ABA application to maize hybrids contrasting for drought tolerance: changes in water parameters and in antioxidant enzyme activity

The objective of this study was to evaluate the effects of abscisic acid (ABA) related to the increase of water-stress tolerance in two drought contrasting maize hybrids: DKB 390 (tolerant) and BRS 1030 (sensitive). The characterization of water status (pre-dawn leaf water potential, Ψ_pd; midday leaf water potential, Ψ_md and stem water potential, Ψ_st) and antioxidant enzyme activity was conducted on greenhouse grown plants. The ABA, hydrogen peroxide (H₂O₂), and malondialdehyde (MDA) contents were also analyzed. Water deficit was imposed for 10 days at the flowering stage and a dosage of 100 μM ABA was applied to plant canopy. Measurements were taken during 10 days after the water recovery. With 5 days of stress, the tolerant hybrid showed lower MDA content, decrease in the water status, and higher activity of the enzymes superoxide dismutase, catalase, ascorbate peroxidase, as well as guaiacol, glutathione reductase, dehydroascorbate reductase, polyphenol oxidase, and l-phenylalanine ammonia-lyase, as compared to the sensitive hybrid. With 10 days of stress, DKB 390 had a decrease in the activity of enzymes whereas BRS 1030 showed a higher activity. In addition, the latter showed greater amounts of H₂O₂ and MDA. ABA application led to a higher tolerance only in DKB 390, due to the increase of water status and the enzymatic activity, mainly the catalase.     

Aerenchyma development in different root zones of maize genotypes under water limitation and different phosphorus nutrition

Root cortical aerenchyma (RCA) is suggested to reduce metabolic cost for root growth, but it might lower water uptake by plants. The objective of this work was to evaluate the effects of drought and phosphorus on the RCA development along the root axis and to elucidate its role in water stress tolerance of two maize genotypes. Plants of drought-tolerant DKB390 and drought-sensitive BRS1010 genotypes were grown in Vermiculite at field capacity of 100, 75, 50, and 25 % and supplied with 0.1, 0.4, and 0.8 mM phosphorus. Growth parameters, RCA, and plant P content were evaluated for all plants. Higher RCA development was observed in DKB390 than in BRS1010. Drought reduced the percentage of RCA in the root-hair zone of both genotypes but increased its development in the root maturation zone. Phosphorus limitation enhanced RCA development only in the DKB390. Under drought stress, DKB390 showed resilient growth whereas growth was inhibited in BRS1010. Higher root P content was related to its higher supply. Therefore, RCA formation was induced either by drought or by phosphorus limitation, while no interaction was evident. The RCA development varied along the root axis in order to balance water and phosphorus uptake and the drought response was genotype dependent.     

Morphophysiology, morphoanatomy, and grain yield under field conditions for two maize hybrids with contrasting response to drought stress

In the northern region of the state of Minas Gerais, lack of rainfall limits crop production in the field, which is possible only with irrigation. Agricultural and physiological practices have been intensively searched to overcome drought effects and consequently increase production. In this context, the objective of this study was to characterize morphophysiological and morphoanatomical changes and evaluate the attributes of grain yield under field conditions in two hybrids contrasting for drought tolerance. The experiment was carried out for 2 years (2010 and 2011) and the water deficit was imposed by stopping irrigation for 22 days at the pre-flowering stage. At the end of the stress treatment, leaf and root anatomy and morphophysiological characteristics (leaf water potential, chlorophyll content, percentage of dry leaves, leaf area, stomatal conductance, chlorophyll fluorescence, and anthesis-silking interval) were evaluated. For a better interpretation of tolerance of the hybrids in the evaluated characteristics, an index was used stress index. Hybrid DKB 390 (tolerant) surpassed hybrid BRS 1030 (sensitive) in grain yield. Furthermore, it presented lower percentage of dry leaves, higher flowering synchronization, higher stomatal conductance, and higher Fv/Fm relationship. In the root, DKB 390 showed higher amount of aerenchyma in the cortex, an increase of exodermis width, and numerous metaxylem with smaller diameter. In the leaf, it presented higher number of stomata and smaller distance between the vascular bundles in the leaf blade. The study concluded that significant morphophysiological and morphoanatomical changes, which are related to drought tolerance, occurred in DKB 390, leading to a higher yield in the field.     

