Foliar Abscisic Acid Concentration Research Articles

Rapid drought-recovery of gas exchange in Caragana species adapted to low mean annual precipitation.

While variation in mean annual precipitation (MAP) of the native habitat of a species has been shown to determine the ability of a species to resist a hydraulic decrease during drought, it remains unknown whether these variations in MAP also influence the ability of a species to recover and survive drought. Leaf hydraulic and gas exchange recovery following drought and the underlying mechanisms of these responses in six Caragana species from habitats along a large precipitation gradient were investigated during rehydration in a common garden. The gas exchange of species from arid habitats recovered more rapidly during rehydration after mild, moderate and severe drought stress treatments than species from humid habitats. The recovery of gas exchange was not associated with foliar abscisic acid concentration, but tightly related to the recovery of leaf hydraulic conductance (Kleaf ). The recovery of Kleaf was associated with the loss of Kleaf during dehydration under mild and moderate drought stress, and to leaf xylem embolism formation under severe drought stress. Results pointed to the different ability to recover in gas exchange in six Caragana species post-drought is associated with the MAP of the species in its native habitat.

Similar soil drying-induced stomatal closure in soybean genotypes varying in abscisic acid accumulation and stomatal sensitivity to abscisic acid.

Different soybean cultivars (Williams 82 , Union , Jindou 21 , Long Huang 1 , Long Huang 2 ) were exposed to drying soil, to investigate whether endogenous abscisic acid (ABA) concentrations and leaf water relations regulated stomatal behaviour. We measured ABA concentrations in xylem and tissue of the first and second trifoliate leaves respectively; stomatal conductance (gs ) and leaf water potential (Ψleaf ) in both leaves; and water content in soil. Cultivar variation in leaf area and g s caused different rates of soil drying, but g s and Ψ leaf declined similarly with soil drying in all cultivars. Variation in leaf xylem ABA concentration better explained stomatal responses than foliar ABA concentration in some cultivars, and was highly correlated with stomatal conductance. Xylem ABA concentration in well-watered soil was highest in Union , and in drying soil was lowest in Jindou 21 and Long Huang 2 , although the latter had the highest foliar ABA concentrations. Jindou 21 accumulated lower xylem ABA concentrations than other cultivars as soil moisture or Ψ leaf decreased, but its stomatal sensitivity to xylem ABA was greater. Because cultivars varied in both ABA accumulation and stomatal sensitivity to ABA, but had similar stomatal sensitivity to Ψ leaf , leaf water relations seem more important in regulating stomatal closure of soybean.

Species-specific variation of photosynthesis and mesophyll conductance to ozone and drought in three Mediterranean oaks.

Mesophyll conductance (g mCO2) is one of the most important components in plant photosynthesis. Tropospheric ozone (O3) and drought impair physiological processes, causing damage to photosynthetic systems. However, the combined effects of O3 and drought on g mCO2 are still largely unclear. We investigated leaf gas exchange during mid‐summer in three Mediterranean oaks exposed to O3 (ambient [35.2 nmol mol−1 as daily mean]; 1.4 × ambient) and water treatments (WW [well‐watered] and WD [water‐deficit]). We also examined if leaf traits (leaf mass per area [LMA], foliar abscisic acid concentration [ABA]) could influence the diffusion of CO2 inside a leaf. The combination of O3 and WD significantly decreased net photosynthetic rate (P N) regardless of the species. The reduction of photosynthesis was associated with a decrease in g mCO2 and stomatal conductance (g sCO2) in evergreen Quercus ilex, while the two deciduous oaks (Q. pubescens, Q. robur) also showed a reduction of the maximum rate of carboxylation (V cmax) and maximum electron transport rate (J max) with decreased diffusive conductance parameters. The reduction of g mCO2 was correlated with increased [ABA] in the three oaks, whereas there was a negative correlation between g mCO2 with LMA in Q. pubescens. Interestingly, two deciduous oaks showed a weak or no significant correlation between g sCO2 and ABA under high O3 and WD due to impaired stomatal physiological behaviour, indicating that the reduction of P N was related to g mCO2 rather than g sCO2. The results suggest that g mCO2 plays an important role in plant carbon gain under concurrent increases in the severity of drought and O3 pollution.

