Maintenance of shoot growth by endogenous ABA: genetic assessment of the involvement of ethylene suppression

Previous work demonstrated that normal levels of endogenous abscisic acid (ABA) are required to maintain shoot growth in well-watered tomato plants independently of effects of hormone status on plant water balance. The results suggested that the impairment of shoot growth in ABA-deficient mutants is at least partly attributable to increased ethylene production. To assess the extent to which ABA maintains shoot growth by ethylene suppression, the growth of ABA-deficient (aba2-1) and ethylene-insensitive (etr1-1) single- and double-mutants of Arabidopsis was examined. To ensure that the results were independent of effects of hormone status on plant water balance, differential relative humidity regimes were used to achieve similar leaf water potentials in all genotypes and treatments. In aba2-1, shoot growth was substantially inhibited and ethylene evolution was doubled compared with the wild type, consistent with the results for tomato. In the aba2-1 etr1-1 double mutant, in which ABA was equally as deficient as in aba2-1 and shoot growth was shown to be insensitive to ethylene, shoot growth was substantially, although incompletely, restored relative to etr1-1. Treatment with ABA resulted in the complete recovery of shoot growth in aba2-1 relative to the wild type, and also significantly increased the growth of aba2-1 etr1-1 such that total leaf area and shoot fresh weight were not significantly lower than in etr1-1. In addition, ABA treatment of aba2-1 etr1-1 restored the wider leaf morphology phenotype exhibited by etr1-1. The results demonstrate that normal levels of endogenous ABA maintain shoot development, particularly leaf expansion, in well-watered Arabidopsis plants, partly by suppressing ethylene synthesis and partly by another mechanism that is independent of ethylene.     

ABA-deficiency results in reduced plant and fruit size in tomato

Abscisic acid (ABA) deficient mutants, such as notabilis and flacca, have helped elucidating the role of ABA during plant development and stress responses in tomato (Solanum lycopersicum L.). However, these mutants have only moderately decreased ABA levels. Here we report on plant and fruit development in the more strongly ABA-deficient notabilis/flacca (not/flc) double mutant. We observed that plant growth, leaf-surface area, drought-induced wilting and ABA-related gene expression in the different genotypes were strongly correlated with the ABA levels and thus most strongly affected in the not/flc double mutants. These mutants also had reduced fruit size that was caused by an overall smaller cell size. Lower ABA levels in fruits did not correlate with changes in auxin levels, but were accompanied by higher ethylene evolution rates. This suggests that in a wild-type background ABA stimulates cell enlargement during tomato fruit growth via a negative effect on ethylene synthesis.     

Manipulation of the apoplastic pH of intact plants mimics stomatal and growth responses to water availability and microclimatic variation

The apoplastic pH of intact Forsythiaxintermedia (cv. Lynwood) and tomato (Solanum lycopersicum) plants has been manipulated using buffered foliar sprays, and thereby stomatal conductance (g(s)), leaf growth rate, and plant water loss have been controlled. The more alkaline the pH of the foliar spray, the lower the g(s) and/or leaf growth rate subsequently measured. The most alkaline pH that was applied corresponds to that measured in sap extracted from shoots of tomato and Forsythia plants experiencing, respectively, soil drying or a relatively high photon flux density (PFD), vapour pressure deficit (VPD), and temperature in the leaf microclimate. The negative correlation between PFD/VPD/temperature and g(s) determined in well-watered Forsythia plants exposed to a naturally varying summer microclimate was eliminated by spraying the plants with relatively alkaline but not acidic buffers, providing evidence for a novel pH-based signalling mechanism linking the aerial microclimate with stomatal aperture. Increasing the pH of the foliar spray only reduced g(s) in plants of the abscisic acid (ABA)-deficient flacca mutant of tomato when ABA was simultaneously sprayed onto leaves or injected into stems. In well-watered Forsythia plants exposed to a naturally varying summer microclimate (variable PFD, VPD, and temperature), xylem pH and leaf ABA concentration fluctuated but were positively correlated. Manipulation of foliar apoplastic pH also affected the response of g(s) and leaf growth to ABA injected into stems of intact Forsythia plants. The techniques used here to control physiology and water use in intact growing plants could easily be applied in a horticultural context.     

