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PubMed2004

Carbon and water relations for developing fruits of Opuntia ficus-indica (L.) Miller, including effects of drought and gibberellic acid.

Growth, gas exchange rates, and carbohydrate content were studied for developing fruits of the cultivated cactus Opuntia ficus-indica (L.) Miller, including effects of drought and exogenous gibberellic acid (GA3). Fruit development required 110 d from the time of bud differentiation to ripening at 80 d after anthesis, when the fruit mass averaged 67 g. Stomatal conductance and net CO2 uptake rates for fruits were higher during the night; they were maximal at 7 d before anthesis and decreased as development progressed. Fruits undergoing drought, imposed by detaching terminal stems bearing fruits, were 50% smaller than the control at 80 d after anthesis and did not ripen. Fruits injected with 2 ml of 500 ppm GA3 were 30% smaller than the control at 80 d after anthesis; they contained a large proportion of aborted seeds that produced a weak sink signal for dry mass accumulation. Gas exchange was higher at 21 d after anthesis for fruits treated with GA3. Total soluble sugars represented 40% of the fruit's dry mass until 45 d after anthesis, when the sugar content rapidly increased, reaching 90% at 73 d after anthesis. Such an increase was not observed for fruits treated with GA3, and the sugar content for fruits undergoing drought remained low throughout development. Starch content increased for developing fruits of O. ficus-indica until 14 d after anthesis and, except for the fruits undergoing drought, decreased thereafter. Fruit development for O. ficus-indica is apparently regulated by water availability as well as hormonal signals originating both within and outside the fruit.

Carbon↗

PubMed2005

Effects of free proline accumulation in petunias under drought stress.

Petunias (Petunia hybrida cv. 'Mitchell') accumulate free proline (Pro) under drought-stress conditions. It is therefore believed that Pro acts as an osmoprotectant in plants subjected to drought conditions. Petunia plants were transformed by Delta(1)-pyrroline-5-carboxylate synthetase genes (AtP5CS from Arabidopsis thaliana L. or OsP5CS from Oryza sativa L.). The transgenic plants accumulated Pro and their drought tolerance was tested. The Pro content amounted to 0.57-1.01% of the total amino acids in the transgenic plants, or 1.5-2.6 times that in wild-type plants grown under normal conditions. The transgenic plant lines tolerated 14 d of drought stress, which confirms that both P5CS transgenes had full functionality. Exogenous L-Pro treatment caused the plants to accumulate Pro; plants treated with 5 mM L-Pro accumulated up to 18 times more free Pro than untreated plants. Exogenous L-Pro restricted the growth of wild-type petunias more than that of Arabidopsis plants. The capacity for free Pro accumulation might depend on the plant species. The growth of petunia plants was influenced not only by the Pro concentration in the plants, but by the ratio of the Pro content to the total amino acids, because the growth of the transgenic petunia plants appeared normal.

Arabidopsis↗

PubMed2006

The effect of transient and continuous drought on yield, photosynthesis and carbon isotope discrimination in sugar beet (Beta vulgaris L.).

Stable carbon isotope discrimination (delta13C), photosynthetic performance (A), dry matter accumulation (DW), and sucrose yield (Y(s)) of sugar beet were evaluated in a glasshouse experiment under transient (TS) and permanent (PS) water stress. A was significantly reduced under drought, to an extent depending on stress duration. The reduced A was strictly associated with a low DW and Y(s), the later being 42% lower in PS than control plants (C). Restoring water steeply increased A and the associated leaf traits (RWC, leaf water potential etc.), but the increase of Y(s) was negligible. Therefore, the negative effects of severe water stress in the early growth period, though reversible on gas-exchange and most leaf traits, can drastically reduce Y(s) of sugar beet. Furthermore, A seems not to be effective in predicting sucrose accumulation, although it was very effective in detecting the occurrence of plant water stress. The A/C(i) model was used to assess the photosynthetic adjustments to continuous or transient drought by calculating the photosynthetic parameters Vcmax and Jmax and then compared with delta13C. Mesophyll conductance (g(m)) was estimated by comparing delta13C measured on soluble sugars and gas-exchange data. This approach confirmed the expectation that g(m) was limiting A and that there was a significant drop in [CO2] from the substomatal cavities and the chloroplast stroma both in favourable and drought conditions. Therefore, the carbon concentration at the carboxylation site was overestimated by 25-35% by conventional gas-exchange measurements, and Vcmax was consistently underestimated when g(m) was not taken into account, especially under severe drought. Root delta13C was found to be strictly related to sucrose content (brix%), Y(s) and root dry weight, and this was especially clear when delta13C was measured on bulk dry matter. By contrast, leaf delta13C measured in soluble sugars (delta(s)) and bulk dry matter (delta(dm)) were found to correlate weakly to brix% and yield, and this was not surprising as the integration time-scale of leaf delta(s) and delta(dm) were found to be shorter than that of root delta13C in bulk dry matter. The effect of water stress on diffusive and biochemical limitations with different integration times ranged from 1 d (leaf delta(s)) to more than 1 month (root delta(dm)).

