Drought- and ABA-Induced Changes in Polypeptide and mRNA Accumulation in Tomato Leaves

Drought stress triggers abscisic acid (ABA) biosynthesis resulting in ABA accumulation. The ABA-deficient tomato mutant, flacca (Lycopersicon esculentum Mill. cv Ailsa Craig), does not synthesize ABA in response to drought stress. This mutant has been used to distinguish polypeptides and in vitro translation products that are synthesized during drought stress in response to elevated ABA levels from those that are induced directly by altered water relations. A set of polypeptides and in vitro translation products was synthesized during drought stress in the wild type. These polypeptides and in vitro translation products were synthesized to a lesser extent in the drought-stressed ABA-deficient mutant. Treatment of flacca with ABA resulted in the synthesis of the drought-stress-induced polypeptides and in vitro translation products. These results support the hypothesis that many of the polypeptides that are synthesized during drought are regulated by alterations in ABA concentration. Similarly, the mRNA population was altered by ABA during drought stress.     

Hormonal changes in papaya seedlings subjected to progressive water stress and re-watering

Changes on abscisic acid (ABA), jasmonic acid (JA) and indole-3-acetic acid (IAA) levels were investigated in papaya seedlings (Carica papaya L.) cv. “Baixinho de Santa Amalia” under progressive water stress and subsequent rehydration. Also, the behaviour of leaf gas exchange and leaf growth was determined under stress condition. The results indicated that ABA and JA differ in their pattern of change under water stress. ABA continuously increased in leaves and roots during the whole period of stress whereas JA showed a sharp increase and a later decrease in both organs. Re-watering reduced rapidly (24 h) leaf and root ABA to control levels whereas the influence on JA levels could not be assessed. Drought and recovery did not alter IAA levels in leaf and root tissues of papaya seedlings. In addition, water stress reduced stomatal conductance, photosynthetic rate, transpiration rate, the percentage of attached leaves and leaf growth. Rehydration reverted in few days the effects of stress on leaf growth and gas exchange parameters. Overall, the data suggest that ABA could be involved in the induction of several progressive responses such as the induction of stomatal closure and leaf abscission to reduce papaya water loss. In addition, the pattern of accumulation of JA is compatible with a triggering signal upstream ABA. The unaltered levels of IAA could suggest a certain adaptive ability of papaya to maintain active physiological processes under progressive drought stress.     

Influence of Exogenous Glycine Betaine and Abscisic Acid on Papaya in Responses to Water-deficit Stress

The effects of exogenous foliar glycine betaine (GB) and abscisic acid (ABA) on papaya responses to water stress were investigated under distinct water regimes. Papaya seedlings (Carica papaya L. cultivar “BH-65”) were pretreated with GB or ABA and subsequently subjected to consecutive periods of drought, rehydration, and a second period of drought conditions. Results indicated that water stress induced ABA, jasmonic acid (JA), and proline accumulation but did not modify malondialdehyde (MDA) concentration. In addition, water deprivation reduced photosynthetic rate, stomatal conductance, relative water content (RWC), leaf fresh weight, and increased leaf abscission. GB applied prior to drought imposition decreased the impact of water stress on ABA, JA, proline accumulation, leaf water status, growth, and photosynthetic performance. However, ABA-pretreated plants did not show alteration of most of these parameters under water stress conditions when compared with non-pretreated plants except a clear induction of JA accumulation. Taken together, the data suggest that GB may modulate ABA, JA, and proline accumulation through the control of stomatal movement and the high availability of compatible solutes, leading to improvement of leaf water status, growth, and photosynthetic machinery function. In contrast, exogenous ABA did not stimulate papaya physiological responses under drought, but interestingly ABA in combination with drought could induce progressive JA synthesis, unlike drought alone, which induces a transitory JA increase and may trigger endogenous ABA accumulation. The data also suggest that irrespective of the pretreatments, papaya did not suffer oxidative damage.     

