干旱胁迫下紫花苜蓿根系形态变化及与水分利用的关系

采用盆栽实验方法研究了紫花苜蓿(品种:陇东和阿尔冈金)根系形态、生物量、蒸腾耗水量等对持续干旱的反应及与水分利用效率(WUE)间的关系,以期揭示紫花苜蓿对干旱胁迫的适应机制。结果表明:干旱胁迫使得紫花苜蓿根系形态特征在年季间、茬次间和品种间发生了显著变化,主要表现为主根伸长生长受到抑制、主根直径变细、侧根和根系总长度伸长生长则被促进、根系表面积和直径≥1mm的侧根数目显著增加、根系生物量下降,这是紫花苜蓿对干旱逆境的适应策略,但这种适应性存在限度。另一方面,干旱胁迫条件下紫花苜蓿草产量和蒸腾耗水量也因生长年限、茬次和品种的不同而呈现不同程度的降低。紫花苜蓿根系形态性状(总根长、根系生物量与根冠比)与植株水分利用效率间具有显著的相关性,其中根重对水分效率的影响是第一位的。WUE在根系形态与冠层水分消耗的协同变化下得到有限提高。对干旱的耐性最终表现为第2年第1年、第1茬和第2茬第3茬、陇东阿尔冈金。     

The responses of Briza media and Koeleria macrantha to drought and re-watering

1. To test whether differences in response to drought can help to explain the differing distributions of Briza media and Koeleria macrantha , changes in their leaf growth and water relations during soil drying were measured. After droughting, plants were re-watered and recovery recorded.
2. Leaf growth of the two species showed a similar sensitivity to drought with respect to duration of soil drying and soil moisture content.
3. In both species tiller base relative water content (RWC) was maintained at similar levels to controls until soil moisture content had fallen to less than 9%. This may aid survival in habitats subject to periodic droughting by preventing damage to the meristems.
4. Briza media did not respond to re-watering. However, plants of K. macrantha re-grew after periods of over 20 days of drought, when tiller base RWCs had fallen as low as 13%. This ability to resurrect may explain the occurrence of K. macrantha in xeric calcicolous grasslands subject to episodes of severe droughting, from which B. media is absent.     

水分亏缺下紫花苜蓿和高粱根系水力学导度 与水分利用效率的关系

利用聚乙二醇(PEG-6000)模拟水分亏缺条件(胁迫水势-0.2MPa,胁迫48h),研究了变水条件下紫花苜蓿(品种:阿尔冈金和陇东)和高粱(品种:抗四)根系水力学导度(Lp_r)、根系活力、根叶相对含水量、水分利用效率等参数的动态变化,以期进一步明确植物水分吸收及散失过程调控的生理生态学基础。结果表明:水分亏缺限制了紫花苜蓿和高粱根系吸水,表现在Lp_r的下降和根系活力的降低;继而调控了其地上部反应,引起气孔导度、光合速率、叶片相对含水量和蒸腾速率等的下降,但限制性的提高了其水分利用效率,尤其在胁迫初期。恢复到正常供水条件后,Lp_r、根系活性、气孔导度等水分利用参数逐渐部分或完全恢复到了胁迫前水平,但恢复程度存在种间和品种间差异,并且根系吸水能力的恢复对于是植株地上部生长状态的恢复至关重要,尤其是水分恢复初期。紫花苜蓿根系中检测到水通道蛋白(AQPs)的存在,水分亏缺对紫花苜蓿Lp_r的影响认为主要是通过影响AQPs的活性实现的。比较紫花苜蓿和高粱水分吸收与利用状况在变水条件下的动态变化,认为紫花苜蓿幼苗对干旱逆境的适应能力相对弱于高粱,品种间陇东适应能力更强。     

