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.     

苗期干旱及复水条件下不同花生品种的光合特性

为探索不同花生(Arachis hypogaea)品种的旱后恢复能力, 研究花生品种耐旱性与光合特性的关系, 通过盆栽土壤水分控制实验, 测定了12个花生品种苗期对干旱胁迫与复水过程的光合响应特征, 并讨论了所测各性状参数与抗旱性强弱的关系, 包括对水分胁迫伤害的修复能力。结果表明, 根据苗期生物量抗旱系数, ‘山花11号’、‘如皋西洋生’、‘A596’、‘山花9号’、‘农大818’的抗旱性较强, 且复水后植株产生超补偿生长效应, 补偿生长能力与抗旱性呈极显著正相关。叶片净光合速率(P_n)、气孔导度(G_s)、胞间CO₂浓度(C_i)、最大光化学效率(F_v/F_m)、实际光化学效率(Φ_PSII)、光化学猝灭系数(q_P)随干旱进程逐渐降低, 复水后逐渐增加, 抗旱性强的花生品种变幅较小。干旱7天, 大多数花生品种的光合参数值未有显著性差异。干旱14天, 抗旱性越强的花生品种光合参数值越高, 不同抗旱性花生品种的光合参数值有显著差异。‘山花11号’、‘如皋西洋生’、‘A596’、‘山花9号’的P_n、G_s、Φ_PSII、F_v/F_m、q_P在复水5天时恢复至对照水平, 复水10天时超过对照, ‘79266’、‘ICG6848’、 ‘白沙1016’、‘花17’在复水10天时仍未达到对照水平, 复水过程中抗旱性强的品种的光合参数显著高于抗旱性弱的品种。相关分析表明, 干旱胁迫14天和复水5天后, 花生的P_n、Φ_PSII、F_v/F_m、q_P与品种抗旱性呈极显著正相关。因此, 可在苗期用40%土壤相对含水量胁迫14天及复水5天时花生的P_n、Φ_PSII、F_v/F_m、q_P鉴定品种的干旱伤害程度及修复能力, ‘山花11号’可作为强干旱适应性鉴定的标准品种。     

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.     

Soybean leaf growth and gas exchange response to drought under carbon dioxide enrichment

This study was conducted to determine the response in leaf growth and gas exchange of soybean (Glycine max Merr.) to the combined effects of water deficits and carbon dioxide (CO₂) enrichment. Plants grown in pots were allowed to develop initially in a glasshouse under ambient CO₂ and well-watered conditions. Four-week old plants were transferred into two different glasshouses with either ambient (360 μmol mol^-1) or elevated (700 μmol mol^-1) CO₂. Following a 2-day acclimation period, the soil of the drought-stressed pots was allowed to dry slowly over a 2-week period. The stressed pots were watered daily so that the soil dried at an equivalent rate under the two CO₂ levels. Elevated [CO₂] decreased water loss rate and increased leaf area development and photosynthetic rate under both well-watered and drought-stressed conditions. There was, however, no significant effect of [CO₂] in the response relative to soil water content of normalized leaf gas exchange and leaf area. The drought response based on soil water content for transpiration, leaf area, and photosynthesis provide an effective method for describing the responses of soybean physiological processes to the available soil water, independent of [CO₂].     

Differential responses of photosystems I and II to seasonal drought in two Ficus species

Hemiepiphytic Ficus species exhibit more conservative water use strategy and are more drought-tolerant compared with their non-hemiepiphytic congeners, but a difference in the response of photosystem I (PSI) and photosystem II (PSII) to drought stress has not been documented to date. The enhancement of non-photochemical quenching (NPQ) and cyclic electron flow (CEF) have been identified as important mechanisms that protect the photosystems under drought conditions. Using the hemiepiphytic Ficus tinctoria and the non-hemiepiphytic Ficus racemosa, we studied the water status and the electron fluxes through PSI and PSII under seasonal water stress. Our results clearly indicated that the decline in the leaf predawn water potential (ψ_pd), the maximum photosynthetic rate (A_max) and the predawn maximum quantum yield of PSII (F_v/F_m) were more pronounced in F. racemosa than in F. tinctoria at peak drought. The F_v/F_m of F. racemosa was reduced to 0.69, indicating net photoinhibition of PSII. Concomitantly, the maximal photo-oxidizable P700 (P_m) decreased significantly in F. racemosa but remained stable in F. tinctoria. The fraction of non-photochemical quenching [Y(NPQ)] and the ratio of effective quantum yield of PSI to PSII [Y(I)/Y(II)] increased for both Ficus species at peak drought, with a stronger increase in F. racemosa. These results indicated that the enhancement of NPQ and the activation of CEF contributed to the photoprotection of PSI and PSII for both Ficus species under seasonal drought, particularly for F. racemosa.     