Effect of drought and ABA on growth, photosynthesis and antioxidant system of Cotinus coggygria seedlings under two different light conditions

We exposed seedlings of Cotinus coggygria var. cinerea to drought and exogenous abscisic acid (ABA) under two different light conditions. Two watering regimes (well-watered and drought), two exogenous ABA applications (no ABA and with ABA) and two light regimes (full sunlight and shade) were employed. Compared with well-watered treatment, drought treatment significantly reduced the relative growth rate, relative water content (RWC), net photosynthesis rate (A) and transpiration (E), but increased chlorophyll a (chla), carbon isotope (δ¹³C), endogenous ABA, malondialdehyde (MDA) and hydrogen peroxide (H₂O₂) contents, and guaiacol peroxidase (POD) and catalase (CAT) activities. There was an apparent alleviation of drought effects by shade, as indicated by the lower relative growth rate, and chlorophyll, MDA and H₂O₂ contents, and increases in indoleacetic acid (IAA) and reduced glutathione (GSH) contents. On the other hand, the exogenous ABA application under shade induced protective effects on drought-stressed seedlings, as visible in RWC, MDA, A, stomatal conductance (g_s), E, δ¹³C, ABA and IAA values. In all, our results suggest that seedlings of C. coggygria are more sensitive to drought under full-light than under shade.     

Chemical and hydraulic signals regulate stomatal behavior and photosynthetic activity in maize during progressive drought

The objectives of this study were to investigate stomatal regulation in maize seedlings during progressive soil drying and to determine the impact of stomatal movement on photosynthetic activity. In well-watered and drought-stressed plants, leaf water potential (Ψ _leaf), relative water content (RWC), stomatal conductance (g _s), photosynthesis, chlorophyll fluorescence, leaf instantaneous water use efficiency (iWUE_leaf), and abscisic acid (ABA) and zeatin-riboside (ZR) accumulation were measured. Results showed that g _s decreased significantly with progressive drought and stomatal limitations were responsible for inhibiting photosynthesis in the initial stages of short-term drought. However, after 5 days of withholding water, non-stomatal limitations, such as damage to the PSII reaction center, became the main limiting factor. Stomatal behavior was correlated with changes in both hydraulic and chemical signals; however, changes in ABA and ZR occurred prior to any change in leaf water status. ABA in leaf and root tissue increased progressively during soil drying, and further analysis found that leaf ABA was negatively correlated with g _s (R ² = 0.907, p < 0.05). In contrast, leaf and root ZR decreased gradually. ZR in leaf tissue was positively correlated with g _s (R ² = 0.859, p < 0.05). These results indicate that ABA could induce stomatal closure, and ZR works antagonistically against ABA in stomatal behavior. In addition, the ABA/ZR ratio also had a strong correlation with g _s, suggesting that the combined chemical signal (the interaction between ABA and cytokinin) plays a role in coordinating stomatal behavior. In addition, Ψ _leaf and RWC decreased significantly after only 3 days of drought stress, also affecting stomatal behavior.     

Reciprocal grafting between clones with contrasting drought tolerance suggests a key role of abscisic acid in coffee acclimation to drought stress

The role of abscisic acid (ABA) in drought tolerance of Coffea canephora is unknown. To determine whether ABA is associated with drought tolerance and if the use of tolerant rootstocks could increase ABA and drought tolerance, we performed reciprocal grafting experiments between clones with contrasting tolerance to drought (clone 109, sensitive; and clone 120, tolerant). Plants were grown in large (120 L) pots in a greenhouse and subjected to drought stress by withholding irrigation. The non-grafted 120 plants and graft treatments with 120 as a rootstock showed a slower reduction of predawn leaf water potential (Ψ_pd) and a lower negative carbon isotopic composition ratio compared with the other grafting combinations in response to drought. The same 120 graft treatments also showed higher leaf ABA concentrations, lower levels of electrolyte leakage, and lower activities of ascorbate peroxidase and catalase under moderate (Ψ_pd?=???1.0 or ??1.5 MPa) and severe (Ψ_pd?=???3.0 MPa) drought. Root ABA concentrations were higher in plants with the 120 rootstocks regardless of watering regime. The 120 shoots could also contribute to drought tolerance because treatment with 120/109 rootstock/scion combination showed postponed dehydration, higher leaf ABA concentration, and lower leaf electrolyte leakage compared with the sensitive clone. We conclude that both the shoot and root systems of the tolerant clone can increase the concentrations of ABA in leaves in response to drought. This further suggests that ABA is associated with a delayed onset of severe water deficit and decreased oxidative damage in C. canephora.     