An abscisic acid-related reduced transpiration promotes gradual embolism repair when grapevines are rehydrated after drought

* Proposed mechanisms of embolism recovery are controversial for plants that are transpiring while undergoing cycles of dehydration and rehydration. * Here, water stress was imposed on grapevines (Vitis vinifera), and the course of embolism recovery, leaf water potential (Psi(leaf)), transpiration (E) and abscisic acid (ABA) concentration followed during the rehydration process. * As expected, Psi(leaf) and E decreased upon water stress, whereas xylem embolism and leaf ABA concentration increased. Upon rehydration, Psi(leaf) recovered in 5 h, whereas E fully recovered only after an additional 48 h. The ABA content of recovering leaves was higher than in droughted controls, both on the day of rewatering and the day after, suggesting that ABA accumulated in roots during drought was delivered to the rehydrated leaves. In recovering plants, xylem embolism in petioles, shoots, and roots decreased during the 24 h following rehydration. * A model is proposed to describe plant recovery after rehydration based on three main points: embolism repair occurs progressively in shoots and further in roots and in petioles, following an almost full recovery of Psi(leaf); hydraulic conductance recovers during diurnal transpiring hours, when formation and repair of embolisms occurs in all plant organs; an ABA residual signal in rehydrated leaves hinders stomatal opening even when water relations have recovered, suggesting that an ABA-induced transpiration control promotes gradual embolism repair in rehydrated grapevines.     

Changes in abscisic acid and its glucose ester in Phaseolus vulgaris L. during chilling and water stress

Levels of free and conjugated abscisic acid (ABA) were determined in leaves and roots of intact bean (Phaseolus vulgaris L., cv. Mondragone) seedlings under chilling (3C) and drought as well as during recovery from stress. Abscisic acid-glucose ester (ABAGE) was the only conjugate releasing free ABA after alkaline hydrolysis of the crude aqueous extracts. During the first 20–30 h chilled plants rapidly dehydrated and wilted without any change in ABA and ABAGE levels. Subsequently, leaf and root ABA levels increased and plants regained turgor. ABAGE concentration showed a slight increase in leaves but not in roots. Upon recovery from chilling a transient, but significant, rise in leaf ABA content was observed, while no appreciable change in ABAGE was found. Drought triggered ABA accumulation in leaves and roots, while a rise in ABAGE content was detected only in leaf tissues. Recovery from stress caused a drop in ABA levels without a correspondent increase in ABAGE concentration. We conclude that ABAGE is not a source of free ABA during either chilling or water stress and that only a small proportion of the ABA produced under stress is metabolised to ABAGE during recovery.Abbreviations ABA = abscisic acid - ABAGE = abscisic acid-glucose ester - DW = dry weight - FW = fresh weight - RIA = radioimmunoassay - RWC = relative water content - w = water potential - o = osmotic potential - p = turgor potential     

Drought- and ABA-induced changes in photosynthesis of barley plants

The changes caused by drought stress and abscisic acid (ABA) on photosynthesis of barley plants (Hordeum vulgare. L. cv. Alfa) have been studied. Drought stress was induced by allowing the leaves to lose 12% of their fresh weight. Cycloheximide (CHI), an inhibitor of stress-induced ABA accumulation, was used to distinguish alterations in photosynthetic reactions that are induced after drought stress in response to elevated ABA levels from those that are caused directly by altered water relations. Four hoars after imposition of drought stress or 2 h after application of ABA, Ihe bulk of the leaf's ABA content measured by enzyme-amplified ELISA, increased 14- and 16-fold, respectively. CHI fully blocked the stress-induced ABA accumulation. Gas exchange measurements and analysis of enzyme activities were used to study the reactions of photosynthesis to drought stress and ABA. Leaf dehydration or ABA treatment led to a noticeable decrease in both the initial slope of the curves representing net photosynthetic rate versus intercellular CO₂ concentration and the maximal rate of photosynthesis; dehydration of CHI-treated plants showed much slower inhibition of the latter. The calculated values of the intercellular CO₂ concentration, CO₂ compensation point and maximal carboxylating efficiency of ribulose 1,5-bisphosphate (RuBP) carboxylase support the suggestion that biochemical factors are involved in the response of photosynthesis to ABA and drought stress. RuBP carboxylase activity was almost unaffected in ABA- and CHI-treated, non-stressed plants. A drop in enzyme activity was observed after leaf dehydration of the control and ABA-treated plants. When barley plants were supplied with ABA, the activity of carbonic anhydrase (CA, EC 4.2.2.1) increased more than 2-fold. Subsequent dehydration caused an over 1.5-fold increase in CA activity of the control plants and a more than 2.5-fold increase in ABA-treated plants. Dehydration of CHI-treated plants caused no change in enzyme activity. It is suggested that increased activity of CA is a photosynthetic response to elevated ABA concentration.     