Beta vulgaris↗

PubMed2006

Acclimation of Rubisco specificity factor to drought in tobacco: discrepancies between in vitro and in vivo estimations.

In studies about the photosynthesis response to environmental stresses, such as drought, the Rubisco specificity factor (tau) is assumed to be constant or derived indirectly from gas exchange measurements. However, an analysis of the acclimation of tau to drought using in vitro determinations is lacking. The aim of the present work was to analyse the acclimation of tau to different drought intensities in tobacco (Nicotiana tabacum L.). Potted tobacco plants were subjected to three different water regimes (100%, 40%, and 15% of field capacity) and new leaves were allowed to develop. When acclimated leaves were fully developed, they were sampled for gas exchange and chlorophyll fluorescence measurements, as well as for the in vitro analysis of Rubisco kinetic properties. Relative water content and gas exchange decreased with increasing water shortage. The apparent Rubisco specificity factor as estimated in vivo by gas exchange decreased with water stress. However, in vitro estimates of tau were identical among treatments, as were Rubisco specific initial activity and activation state. The reasons for the observed discrepancy between in vitro and in vivo estimates are profusely discussed. It is suggested that the Rubisco specificity factor does not acclimate to water stress in the short term (weeks or months) in tobacco, and the validity of the so-called Laisk gas exchange method to estimate tau under drought is questioned.

Acclimatization↗

PubMed2006

Osmotic adjustment in chickpea (Cicer arietinum L.) results in no yield benefit under terminal drought.

Variation in osmotic adjustment (OA) among chickpea (Cicer arietinum L.) cultivars has been observed when exposed to terminal drought, but some studies suggest that this benefits yield while others suggest it does not benefit yield in water-limited environments. In the present study, parents differing in OA were crossed and a set of advanced breeding lines (ABLs) developed for yield testing. The variation in OA during podding was measured under terminal drought in the F(2), F(3), F(7), and F(8) progeny and in the parents by either rehydrating the leaves before sampling for osmotic potential (OP) or by measuring the relative water content (RWC) and OP on adjacent leaves for the calculation of the OP at full turgor. Yields were measured in the F(8) progeny under terminal drought in Australia and India. While differences in OA were measured in the chickpea lines and parents, OA varied from year to year and did not consistently benefit yield when measured in the field under terminal drought. In Australia, differences in OA were not associated with any yield benefit in any year, while in India early flowering resulted in higher yields at three of the four sites, and OA had an inconsistent effect on seed yields. A comparison of OP at full turgor measured after rehydration and from measurements of RWC and OP showed that the rehydration technique underestimated OA. The lack of contribution of OA to yield of chickpea is discussed.

Australia↗

PubMed2007

Overexpression of wheat Na+/H+ antiporter TNHX1 and H+-pyrophosphatase TVP1 improve salt- and drought-stress tolerance in Arabidopsis thaliana plants.