Sap flow as an indicator of transpiration and the water status of young apricot trees

The relationship between water loss via transpiration and stem sap flow in young apricot trees was studied under different environmental conditions and different levels of soil water status. The experiment was carried out in a greenhouse over a 2-week period (November 2–14, 1997) using three-year-old apricot trees (Prunus armeniaca cv. Búlida) growing in pots. Diurnal courses of leaf water potential, leaf conductance and leaf turgor potential also were recorded throughout the experiment. Data from four days of different enviromental conditions and soil water availability have been selected for analysis. On each of the selected days the leaf water potential and the mean transpiration rates were well correlated. The slope of the linear regression of this correlation, taken to indicate the total hydraulic resistance of the tree, confirmed an increasing hydraulic resistance under drought conditions. When the trees were not drought stressed the diurnal courses of sap flow and transpiration were very similar. However, when the trees were droughted, measured of sap flow slightly underestimated actual transpiration. Our heat-pulse measurements suggest the amount of readily available water stored in the stem and leaf tissues of young apricot trees is sufficient to sustain the peak transpiration rates for about 1 hour. This revised version was published online in June 2006 with corrections to the Cover Date.     

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.     

Differences in osmotic adjustment,foliar abscisic acid dynamics,and stomatal regulation between an isohydric and anisohydric woody angiosperm during drought

Species are often classified along a continuum from isohydric to anisohydric, with isohydric species exhibiting tighter regulation of leaf water potential through stomatal closure in response to drought. We investigated plasticity in stomatal regulation in an isohydric (Eucalyptus camaldulensis) and an anisohydric (Acacia aptaneura) angiosperm species subject to repeated drying cycles. We also assessed foliar abscisic acid (ABA) content dynamics, aboveground/belowground biomass allocation and nonstructural carbohydrates. The anisohydric species exhibited large plasticity in the turgor loss point (Ψ_TLP), with plants subject to repeated drying exhibiting lower Ψ_TLP and correspondingly larger stomatal conductance at low water potential, compared to plants not previously exposed to drought. The anisohydric species exhibited a switch from ABA to water potential‐driven stomatal closure during drought, a response previously only reported for anisohydric gymnosperms. The isohydric species showed little osmotic adjustment, with no evidence of switching to water potential‐driven stomatal closure, but did exhibit increased root:shoot ratios. There were no differences in carbohydrate depletion between species. We conclude that a large range in Ψ_TLP and biphasic ABA dynamics are indicative of anisohydric species, and these traits are associated with exposure to low minimum foliar water potential, dense sapwood and large resistance to xylem embolism.     

Leaf hydraulics and drought stress: response, recovery and survivorship in four woody temperate plant species

Efficient conduction of water inside leaves is essential for leaf function, yet the hydraulic-mediated impact of drought on gas exchange remains poorly understood. Here we examine the decline and subsequent recovery of leaf water potential ( Ψ _leaf), leaf hydraulic conductance ( K _leaf), and midday transpiration ( E ) in four temperate woody species exposed to controlled drought conditions ranging from mild to lethal. During drought the vulnerability of K _leaf to declining Ψ _leaf varied greatly among the species sampled. Following drought, plants were rewatered and the rate of E and K _leaf recovery was found to be strongly dependent on the severity of the drought imposed. Gas exchange recovery was strongly correlated with the relatively slow recovery of K _leaf for three of the four species, indicating conformity to a hydraulic-stomatal limitation model of plant recovery. However, there was also a shift in the sensitivity of stomata to Ψ _leaf suggesting that the plant hormone abscisic acid may be involved in limiting the rate of stomatal reopening. The level of drought tolerance varied among the four species and was correlated with leaf hydraulic vulnerability. These results suggest that species-specific variation in hydraulic properties plays a fundamental role in steering the dynamic response of plants during recovery.     

不同西瓜基因型对干旱胁迫的生理响应及其抗旱性评价

以来源于不同地区的12个西瓜基因型为试材,采用盆栽控水的方式进行持续干旱处理,研究干旱胁迫对西瓜幼苗株高、根长、鲜质量及干物质积累等生长状况的影响,比较不同基因型材料对干旱胁迫的生理响应差异,同时依据旱害指数对其抗旱性进行直接评价,并采用隶属函数法进一步验证和综合评价.结果表明: 干旱处理下,12个西瓜基因型对干旱胁迫的耐受能力存在明显差异,各基因型开始出现旱害症状的时间和发生旱害的程度各不相同.干旱胁迫降低了西瓜幼苗的株高、地上、地下鲜质量和地上干物质积累量,普遍提高了根冠比,而对根长和地下干质量存在正向和负向两种不同的影响.干旱处理后,西瓜叶片的相对含水量和叶绿素含量均不同程度降低,丙二醛、过氧化氢和超氧阴离子含量增加,脯氨酸大量积累,而可溶性蛋白含量以及抗氧化酶活性则因基因型的不同而表现出不同的结果.分析认为,3个野生型材料M20、KY-3和Y-2为抗旱性强的西瓜种质,Y34、金美人和04-1-2为敏感种质,而其余基因型为中抗种质.     