Alternate wetting and drying irrigation increases water and phosphorus use efficiency independent of substrate phosphorus status of vegetative rice plants

Sustainable approaches to rice cultivation that apply less irrigation and chemical fertilisers are required to increase crop resource use efficiency. Although alternate wetting and drying (AWD) has been widely promoted as a water-saving irrigation technique, its interactions with phosphorus (P) nutrition have attracted little attention. Vegetative rice plants were grown with two phosphorus levels, fertilised (HP) or un-fertilised (LP), and either continuous flooding (CF) or AWD irrigation. Treatment effects on substrate P bioavailability (measured by Diffusive Gradients in Thin films – DGT-P), plant and substrate water relations, and foliar phytohormone status, were assessed along with P partitioning in planta. Shoot biomass and leaf area under different irrigation treatments depended on substrate P status (significant P x irrigation interaction), since LP decreased these variables under CF, but had no significant effect on plants grown under AWD. AWD maintained DGT-P concentrations and increased maximal root length, but decreased root P concentrations and P offtake. Substrate drying decreased stomatal conductance (gs) and leaf water potential (Ψleaf) but re-flooding increased gs. AWD increased foliar abscisic acid (ABA), isopentenyl adenine (iP) and 1-aminocyclopropane-1-carboxylic acid (ACC) concentrations, but decreased trans-zeatin (tZ) and gibberellin A1 (GA1) concentrations. Low P increased ACC and jasmonic acid (JA) concentrations but decreased gibberellin A4 (GA4) concentrations. Across all treatments, stomatal conductance was negatively correlated with foliar ABA concentration but positively correlated with GA1 concentration. Changes in shoot phytohormone concentrations were associated with increased water and phosphorus use efficiency (WUE and PUE) of vegetative rice plants grown under AWD.

Deep root growth, ABA adjustments and root water uptake response to soil water deficit in giant reed.

Giant reed (Arundo donax L.) is a deep-rooted crop that can survive prolonged dry periods probably as a result of its capacity to uptake water from below ground, but specific information on the functioning of deep/shallow roots is missing. The objective of this study was to understand the dynamic interrelationships of root water acquisition, canopy water conservation and abscisic acid (ABA) signals from both shallow and deep roots. In transparent split top-bottom rhizotron systems (1-m-high columns), where hydraulically isolated and independently watered layers were created with the aid of calibrated soil moisture sensors, water uptake trends were monitored. Rooting patterns were traced on the walls of the rhizotrons. Leaf gas exchange was determined using a portable infrared gas analyser. Leaf and root ABA concentrations were monitored. Under well-watered conditions, water uptake from both upper and deeper soil layers was similar. Water uptake from deeper soil layers increased gradually by up to 2.2-fold when drought stress was imposed to upper layers compared to the control conditions. Despite the significant increase in water uptake from deeper layers, surface root length density of drought-treated plants remained unchanged, suggesting increased root water uptake efficiency by these roots. However, these adjustments were not sufficient to sustain photosynthesis and therefore biomass accumulation, which was reduced by 42 %. The ABA content in shallower drought-treated roots increased 2.6-fold. This increase closely and positively correlated with foliar ABA concentration, increased intrinsic water use efficiency and leaf water potential (LWP). Giant reed is able to change its water sources depending on water availability and to maximize water uptake efficiency to satisfy canopy evapotranspirative demands. The regulation of deep root functioning and distribution, adjustment of canopy size, and root/foliar synthesized ABA play a central role in controlling LWP and leaf transpiration efficiency.