Root-derived cytokinins as long-distance signals for NO3--induced stimulation of leaf growth

Leaf growth of many plant species shows rapid changes in response to alterations of the form and the level of N supply. In hydroponically-grown tomato (Lycopersicon esculentum L.), leaf growth was rapidly stimulated by NO(3)(-) application to NH(4)(+) precultured plants, while NH(4)(+) supply or complete N deprivation to NO(3)(-) precultured plants resulted in a rapid inhibition of leaf growth. Just 10 microM NO(3)(-) supply was sufficient to stimulate leaf growth to the same extent as 2 mM. Furthermore, continuous NO(3)(-) supply induced an oscillation of leaf growth rate with a 48 h interval. Since changes in NO(3)(-) levels in the xylem exudate and leaves did not correlate with NO(3)(-)-induced alterations of leaf growth rate, additional signals such as phytohormones may be involved. Levels of a known inhibitor of leaf growth, abscisic acid (ABA), did not consistently correspond to leaf growth rates in wild-type plants. Moreover, leaf growth of the ABA-deficient tomato mutant flacca was inhibited by NH(4)(+) without an increase in ABA concentration and was stimulated by NO(3)(-) despite its excessive ethylene production. These findings suggest that neither ABA nor ethylene are directly involved in the effects of N form on leaf growth. However, under all experimental conditions, stimulation of leaf growth by NO(3)(-) was consistently associated with increased concentration of the physiologically active forms of cytokinins, zeatin and zeatin riboside, in the xylem exudate. This indicates a major role for cytokinins as long-distance signals mediating the shoot response to NO(3)(-) perception in roots.     

Auxin-induced ethylene triggers abscisic acid biosynthesis and growth inhibition

The growth-inhibiting effects of indole-3-acetic acid (IAA) at high concentration and the synthetic auxins 7-chloro-3-methyl-8-quinolinecarboxylic acid (quinmerac), 2-methoxy-3,6-dichlorobenzoic acid (dicamba), 4-amino-3,6, 6-trichloropicolinic acid (picloram), and naphthalene acetic acid, were investigated in cleavers (Galium aparine). When plants were root treated with 0.5 mM IAA, shoot epinasty and inhibition of root and shoot growth developed during 24 h. Concomitantly, 1-aminocyclopropane-1-carboxylic acid (ACC) synthase activity, and ACC and ethylene production were transiently stimulated in the shoot tissue within 2 h, followed by increases in immunoreactive (+)-abscisic acid (ABA) and its precursor xanthoxal (xanthoxin) after 5 h. After 24 h of treatment, levels of xanthoxal and ABA were elevated up to 2- and 24-fold, relative to control, respectively. In plants treated with IAA, 7-chloro-3-methyl-8-quinolinecarboxylic acid, naphthalene acetic acid, 2-methoxy-3,6-dichlorobenzoic acid, and 4-amino-3,6,6-trichloropicolinic acid, levels of ethylene, ACC, and ABA increased in close correlation with inhibition of shoot growth. Aminoethoxyvinyl-glycine and cobalt ions, which inhibit ethylene synthesis, decreased ABA accumulation and growth inhibition, whereas the ethylene-releasing ethephon promoted ABA levels and growth inhibition. In accordance, tomato mutants defective in ethylene perception (never ripe) did not produce the xanthoxal and ABA increases and growth inhibition induced by auxins in wild-type plants. This suggests that auxin-stimulated ethylene triggers ABA accumulation and the consequent growth inhibition. Reduced catabolism most probably did not contribute to ABA increase, as indicated by immunoanalyses of ABA degradation and conjugation products in shoot tissue and by pulse experiments with [(3)H]-ABA in cell suspensions of G. aparine. In contrast, studies using inhibitors of ABA biosynthesis (fluridone, naproxen, and tungstate), ABA-deficient tomato mutants (notabilis, flacca, and sitiens), and quantification of xanthophylls indicate that ABA biosynthesis is influenced, probably through stimulated cleavage of xanthophylls to xanthoxal in shoot tissue.     

