重、相对株高和气孔导度则可以作为谷子全生育期抗旱性鉴定的参考指标;在供试的品种中,红根谷和大齐头白表现了良好的综合抗旱性。本文还讨论了抗旱性的复杂性以及不同性状对干旱胁迫的反应,所建立的指标对谷子全生育期抗旱鉴定具有指导意义。     

作物干旱指标对西北半干旱区春小麦缺水特征的反映

针对作物水分胁迫较为严重的西北半干旱区,应用CI301-PS光合作用仪对春小麦开花到乳熟期间的生理特征和环境因子进行了近1个月的观测, 并研究分析了3种作物干旱指标叶水势、作物水分胁迫指数以及气孔导度随时间变化和对气象因子的响应.发现干旱胁迫增加时,叶片水分减少,作物水分胁迫指数增大,叶水势降低,气孔导度有所减小.因此,气孔下腔的CO2浓度降低,作物净光合速率有所减小,不利于半干旱区小麦生物量的累积;三者相比,叶水势是反应西北半干旱区作物干旱最敏感的指标;受半干旱区逆湿现象的影响,9:00或之后一段时间观测叶水势和气孔导度对小麦等作物缺水状况反映得更客观.     

三种锦鸡儿属植物水力结构特征及其干旱适应策略

水分胁迫是干旱半干旱区限制植物生长的主要因素。以干旱半干旱区的3种锦鸡儿属植物为研究对象,从生态适应策略角度来分析3种锦鸡儿植物产生生态分离的原因。对三种锦鸡儿属植物茎干叶片的显微结构、生理功能(导水率、光合速率以及水分利用效率)进行测定,并统计了3种锦鸡儿植株的形态特征,如一、二级枝的直径、长度、末端叶面积。结果表明:三种锦鸡儿属植物都能形成较小的导管直径来适应旱生环境,但是在导水结构上又表现出一定的差异性。中间锦鸡儿的导管直径最小,次脉密度和最大净光合速率最大;柠条锦鸡儿的导管直径、叶片厚度和比叶重(LMA)最大。小叶锦鸡儿在导水率下降50%时的水势(P_(50))最大,水分胁迫时极易发生栓塞,但正是由于导管的栓塞降低了水分运输效率,使其在旱生环境中能够通过减少水分的供应来降低水分的丧失,从而保证自身生长的水分需求;而中间锦鸡儿则主要通过减小导管直径来适应旱生环境;柠条锦鸡儿的水分利用效率最高,抗栓塞能力最强,抗旱性最好,同时柠条锦鸡儿可以通过减少蒸腾面积来减少水分的丧失。植物的导管直径大小、叶片厚度、LMA、叶脉密度对植物导水速率、光合速率等生理功能都有一定的影响。     

Adjusting Nitrogen Application in Accordance with Soil Water Availability Enhances Yield and Water Use by Regulating Physiological Traits of Maize under Drip Fertigation

Knowledge of the interactive effects of water and nitrogen (N) on physio-chemical traits of maize (Zea mays L.) helps to optimize water and N management and improve productivity. A split-plot experiment was conducted with three soil water conditions (severe drought, moderate drought, and fully water supply referring to 45%–55%, 65%–75%, and 85%–95% field capacity, respectively) and four N application rates (N₀, N₁₅₀, N₂₄₀, and N₃₃₀ referring to 0, 150, 240, 330 kg N ha^–1 respectively) under drip fertigation in 2014 and 2015 in the Huang-Huai-Hai Plain of China. The results indicated that drought stress inhibited physiological activity of plants (leaf relative water content, root bleeding sap, and net photosynthetic rate), resulting in low dry matter accumulation after silking, yield, and N uptake, whereas increased WUE and NUE. N application rates over than 150 kg ha^–1 aggravated the inhibition of physiological activity under severe drought condition, while it was offset under moderate drought condition. High N application rates (N₃₃₀) still revealed negative effects under moderate drought condition, as it did not consistently enhance plant physiological activity and significantly reduced N uptake as compared to the N₂₄₀ treatment. With fully water supply, increasing N application rates synergistically enhanced physiological activity, promoted dry matter accumulation after silking, and increased yield, WUE, and N uptake. Although the N₂₄₀ treatment reduced yield by 5.4% in average, it saved 27.3% N under full water supply condition as compared with N₃₃₀ treatment. The results indicated that N regulated growth of maize in aspects of physiological traits, dry matter accumulation, and yield as well as water and N use was depended on soil water status. The appropriate N application rates for maize production was 150 kg ha^–1 under moderate drought or 240 kg ha^–1 under fully water supply under drip fertigation, and high N supply (>150 kg ha^–1) should be avoided under severe drought condition.     