AM1 is a potential ABA substitute for drought tolerance as revealed by physiological and ultra-structural responses of oilseed rape

Abscisic acid (ABA) is an important signaling molecule for plants under drought tolerance. However, ABA itself has many limitations to be used in agriculture practically. Recently, AM1 (ABA-mimicking ligand) has been found to replace ABA. In this study, we have investigated AM1’s potential role for drought tolerance by growing two contrasting rapeseed (Brassica napus L.) genotypes: Qinyou 8 (drought sensitive) and Q2 (drought resistant) with exogenous ABA or AM1 application under well-watered and drought-stressed conditions. Results demonstrate that drought stress has hampered plant growth (relative height growth rate, plant biomass, leaf area), plant water status (leaf relative water content, root moisture content, leaf water potential), photosynthetic gas exchange attributes like net photosynthesis rate (Pn), stomatal conductance (Gs), intercellular CO₂ concentration (Ci), transpiration rate (E); chlorophyll fluorescence parameters like photosynthetic efficiency (Fv/Fm), effective quantum yield of PSII (Φ _PSII ), photochemical quenching coefficient (qL), electron transport rate (ETR) and chlorophyll content, especially for Qinyou 8 significantly compared to well-watered plants. Whereas increased root/shoot ratio (R/S), water use efficiency (WUE) and non-photochemical quenching (NPQ) was recorded in both genotypes under drought stress. On the other hand, exogenous ABA or AM1 treatment has regulated all the above parameters in a rational way to avoid drought stress. Chloroplast transmission electron microscope images, especially for Qinyou8, have revealed that oxidative stress induced by drought has blurred the grana thylakoids, increased the size or number of plastoglobules due to lipid peroxidation, and the presence of starch granules depict weak capacity to convert them into simple sugars for osmotic adjustment. However, intact grana thylakoid, few plastoglobules with no or very few starch granules were observed in the chloroplast from ABA- or AM1-treated plants under drought. More importantly, AM1-treated plants under drought stress have responded in an extremely similar way like ABA-treated ones. Finally, it is suggested that AM1 is a potential ABA substitute for plant drought tolerance.     

Water stress and abscisic acid treatments induce the CAM pathway in the epiphytic fern Vittaria lineata (L.) Smith

Among various epiphytic ferns found in the Brazilian Atlantic Forest, we studied Vittaria lineata (L.) Smith (Polypodiopsida, Pteridaceae). Anatomical characterization of the leaf was carried out by light microscopy, fluorescence microscopy, and scanning electron microscopy. V. lineata possesses succulent leaves with two longitudinal furrows on the abaxial surface. We observed abundant stomata inside the furrows, glandular trichomes, paraphises, and sporangia. We examined malate concentrations in leaves, relative water content (RWC), photosynthetic pigments, and chlorophyll (Chl) a fluorescence in control, water-deficient, and abscisic acid (ABA)-treated plants. Plants subjected to drought stress (DS) and treated by exogenous ABA showed significant increase in the malate concentration, demonstrating nocturnal acidification. These findings suggest that V. lineata could change its mode of carbon fixation from C₃ to the CAM pathway in response to drought. No significant changes in RWC were observed among treatments. Moreover, although plants subjected to stress treatments showed a significant decline in the contents of Chl a and b, the concentrations of carotenoids were stable. Photosynthetic parameters obtained from rapid light curves showed a significant decrease after DS and ABA treatments.     

Barley strigolactone signalling mutant hvd14.d reveals the role of strigolactones in abscisic acid-dependent response to drought

Strigolactones (SLs) are a group of plant hormones involved in many aspects of plant development and stress adaptation. Here, we investigated the drought response of a barley (Hordeum vulgare L.) mutant carrying a missense mutation in the gene encoding the SL-specific receptor HvD14. Our results clearly showed that hvd14.d mutant is hyper-sensitive to drought stress. This was illustrated by a lower leaf relative water content (RWC), impaired photosynthesis, disorganization of chloroplast structure, altered stomatal density and slower closure of stomata in response to drought in the mutant compared to the wild type parent cultivar Sebastian. Although the content of abscisic acid (ABA) and its derivatives remained unchanged in the mutant, significant differences in expression of genes related to ABA biosynthesis were observed. Moreover, hvd14.d was insensitive to ABA during seed germination. Analysis of Arabidopsis thaliana mutant atd14-1 also demonstrated that mutation in the SL receptor resulted in increased sensitivity to drought. Our results indicate that the drought-sensitive phenotype of barley SL mutant might be caused by a disturbed ABA metabolism and/or signalling pathways. These results together uncovered a link between SL signalling and ABA-dependent drought stress response in barley.     

