Effect of soil compaction on barley (Hordeum vulgare L.) growth: I. Possible role for ABA as a root-sourced chemical signal

Wild-type (Steptoe) and abscisic acid (ABA)-deficient mutant(Az34) genotypes of barley were grown in compacted soil to examinethe potential role of ABA as a root-to-shoot signal. Root andshoot growth and leaf conductance were all reduced when plantswere grown in compacted soil with a bulk density of 1.7g cm^–3,relative to uncompacted control plants (1.1 g cm^–3. Theseeffects occurred in the absence of detectable changes in leafwater status or foliar abscisic acid (ABA) content. Analysisof Steptoe and Az34 xylem sap showed that the ABA concentrationwas greatly increased at 6 d after emergence (6 DAE) when seedlingswere grown in compacted soil (1.7 g cm^–3); however, ABAconcentrations were never as high in the mutant as in the wildtype. The increase in xylem sap ABA concentration observed athigh bulk density was closely correlated with reductions inleaf conductance, but not leaf area. These increases were transitory,and xylem sap ABA concentrations subsequently decreased towardsthe control level by 18 DAE in both genotypes. The ABA-deficient mutant, Az34, produced a much lower leaf areathan Steptoe at a bulk density of 1.6 g cm^–3. Examinationof epidermal characteristics indicates that this effect resultedmainly from reductions in cell expansion rather than cell division,suggesting that the higher ABA concentrations detected in xylemsap from the wild-type Steptoe may have exerted a positive rolein maintaining leaf expansion in this treatment. The possibleinvolvement of ABA as a root-to-shoot signal mediating the effectsof compaction stress is discussed. Key words: Soil compaction, bulk density, ABA, ABA-deficient mutant, leaf growth     

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.     

Does an antagonistic relationship between ABA and ethylene mediate shoot growth when tomato (Lycopersicon esculentum Mill.) plants encounter compacted soil?

This study tested the hypothesis that antagonistic interactions between abscisic acid (ABA) and ethylene mediate the effects of soil compaction on shoot growth. Isogenic wild‐type (Ailsa Craig), ABA‐deficient (notabilis) and a transgenic (ACO1_AS) tomato genotype with a reduced capacity to synthesize ethylene were examined. Exogenous ABA was also applied. Leaf area was comparable when Ailsa Craig and ACO1_AS were grown in uncompacted (1·1 g cm^?3) or compacted (1·5 g cm^?3) soil, but was lower in notabilis. However, a 1·1/1·5 g cm^?3 split‐pot treatment invoked marked genotypic differences, whereby leaf area was comparable to 1·1 g cm^?3 control plants in ACO1_AS but was intermediate between the 1·1 and 1·5 g cm^?3 treatments in Ailsa Craig and notabilis. ABA may be discounted as the root‐sourced signal responsible for reducing leaf area when the roots encountered compacted soil as Ailsa Craig and ACO1_AS showed differing responses despite similar increases in xylem sap ABA concentration; leaf area was invariably lower in notabilis. These genotypic differences were correlated with ethylene evolution; thus the greater leaf area in ACO1_AS was associated with its reduced ability to synthesize ethylene, whereas the reductions in leaf expansion observed when Ailsa Craig and notabilis encountered compacted soil were accompanied by increased ethylene production. Application of ABA had little effect on ACO1_AS, but promoted a recovery of leaf expansion in notabilis, and more surprisingly in Ailsa Craig. These results suggest that antagonistic interactions between ABA and ethylene may regulate leaf expansion when the root system simultaneously encounters uncompacted and compacted soil.     

Effect of soil compaction on barley (Hordeum vulgare L.) growth: II Are increased xylem sap ABA concentrations involved in maintaining leaf expansion in compacted soils?