不同花生品种响应干旱胁迫后叶片内ABA与AhNCEDl的分布

以粤油7号和汕优523两个不同抗旱性品种为材料,研究响应干旱胁迫后叶片ABA（abscisicacid,脱落酸）和AhNCEDl（Arachishypogaeanine-cis-epoxycarotenoiddioxygenase）的分布以及含量变化。结果表明,两种花生品种响应干旱胁迫后叶片的维管组织中ABA分布增强且含量增加,AhNCEDl蛋白分布也增强;且在水分胁迫初期粤油7号花生AhNCEDl蛋白分布强于汕优523,其体内ABA分布水平也高于汕优523;经ABA生物合成抑制剂N印roxen处理后,两种花生叶片ABA分布减弱,但粤油7号叶片维管组织ABA分布水平仍高于汕优523。结果表明维管组织是干旱胁迫下花生叶片中ABA和AhNCEDl分布的主要区域,且粤油7号花生抗旱性强可能与其体内AhNCEDl和ABA的分布量较高有关。     

外源脱落酸对干旱胁迫下滇润楠幼苗生长及生理特性的影响

以滇润楠一年生实生苗为试验材料,研究在良好水分条件(土壤含水量为70%～75%田间持水量)、轻度干旱胁迫及重度干旱胁迫处理下(50%～55%和30%～35%田间持水量)进行外源脱落酸(ABA)喷施对其生长及生理特性的影响。结果表明: 干旱胁迫使得滇润楠幼苗叶片的相对含水量、株高和生物量显著下降,净光合速率及叶绿素荧光参数(PSⅡ最大光化学效率,F_v/F_m)有不同程度的下降,而根冠比、膜脂过氧化产物丙二醛(MDA)含量显著增加。外源ABA的喷施可提高干旱胁迫下滇润楠幼苗的适应性,尤其是重度干旱下,外源ABA显著提高了叶片相对含水量21.0%,同时增加了植株株高和生物量的累积,提高了根冠比,为良好水分条件的2.1倍;减少了干旱下膜脂过氧化产物MDA的累积,提高了抗氧化酶过氧化氢酶、超氧化物岐化酶的活性,显著增加了脯氨酸的含量,为良好水分条件的7.7倍。外源ABA的喷施显著缓解了干旱胁迫对植株光合器官的不利影响,减少干旱引起的叶片净光合速率及气孔导度的下降,并且减轻了PSⅡ受到干旱的伤害程度,重度干旱下喷施ABA的植株的F_v/F_m显著高于未喷施ABA的植株。外源ABA的喷施可以减轻干旱对滇润楠植株的伤害,提高其抗旱性。     

Abscisic acid, indole-3-acetic acid and mineral–nutrient changes induced by drought and salinity in longan (Dimocarpus longan Lour.) plants

Longan species (Dimocarpus longan Lour.) exhibit a high agronomic potential in many subtropical regions worldwide; however, little is known about its responses to abiotic stress conditions. Drought and salinity are the most environmental factors inducing negative effects on plant growth and development. In order to elucidate the responses of longan to drought and salinity, seedlings were grown under conditions of drought and salt stresses. Drought was imposed by suspending water supply leading to progressive soil dehydration, and salinity was induced using two concentrations of NaCl, 100 and 150 mM in water solution, for 64 days. Data showed that salt concentrations increased foliar abscisic acid (ABA) and only 150 mM NaCl reduced indole-3-acetic acid (IAA) and increased proline levels. NaCl treatments also increased Na⁺ and Cl^? content in plant organs proportionally to salt concentration. Drought increased leaf ABA but did not change IAA concentrations, and also increased proline synthesis. In addition, drought and salt stresses reduced the photosynthesis performance; however, only drought decreased leaf growth and relative leaf water content. Overall, data indicate that under severe salt stress, high ABA accumulation was accompanied by a reduction of IAA levels; however, drought strongly increased ABA but did not change IAA concentrations. Moreover, drought and high salinity similarly increased (or maintained) ion levels and proline synthesis. Data also suggest that ABA accumulation may mitigate the impact of salt stress through inducing stomatal closure and delaying water loss, but did not mediate the effects of long-term drought conditions probably because leaves reached a strong dehydration and the role of ABA at this stage was not effective to detain leaf injuries.     