Hydrogen peroxide is involved in abscisic acid-induced adventitious rooting in cucumber (<Emphasis Type="Italic">Cucumis sativus</Emphasis> L.) under drought stress

Abscisic acid (ABA) and hydrogen peroxide (H₂O₂) are important regulatory factors involved in plant development under adversity stress. Here, the involvement of H₂O₂ in ABA-induced adventitious root formation in cucumber (Cucumis sativus L.) under drought stress was determined. The results indicated that exogenous ABA or H₂O₂ promoted adventitious rooting under drought stress, with a maximal biological response at 0.5 μM ABA or 800 μM H₂O₂. The promotive effects of ABA-induced adventitious rooting under drought stress were suppressed by CAT or DPI, suggesting that endogenous H₂O₂ might be involved in ABA-induced adventitious rooting. ABA increased relative water content (RWC), leaf chlorophyll content, chlorophyll fluorescence parameters (Fv/Fm, ΦPS II and qP), water soluble carbohydrate (WSC) and soluble protein content, and peroxidase (POD), polyphenol oxidase (PPO) and indoleacetate oxidase (IAAO) activities, while decreasing transpiration rate. However, the effects of ABA were inhibited by H₂O₂ scavenger CAT. Therefore, H₂O₂ may be involved in ABA-induced adventitious root development under drought stress by stimulating water and chlorophyll content, chlorophyll fluorescence, carbohydrate and nitrogen content, as well as some enzyme activities.     

Interplay Between Abscisic Acid and Jasmonic Acid and its Role in Water-oxidative Stress in Wild-type, ABA-deficient, JA-deficient, and Ascorbate-deficient Arabidopsis Plants

The interplay between jasmonic acid (JA) and abscisic acid (ABA) in plant responses to water stress and in water-stress-enhanced oxidative stress was investigated in Arabidopsis thaliana plants subjected to water stress by water deprivation. For this purpose a drought assay was conducted using Arabidopsis mutants impaired in ABA (aba2), JA (aos), and ascorbate (vtc1) biosynthesis. Our results show an interaction between ABA and JA during their biosynthesis. Moreover, the coordinated action of ABA and JA protected wild-type, aba2, and aos plants from the effects of stress. However, this effect was not observed in the vtc1 mutant, which showed a distinct decrease in the F _v/F _m ratio, concomitant with a marked fall in relative water content (RWC), despite high endogenous concentrations of JA and ABA. This finding indicates the relevance of ascorbate metabolism in plant acclimation to stress. Despite the interaction between the two phytohormones, drought-associated stomatal closure is regulated mainly by ABA and weakly by JA, whereas JA plays a role in the formation of antioxidants regulating ascorbate and glutathione metabolism. A time course analysis revealed the relevance of plant age and stress duration in the responses of the mutants compared to wild-type plants. Here we discuss the relationship between ABA, JA, ascorbate, and glutathione in plants under water stress.     

Overproduction of the Membrane-bound Receptor-like Protein Kinase 1, RPK1, Enhances Abiotic Stress Tolerance in Arabidopsis