Transgenic Arabidopsis plants overexpressing the wheat vacuolar Na(+)/H(+) antiporter TNHX1 and H(+)-PPase TVP1 are much more resistant to high concentrations of NaCl and to water deprivation than the wild-type strains. These transgenic plants grow well in the presence of 200 mM NaCl and also under a water-deprivation regime, while wild-type plants exhibit chlorosis and growth inhibition. Leaf area decreased much more in wild-type than in transgenic plants subjected to salt or drought stress. The leaf water potential was less negative for wild-type than for transgenic plants. This could be due to an enhanced osmotic adjustment in the transgenic plants. Moreover, these transgenic plants accumulate more Na(+) and K(+) in their leaf tissue than the wild-type plants. The toxic effect of Na(+) accumulation in the cytosol is reduced by its sequestration into the vacuole. The rate of water loss under drought or salt stress was higher in wild-type than transgenic plants. Increased vacuolar solute accumulation and water retention could confer the phenotype of salt and drought tolerance of the transgenic plants. Overexpression of the isolated genes from wheat in Arabidopsis thaliana plants is worthwhile to elucidate the contribution of these proteins to the tolerance mechanism to salt and drought. Adopting a similar strategy could be one way of developing transgenic staple crops with improved tolerance to these important abiotic stresses.

Arabidopsis↗

PubMed2000

Responses of wild watermelon to drought stress: accumulation of an ArgE homologue and citrulline in leaves during water deficits.

Wild watermelon from the Botswana desert had an ability to survive under severe drought conditions by maintaining its water status (water content and water potential). In the analysis by two-dimensional electrophoresis of leaf proteins, seven spots were newly induced after watering stopped. One with the molecular mass of 40 kilodaltons of the spots was accumulated abundantly. The cDNA encoding for the protein was cloned based on its amino-terminal sequence and the amino acid sequence deduced from the determined nucleotide sequences of the cDNA exhibited homology to the enzymes belong to the ArgE/DapE/Acy1/Cpg2/YscS protein family (including acetylornithine deacetylase, carboxypeptidase and aminoacylase-1). This suggests that the protein is involved in the release of free amino acid by hydrolyzing a peptidic bond. As the drought stress progressed, citrulline became one of the major components in the total free amino acids. Eight days after withholding watering, although the lower leaves wilted significantly, the upper leaves still maintained their water status and the content of citrulline reached about 50% in the total free amino acids. The accumulation of citrulline during the drought stress in wild watermelon is an unique phenomenon in C3-plants. These results suggest that the drought tolerance of wild watermelon is related to (1) the maintenance of the water status and (2) a metabolic change to accumulate citrulline.

Amidohydrolases↗

PubMed2004

The role of aquaporin RWC3 in drought avoidance in rice.

Although the discovery of aquaporins in plants has resulted in a paradigm shift in the understanding of plant water relations, the relationship between aquaporins and drought resistance still remains elusive. From an agronomic viewpoint, upland rice is traditionally considered as showing drought avoidance. In the investigation of different morphological and physiological responses of upland rice (Oryza sativa L. spp indica cv. Zhonghan 3) and lowland rice (O. sativa L. spp japonica cv. Xiushui 63) to water deficit, we observed young leaf rolling and the remarkable decline of cumulative transpiration in the upland rice. The expression of water channel protein RWC3 mRNA was increased in upland rice at the early response (up to 4 h) to the 20% polyethylene glycol (PEG) 6000 treatment, whereas there was no significant expression changes in lowland rice. Protein levels were increased in upland rice and decreased in lowland rice at 10 h after the water deficit. The up-regulation of RWC3 in upland rice fits well with the knowledge that upland rice adopts the mechanism of drought avoidance. The physiological significance of this RWC3 up-regulation was then explored with the over-expression of RWC3 in transgenic lowland rice (O. sativa L. spp japonica cv. Zhonghua 11) controlled by a stress-inducible SWPA2 promoter. Compared to the wild-type plant, the transgenic lowland rice exhibited higher root osmotic hydraulic conductivity (Lp), leaf water potential and relative cumulative transpiration at the end of 10 h PEG treatment. These results indicated that RWC3 probably played a role in drought avoidance in rice.