The abscisic acid (ABA)-deficient mutant of barley, Az34, exhibiteda much reduced rate of leaf expansion at a bulk density of 1.6g cm^–3 as compared to the isogenic wild-type variety,Steptoe. Az34 had a consistently lower xylem sap ABA concentrationat 7 d and 14 d after emergence (DAE). The xylem sap data suggestthat ABA present at Steptoe concentrations may have a directrole in maintaining leaf expansion at the sub-critical bulkdensity (1.6 g cm^–3 To test this hypothesis, additionof synthetic ABA either to the rooting environment (100 nM)or directly to the xylem sap (5 pg µl^–1 to reproducethe xylem sap ABA concentrations found in Steptoe, increasedleaf expansion in Az34 to the wild-type level. Furthermore,feeding Steptoe xylem sap to Az34 produced similar effects.These experiments provide direct evidence of a positive rolefor ABA as a root-to-shoot signal which assists in maintainingleaf growth in plants experiencing subcritical levels of compactionstress. Key words: ABA-deficient mutant, leaf expansion, xylem sap, ABA     

Soil compaction. A role for ethylene in regulating leaf expansion and shoot growth in tomato?

The role of ethylene in regulating growth in tomato (Lycopersicon esculentum Mill.) during compaction stress was examined using wild-type (cv Ailsa Craig) and transgenic (ACO1_AS) genotypes; the latter has a reduced capacity to produce ethylene. Ethephon or silver ions were applied to increase ethylene production or block its action. Shoot growth in both genotypes was comparable in uncompacted (1.1 g cm⁻³) and uniformly compacted soil (1.5 g cm⁻³). However, a 1.1/1.5-g cm⁻³ split-pot treatment invoked marked genotypic differences: growth was reduced in cv Ailsa Craig but was comparable to uncompacted control plants in ACO1_AS. As xylem sap abscisic acid levels were similar, abscisic acid was not responsible for inhibiting growth in cv Ailsa Craig. These genotypic differences in growth were accompanied by increased ethylene evolution in cv Ailsa Craig, suggesting that the ability of ACO1_AS to maintain growth in the split-pot treatment reflected its lower ethylene levels, a view supported by the observation that excising the roots in the compacted compartment reduced ethylene evolution and restored shoot growth in cv Ailsa Craig. Treatment with silver restored shoot growth in cv Ailsa Craig, whereas treatment with ethephon reduced growth in ACO1_AS. Thus, ethylene apparently has a key role in determining growth when tomato plants encounter differential soil compaction.     

Abscisic acid introduced into the transpiration stream accumulates in the guard-cell apoplast and causes stomatal closure

Petioles of water‐sufficient intact Vicia faba L. plants were infused with 1 µm abscisic acid (ABA) to simulate the import of root‐source ABA. This protocol permitted quantitative ABA delivery, up to 300 pmol ABA over 60 min, to the leaf without ambiguities associated with perturbations in plant–water status. The ABA concentrations in whole‐leaf samples and in apoplastic sap increased with the amount infused; ABA degradation was not detected. The ABA concentration in apoplastic sap was consistent with uptake of imported ABA into the leaf symplast, but this interpretation is qualified. Our focus was quantitative cellular compartmentation of imported ABA in guard cells. Unlike when leaves are stressed, the guard‐cell symplast ABA content did not increase because of ABA infusion (P = 0·48; 3·0 ± 0·5 versus 4·0 ± 1·2 fg guard‐cell‐pair^?1). However, the guard‐cell apoplast ABA content increased linearly (R² = 0·98) from ?0·2 ± 0·5 to 3·1 ± 1·3 fg guard‐cell‐pair^?1 (≈ 3·1 µm ) and was inversely related to leaf conductance (R² = 0·82). Apparently, xylem ABA accumulates in the guard‐cell wall as a result of evaporation of the apoplast solution. This mechanism provides for integrating transpiration rate and ABA concentration in the xylem solution.     