Optimized potassium nutrition improves plant-water-relations of barley under PEG-induced osmotic stress

Water use efficiency (WUE) of crop plants is an important plant trait for maintaining high yield in water limited areas. By influencing osmoregulation of plants, potassium (K) plays a critical role in stress avoidance and adaptation. However, whole plant physiological mechanisms modulated by K supply in respect of plant drought tolerance and water use efficiency are not well understood. In the present study, growth, development and transpiration dynamics of two barley cultivars were evaluated with and without PEG-induced osmotic stress using an automated balance system and image based leaf area determination. Experiments were conducted to study the effects of varied K supply under different osmotic stress treatments on a wide range of morphological, biochemical and physiological characteristics of barley plants such as leaf area development, daily whole plant transpiration rate (DTR), stomatal conductance (gs), assimilation rate (AN), biomass and leaf water use efficiency (WUE) as well as foliar abscisic acid (ABA) concentrations. Two barley cultivars (cv. Sahin-91 and cv. Milford) were treated with two K supply levels (0.04 and 0.8 mM K) and osmotic stress induced by polyethylene glycol 6000 (PEG) for a period of 9 days (in total 48 days experiment) in the hydroponic plant culture (non-PEG and + 20% PEG ). Without PEG, low-K supply depressed dry matter (DM) by almost 60% averaged across both cultivars. Under osmotic stress (+PEG), total leaf area was reduced by almost 70% in low-K compared to adequate-K plants. Low K concentration under PEG stress was correlated with higher ABA concentration and was correlated with lower leaf- and whole plant transpiration rate. Biomass-WUE under low K supply decreased significantly in both barley cultivars, to a greater extent in cv. Milford under osmotic stress. However, leaf-WUE was not affected by K supply in the absence of osmotic stress. It was suggested that reduced biomass-WUE in low-K treated barley plants was not related to inefficient stomatal control under K deficiency, but instead due to reduced assimilation rate. It was further hypothesized that under low K supply, a number of energy consuming activities reduce biomass-WUE, which are not distinguished by measuring leaf-WUE. This study showed that low K supply under osmotic stress increases foliar ABA concentration thereby decreasing plant transpiration.

Long-distance ABA transport can mediate distal tissue responses by affecting local ABA concentrations.

Environmental stresses that perturb plant water relations influence abscisic acid (ABA) concentrations, but it is unclear whether long-distance ABA transport contributes to changes in local ABA levels. To determine the physiological relevance of ABA transport, we made reciprocal- and self-grafts of ABA-deficient flacca mutant and wild-type (WT) tomato plants, in which low phosphorus (P) conditions decreased ABA concentrations while salinity increased ABA concentrations. Whereas foliar ABA concentrations in the WT scions were rootstock independent under conditions, salinity resulted in long-distance transport of ABA: flacca scions had approximately twice as much ABA when grafted on WT rootstocks compared to flacca rootstocks. Root ABA concentrations were scion dependent: both WT and flacca rootstocks had less ABA with the flacca mutant scion than with the WT scion under conditions. In WT scions, whereas rootstock genotype had limited effects on stomatal conductance under conditions, a flacca rootstock decreased leaf area of stressed plants, presumably due to attenuated root-to-shoot ABA transport. In flacca scions, a WT rootstock decreased stomatal conductance but increased leaf area of stressed plants, likely due to enhanced root-to-shoot ABA transport. Thus, long-distance ABA transport can affect responses in distal tissues by changing local ABA concentrations.

Dissecting the role of isoprene and stress-related hormones (ABA and ethylene) in Populus nigra exposed to unequal root zone water stress.

Isoprene is synthesized through the 2-C-methylerythritol-5-phosphate (MEP) pathway that also produces abscisic acid (ABA). Increases in foliar free ABA concentration during drought induce stomatal closure and may also alter ethylene biosynthesis. We hypothesized a role of isoprene biosynthesis in protecting plants challenged by increasing water deficit, by influencing ABA production and ethylene evolution. We performed a split-root experiment on Populus nigra L. subjected to three water treatments: well-watered (WW) plants with both root sectors kept at pot capacity, plants with both root compartments allowed to dry for 5 days (DD) and plants with one-half of the roots irrigated to pot capacity, while the other half did not receive water (WD). WD and WW plants were similar in photosynthesis, water relations, foliar ABA concentration and isoprene emission, whereas these parameters were significantly affected in DD plants: leaf isoprene emission increased despite the fact that photosynthesis declined by 85% and the ABA-glucoside/free ABA ratio decreased significantly. Enhanced isoprene biosynthesis in water-stressed poplars may have contributed to sustaining leaf ABA biosynthesis by keeping the MEP pathway active. However, this enhancement in ABA was accompanied by no change in ethylene biosynthesis, likely confirming the antagonistic role between ABA and ethylene. These results may indicate a potential cross-talk among isoprene, ABA and ethylene under drought.