旱作条件下不同苜蓿品种光合作用的日变化

晴天,利用LI-6400光合仪研究了旱作条件下4年生紫花苜蓿新疆大叶、巨人201、牧歌401和路宝再生草初花期（2004年6月23日）的光合作用日变化特征.结果表明：①4个苜蓿品种Pn、Tr和Gs的日变化曲线均呈“双峰”型,12：00左右存在明显的光合＂午休＂现象,但不同品种Pn、Tr和Gs的高峰和低谷出现的时刻和高低不同.WUE日进程,除路宝呈“单峰”型外,新疆大叶、巨人201和牧歌401呈“双峰”型,4个品种在8：00左右的WUE均达到全天的最高值,14：00左右的峰值不明显.②根据Pn、Ci、Ls的变化方向,推测4个品种的光合“午休”主要受气孔因素限制.③国外苜蓿品种巨人201、牧歌401和路宝日平均Pn、Tr和WUE均优于国内地方品种新疆大叶,尤其是巨人201是一种高光合、高蒸腾、高水分利用率的品种.④相关分析结果表明,对Pn影响最显著的因子是PAR,其次是Gs、RH、Ta和VPD;对蒸腾速率影响最显著的因子为PAR,其次是RH、Ta、Gs和VPD.     

紫花苜蓿(Medicago sativa)对干旱胁迫的光合生理响应

紫花苜蓿是重要的豆科牧草,具有较强的抗旱性,然而干旱仍是制约紫花苜蓿生产的主要逆境因子。通过盆栽试验,以抗旱性强弱不同的两种紫花苜蓿为试验材料,对干旱胁迫下紫花苜蓿的光合生理进行较为系统的研究,结果表明：（1）干旱胁迫下两种紫花苜蓿叶片净光合速率（Pn）、蒸腾速率（E）、气孔导度（Gs）、叶绿素含量（Chl）都有不同幅度的下降;叶绿体超微结构遭到破坏。相对于抗旱性弱的苜蓿,抗旱性强的苜蓿随干旱胁迫程度的加深,净光合速率下降较慢,叶绿体的外形及基粒结构受到的影响较小。（2）轻度干旱胁迫下气孔限制是两种紫花苜蓿P。降低的主要因素,中度和重度干旱胁迫下非气孔限制是Pn降低的主要因素。（3）对叶绿素荧光参数的研究表明：干旱胁迫下两种紫花苜蓿PSⅡ反应中心光化学效率（F／F=）、PSⅡ潜在活性（Fv／Fo）降低。总体上抗旱性强的紫花苜蓿Fv／Fm和Fv／Fo下降幅度小,PSⅡ利用光能的能力及PSⅡ的潜在活性均较强。PsⅡ光化学淬灭系数（qP）、非光化学淬灭系数（qN）的变化表现为干旱胁迫下两种紫花苜蓿qP值降低、qN值升高,总体上抗旱性强的紫花苜蓿qP降低的幅度低且qN升高幅度大,表明抗旱性强的紫花苜蓿PSⅡ反应中心电子传递活性受到的影响小,光合机构的损伤程度低。     

紫花苜蓿(Medicago sativa)对干旱胁迫的光合生理响应

紫花苜蓿是重要的豆科牧草,具有较强的抗旱性,然而干旱仍是制约紫花苜蓿生产的主要逆境因子。通过盆栽试验,以抗旱性强弱不同的两种紫花苜蓿为试验材料,对干旱胁迫下紫花苜蓿的光合生理进行较为系统的研究,结果表明：（1）干旱胁迫下两种紫花苜蓿叶片净光合速率（Pn）、蒸腾速率(E)、气孔导度(Gs)、叶绿素含量(Chl)都有不同幅度的下降;叶绿体超微结构遭到破坏。相对于抗旱性弱的苜蓿,抗旱性强的苜蓿随干旱胁迫程度的加深,净光合速率下降较慢,叶绿体的外形及基粒结构受到的影响较小。（2）轻度干旱胁迫下气孔限制是两种紫花苜蓿Pn降低的主要因素,中度和重度干旱胁迫下非气孔限制是Pn降低的主要因素。（3）对叶绿素荧光参数的研究表明：干旱胁迫下两种紫花苜蓿PSⅡ反应中心光化学效率(Fv/Fm)、PSⅡ潜在活性(Fv/Fo)降低。总体上抗旱性强的紫花苜蓿Fv/Fm和Fv/Fo下降幅度小,PSⅡ利用光能的能力及PSⅡ的潜在活性均较强。PSⅡ光化学淬灭系数（qP）、非光化学淬灭系数（qN）的变化表现为干旱胁迫下两种紫花苜蓿qP值降低、qN值升高,总体上抗旱性强的紫花苜蓿qP降低的幅度低且qN升高幅度大,表明抗旱性强的紫花苜蓿PSⅡ反应中心电子传递活性受到的影响小,光合机构的损伤程度低。     