水分和磷素对木荷不同种源苗木生长和磷效率的影响

以用木荷中心产区的浙江龙泉、福建建瓯、尤溪和江西吉安4个代表性优良种源为试验材料,以广西产红荷为对照,设置不同水分处理和磷素水平的盆栽试验,研究水分和磷素对木荷种源苗木生长和生理指标的影响.结果表明：在不同的水分和磷素处理下,4个参试种源的苗木生长、根系形态参数和磷素吸收效率等均存在显著的遗传差异,福建建瓯和浙江龙泉种源苗木生长量大、根系发达、磷素吸收效率高,生长表现明显优于福建尤溪和江西吉安种源,而广西产红荷则保持其原产地速生、抗旱、耐瘠的特性.土壤水分和磷素对木荷种源苗木生长影响显著.适宜水分条件下,种源苗木径生长、干物质量、根系参数和磷素吸收效率较干旱胁迫条件下高18.5%～105.6%,高磷水平下种源苗木上述性状较低磷处理高37.5%～286.2%.但在干旱和低磷胁迫下,木荷将光合产物更多地分配至地下根系,磷素利用效率也较高,这可能是木荷适应干旱和低磷胁迫的重要生理机制.相对于显著的种源、水分和磷素主效应,其间的交互作用则可以忽略.     

硅和干旱胁迫对水稻叶片光合特性和矿质养分吸收的影响

硅被认为是植物生长的有益元素,它能增强植物对非生物逆境和生物逆境胁迫的抗性。以抗旱性不同的一对水稻近等基因系w-14-和w-20为实验材料,采用盆栽实验,研究了干旱胁迫下硅处理对水稻生长性状、光合生理特性和矿质养分吸收的影响。结果表明,在正常水分条件下硅处理对水稻的生长及生理特性没有明显影响。干旱胁迫显著降低水稻植株的生长,叶绿素含量、叶绿素荧光参数Fv/Fm及Fv/F0值显著降低,光合作用受到明显抑制。加硅能提高干旱胁迫条件下水稻植株的生物量、水分利用效率、叶片叶绿素含量、净光合速率和蒸腾速率,而气孔导度和细胞间隙CO2浓度则下降。无论干旱与否,施硅后水稻的叶片硅含量均显著上升。两个水稻品系叶片的无机离子含量在干旱胁迫条件下均呈显著增加的趋势,而硅处理后材料w-14的叶片K+、Na+、Ca2+、Mg2+、Fe3+含量分别降低16.38%,24.50%,19.70%,21.52%,18.58%,w-20则分别降低11.64%,12.11%,16.06%,11.11%和19.15%,并使之回复到与对照更接近的水平。研究结果表明了硅提高水稻植株的抗旱性与光合作用的改善和矿质养分的调节有关。     

干旱胁迫对大豆苗期生长发育和生理生态特征的影响

对土壤逐步干旱下大豆幼苗形态特征和生理生态特征的变化研究表明:随着胁迫强度的增加,大豆苗期的光合速率、蒸腾速率、气孔导度降低,且相互之间呈显著正相关;随着胁迫时间的延长,叶绿素含量和叶片含水量呈低-高-低的变化趋势;在胁迫初期,对干物质分配率的影响很小,随着胁迫时间的增加,差异越来越大;随着干旱胁迫天数和强度的增加,游离脯氨酸含量呈逐渐递增的趋势,尤其是重度胁迫下积累能力更强。     

Evaluation of physiological traits for improving drought tolerance in faba bean (<Emphasis Type="Italic">Vicia faba</Emphasis> L.)