Stomatal control in tomato with ABA-deficient roots: response of grafted plants to soil drying

The hypothesis that ABA produced by roots in drying soil is responsible for stomatal closure was tested with grafted plants constructed from the ABA-deficient tomato mutants, sitiens and flacca and their near-isogenic wild-type parent. Three types of experiments were conducted. In the first type, reciprocal grafts were made between the wild type and sitiens or flacca. Stomatal conductance accorded with the genotype of the shoot, not the root. Stomates closed in all of the grafted plants in response to soil drying, regardless of the root genotype, i.e. regardless of the ability of the roots to produce ABA. In the second type of experiment, wild-type shoots were grafted onto a split-root system consisting of one wild-type root grafted to one mutant (flacca or sitiens) root. Water was withheld from one root system, while the other was watered well so that the shoots did not experience any decline in water potential or loss of turgor. Stomates closed to a similar extent when water was withheld from the mutant roots or the wild-type roots. In the third type of experiment, grafted plants with wild-type shoots and either wild-type or sitiens roots were established in pots that could be placed inside a pressure chamber, and the pressure increased as the soil dried so that the shoots remained fully turgid throughout. Stomates closed as the soil dried, regardless of whether the roots were wild type or sitiens. These experiments demonstrate that stomatal closure in response to soil drying can occur in the absence of leaf water deficit, and does not require ABA production by roots. A chemical signal from roots leading to a change in apoplastic ABA levels in leaves may be responsible for the stomatal closure.     

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.     

Plant Responses to Drought Stress and Exogenous ABA Application are Modulated Differently by Mycorrhization in Tomato and an ABA-deficient Mutant (<Emphasis Type="Italic">Sitiens</Emphasis>)

The aims of the present study are to find out whether the effects of arbuscular mycorrhizal (AM) symbiosis on plant resistance to water deficit are mediated by the endogenous abscisic acid (ABA) content of the host plant and whether the exogenous ABA application modifies such effects. The ABA-deficient tomato mutant sitiens and its near-isogenic wild-type parental line were used. Plant development, physiology, and expression of plant genes expected to be modulated by AM symbiosis, drought, and ABA were studied. Results showed that only wild-type tomato plants responded positively to mycorrhizal inoculation, while AM symbiosis was not observed to have any effect on plant development in sitiens plants grown under well-watered conditions. The application of ABA to sitiens plants enhanced plant growth both under well-watered and drought stress conditions. In respect to sitiens plants subjected to drought stress, the addition of ABA had a cumulative effect in relation to that of inoculation with G. intraradices. Most of the genes analyzed in this study showed different regulation patterns in wild-type and sitiens plants, suggesting that their gene expression is modulated by the plant ABA phenotype. In the same way, the colonization of roots with the AM fungus G. intraradices differently regulated the expression of these genes in wild-type and in sitiens plants, which could explain the distinctive effect of the symbiosis on each plant ABA phenotype. This also suggests that the effects of the AM symbiosis on plant responses and resistance to water deficit are mediated by the plant ABA phenotype.     

Evidence that abscisic acid does not regulate a centralized whole-plant response to low soil-resource availability

It has been suggested that abscisic acid (ABA) regulates a centralized response of plants to low soil resource availability that is characterized by decreased shoot growth relative to root growth, decreased photosynthesis and stomatal conductance, and decreased plant growth rate. The hypothesis was tested that an ABA-deficient mutant of tomato (flacca; flc) would not exhibit the same pattern of down-regulation of photosynthesis, conductance, leaf area and growth, as well as increased root/shoot partitioning, as its near isogenic wild-type in response to nitrogen or water deficiency, or at least not exhibit these responses to the same degree. Plants were grown from seed in acid-washed sand and exposed to control, nutrient stress, or water stress treatments. Additionally, exogenous ABA was sprayed onto the leaves of a separate group of flc individuals in each treatment. Growth analysis, based on data from frequent harvests of a few individuals, was used to assess the growth and partitioning responses of plants, and gas exchange characteristics were measured on plants throughout the experiment to examine the response of photosynthesis and stomatal conductance. Differences in growth, partitioning and gas exchange variables were found between flc and wild-type individuals, and both nutrient and water treatments caused significant reductions in relative growth rate (RGR) and changes in biomass partitioning. Only the nutrient treatment caused significant reductions in photosynthetic rates. However, flc and wild-type plants responded identically to nutrient and water stress for all but one of the variables measured. The exception was that flc showed a greater decrease in the relative change in leaf area per unit increase of plant biomass (an estimate of the dynamics of leaf area ratio) in response to nutrient stress—a result that is opposite to that predicted by the centralized stress response model. Furthermore, addition of exogenous ABA to flc did not significantly alter any of the responses to nutrient and water stress that we examined. Although it was clear that ABA regulated short-term stomatal responses, we found no evidence to support a pivotal role for ABA, at least absolute amounts of ABA, in regulating a centralized whole-plant response to low soil resource availability.     