Different responses of two contrasting wheat genotypes to abscisic acid application

Purpose of this study was to investigate different responses of two wheat genotypes (Triticum aestivum L.) from the wet and dry climate regions to exogenous abscisic acid (ABA) application under well-watered and water-stressed conditions. Exogenous ABA was applied to the leaves by spraying and changes in dry matter accumulation and allocation, endogenous ABA content and carbon isotope ratio (δ¹³C) were monitored. The ABA application significantly decreased stem height, total biomass, total leaf area, total grain mass and leaf area/mass ratio, and significantly increased root/aboveground biomass ratio, endogenous ABA content and δ¹³C under well-watered and water-stressed conditions. Compared with the wet climate genotype, the dry climate genotype was more responsive to exogenous ABA application, resulting in lower stem height, total biomass, total leaf area, total grain mass and leaf area/mass ratio, and higher root/aboveground biomass ratio, endogenous ABA content and δ¹³C under all experimental treatments.The research was supported by the Program of “100 Distinguished Young Scientists” and “ Knowledge Innovation Engineering” of the Chinese Academy of Sciences (No. KSCX2-SW-115).     

Physiological and growth responses of two dogwoods to short-term drought stress and re-watering

Cornus florida and its cultivars have attracted many attentions by its colorful ornamental features. Suitable moisture condition is a major factor in the success of introduction. However, little is known about dogwoods drought adaptation to seasonal water deficit, and recovery potential from the following rainfall. In this paper, treatment of continuous drought lasted 19 days, followed by re-watering for 7 days was performed on 10-month-old seedlings of C. florida, comparing with native C. kousa. Meantime, well-watered seedlings of two species were regarded as controls. Soil relative water content (SRWC) in stressed seedlings of both dogwoods decreased significantly with drought stress prolonged, and recovered to the normal level after re-watering. As the response to drought stress, significant decline in internal carbon dioxide concentration (C_i), remarkable increment in intrinsic water use efficiency (WUE_i) in C. florida, significant increment in chlorophyll content in C. kousa, and notable decline in leaf relative water content (LRWC), maximum quantum efficiency of PSII photochemistry (F_v/F_m), photochemical quenching (qP), as well as significant increment in malondialdehyde (MDA) content, superoxide dismutase (SOD) activity, soluble sugar content (SSC) in both dogwoods were observed. However, most of physiological variables recovered to the level of control after re-watering. Furthermore, drought stress promoted root volume, root area, root biomass, whereas inhibited seedling height, basal diameter, aboveground biomass, resulting in increase of root/shoot ratio. Our findings indicate that, although C. florida has a weaker performance than C. kousa under drought stress, it can recover to the normal level after re-watering. These results suggest that C. florida and its cultivars possess drought adaptive potential for introducing to southern China.     

Photosynthesis, photochemistry and antioxidative defence in response to two drought severities and with re-watering in Allocasuarina luehmannii

Gas exchange, chlorophyll fluorescence and water potentials, together with ascorbate and glutathione concentrations, were studied during moderate and severe drought stress and in response to re-watering in Allocasuarina luehmannii seedlings. Moderate drought stress (MS) decreased stomatal conductance (g_s) and net CO₂ assimilation rates (A) to ∼40% and ∼60% of control values, respectively, and caused decreases in internal CO₂ concentration (C_i) and maximum light use efficiency of light-acclimated photosystem II (PSII) centres (Fv'/Fm'). Severe drought stress (SS) decreased g_s and A to ∼5% and ∼15% of the control values, respectively, and caused increases in C_i and PSII excitation pressure (1 − qP), as well as decreases in water potentials, effective quantum yield of PSII (ΦPSII), maximum efficiency of PSII (Fv/Fm) and Fv'/Fm'. Ascorbate and glutathione concentrations remained unaffected by drought treatments, but ascorbate became more oxidised under severe stress. MS seedlings recovered within 1 day (C_i, Fv'/Fm') to 1 week (A, g_s) of re-watering. In comparison, SS seedlings had longer-lasting after-stress effects, with recovery of many variables (g_s, water potentials, Fv/Fm, ΦPSII, Fv'/Fm') taking between 1 and 3 weeks from re-watering. We found no indication that interaction with antioxidants played a significant role in recovery. In conclusion, A. luehmannii seedlings appear to function normally under moderate drought, but do not seem to have particular metabolic tolerance mechanisms to endure severe drought, which may have implications for its persistence under climate change at the drier margins of its distribution.     