RPK1 (receptor-like protein kinase 1) localizes to the plasma membrane and functions as a regulator of abscisic acid (ABA) signaling in Arabidopsis. In our current study, we investigated the effect of RPK1 disruption and overproduction upon plant responses to drought stress. Transgenic Arabidopsis overexpressing the RPK1 protein showed increased ABA sensitivity in their root growth and stomatal closure and also displayed less transpirational water loss. In contrast, a mutant lacking RPK1 function, rpk1-1, was found to be resistant to ABA during these processes and showed increased water loss. RPK1 overproduction in these transgenic plants thus increased their tolerance to drought stress. We performed microarray analysis of RPK1 transgenic plants and observed enhanced expression of several stress-responsive genes, such as Cor15a, Cor15b, and rd29A, in addition to H₂O₂-responsive genes. Consistently, the expression levels of ABA/stress-responsive genes in rpk1-1 had decreased compared with wild type. The results suggest that the overproduction of RPK1 enhances both the ABA and drought stress signaling pathways. Furthermore, the leaves of the rpk1-1 plants exhibit higher sensitivity to oxidative stress upon ABA-pretreatment, whereas transgenic plants overproducing RPK1 manifest increased tolerance to this stress. Our current data suggest therefore that RPK1 overproduction controls reactive oxygen species homeostasis and enhances both water and oxidative stress tolerance in Arabidopsis.     

Abscisic acid cross-talking with hydrogen peroxide and osmolyte compounds may regulate the leaf rolling mechanism under drought

Leaf rolling observed in some crops such as maize, rice, wheat and sorghum is an indicator of decreased water status. Moderate leaf rolling not tightly or early increases the photosynthesis and grain yield of crop cultivars under environmental stresses. Moreover, the effects of exogenous abscisic acid (ABA) on stomatal conductance, water status and synthesis of osmotic compounds are a well-known issue in plants subjected to water deficit. However, it is not clear how the cross-talk of ABA with H₂O₂ and osmolyte compounds affects the leaf rolling mechanism. Regulation mechanism of leaf rolling by ABA has been first studied in maize seedlings under drought stress induced by polyethylene glycol 6000 (PEG 6000) in this study. ABA treatment under drought stress reduced hydrogen peroxide (H₂O₂) content and the degree of leaf rolling (%) while the treatment-induced ABA synthesis, osmolyte levels (proline, polyamine and total soluble sugars) and some antioxidant enzyme activities in comparison to the plants that were not treated with ABA. Furthermore, exogenous ABA up-regulated the expression levels of arginine decarboxylase (ADC) and pyrroline-5-carboxylate synthase (P5CS) genes and down-regulated polyamine oxidase (PAO), diamine oxidase (DAO) and proline dehydrogenase (ProDH) gene expressions. When endogenous ABA content was decreased by the treatment of fluoridone (FLU) that is an ABA inhibitor, leaf rolling degree (%), H₂O₂ content and antioxidant enzyme activities increased, but osmolyte levels, ADC and P5CS gene expressions decreased. Finally, the treatment of ABA to maize seedlings exposed to drought stress resulted in the stimulation of the antioxidant system, osmotic adjustment and reduction of leaf rolling. We concluded that ABA can be a signal compound cross-talking H₂O₂, proline and polyamines and thus involved in the leaf rolling mechanism by providing osmotic adjustment. The results of this study can be used to provide data for the molecular breeding of maize hybrids with high grain yield by means of moderately rolled leaves.     

Leaf gas exchange and multiple enzymatic and non-enzymatic antioxidant strategies related to drought tolerance in two oil palm hybrids

Key message

The drought tolerance in young oil palm plants is related to greater efficiency in preventing oxidative damage by activating enzymatic and non-enzymatic antioxidant strategies simultaneously.

Abstract

Drought is a major environmental constraint limiting growth and yield of oil palm trees. In this study, two oil palm hybrids (BRS Manicoré and BRS C 2501) were grown in large containers and subjected to a water deficit during 57 days. Leaf gas exchange analysis was combined with an in-depth assessment of the antioxidant system over the drought imposition. Under drought, leaf water potential at predawn (Ψ _pd) decreased similarly in both hybrids. In parallel, there were decreases in the net CO₂ assimilation rate (A), chlorophyll concentrations and Rubisco total activity. Overall, these decreases were more pronounced in BRS C 2501 than in BRS Manicoré. BRS C 2501 plants triggered more markedly its enzymatic antioxidant system earlier (Ψ _pd = ?2.1 MPa) than did BRS Manicoré, but these responses were accompanied by higher concentrations of H₂O₂ and malondialdehyde in BRS C 2510 than in BRS Manicoré. With the progress of drought stress (Ψ _pd = ?2.9 MPa and below), BRS Manicoré was better able to cope with oxidative stress through a more robust antioxidant system. In addition, significant decreases in drought-induced NAD⁺-malate dehydrogenase activities were only observed in stressed BRS C 2501 plants. Regardless of watering regimes, the total carotenoid, ascorbate and glutathione concentrations were higher in BRS Manicoré than in BRS C 2501. In conclusion, BRS Manicoré is better able to tolerate drought than BRS C 2501 by triggering multiple antioxidant strategies involved both in reactive oxygen species scavenging and dissipation of excess energy and/or reducing equivalents particularly under severe drought stress.