Rapid Effects of Abscisic Acid on Ion Uptake in Sunflower Roots

Short-term effects of ABA, ABA + kinetin and kinetin on ion (⁸⁶Rb-potassium and phosphate) and water uptake in sunflower plants (Helianthus annuus var. californicus) were examined with a continuous-recording technique. Ion uptake in the roots and transport to the shoots were also investigated by conventional tracer uptake experiments and by sap bleeding experiments with excised roots. After addition of 5 × 10^?6-4 × 10^?5M ABA to the root medium there was an immediate decrease (30–70%) in the rate of ion uptake which lasted 30–70 min. The rate of water uptake was not significantly affected as measured with this method. Ion transport to the shoots and to the bleeding sap of excised roots was decreased by ABA. ABA-induced inhibition of ion uptake was abolished by the presence of kinetin, and uptake was slightly stimulated by 2 × 10^?5M kinetin alone. We suggest that concentration gradients of ABA or rapid changes in the ABA-kinetin balance in the roots affect ion uptake and transport.     

Abscisic Acid is not the only stomatal inhibitor in the transpiration stream of wheat plants

Xylem sap was collected from the transpiration stream of wheat (Triticum aestivum L.) plants and assayed for the presence of an inhibitor of transpiration using leaves detached from well-watered plants. Transpiration of detached leaves was reduced by nearly 60% by sap collected from plants in drying soil, and to a lesser extent (about 25%) by sap from plants in well-watered soil. As the soil dried the abscisic acid (ABA) concentration in the sap increased by about 50 times to 5 × 10⁻⁸ molar. However, the ABA in the sap did not cause its inhibitory activity. Synthetic ABA of one hundred times this concentration was needed to reduce transpiration rates of detached leaves to the same extent. Furthermore, inhibitory activity of the sap was retained after its passage through an immunoaffinity column to remove ABA. Xylem sap was also collected by applying pressure to the roots of plants whose leaf water status was kept high as the soil dried. Sap collected from these plants reduced transpiration to a lesser extent than sap from nonpressurised plants. This suggests that the inhibitory activity was triggered partly by leaf water deficit and partly by root water deficit.     

The effects of exogenous ABA applied to maize (<Emphasis Type="Italic">Zea mays</Emphasis> L.) roots on plant responses to chilling stress

This work aimed to discuss the effects of exogenous abscisic acid (ABA) on the root growth regulation of maize seedlings under chilling stress. The roots of the maize cultivar Zhengdan 958 were irrigated with ABA (10^?7, 10^?6, 10^?5 and 10^?4 M) at the third true leaf stage under chilling duration (0, 2, 4, 6, and 8 days). The biomass, the phenylalanine ammonia lyase (PAL), and polyphenol oxidase (PPO) enzyme activities, total phenolic and flavonoid contents, the ferric reducing ability of plasma (FRAP) antioxidant capacity, and 2,2-azinobis (3-ethlbenzothiazo-line-6-sulfonic acid) diammonium salt radical (ABTS^·+) scavenging capacity of the roots of maize seedlings were measured after the treatment. The results showed that appropriate concentrations of exogenous ABA effectively enhanced root biomass, increased PAL and PPO enzyme activities, and significantly increased total phenolic contents and flavonoid contents. Moreover, the ABA markedly improved the FRAP antioxidant capacity and ABTS^·+ scavenging capacity under low-temperature stress. These results indicate that ABA-treated maize seedlings are resistant to chilling stress and that the optimum concentration of ABA is 10^?5 M. Exogenous applications of ABA have a concentration effect in alleviating chilling stress, in which low concentrations have a promoting effect and high concentrations have an inhibiting effect.     