Sugar accumulation in roots of two grape varieties with contrasting response to water stress

Root sugar accumulation was studied in two grapevine varieties contrasting in tolerance to water stress. During a 10‐day water withholding treatment, the drought‐tolerant variety, Grenache, sustained less negative predawn and midday leaf water potentials as well as root water potential compared with the sensitive variety, Semillon. Grenache vines also maintained lower stomatal conductance and transpiration than Semillon vines throughout the drying period. In both varieties there was accumulation of sucrose in the roots and concentrations were inversely correlated to leaf and root water status. In both Grenache and Semillon, elevated root osmolality was associated with decreased soil moisture indicating that sugar accumulation may play a role in osmotic protection. Petiole xylem sap abscisic acid (ABA) concentrations increased with water deficit in both varieties and were highest for vines with the most negative root and predawn leaf water potentials. Furthermore, root sucrose concentrations were positively correlated with leaf xylem sap ABA concentrations, indicative of integration between carbohydrate metabolism and the ABA signalling system. Similar root sugar accumulation patterns between the two varieties, however, demonstrate that other factors are likely influencing the ability of the drought‐tolerant variety to remain hydrated.     

Global regulation of reactive oxygen species scavenging genes in alfalfa root and shoot under gradual drought stress and recovery

Reactive oxygen species (ROS) production and scavenging in plants under drought stress have been studied intensively in recent years. Here we report a global analysis of gene expression for the major ROS generating and scavenging proteins in alfalfa root and shoot under gradual drought stress followed by one-day recovery. Data from two alfalfa varieties, one drought tolerant and one drought sensitive, were compared and no qualitative differences in ROS gene regulation between the two were found. Conserved, tissue-specific patterns of gene expression in response to drought were observed for several ROS-scavenging gene families, including ascorbate peroxidase, monodehydroascorbate reductase, and peroxiredoxin. In addition, differential gene expression within families was observed. Genes for the ROS-generating enzyme, NADPH oxidase were generally induced under drought, while those for glycolate oxidase were repressed. Among the ROS-scavenging protein genes, Ferritin, Cu/Zn superoxide dismutase (SOD), and the majority of the glutathione peroxidase family members were induced under drought in both roots and shoots of both alfalfa varieties. In contrast, Fe-SOD, CC-type glutaredoxins, and thoiredoxins were downregulated.     

不同土壤水分条件下硅对紫花苜蓿水分利用效率及产量构成要素的影响

硅是地壳中含量仅次于氧的元素,植物不可能在无硅的环境中生长.通过盆栽试验研究了不同土壤水分条件下硅对紫花苜蓿(Medicago sativa)水分利用效率及产量构成要素的影响.结果表明,在土壤含水量为田间最大持水量的35%和80%的条件下,硅对紫花苜蓿水分利用效率和生物量没有显著影响,而在土壤含水量为田间最大持水量的50%和65%的条件下,硅显著提高了紫花苜蓿水分利用效率和生物量(p<0.05),紫花苜蓿水分利用效率的增幅分别为35%和20%,主要途径为降低叶片蒸腾速率;紫花苜蓿生物量增幅分别为41%和14%,主要通过促进分枝和株高生长,而不受单枝生物量的影响.因此硅对紫花苜蓿水分利用效率和生物量的有益作用与其生长环境中的土壤水分条件密切相关.     