Gas exchange recovery following natural drought is rapid unless limited by loss of leaf hydraulic conductance: evidence from an evergreen woodland.

Drought can cause major damage to plant communities, but species damage thresholds and postdrought recovery of forest productivity are not yet predictable. We used an El Niño drought event as a natural experiment to test whether postdrought recovery of gas exchange could be predicted by properties of the water transport system, or if metabolism, primarily high abscisic acid concentration, might delay recovery. We monitored detailed physiological responses, including shoot sapflow, leaf gas exchange, leaf water potential and foliar abscisic acid (ABA), during drought and through the subsequent rehydration period for a sample of eight canopy and understory species. Severe drought caused major declines in leaf water potential, elevated foliar ABA concentrations and reduced stomatal conductance and assimilation rates in our eight sample species. Leaf water potential surpassed levels associated with incipient loss of leaf hydraulic conductance in four species. Following heavy rainfall gas exchange in all species, except those trees predicted to have suffered hydraulic impairment, recovered to prestressed rates within 1d. Recovery of plant gas exchange was rapid and could be predicted by the hydraulic safety margin, providing strong support for leaf vulnerability to water deficit as an index of damage under natural drought conditions.

Inhibition of tomato shoot growth by over‐irrigation is linked to nitrogen deficiency and ethylene

Although physiological effects of acute flooding have been well studied, chronic effects of suboptimal soil aeration caused by over-irrigation of containerized plants have not, despite its likely commercial significance. By automatically scheduling irrigation according to soil moisture thresholds, effects of over-irrigation on soil properties (oxygen concentration, temperature and moisture), leaf growth, gas exchange, phytohormone [abscisic acid (ABA) and ethylene] relations and nutrient status of tomato (Solanum lycopersicum Mill. cv. Ailsa Craig) were studied. Over-irrigation slowly increased soil moisture and decreased soil oxygen concentration by 4%. Soil temperature was approximately 1°C lower in the over-irrigated substrate. Over-irrigating tomato plants for 2 weeks significantly reduced shoot height (by 25%) and fresh weight and total leaf area (by 60-70%) compared with well-drained plants. Over-irrigation did not alter stomatal conductance, leaf water potential or foliar ABA concentrations, suggesting that growth inhibition was not hydraulically regulated or dependent on stomatal closure or changes in ABA. However, over-irrigation significantly increased foliar ethylene emission. Ethylene seemed to inhibit growth, as the partially ethylene-insensitive genotype Never ripe (Nr) was much less sensitive to over-irrigation than the wild type. Over-irrigation induced significant foliar nitrogen deficiency and daily supplementation of small volumes of 10 mM Ca(NO3 )2 to over-irrigated soil restored foliar nitrogen concentrations, ethylene emission and shoot fresh weight of over-irrigated plants to control levels. Thus reduced nitrogen uptake plays an important role in inhibiting growth of over-irrigated plants, in part by stimulating foliar ethylene emission.

Water deficit affects plant and soil water status, plant growth, and ginsenoside contents in American ginseng