土壤干旱条件下氮素营养对玉米内源激素含量影响

在田间持水量分别保持于35％、55％和75％±5％的土壤水分条件下,利用盆栽实验研究了土壤干旱和氮素营养对玉米内源激素和气孔导度的影响．结果表明,土壤干旱下氮素营养明显降低了玉米根系木质部汁液ABA浓度,而正常供水下施氮处理间则无显著差异(施氮处理仍较低),同时测定的叶片ABA浓度则呈相反的变化趋势,表现为干旱下施氮处理要高于不施氮处理;施氮处理木质部汁液中ZRs浓度应低于相应的不施氮处理,在调控气孔行为方面并未表现拮抗ABA作用;3种土壤水分条件下,施氮玉米叶片的气孔导度均高于不施氮处理,与木质部汁液ABA浓度呈负相关,说明施氮处理较低的根源ABA浓度是导致其气孔导度较大的主要原因．     

The relations of stomatal closure and reopening to xylem ABA concentration and leaf water potential during soil drying and rewatering

Two tropical tree species, Acacia confusa and Leucaena leucocephala, were used to study the relationships among stomatal conductance, xylem ABA concentration and leaf water potential during a soil drying and rewatering cycle. Stomatal conductance of both A. confusa and L. leucocephala steadily decreased with the decreases in soil water content and pre-dawn leaf water potential. Upon rewatering, soil water content and pre-dawn leaf water potential rapidly returned to the control levels, whereas the reopening of stomata showed an obvious lag time. The length of this lag time was highly dependent not only upon the degree of water stress but also on plant species. The more severe the water stress, the longer the lag time. When A. confusa and L. leucocephala plants were exposed to the same degree of water stress (around –2.0 MPa in pre-dawn leaf water potential), the stomata of A. confusa reopened to the control level 6 days after rewatering. However, it took L. leucocephala about 14 days to reopen fully. A very similar response of leaf photosynthesis to soil water deficit was also observed for both species. Soil drying resulted in a significant increase in leaf and xylem ABA concentrations in both species. The more severe the water stress, the higher the leaf and xylem ABA concentrations. Both leaf ABA and xylem ABA returned to the control level following relief from water deficit and preceded the full recovery of stomata, suggesting that the lag phase of stomatal reopening was not controlled by leaf and/or xylem ABA. In contrast to drying the whole root system, drying half of the root system did not change the leaf water relations, but caused a significant increase in xylem ABA concentration, which could fully explain the decrease of stomatal conductance. After rewatering, the stomatal conductance of plants in which half of the roots were dried recovered more rapidly than those of whole-root dried plants, indicating that the leaf water deficit that occurred during the drying period was related to the post-stress stomatal inhibition. These results indicated that the decrease in stomatal conductance caused by water deficit was closely related to the increase in xylem ABA, but xylem ABA could not fully explain the reopening of stomata after relief of water stress, neither did the leaf ABA. Some unknown physiological and/or morphological processes in the guard cells may be related to the recovery process.     

Activation of abscisic acid biosynthesis in the leaves of <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> in response to water deficit

It is well known that endogenous abscisic acid (ABA) levels increase rapidly in response to drought stress and that this induces stomatal closure. In Arabidopsis thaliana, ABA levels increased rapidly in the leaves and roots when intact wild-type whole plants were exposed to drought stress. However, if the leaves and roots were separated and exposed to drought independently, the ABA level increased only in the leaves. These results suggest that, under our experimental conditions, ABA is synthesized mainly in the leaves in response to drought stress and that some of the ABA accumulated in the leaves is transported to the roots. Tracer experiments using isotope-labeled ABA indicate that the movement of ABA from leaves to roots is activated by water deficit in the roots. We also demonstrate that the endogenous ABA level in the leaves increased only when the leaves themselves were exposed to drought stress, suggesting that leaves play a major role in the production of ABA in response to acute water shortage.     

Photo- and antioxidative protection,and a role for salicylic acid during drought and recovery in field-grown<Emphasis Type="Italic"> Phillyrea angustifolia</Emphasis> plants