Animals↗

PubMed1986

Relationships among drought resistance, transpiration rates, and abscisic acid levels in three northern conifers.

Water potentials, transpiration rates and abscisic acid (ABA) levels in shoots of black spruce (Picea mariana (Mill.) BSP), white spruce (Picea glauca (Moench) Voss) and jack pine (Pinus banksiana Lamb.) seedlings were monitored during periods of drought and recovery from drought. Abscisic acid contents of shoots increased during the period of drought as water potentials decreased. The increase in levels of ABA was closely associated with a decrease in rates of transpiration. In the spruces, the levels of ABA peaked and then fell while plant water potentials continued to decrease, whereas in jack pine, the level of ABA rose throughout the drought treatment. After rewatering, the levels of ABA in all three conifers fell concurrent with a rise in transpiration rates. At the end of the three-day recovery period, ABA levels and transpiration rates in the spruces were either at or near control levels, whereas the concentration of ABA in jack pine remained approximately twice the control level, and transpiration was only 60% of the control rate. A compound tentatively identified as phaseic acid followed trends similar to those for ABA.

Journal Article↗

PubMed1986

Free proline changes in Pinus taeda L. callus in response to drought stress.

The amino acid, proline (PRO), may have an important role in plant adaptation to drought stress. To study the effect of drought stress on PRO content of pine tissues, we measured free PRO by high pressure liquid chromatography in control ( -0.4 MPa) and drought-stressed ( -0.8, -1.0, -1.3, -1.6 MPa) callus tissue of Pinus taeda L. after eight weeks of growth. Drought stress was induced by adding polyethylene glycol (PEG) to the nutrient media. The relation between PRO accumulation and water potential was influenced by composition of the medium and temperature. Callus growing in media with water potentials of -0.8 MPa showed no increase in PRO compared to control callus in media at -0.4 MPa. When callus tissue was subjected to low water potentials (-1.6 MPa), endogenous PRO concentration increased 40-fold, while callus fresh weight decreased by more than 90%. When exogenous PRO was added to the nutrient media, endogenous PRO was directly proportional to the exogenous PRO concentration rather than reduction in callus growth. Thus low water potential in callus results in endogenous PRO accumulation and large reductions in callus fresh weight growth. Proline accumulation in pine cells appears to be related to mechanisms associated with tolerance to desiccation.

Journal Article↗

PubMed1993

Rapid response of large, drought-stressed beech trees to irrigation.

Large, declining beech (Fagus sylvatica L.) trees (diameter at breast height = 50 cm), growing on heavy clay soils in the highlands near Zurich, Switzerland, were amply irrigated in late summer. During irrigation, the xylem sap flow rate, Q(wt), was measured by the stem-tissue heat balance method with internal heating and sensing. Only a gradual and slight increase in Q(wt) in response to irrigation was observed in the control trees, whereas Q(wt) in the declining trees, whose transpiration rates were only 2-20% those of the control trees, increased 2-5 times within minutes. This suggests, that severe local drought was the major factor limiting tree growth at the site. The extent of the response permits estimation of the supply-limited (soil water) and demand-limited (tree structure) components of stress. Drought caused a decline in Q(wt) in the trees with short crowns and limited root systems that had originally been growing in dense canopies and had become suddenly exposed to full illumination as a result of a severe wind storm and thinning. Trees with deep, narrow, dense crowns, growing in more open places and adapted over a long period to high irradiance remained healthy during drought. Prolonged, periodic water shortage reduced the amount of foliage up to 90% but during drought stimulated the growth of fine roots in the surface and upper soil layers. The stem conductive systems of the declining trees were still partially functional.

Journal Article↗

PubMed1995

Gas exchange, leaf structure and nitrogen in contrasting successional tree species growing in open and understory sites during a drought.