Growth,graviresponsiveness and abscisic-acid content of Zea mays seedlings treated with Fluridone

Ten-d-old seedlings of Zea mays L. cv. Tx 5855 treated with 1-methyl-3-phenyl-5-(3-[trifluoromethyl]phenyl)-4-(1H)-pyridinone (Fluridone) were analyzed for abscisic acid (ABA) content using high-performance liquid chromatography with an analysis sensitivity of 2.5 ng ABA g^-1 fresh weight (FW). Seedlings were divided into three portions: leaves, detipped roots, and root tips (terminal 1.5 mm). Control plants (water treatment only; no Fluridone) were characterized by the following amounts of ABA: leaves, 0.114±0.024 (standard deviation) g ABA g^-1 FW; detipped roots, 0.260±0.039±g ABA g^-1 FW; root tips, no ABA detected. We did not detect any ABA in tissues of Fluridone-treated plants. Primary roots of treated and untreated seedlings were strongly graviresponsive, with no significant differences between the curvatures or the growth rates of primary roots of Fluridone-treated and control seedlings. These results indicate that 1) Fluridone completely inhibits ABA synthesis, and 2) ABA is not necessary for positive gravitropism by primary roots of Zea mays.Abbreviations ABA abscisic acid - Fluridone 1-methyl-3-phenyl-5-(3-[trifluoromethyl]phenyl)-4-(1H)-pyridinone - FW fresh weight - SD standard deviation     

Effects of surgically implanted acoustic transmitters >2% of body mass on the swimming performance, survival and growth of juvenile sockeye and Chinook salmon

The influence of surgical implantation of an acoustic transmitter on the swimming performance, growth and survival of juvenile sockeye salmon Oncorhynchus nerka and Chinook salmon Oncorhynchus tshawytscha was examined. The transmitter had a mass of 0·7 g in air while sockeye salmon had a mass of 7·0–16·0 g and Chinook salmon had a mass of 6·7–23·1 g (a transmitter burden of 4·5–10·3% for sockeye salmon and 3·1–10·7% for Chinook salmon). Mean critical swimming speeds (U_crit) for Chinook salmon ranged from 47·5 to 51·2 cm s^?1 [4·34–4·69 body lengths (fork length, L_F) s^?1] and did not differ among tagged, untagged and sham‐tagged groups. Tagged sockeye salmon, however, did have lower U_crit than control or sham fish. The mean U_crit for tagged sockeye salmon was 46·1 cm s^?1 (4·1 L_F s^?1), which was c. 5% less than the mean U_crit for control and sham fish (both groups were 48·6 cm s^?1 or 4·3 L_F s^?1). A laboratory evaluation determined that there was no difference in L_F or mass among treatments (control, sham or tag) either at the start or at the end of the test period, suggesting that implantation did not negatively influence the growth of either species. None of the sockeye salmon held under laboratory conditions died from the influence of surgical implantation of transmitters. In contrast, this study found that the 21 day survival differed between tagged and control groups of Chinook salmon, although this result may have been confounded by the poor health of Chinook salmon treatment groups.     

The Effects of Partially Drying Part of the Root System of Helianthus annuus on the Abscisic Acid Content of the Roots, Xylem Sap and Leaves

Plants of Helianthus annuus were grown in soil in pots suchthat approximately 30% of the root system protruded throughthe base of the pot. After 7 d further growth in aerated nutrientsolution, the attached, protruding roots were air-dried for10–15 min and thereafter surrounded with moist still air,in the dark, for 49 h, whilst the soil was kept at field capacity.The roots of the control plants remained in the nutrient solutionthroughout the experiment. This treatment rapidly reduced the water content of protrudingroots from 20.5 to 17.8 g g^–1 dry mass (DM), which remainedless than that of the control roots for the rest of the experiment.This treatment also reduced root turgor and water potential.The abscisic acid (ABA) concentrations in the protruding roots,xylem sap and leaves of the treated plants increased significantly,compared to values recorded for control plants. In treated roots, the ABA concentration was significantly increased4 h after treatment, with a maximum of 4.4+0.1 nmol g^–1(DM) after 25 h. The ABA concentration in the xylem sap of thetreated plants was significantly greater than in the controls25 h, 30 h, and 49 h after the partial drying of the roots,with a maximum concentration of approximately 970 pmol ABA cm-3at 49 h. Initially, the ABA concentration in the leaves was0.45 nmol g^–1 (DM) which increased significantly to 1.1±0.1 nmol g^–1 at 25 h, to 1.7±0.3 nmol g^–1at 49 h. Leaf conductance was significantly less in plants with air-driedroots than in the controls 8 h after the start of the treatmentand thereafter. The water relations of the leaves of the treatedplants did not differ from those of the control plants. These results confirm previous reports that ABA is rapidly generatedin partially-dried and attached root systems and demonstratesa concomitant large increase in the ABA content of the xylemsap. It is suggested that partial dehydration of some of theroots of Helianthus annuus, increases ABA concentration in thetranspiration stream and decreases leaf conductance in the absenceof changes in leaf water status. As these responses were initiatedin free-growing roots the stimulus is independent of any increasesin soil shear strength that are associated with soil drying. Key words: Soil drying, roots, ABA, leaf conductance, water relations     