The effect of competition from neighbours on stomatal conductance in lettuce and tomato plants

Competition decreased transpiration from young lettuce plants after 2 days, before any reductions in leaf area became apparent, and stomatal conductance (g(s) ) of lettuce and tomato plants was also reduced. Stomatal closure was not due to hydraulic signals or competition for nutrients, as soil water content, leaf water status and leaf nitrate concentrations were unaffected by neighbours. Competition-induced stomatal closure was absent in an abscisic acid (ABA)-deficient tomato mutant, flacca, indicating a fundamental involvement of ABA. Although tomato xylem sap ABA concentrations were unaffected by the presence of neighbours, ABA/pH-based stomatal modulation is still likely to underlie the response to competition, as soil and xylem sap alkalization was observed in competing plants. Competition also modulated leaf ethylene production, and treatment of lettuce plants with an ethylene perception inhibitor (1-methylcyclopropene) diminished the difference in g(s) between single and competing plants grown in a controlled environment room, but increased it in plants grown in the greenhouse: ethylene altered the extent of the stomatal response to competition. Effects of competition on g(s) are discussed in terms of the detection of the absence of neighbours: increases in g(s) and carbon fixation may allow faster initial space occupancy within an emerging community/crop.     

Long-distance signals regulating stomatal conductance and leaf growth in tomato (Lycopersicon esculentum) plants subjected to partial root-zone drying

Tomato (Lycopersicon esculentum Mill. cv. Ailsa Craig) plants were grown with roots split between two soil columns. After plant establishment, water was applied daily to one (partial root-zone drying-PRD) or both (well-watered control-WW) columns. Water was withheld from the other column in the PRD treatment, to expose some roots to drying soil. Soil and plant water status were monitored daily and throughout diurnal courses. Over 8 d, there were no treatment differences in leaf water potential (psileaf) even though soil moisture content of the upper 6 cm (theta) of the dry column in the PRD treatment decreased by up to 70%. Stomatal conductance (gs) of PRD plants decreased (relative to WW plants) when of the dry column decreased by 45%. Such closure coincided with increased xylem sap pH and did not require increased xylem sap abscisic acid (ABA) concentration ([X-ABA]). Detached leaflet ethylene evolution of PRD plants increased when of the dry column decreased by 55%, concurrent with decreased leaf elongation. The physiological significance of enhanced ethylene evolution of PRD plants was examined using a transgenic tomato (ACO1AS) with low stress-induced ethylene production. In response to PRD, ACO1AS and wild-type plants showed similar xylem sap pH, [X-ABA] and gs, but ACO1AS plants showed neither enhanced ethylene evolution nor significant reductions in leaf elongation. Combined use of genetic technologies to reduce ethylene production and agronomic technologies to sustain water status (such as PRD) may sustain plant growth under conditions where yield would otherwise be significantly reduced.     

Differential sensitivity of canola (Brassica napus) seedlings to ultraviolet-B radiation,water stress and abscisic acid

Responses of canola (Brassica napus L.) seedlings to three ultraviolet (UV)-B levels [0 (zero), 5 (ambient) and 10 (enhanced) kJ m^?2 d^?1], two watering regimes (well-watered and water-stressed), and two abscisic acid (ABA) levels (with and without application) were investigated. Overall, enhanced UVB and water stress negatively affected plant growth and physiology, but ABA had very little effect. Enhanced UVB decreased stem height, leaf area, plant dry matter, water use efficiency and wax content, but increased concentrations of chlorophyll a, carotenoids and flavonoids, and ethylene evolution. Water stress reduced stem height and diameter, leaf area, plant dry matter, leaf weight ratio and shoot:root weight ratio under zero and ambient UVB. Water stress also reduced chlorophyll a and carotenoids in plants exposed to enhanced UVB. ABA with watering regime had significant interactive effects only on leaf dry matter and wax content. We found that enhanced UVB and water stress adversely affected B. napus seedlings. Interaction between these two factors affected plant performance. In this interaction, ABA had little significant role. Also, optimum vegetative growth and biomass were achieved under ambient UVB.     