Mechanisms of photo- and antioxidative protection, the extent of oxidative stress, and salicylic acid accumulation in leaves of Phillyrea angustifolia L. (Oleaceae) plants exposed to drought and recovery in Mediterranean field conditions were studied. The amounts of alpha-tocopherol increased up to 4-fold and those of zeaxanthin increased up to 3-fold at relative leaf water contents (RWCs) of ca. 60%, which caused up to 70% increases in the de-epoxidation state of the xanthophyll cycle (DPS). While alpha-tocopherol increased further in severe drought, zeaxanthin levels and DPS remained constant, beta-carotene decreased and malondialdehyde (MDA) levels increased at RWCs below 50%. Though this was associated with significant decreases in the maximum efficiency of photosystem II photochemistry (F(v)/ F(m)), the same leaves that suffered from drought recovered after rainfalls, and similar MDA levels and F(v)/ F(m) ratios to those observed before drought were attained. During recovery (i) the F(v)/ F(m) ratio and beta-carotene levels increased slowly, (ii) alpha-tocopherol levels decreased sharply, to increase again, and (iii) MDA levels in leaves increased to values 35% higher than those observed at maximum drought, and decreased later. Salicylic acid (SA) levels showed a strong negative correlation (r(2)=0.857) with the RWC, and increased progressively up to 5-fold, during drought. During recovery, SA levels decreased, but remained slightly higher than those observed before drought. SA levels were positively correlated with those of alpha-tocopherol during drought (r(2)=0.718), but not during recovery (r(2)=0.221). We conclude that (i) P. angustifolia plants activate several mechanisms of photo- and antioxidative protection to withstand drought stress during a Mediterranean summer, (ii) endogenous SA levels increase in leaves of drought-stressed plants, thus suggesting a role for SA in plant responses to drought, and (iii) plants suffer oxidative stress during recovery, and this stress is more severe as the previous drought is more intense.     

Effect of drought and high solar radiation on 1-aminocyclopropane-1-carboxylic acid and abscisic acid concentrations in Rosmarinus officinalis plants

The endogenous concentrations of ACC and ABA were measured, at predawn and at maximum solar radiation, during a summer drought, and recovery after autumn rainfalls, in rosemary (Rosmarinus officinalis L.), a drought-tolerant species, growing under Mediterranean field conditions. During the summer, plants were subjected to both water deficit and high solar radiation. Plants showed severe reductions in shoot water potential to -3 MPa, which were associated with drastic stomatal closure (73%), a decrease in net photosynthesis, reaching almost zero, and a severe chlorophyll loss (74%). Despite the severity of the stress, plants recovered after the autumn rainfalls. The concentration of ACC was not enhanced by drought, and at predawn these concentrations remained constant at approximately 600 pmol ACC-1 DW throughout the experiment. Thus, ethylene did not regulate the response of rosemary to drought. However, a sharp increase in ACC levels between predawn and midday was observed. This increase was positively correlated to the intensity of the incident solar radiation. ACC levels recorded in June at midday reached 16 000 pmol g DW and in October values of 1000 pmol g-1 DW were observed. In contrast, in drought-stressed plants predawn concentrations of ABA were up to 130-fold those of recovered plants, and the levels of ABA scored at midday were double of those scored at predawn. In conclusion, although drought-stressed rosemary plants showed a relatively moderate ABA accumulation (approximately 500 pmol g-1 DW#, at predawn), it seems to be an essential factor for the regulation of the plant response to stress, thereby enabling a rapid recovery after stress release, although other mechanisms can not be excluded. As drought stress did not induce ACC accumulation, it was concluded that ethylene production was not a major factor in the drought stress resistance of rosemary plants. The increased ACC and ABA concentrations at midday were correlated with day length and light intensity and not with the water status of the plant.     

Overexpression of Arabidopsis Molybdenum Cofactor Sulfurase Gene Confers Drought Tolerance in Maize (Zea mays L.)

Abscisic acid (ABA) is a key component of the signaling system that integrates plant adaptive responses to abiotic stress. Overexpression of Arabidopsis molybdenum cofactor sulfurase gene (LOS5) in maize markedly enhanced the expression of ZmAO and aldehyde oxidase (AO) activity, leading to ABA accumulation and increased drought tolerance. Transgenic maize (Zea mays L.) exhibited the expected reductions in stomatal aperture, which led to decreased water loss and maintenance of higher relative water content (RWC) and leaf water potential. Also, transgenic maize subjected to drought treatment exhibited lower leaf wilting, electrolyte leakage, malondialdehyde (MDA) and H₂O₂ content, and higher activities of antioxidative enzymes and proline content compared to wild-type (WT) maize. Moreover, overexpression of LOS5 enhanced the expression of stress-regulated genes such as Rad 17, NCED1, CAT1, and ZmP5CS1 under drought stress conditions, and increased root system development and biomass yield after re-watering. The increased drought tolerance in transgenic plants was associated with ABA accumulation via activated AO and expression of stress-related gene via ABA induction, which sequentially induced a set of favorable stress-related physiological and biochemical responses.