American ginseng (Panax quinquefolius L.) produces pharmacologically active secondary compounds known as ginsenosides which have been shown to be influenced by both genetic and environmental factors. In a greenhouse experiment, effects of water deficit on ginseng plant growth, predawn leaf water potential (ΨLeaf), soil water potential (ΨSoil), leaf abscisic acid (ABA) concentration, and root ginsenoside contents as well as photosynthesis-related physiological responses were studied. Three-year-old seedlings, grown in 200 mL volume of plastic pots, were well watered for 45 days prior to the initiation of water deficit treatments. Plants in the water deficit treatments were irrigated every 10 or 20 days for the mild and severe water deficit treatments, respectively, while the control plants were watered every 4 days. The experiment was terminated after 15, 6, and 3 dry down cycles (60 days) for the control, mild, and severe water deficit treatments, respectively. As water deficit progressed, both ΨSoil and ΨLeaf decreased, but foliar ABA concentration increased. Other physiological responses to water deficit, including transpiration rate, stomatal conductance, and CO2 assimilation rate, were decreased. Water deficit decreased root growth, but unaffected shoot growth. Foliar chlorophyll content was also decreased in the water deficit treatments. The contents of individual ginsenosides Re, Rb1, Rc and Rd, and total ginsenosides were increased in the storage roots of water deficit-treated plants as compared with well-watered controls. Rootlet fresh weight before transplanting (RFWBT) as a covariate had a significant effect on the contents of ginsenoside Rb1, Rc, and Rb2. Overall, the results indicate that water deficit could contribute not only to reducing plant performance but also increasing the levels of ABA and certain ginsenoisdes.

Isoprene emission protects photosynthesis but reduces plant productivity during drought in transgenic tobacco (Nicotiana tabacum) plants

Isoprene protects the photosynthetic apparatus of isoprene-emitting plants from oxidative stress. The role of isoprene in the response of plants to drought is less clear. Water was withheld from transgenic isoprene-emitting and non-emitting tobacco (Nicotiana tabacum) plants, to examine: the response of isoprene emission to plant water deficit; a possible relationship between concentrations of the drought-induced phytohormone abscisic acid (ABA) and isoprene; and whether isoprene affected foliar reactive oxygen species (ROS) and lipid peroxidation levels. Isoprene emission did not affect whole-plant water use, foliar ABA concentration or leaf water potential under water deficit. Compared with well-watered controls, droughted non-emitting plants significantly increased ROS content (31-46%) and lipid peroxidation (30-47%), concomitant with decreased operating and maximum efficiencies of photosystem II photochemistry and lower leaf and whole-plant water use efficiency (WUE). Droughted isoprene-emitting plants showed no increase in ROS content or lipid peroxidation relative to well-watered controls, despite isoprene emission decreasing before leaf wilting. Although isoprene emission protected the photosynthetic apparatus and enhanced leaf and whole-plant WUE, non-emitting plants had 8-24% more biomass under drought, implying that isoprene emission incurred a yield penalty.

Root‐to‐shoot signalling when soil moisture is heterogeneous: increasing the proportion of root biomass in drying soil inhibits leaf growth and increases leaf abscisic acid concentration

To determine whether root-to-shoot signalling of soil moisture heterogeneity depended on root distribution, wild-type (WT) and abscisic acid (ABA)-deficient (Az34) barley (Hordeum vulgare) plants were grown in split pots into which different numbers of seminal roots were inserted. After establishment, all plants received the same irrigation volumes, with one pot watered (w) and the other allowed to dry the soil (d), imposing three treatments (1 d: 3 w, 2 d: 2 w, 3 d: 1 w) that differed in the number of seminal roots exposed to drying soil. Root distribution did not affect leaf water relations and had no sustained effect on plant evapotranspiration (ET). In both genotypes, leaf elongation was less and leaf ABA concentrations were higher in plants with more roots in drying soil, with leaf ABA concentrations and water potentials 30% and 0.2 MPa higher, respectively, in WT plants. Whole-pot soil drying increased xylem ABA concentrations, but maximum values obtained when leaf growth had virtually ceased (100 nm in Az34, 330 nm in WT) had minimal effects (<40% leaf growth inhibition) when xylem supplied to detached shoots. Although ABA may not regulate leaf growth in vivo, genetic variation in foliar ABA concentration in the field may indicate different root distributions between upper (drier) and lower (wetter) soil layers.