Interaction with ethylene: changing views on the role of abscisic acid in root and shoot growth responses to water stress

Shoot and root growth are differentially sensitive to water stress. Interest in the involvement of hormones in regulating these responses has focused on abscisic acid (ABA) because it accumulates in shoot and root tissues under water-limited conditions, and because it usually inhibits growth when applied to well-watered plants. However, the effects of ABA can differ in stressed and non-stressed plants, and it is therefore advantageous to manipulate endogenous ABA levels under water-stressed conditions. Studies utilizing ABA-deficient mutants and inhibitors of ABA synthesis to decrease endogenous ABA levels, and experimental strategies to circumvent variation in plant water status with ABA deficiency, are changing the view of the role of ABA from the traditional idea that the hormone is generally involved in growth inhibition. In particular, studies of several species indicate that an important role of endogenous ABA is to limit ethylene production, and that as a result of this interaction ABA may often function to maintain rather than inhibit shoot and root growth. Despite early speculation that interaction between these hormones may influence many of the effects of water deficit, this topic has received little attention until recently.     

ABA and polyamines act independently in primary leaves of cold-stressed tomato (Lycopersicon esculentum)

The effects of ABA and putrescine, a polyamine, on cold-induced membrane leakage were investigated using primary leaves of wild-type and an ABA-deficient mutant, flacca , of tomato ( Lycopersicon esculentum Mill.). The amount of chilling-induced electrolyte leakage from flacca leaves was much higher than that from the wild-type leaves. When applied exogenously ABA reduced cold-induced electrolyte leakage from leaves of both wild-type and the flacca mutant. However, the cold-induced electrolyte leakage from flacca leaves was not as pronounced as in the wild-type indicating that ABA is an important mediator in response to cold stress in the leaves. Putrescine reduced cold-induced electrolyte leakage from both wild-type and flacca leaves. Synthesis of putrescine in the leaves was increased by cold treatment. DFMO, a biosynthetic inhibitor of the polyamine, increased electrolyte leakage from cold-treated leaves, and exogenously applied putrescine decreased the enhanced leakage to the control level. Therefore, this polyamine is thought also to be involved in the response to cold stress of tomato leaves. Both ABA and putrescine were protective against cold stress, but exogenously applied ABA decreased the endogenous level of putrescine in the leaves. Furthermore, the DMFO-increased electrolyte leakage in cold-stressed leaves was completely abolished by the application of ABA. These results suggest that ABA is a major regulator in the response to cold stress in tomato leaves and that it does not exert its role via putrescine in the response to cold stress.     

Abscisic acid mimics effects of dehydration on area expansion and photosynthetic partitioning in young soybean leaves

Abstract. Leaf area expansion, photosynthetic carbon dioxide uptake and leaf dry mass accumulation were compared for expanding leaves of well-watered soybean ( Glycine max [L.] Merr.) plants, mildly dehydrated plants and well-watered plants treated with abscisic acid (ABA). Both ABA treatment and dehydration reduced area expansion in the light and over a 24 h period without decreasing the photosynthetic rates of expanding leaves. Dry mass accumulation during the light was less in ABA-treated and water-stressed leaves than in control leaves, with no differences among treatments in leaf mass per unit of area. ABA treatment and water stress both increased export of carbon from expanding leaves in the light. ABA applied near the end of the light period also increased export of carbon during the following dark period. However, it is unlikely that decreased availability of photosynthate caused slow expansion in the ABA and dehydration treatments, because expansion rates were not slowed in plants kept in dim light, even though photosynthetic rates and dry mass accumulation rates were greatly reduced. The data suggest that ABA may mediate the effects of mild dehydration on leaf area expansion and partitioning of photosynthate.     