Among grain legumes, faba bean is becoming increasingly popular in European agriculture due to recent economic and environmental interests. Faba bean can be a highly productive crop, but it is sensitive to drought stress and yields can vary considerably from season to season. Understanding the physiological basis of drought tolerance would indicate traits that can be used as indirect selection criteria for the development of cultivars adapted to drought conditions. To assess genotypic variation in physiological traits associated with drought tolerance in faba bean and to determine relationships among these attributes, two pot experiments were established in a growth chamber using genetic materials that had previously been screened for drought response in the field. Nine inbred lines of diverse genetic backgrounds were tested under adequate water supply and limited water conditions. The genotypes showed substantial variation in shoot dry matter, water use, stomatal conductance, leaf temperature, transpiration efficiency, carbon isotope discrimination (Δ¹³C), relative water content (RWC) and osmotic potential, determined at pre-flowering vegetative stage. Moisture deficits decreased water usage and consequently shoot dry matter production. RWC, osmotic potential, stomatal conductance and Δ¹³C were lower, whereas leaf temperature and transpiration efficiency were higher in stressed plants, probably due to restricted transpirational cooling induced by stomatal closure. Furthermore, differences in stomatal conductance, leaf temperature, Δ¹³C and transpiration efficiency characterized genotypes that were physiologically more adapted to water deficit conditions. Correlation analysis also showed relatively strong relationships among these variables under well watered conditions. The drought tolerant genotypes, ILB-938/2 and Melodie showed lower stomatal conductance associated with warmer leaves, whereas higher stomatal conductance and cooler leaves were observed in sensitive lines (332/2/91/015/1 and Aurora/1). The lower value of Δ¹³C coupled with higher transpiration efficiency in ILB-938/2, relative to sensitive lines (Aurora/1 and Condor/3), is indeed a desirable characteristic for water-limited environments. Finally, the results showed that stomatal conductance, leaf temperature and Δ¹³C are promising physiological indicators for drought tolerance in faba bean. These variables could be measured in pot-grown plants at adequate water supply and may serve as indirect selection criteria to pre-screen genotypes.     

Growth and Photosynthetic Characteristics in Pearl Millet under Water Stress and Different Potassium Supply

Influence of supra-optimal concentrations of K on growth, water relations, and photosynthetic capacity in pearl millet under severe water deficit conditions was assessed in a glasshouse. Nineteen-days-old plants of two lines, ICMV-94133 and WCA-78, of Pennisetum glaucum (L.) R.Br. were subjected for 30 d to 235.0, 352.5, and 470.0 mg(K) kg^–1(soil) and two water regimes (100 and 30 % field capacity). Increasing K supply did not alleviate the effect of water deficit on the growth of two lines of pearl millet since additional amount of K in the growth medium had no effect on shoot dry mass, relative growth rate, plant leaf area, net assimilation rate, or leaf area ratio, although there was significant effect of drought stress on these variables. Soil moisture had a significant effect on net photosynthetic rate (P _N), transpiration rate, stomatal conductance, and water use efficiency of both pearl millet lines, but there was no significant effect of varying K supply on these variables. In WCA-78 an ameliorative effect of increasing supply of K on P _N was observed under water deficit. Chlorophyll (Chl) a and b contents increased significantly in both lines with increase in K supply under well watered conditions, but under water deficit they increased only in ICMV-94133. Chl a/b ratios were reduced significantly in WCA-78 with increasing K supply under both watering regimes, but by contrast, in ICMV-94133 this variable was decreased only under water stress. Leaf water potential and osmotic potential of both lines decreased significantly with the imposition of drought. Leaf pressure potential in both lines increased with increase in K supply under water stress. Contents of total free amino acids in the leaves of both pearl millet lines increased significantly with increase in K supply under water stress. Potassium supply had no effect on leaf soluble sugars or soluble proteins. Considerable osmotic adjustment occurred in pearl millet plants experiencing water deficit under high K supply.     