     

Time course analysis of ABA and non-ionic osmotic stress-induced changes in water status,chlorophyll fluorescence and osmotic adjustment in Arabidopsis thaliana wild-type (Columbia) and ABA-deficient mutant (aba2)

In the present study, we investigated time course changes of water status including relative water content (RWC), leaf osmotic potential (Ψ_Π), stomatal conductance (g_s), proline (Pro), chlorophyll fluorescence (F_v/F_m) and total chlorophyll content in the Arabidopsis thaliana under PEG-induced drought stress after exogenous ABA treatment. To a better explanation for the role of ABA in the water status of A. thaliana to drought stress, wild-type (Columbia) and ABA-deficient mutant (aba2) of A. thaliana were used in the present study. Moreover, three weeks old Arabidopsis seedlings were applied exogenously with 50 μM ABA and exposed to drought stress induced by 40% PEG8000 (−0.73 MPa) for 6 h, 12 h and 24 h (hours). Our findings indicate that RWC of wild-type and aba2 started to decrease in the first 12 h and 6 h of PEG-induced drought stress, respectively. However, exogenous treatment of 50 μM ABA increased their RWC under drought stress. On the other hand, while Ψ_Π of both genotypes started to decrease in the first 6 h of drought stress, these declines in Ψ_Π were prevented by ABA treatment under stress throughout the experiment; it was more pronounced in aba2 at 24 h. While the highest increase in g_s was obtained in aba2 after 24 h stress, ABA-induced highest decrease in g_s was obtained in the same genotype during 12 h, as compared to PEG-treated group alone. On the other hand, Pro content increased in all treatment groups of ABA-deficient mutant aba2 at 12 h and 24 h. However, Pro content in ABA + PEG treated aba2 plants was higher than in PEG- and ABA-treated plants alone at the end of the 24 h. Drought stress decreased F_v/F_m and total chlorophyll contents of both genotypes while 50 μM ABA alleviated these reductions during drought stress, as compared to PEG stressed plants. On the other hand, 50 μM ABA treatment alone did not create any remarkable effect on F_v/F_m and total chlorophyll contents.These findings indicate that exogenous ABA showed an alleviative effect against damage of drought stress on relative water content, osmotic potential, stomatal conductance, proline, chlorophyll fluorescence and total chlorophyll content of both genotypes during 24 h of drought stress treatment.     

Effect of drought stress,abscisic acid,and abscisic acid analogues on the efficacy of diclofop-methyl and tralkoxydim

The effects of drought stress, abscisic acid (ABA), and four ABA analogues on diclofop-methyl and tralkoxydim efficacy were investigated in oat (Avenu sativa). Drought stress conditions (6% soil moisture content) reduced the efficacy of diclofop-methyl at 350 g ha^–1, but not at 700 g ha^–1. Similarly, tralkoxydim efficacy was reduced by drought stress at 62.5 and 125 g ha^–1, but not at 250 g ha^–1. ABA (100 m), applied as a root drench 2 days before the herbicide, protected oat plants against all rates of diclofop-methyl and against low rates of tralkoxydim. Two ABA analogues protected oat plants from diclofop-methyl injury, whereas two others had no effect. Foliage applications of ABA were much less effective than root applications in protecting against herbicide injury. Protection by ABA and the two active analogues was dependent on the relative time of application with respect to the herbicides. Optimal protection by ABA and analogue I was obtained when they were applied between 2 days before and 1 day after diclofop-methyl application. Analogue IV protected plants when applied between 3 days before and 1 day after diclofop-methyl application. Partial protection against tralkoxydim activity by ABA was observed when it was applied between 1 day before and 1 day after herbicide application. Analogue I did not afford any protection against tralkoxydim, and analogue IV afforded partial protection when applied the same day or 1 day after tralkoxydim. The results indicate that protection against these postemergence herbicides, similar to that conferred by water stress, can be induced by ABA and structural analogues that apparently mimic the action of ABA.Abbreviations SMC soil moisture content - ABA abscisic acid