Seasonal ecophysiology, leaf structure and nitrogen were measured in saplings of early (Populus grandidentata Michx. and Prunus serotina J.F. Ehrh.), middle (Fraxinus americana L. and Carya tomentosa Nutt.) and late (Acer rubrum L. and Cornus florida L.) successional tree species during severe drought on adjacent open and understory sites in central Pennsylvania, USA. Area-based net photosynthesis (A) and leaf conductance to water vapor diffusion (g(wv)) varied by site and species and were highest in open growing plants and early successional species at both the open and understory sites. In response to the period of maximum drought, both sunfleck and sun leaves of the early successional species exhibited smaller decreases in A than leaves of the other species. Shaded understory leaves of all species were more susceptible to drought than sun leaves and had negative midday A values during the middle and later growing season. Shaded understory leaves also displayed a reduced photosynthetic light response during the peak drought period. Sun leaves were thicker and had a greater mass per area (LMA) and nitrogen (N) content than shaded leaves, and early and middle successional species had higher N contents and concentrations than late successional species. In both sunfleck and sun leaves, seasonal A was positively related to predawn leaf Psi, g(wv), LMA and N, and was negatively related to vapor pressure deficit, midday leaf Psi and internal CO(2). Although a significant amount of plasticity occurred in all species for most gas exchange and leaf structural parameters, middle successional species exhibited the largest degree of phenotypic plasticity between open and understory plants.

Journal Article↗

PubMedDate not supplied

Non-nutritional stress acclimation of mycorrhizal woody plants exposed to drought.

Mycorrhizal enhancement of drought resistance of two woody plant species, loblolly pine (Pinus taeda L.) and rose (Rosa hybrida L. cv. Ferdy), occurred independently of phosphorus nutrition. Mycorrhizae tended to alter root morphology and carbon allocation patterns of shoots and roots. Increased drought resistance of mycorrhizal plants was in part attributed to drought-induced colonization by mycorrhizae and the ability of the mycorrhizal plants to maintain high transpiration rates as a result of greater lateral root formation and lower shoot mass (in ectomycorrhizal loblolly pine), and a higher root/shoot ratio and leaf abscission (in endomycorrhizal roses). Neither the endo- nor ectomycorrhizal symbionts affected osmotic adjustment of droughted plants.

Journal Article↗

PubMed1997

Carbon isotopic composition, gas exchange, and growth of three populations of ponderosa pine differing in drought tolerance.

Effects of water supply on gas exchange, carbon isotopic composition, and relative growth rate were compared among seedlings from three populations of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) grown in a controlled environment chamber. The three populations were chosen to represent high, moderate and low drought tolerance. There was no indication that drought tolerance was related to high water-use efficiency. Populations differed (P < 0.05) in relative growth rate (RGR), but did not differ (P > 0.10) in gas exchange variables or carbon isotope ratio (delta(13)C). Well-watered seedlings had significantly higher RGR, xylem pressure potential (Psi(xpp)), net photosynthesis (A), stomatal conductance to water vapor (g), and lower delta(13)C and instantaneous water-use efficiency than water-stressed seedlings. With decreasing Psi(xpp), A decreased linearly, whereas g decreased exponentially. Seedlings of the highly drought-tolerant population were more sensitive to water availability than seedlings from the other populations; they used water quickly when water was available, but closed their stomata in response to water stress. We conclude that, in ponderosa pine, the drought avoidance mechanism is more important for survival and growth in arid and semiarid environments than the efficient use of water.

Journal Article↗

PubMed1998

Drought tolerance, xylem sap abscisic acid and stomatal conductance during soil drying: a comparison of canopy trees of three temperate deciduous angiosperms.

Patterns of water relations, xylem sap abscisic acid concentration ([ABA]) and stomatal aperture were characterized and compared in drought-sensitive black walnut (Juglans nigra L.), less drought-sensitive sugar maple (Acer saccharum Marsh.) and drought-tolerant white oak (Quercus alba L.) trees co-occurring in a second-growth forest in Missouri, USA. There were strong correlations among reduction in predawn leaf water potential, increased xylem sap [ABA] and stomatal closure in all species. Stomatal conductance was more closely correlated with xylem sap ABA concentration than with ABA flux or xylem sap pH and cation concentrations. In isohydric black walnut, increased concentrations of ABA in the xylem sap appeared to be primarily of root origin, causing stomatal closure in response to soil drying. In anisohydric sugar maple and white oak, however, there were reductions in midday leaf water potential associated with stomatal closure, making it uncertain whether drought-induced xylem sap ABA was of leaf or root origin. The role of root-originated xylem sap ABA in these species as a signal to the shoot of the water status of the roots is, therefore, less certain.