Sex-specific physiological and growth responses to water stress in <Emphasis Type="Italic">Hippophae rhamnoides</Emphasis> L. populations

Dioecious plant species and those occupying diverse habitats may present special analytical problems to determine effects of environmental stress. Here, sex-specific physiological and growth responses of two contrasting sea buckthorn (Hippophae rhamnoides L.) populations were recorded after exposure to different watering regimes. The populations used were from wet and dry climate regions in China, respectively. In the semi-controlled environmental study, the well-watered and water-deficiency plants which were watered to 100 % and 50 % field capacity were used, respectively. Sexual differences in height growth (HT), dry matter accumulation (DMA), root/shoot ratio (RS), specific leaf area (SLA), net photosynthesis (A), transpiration (E), instantaneous water use efficiency (WUE_i) and carbon isotope composition (δ¹³C) between the male and female individuals were detected under water-deficiency treatment in both populations tested. However, these sexual differences were not detected under well-watered treatment. On the other hand, compared with the wet climate population, the dry climate population showed lower HT, DMA, SLA, A and E, and higher RS under both watering regimes. The dry climate population also showed higher WUE_i and δ¹³C as affected by water deficit than the wet climate population. These morphological and physiological responses to drought showed that the different populations and the different sexual individuals may employ different survival strategies under environmental stress. The male individuals and the dry climate population would have a conservative water-use strategy in response to drought stress.     

不同水氮条件下骆驼刺幼苗生长及氮效率变化特征

在荒漠生态系统氮沉降背景下,研究退化植被幼苗对水分和氮素变化的响应特征,对实现植被恢复和重建具有重要意义。在塔里木盆地南缘对骆驼刺（Alhagi sparsifolia）幼苗通过2年的水分（干旱、中水和湿润）和氮素（不施氮、低氮（51 mg/kg）、中氮（102 mg/kg）和高氮（306 mg/kg））添加试验,研究骆驼刺幼苗干物质累积、生物固氮和氮效率对水氮条件变化的响应。结果表明,骆驼刺幼苗不同器官的干物质累积和吸氮效率对水氮条件变化的响应因生长年份而异,但幼苗整株干物质累积和吸氮效率在2个生长年份的变化趋势却相似。在干旱条件下,骆驼刺幼苗的干物质量、吸氮效率和生物固氮量均在低氮处理下显著增加,之后随施氮量增加而降低。水氮交互可显著提高幼苗干物质累积、吸氮效率和生物固氮量,其中以中水中氮处理的效果最好。水氮添加有降低骆驼刺幼苗氮素利用效率（NUE）的趋势,但在干旱和中水条件下施氮可显著提高幼苗的生物固氮比例,然而生物固氮比例与NUE仅在第2个生长年份呈显著负相关。在2个生长年份,骆驼刺幼苗干物质量与吸氮效率和生物固氮量呈极显著正相关关系,但与NUE和生物固氮比例并无明显相关性。这表明骆驼刺幼苗主要是通过调节吸氮效率和生物固氮量来适应水氮条件变化,进而影响幼苗干物质累积。     

Adaptive responses of Populus kangdingensis to drought stress

We measured dry matter accumulation and allocation, photosynthesis, lipid peroxidation, osmotic adjustment, antioxidative defences and ABA content of Populus kangdingensis C. Wang et Tung under three different watering regimes (100%, 50% and 25% of the field capacity) to characterize the morphological, physiological and biochemical basis of drought resistance in woody plants. The results showed that drought stress caused pronounced inhibition of the growth and photosynthesis rate, and that the stomatal limitation to photosynthesis was dominant. The decrease in stomatal conductance effectively controlled water loss and increased water use efficiency. Drought also affected many physiological and biochemical processes, including increases in free proline, malondialdehyde and ABA content, and superoxide dismutase activity. On the other hand, the ABA content of leaves was significantly higher than that of stem and roots under all watering regimes; the high level of ABA in the leaf may result from the large import of ABA to leaves from other organs. These results demonstrate that there are a large set of parallel changes in the morphological, physiological and biochemical responses when plants are exposed to drought stress; these changes may enhance the capability of plants to survive and grow during drought periods.