The Relationship between the Methyl-Erythritol Phosphate Pathway Leading to Emission of Volatile Isoprenoids and Abscisic Acid Content in Leaves

It was investigated whether the methyl-erythritol phosphate (MEP) pathway that generates volatile isoprenoids and carotenoids also produces foliar abscisic acid (ABA) and controls stomatal opening. When the MEP pathway was blocked by fosmidomycin and volatile isoprenoid emission was largely suppressed, leaf ABA content decreased to about 50% and leaf stomatal conductance increased significantly. No effect of fosmidomycin was seen in leaves with constitutively high rates of stomatal conductance and in plant species with low foliar ABA concentration. In all other cases, isoprene emission was directly associated with foliar ABA, but ABA reduction upon MEP pathway inhibition was also observed in plant species that do not emit isoprenoids. Stomatal closure causing a midday depression of photosynthesis was also associated with a concurrent increase of isoprene emission and ABA content. It is suggested that the MEP pathway generates a labile pool of ABA that responds rapidly to environmental changes. This pool also regulates stomatal conductance, possibly when coping with frequent changes of water availability. MEP pathway inhibition by leaf darkening, and its down-regulation by exposure to elevated CO2, was also associated with a reduction of foliar ABA content. However, stomatal conductance was reduced, indicating that stomatal aperture is not regulated by the MEP-dependent foliar ABA pool, under these specific cases.

Does root‐sourced ABA have a role in mediating growth and stomatal responses to soil compaction in tomato (Lycopersicon esculentum)?

Isogenic wild‐type (Ailsa Craig) and abscisic acid (ABA)‐deficient mutant (flacca) genotypes of tomato were used to examine the role of root‐sourced ABA in mediating growth and stomatal responses to compaction. Plants were grown in uniform soil columns providing low to moderate bulk densities (1.1–1.5 g cm−3), or in a split‐pot system, which allowed the roots to divide between soils of the same or differing bulk density (1.1/1.5 g cm−3). Root and shoot growth and leaf expansion were reduced when plants were grown in compacted soil (1.5 g cm−3) but leaf water status was not altered. However, stomatal conductance was affected, suggesting that non‐hydraulic signal(s) transported in the transpiration stream were responsible for the observed effects. Xylem sap and foliar ABA concentrations increased with bulk density for 10 and 15 days after emergence (DAE), respectively, but were thereafter poorly correlated with the observed growth responses. Growth was reduced to a similar extent in both genotypes in compacted soil (1.5 g cm−3), suggesting that ABA is not centrally involved in mediating growth in this severely limiting ‘critical’ compaction stress treatment. Growth performance in the 1.1/1.5 g cm−3 split‐pot treatment of Ailsa Craig was intermediate between the uniform 1.1 and 1.5 g cm−3 treatments, whereas stomatal conductance was comparable to the compacted 1.5 g cm−3 treatment. In contrast, shoot dry weight and leaf area in the split‐pot treatment of flacca were similar to the 1.5 g cm−3 treatment, but stomatal conductance was comparable to uncompacted control plants. These results suggest a role for root‐sourced ABA in regulating growth and stomatal conductance during ‘sub‐critical’ compaction stress, when genotypic differences in response are apparent. The observed genotypic differences are comparable to those previously reported for barley, but occurred at a much lower bulk density, reflecting the greater sensitivity of tomato to compaction. By alleviating the severe growth reductions induced when the entire root system encounters compacted soil, the split‐pot approach has important applications for studies of the role of root‐sourced signals in compaction‐sensitive species such as tomato.

Abscisic acid relations and the response of Populus trichocarpa stomata to leaf water potential.

The role of abscisic acid (ABA) in the mediation of stomatal responses to low leaf water potential was examined with intact plants and epidermal strips of Populus trichocarpa Torr. & A. Gray. Clones of this species grown under well-watered conditions maintain a high leaf conductance when the foliage wilts. However, foliar ABA concentration in P. trichocarpa increased manyfold in response to water stress as it did also in P. deltoides Bartr. ex Marsh. and P. trichocarpa x deltoides hybrids. Application of ABA to epidermal strips appeared to cause solute leakage, however stomata of P. trichocarpa remained partially open even when the guard cells were plasmolyzed. Foliar application of ABA induced closure of stomata in young expanding leaves, but not in fully expanded foliage. Ten days after ABA application, stomata on young leaves were open at high water potential but closed at low water potential. These characteristics are discussed with respect to wilty mutants of tomato and potato, which also have stomata unresponsive to leaf wilting.