Coupled biogeochemical and hydrological responses of streams and rivers to drought

• 1 Severe or extreme droughts occurred about 10% of the time over a 105‐year record from central New Mexico, U.S.A., based on the Palmer Drought Severity Index.
• 2 Drought lowers water tables, creating extensive areas of groundwater recharge and fragmenting reaches of streams and rivers. Deeper groundwater inputs predominate as sources of surface flows during drought. Nutrient inputs to streams and rivers reflect the biogeochemistry of regional ground waters with longer subsurface residence times.
• 3 Inputs of bioavailable dissolved organic carbon to surface waters decrease during drought, with labile carbon limitation of microbial metabolism a byproduct of drought conditions.
• 4 Decreased inputs of organic forms of carbon, nitrogen and phosphorus and a decrease in the organic : inorganic ratio of nutrient inputs favours autotrophs over heterotrophs during drought.
• 5 The fate of autotrophic production during drought will be strongly influenced by the structure of the aquatic food web within impacted sites.

     

Subsequent drought and re-watering events cause a shift in productivity and nutritive value of alfalfa and hybrid fescue plant

Global climate change leads to increasing frequency of droughts, threatening the productivity and quality of forage plants. Therefore, this glasshouse experiment was conducted to investigate the effects of reoccurring droughts on productivity and nutritive value of alfalfa and hybrid fescue plants. Starting from 63 days after sowing, seedlings of plants were exposed to double drought-recovery treatments, each separate period lasting for 1 week. At the end of each treatment, growth of above- and below-ground dry weight, lipid peroxidation, total antioxidative capacity, content of polyphenols, soluble and insoluble carbohydrates and proteins, as well as elemental composition of shoots were investigated. The findings confirmed our hypothesis, as growth of both plant species were mostly insignificantly affected by subsequent drought events. Whereas effect on leave redox homeostasis and plant nutritive value was much stronger, highly depending on both plant species and the order of stress event. We found significant drought-induced increase in the content of water-soluble carbohydrates (WSCs), starch, soluble and insoluble proteins, and polyphenols. Water shortage tended to decrease and increase the content of most macro- and microelements in alfalfa and fescue, respectively. These effects were reversed by re-watering with several exceptions, such as constant reduction of Ca content in alfalfa. To conclude, alfalfa was found to be more sensitive than fescue to the first drought, but able to recover after both stress events. The second drought-initiated prolonged accumulation of WSC implies a shift in plants metabolism towards increased resistance and herewith nutritive value with respect to soluble sugars.     

Reflectance and biochemical responses of maize plants to drought and re‐watering cycles

The ability to recover from drought stress after re‐watering is an important feature that will enable plants to cope with the predicted increase in episodic drought. The effects of pre‐drought and re‐watering conditions on leaf spectral properties and their relationships with the biochemical processes that underlie the recovery from pre‐drought conditions should be better understood. The reflectance spectra, 10 spectral reflectance indices (SRIs) and biochemical characteristics of maize (Zea mays) leaves were monitored 7, 14, 21 and 28 days after the initiation of soil drought stress during two successive cycles of drought and re‐watering periods. The leaf reflectance of the two inbred maize lines increased under the drought stress, especially in the visible spectral range. In addition, an obvious recovery of the leaf reflectance was only observed in the first re‐watering period, and its value remained higher than that of the control plants during the second recovery period. A recovery lag in the pigment contents was also observed during the second cycle. The recovery variations in the pattern and magnitude of the SRIs and the total contents of C, N and P that were measured in response to the re‐watering during both cycles were diverse and complex; both full and partial recoveries were observed. The SRIs representing different physiological attributes of plant growth, including the water index, red edge position, photochemical reflectance index and near‐infrared reflectance at 800 nm, showed strong linear relationships (P < 0.01 or 0.05) with the growth and biochemical traits across the successive drought and re‐watering cycles. The results suggest that maize plants can adjust their leaf reflectance properties and employ growth and biochemical strategies to adapt to cyclic drought stress and recover from drought stress after re‐watering.     

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.     

春小麦对骤旱的响应特征及其阈值分析

与缓慢发展的干旱过程不同,骤旱具有发生速度快,短期内可致害的特点。目前,关于作物骤旱致害的临界阈值及其调控机制尚不清楚。以春小麦为供试作物,通过桶栽试验,模拟研究骤旱过程中小麦受旱致害的过程特征及其控制因素。结果发现,发生骤旱时土壤含水量下降呈先快后慢的变化趋势,叶片水分和叶水势则呈先慢后快的指数变化趋势。叶片光合生理指标对土壤水分的下降存在明显的阈值响应,且不同生理指标的阈值并不完全相同,其中净光合速率与表征叶片光合能力的指标（最大羧化速率）对土壤有效含水量的响应阈值为0.4,气孔导度和蒸腾速率对土壤有效含水量的响应阈值分别为0.5和0.4。而小麦光合生理指标对叶片水分和叶水势的阈值响应并不明显。同时依据各生理指标相关和通径分析结果得出,骤旱发生时引起小麦叶片净光合速率快速降低的主导因子为非气孔因素,而并不是以往作物受旱研究中的气孔因素。本研究结果有望丰富干旱影响认知,并可为科学应对干旱提供依据。