Journal Article↗

PubMed2000

Quantifying stomatal and non-stomatal limitations to carbon assimilation resulting from leaf aging and drought in mature deciduous tree species.

Gas exchange techniques were used to investigate light-saturated carbon assimilation and its stomatal and non-stomatal limitations over two seasons in mature trees of five species in a closed deciduous forest. Stomatal and non-stomatal contributions to decreases in assimilation resulting from leaf age and drought were quantified relative to the maximum rates obtained early in the season at optimal soil water contents. Although carbon assimilation, stomatal conductance and photosynthetic capacity (V(cmax)) decreased with leaf age, decreases in V(cmax) accounted for about 75% of the leaf-age related reduction in light-saturated assimilation rates, with a secondary role for stomatal conductance (around 25%). However, when considered independently from leaf age, the drought response was dominated by stomatal limitations, accounting for about 75% of the total limitation. Some of the analytical difficulties associated with computing limitation partitioning are discussed, including path dependence, patchy stomatal closure and diffusion in the mesophyll. Although these considerations may introduce errors in our estimates, our analysis establishes some reasonable boundaries on relative limitations and shows differences between drought and non-drought years. Estimating seasonal limitations under natural conditions, as shown in this study, provides a useful basis for comparing limitation processes between years and species.

Journal Article↗

PubMed2003

Enhanced photo- and antioxidative protection, and hydrogen peroxide accumulation in drought-stressed Cistus clusii and Cistus albidus plants.

Mechanisms of drought stress resistance were studied in Cistus clusii Dunal and Cistus albidus L., two native Mediterranean shrubs that can withstand severe summer drought. While water deficit, solar radiation and temperature increased from winter to summer in the field, C. clusii and C. albidus reduced leaf area, increased root mass per leaf area, and showed diurnal changes in stomatal conductance to minimize water loss. In both species, the consequent reductions in CO2 assimilation were accompanied by reduced efficiency of photosystem II photochemistry, and protection against stress was afforded by enhanced de-epoxidation of violaxanthin in the xanthophyll cycle and increases in alpha-tocopherol and beta-carotene. In addition, hydrogen peroxide (H2O2) accumulation was observed in mesophyll cell walls of both species during the first stages of drought, although no accumulation of H2O2 was observed in chloroplasts or other organelles during the study. Despite these common responses, C. albidus and C. clusii differed in the extent of photo- and antioxidative protection. In response to drought, C. clusii showed a higher de-epoxidation state of the xanthophyll cycle and higher alpha-tocopherol and beta-carotene concentrations than C. albidus. We conclude that several structural and biochemical mechanisms underlie stress resistance in C. clusii and C. albidus, and are indicative of the different degrees of stress resistance of these shrubs.

Antioxidants↗

PubMed2004

Xylem sap flow and drought stress of Fraxinus excelsior saplings.

We report on diurnal and seasonal variations in sap flow rate and stem water potential of Fraxinus excelsior L. saplings growing at the edge of a Fraxino-Aceretum forest in western Germany. Because of shallow soil, the trees were subjected to drought in summer. When soil water availability was not limiting, sap flow rate was related to changes in solar radiation and vapor pressure deficit. Maximum transpiration rates per leaf area were 3.5-7.4 mmol m-2 s-1, and maximum daily totals were 1.7-3.3 kg m-2 day-1. Under drought conditions, stem water potential dropped to midday minima of -2.6 to -3.5 MPa and sap flow rate was strongly related to this parameter. After the drought period, reduced apparent (whole-plant) hydraulic conductance was observed, which was attributed to a continued reduction in stomatal conductance after the drought stress had ceased. A model was developed that linked sap flow rate directly to climatic variables and stem water potential. Good correlation between measured and simulated sap flow rates allowed the model to be used for data interpretation.

Circadian Rhythm↗