干旱胁迫对不同根型苜蓿根系构型的影响

采用盆栽试验,比较播种当年及生长第2年对照、中度、重度干旱胁迫下根茎型、直根型、根蘖型苜蓿根系平面、立体几何构型和分形特征差异,以探究不同根型苜蓿对干旱的适应策略。结果表明：随干旱胁迫增加,各根型苜蓿根系总长度、总表面积、总体积和平均直径均减小;各根型苜蓿的根系拓扑指数较小,根系分支模式为叉状分支结构;各根型苜蓿的根系具有较好的分形特征,其分形维数较小,而分形丰度和平均连接长度均较大,表明其根系分支相对简单,但空间拓展能力强,有利于对营养空间的占有;各根型苜蓿的总分支较小、逐级分支率较大,有助于资源的快速获取和促进主根的向下生长。对根系构型参数进行主成分分析,根系总表面积、分形丰度、根长、根体积、分形维数和比根长6个根系构型参数能较好反映不同根型苜蓿根系构型特征。重度胁迫下,根茎型、根蘖型苜蓿的根系总长度、总表面积、总体积、比根长、比表面积、平均连接长度均大于直根型苜蓿,表明根茎型、根蘖型苜蓿适应干旱环境的能力强于直根型苜蓿。     

塔里木沙漠公路防护林3种植物光合特性对干旱胁迫的响应

利用Li-6400光合作用系统在沙漠腹地测定分析塔里木沙漠公路防护林植物梭梭(Haloxylon ammodendron)、多枝柽柳(Tamarix ramosissima)和乔木状沙拐枣(Calligonum arborescens)光合特性对干旱胁迫的响应,探讨了水分亏缺对防护林植物光合积累的影响作用。结果显示:沙拐枣的净光合速率、蒸腾速率、水分利用效率、光能利用效率及潜在光合作用能力对水分亏缺最敏感,而柽柳则最不敏感;不同处理下3种植物光合特性变化的差异表明,不同植物对水分亏缺有着不同的响应变化和适应方式;此外,干旱胁迫未提高3种植物的水分利用效率,反而降低了其光能利用效率;C3植物多枝柽柳在干旱高温条件下保持着比C4植物梭梭和沙拐枣更为稳定的光合积累和水分利用效率,说明部分C3植物具备不弱于C4荒漠植物的干旱耐受适应能力;虽然水分亏缺对3种植物的光合作用能力均存在不同程度抑制作用,但未对其光合积累造成不可逆转的影响。可见3种植物都有很强的干旱适应与耐受能力,这种能力表明沙漠公路防护林的灌溉管理还有进一步的节水空间。     

覆盖材料和沟垄比对土壤水分和紫花苜蓿干草产量的影响

为寻求半干旱黄土高原区种植紫花苜蓿的适宜覆盖材料和最佳沟垄比,采用完全随机设计布置大田试验,以传统平作为对照,研究不同垄覆盖材料(土壤结皮、生物可降解地膜和普通地膜)和不同沟垄比(沟宽:垄宽分别为60∶30、60∶45和60∶60,单位是cm)对土壤水分和紫花苜蓿干草产量等的影响。结果表明:通过对2012年和2013年紫花苜蓿生育期降雨量统计,2a平均值显示,无效降雨次数(53次)大于有效降雨次数(27次),无效降雨对总降雨量的贡献率(19%)小于有效降雨(81%)。就紫花苜蓿全生育期而言,与平作相比,SR_(30)、SR_(45)、SR_(60)、BMR_(30)、BMR_(45)、BMR_(60)、CMR_(30)、CMR_(45)和CMR_(60)(SR、BMR和CMR分别代表土垄、生物可降解膜垄和普通膜垄,下标分别表示垄宽为30、45cm和60cm)连续2a的平均根层(0—140 cm)土壤贮水量分别提高12.8、19.2、24.4、26.0、30.7、40.5、29.9、37.1 mm和47.7 mm。垄沟集雨种植第1年龄和第2年龄紫花苜蓿根层没有出现明显干层。与平作相比,SR_(30)、SR_(45)和SR_(60)的连续2a紫花苜蓿平均实际干草产量分别降低3%、8%和13%,WUE分别提高52%、58%和55%;BMR_(30)、BMR_(45)、BMR_(60)、CMR_(30)、CMR_(45)和CMR_(60)的连续2a紫花苜蓿平均实际干草产量分别提高14%、12%、7%、17%、19%和9%,WUE分别提高49%、62%、59%、51%、67%和56%。当紫花苜蓿生育期降雨量为380.7—427.6 mm和沟垄比为60 cm∶35—36 cm时,生物可降解膜垄和普通膜垄的紫花苜蓿实际干草产量达到最大值,为该地区垄沟集雨种植紫花苜蓿提供参考。     