Overproduction of abscisic acid in tomato increases transpiration efficiency and root hydraulic conductivity and influences leaf expansion

Overexpression of genes that respond to drought stress is a seemingly attractive approach for improving drought resistance in crops. However, the consequences for both water-use efficiency and productivity must be considered if agronomic utility is sought. Here, we characterize two tomato (Solanum lycopersicum) lines (sp12 and sp5) that overexpress a gene encoding 9-cis-epoxycarotenoid dioxygenase, the enzyme that catalyzes a key rate-limiting step in abscisic acid (ABA) biosynthesis. Both lines contained more ABA than the wild type, with sp5 accumulating more than sp12. Both had higher transpiration efficiency because of their lower stomatal conductance, as demonstrated by increases in delta(13)C and delta(18)O, and also by gravimetric and gas-exchange methods. They also had greater root hydraulic conductivity. Under well-watered glasshouse conditions, mature sp5 plants were found to have a shoot biomass equal to the wild type despite their lower assimilation rate per unit leaf area. These plants also had longer petioles, larger leaf area, increased specific leaf area, and reduced leaf epinasty. When exposed to root-zone water deficits, line sp12 showed an increase in xylem ABA concentration and a reduction in stomatal conductance to the same final levels as the wild type, but from a different basal level. Indeed, the main difference between the high ABA plants and the wild type was their performance under well-watered conditions: the former conserved soil water by limiting maximum stomatal conductance per unit leaf area, but also, at least in the case of sp5, developed a canopy more suited to light interception, maximizing assimilation per plant, possibly due to improved turgor or suppression of epinasty.     

Cytokinin-producing, plant growth-promoting rhizobacteria that confer resistance to drought stress in Platycladus orientalis container seedlings

One of the proposed mechanisms through which plant growth-promoting rhizobacteria (PGPR) enhance plant growth is the production of plant growth regulators, especially cytokinin. However, little information is available regarding cytokinin-producing PGPR inoculation on growth and water stress consistence of forest container seedlings under drought condition. This study determined the effects of Bacillus subtilis on hormone concentration, drought resistance, and plant growth under water-stressed conditions. Although no significant difference was observed under well-watered conditions, leaves of inoculated Platycladus orientalis (oriental thuja) seedlings under drought stress had higher relative water content and leaf water potential compared with those of noninoculated ones. Regardless of water supply levels, the root exudates, namely sugars, amino acids and organic acids, significantly increased because of B. subtilis inoculation. Water stress reduced shoot cytokinins by 39.14 %. However, inoculation decreased this deficit to only 10.22 %. The elevated levels of cytokinins in P. orientalis shoot were associated with higher concentration of abscisic acid (ABA). Stomatal conductance was significantly increased by B. subtilis inoculation in well-watered seedlings. However, the promoting effect of cytokinins on stomatal conductance was hampered, possibly by the combined action of elevated cytokinins and ABA. B. subtilis inoculation increased the shoot dry weight of well-watered and drought seedlings by 34.85 and 19.23 %, as well as the root by 15.445 and 13.99 %, respectively. Consequently, the root/shoot ratio significantly decreased, indicative of the greater benefits of PGPR on shoot growth than root. Thus, inoculation of cytokinin-producing PGPR in container seedlings can alleviate the drought stress and interfere with the suppression of shoot growth, showing a real potential to perform as a drought stress inhibitor in arid environments.     

Growth rate, plant development and water relations of the ABA-deficient tomato mutant sitiens

Given the close relationship between a plant's growth rate and its pattern of biomass allocation and the effects of abscisic acid (ABA) on biomass allocation, we studied the influence of ABA on biomass allocation and growth rate of wildtype tomato ( Lycopersicon esculentum Mill. cv. Moneymaker) plants and their strongly ABA-deficient mutant sitiens. The relative growth rate of sitiens was 22% lower than that of the wildtype, as the result of a decreased specific leaf area. The net assimilation rate and the leaf weight ratio were not affected. The mutant showed a much higher transpiration rate and lower hydraulic conductance of the roots. These two factors resulted in sitiens having a significantly lower leaf water potential and turgor. resulting in reduced leaf expansion and, consequently, a lower specific leaf area relative to the wildtype. Addition of ABA to the sitiens roots resulted in phenotypic reversion to the wildtype. We conclude that the influence of ABA-deficiency on biomass allocation and relative growth rate is the result of altered water relations in the plants, rather than of a direct effect on sink strength of different plant organs.