水分胁迫下紫花苜蓿和高粱种子萌发特性及幼苗耐旱性

利用PEG溶液(水势梯度:-0.1～-0.5MPa)模拟水分胁迫,研究了紫花苜蓿(品种:阿尔冈金和陇东)和高粱(品种:抗四)种子的萌发能力的变化及对萌发环境的最低水分需求,并进行种间差异比较.结果显示:PEG水分胁迫通过限制种子有效水分的吸收而抑制了其萌发,且随着胁迫强度的增加,萌发能力减弱,主要表现在:萌发率、吸水速率、萌发活力、萌发胁迫指数等随胁迫强度的增加而下降,根芽比则随之增加.另一方面,种子群体萌动、萌发和出苗达50%概率时间随胁迫强度的增加而越发延迟,且各阶段对环境临界水势的需求不同,出苗阶段最为严格,说明种子出苗过程对环境水分胁迫最为敏感,耐旱能力最弱.相比之下,在同等胁迫条件下,高粱种子的萌发能力较苜蓿种子受到影响较小,各个阶段对环境水势的需求也相对较为宽松.因此,苜蓿苗期对干旱胁迫的忍耐能力不及高粱,且出苗过程中对环境水分条件的需求存在品种间差异.     

甘薯愈伤组织对干旱胁迫和盐胁迫的生理反应对比

研究干旱胁迫和盐胁迫对“芦选一号”。日‘薯愈伤组织可溶性蛋白、可溶性糖、脯氨酸含量、SOD活性等的影响,从而在细胞水平上探讨甘薯抵御渗透胁迫的生理机制。并分析甘薯细胞对干旱处理（PEG-6000）和盐处理（NaCl）的反应差异。结果表明,可溶性蛋白质含量在干旱胁迫下缓慢升高,在轻度和中度盐胁迫的生长前期和中期有较大幅度的上升。但后期下降,表明短时间盐胁迫下,Na^＋可能促进可溶性蛋白的合成;MDA在重度干旱胁迫下的含量显著低于重度盐胁迫,而SOD活性显著高于盐胁迫。表明在盐胁迫下细胞膜透性增加的主要原网是膜脂过氧化作用。干旱处理则是PEG-6000脱水的直接结果;重度干旱胁迫下,可溶性糖含量在短期内迅速升高,然后下降,而脯氨酸含量则在胁迫中后期迅速上升。脯氨酸可能有补偿可溶性糖含量降低的作用。     

Physiological responses to drought and shade in two contrasting Picea asperata populations

The responses of photosynthetic gas exchange, chlorophyll fluorescence, activities of antioxidant enzymes and lipid membrane peroxidation of two contrasting Picea asperata Mast. populations to 30% of full sunlight (shade) and full sunlight (sun) were investigated under well-watered and drought conditions. Two contrasting populations were from the wet and dry climate regions in China, respectively. For both populations tested, drought resulted in lower needle relative water content (RWC), CO₂ assimilation rate ( A ), stomatal conductance ( gs ) and effective PSII quantum yield ( Y ), and higher non-photochemical quenching ( qN ), superoxide dismutase (SOD), ascorbate peroxidase (APX) activities as well as malondialdehyde (MDA) levels and electrolyte leakage in sun plants, whereas these changes were not significant in shade plants. For the wet climate population, shade plants showed higher chlorophyll contents (Chl a , Chl b and Chl a + b ) than sun plants under both well-watered and drought conditions. Our study results implied that shade, applied together with drought, ameliorated the detrimental effects of drought. On the other hand, compared with the wet climate population, the dry climate population was more tolerant to drought in the sun treatment, as indicated by less decreases in A and mass-based leaf nitrogen content ( N _mass), more responsive stomata, greater capacity for non-radiative dissipation of excitation energy as heat (analysed by qN ), and higher level of antioxidant enzyme activities as well as lower MDA content and electrolyte leakage. These results demonstrated that the different physiological strategies were employed by the P. asperata populations from contrasting climate regions when the plants were exposed to drought and shade.