Oxygen Diffusion from the Roots of Rice Grown under Non-waterlogged Conditions

Three rice varieties, cv. Norin 36, cv. Norin 37 and cv. Yubae, were grown in a loam with a 20 cm water-table which gave aerobic conditions to a depth of not less than 15 to 17 cm. Under these conditions Norin 36 grew more vigorously and tillered more frequently than the other two varieties. The rates of oxygen diffusion at 23°C from roots up to 11 cm in length were however appreciably lower for Norin 36 (4.3 × 10^?8g · cm^?2 of root surface · min^?1) than for Norin 37 or Yubae (c. 7.8 × 10^?8g). A considerable increase (up to 200 %) in the oxygen diffusion rate (ODR) from the roots occurred if they were cooled to 3°C, and at this temperature differences in ODR between the varieties were not significant. For a purely physical system, because of the decrease in the diffusion coefficient of oxygen in water, and, the increase in oxygen solubility, a drop of c. 20 % in ODR should accompany the above 20°C drop in temperature. A 16 % drop was recorded for artificial ‘roots’ under these conditions. It was concluded that respiratory activity at the higher temperature must have been responsible for the low readings and intervarietal differences observed at 23°C. By increasing the 3°C values by 25 % a mean value of 14.2 × 10^?8g · cm^?2 of root surface · min^?1 was recorded for the three varieties, being the probable ODR at 23°C in the absence of a respiratory factor. Calculations show that respiratory activity removed enough oxygen to reduce the ODR for Norin 36 by more than 9 × 10^?8g, and for Norin 37 and Yubae by c. 6.7 × 10^?8g · cm^?2 of root surface · min^?1. Anatomical investigations showed that cortical breakdown was always extensive at 4 to 4.5 cm from the apex of the roots. In some cases however breakdown had not occurred in the basal segment of the root. No opinion could be formed as to whether differences in the amount of cortical breakdown between the varieties might have occasioned the respiratory differences observed. An interesting feature of the root anatomy was the failure of breakdown in those regions of the roots through which lateral roots emerged.     

Hormonal responses to partial drying of the root system of Helianthus annuus

Plants of Helianthus annuus were pot-grown in soil, with approximately30% of the root system protruding through the base. After 7d, the upper part of the root system of half of the plants wasexposed to drought (internal roots) while the lower part waskept in aerated nutrient solution (protruding root). The treatmentrapidly reduced the internal roots' water content from 26.1to 21.9 g g^–1 dry weight (DW), while in protruding rootsof stressed plants it slowly and continuously decreased from31.9 to 25.2 g g^–1 DW. Leaf water content rapidly decreasedin treated plants from 7.4 to 6.4 g g^–1 DW in the first2d and then reached a plateau. In stressed plants leaf stomatalresistance was significantly higher in the first 3 d while leafwater potential was lower only on the last day. Abscisic acid (ABA) concentration in treated plants increasedsignificantly compared to the controls. In treated internalroots, ABA rose from the first day, reaching a maximum of 1.48±0.49nmol g^–1 DW after 3 d. In treated protruding roots a maximumof 0.99±0.09 nmol g^–1 DW was reached after 1 d.ABA concentration in the xylem sap increased 2 d and 3 d afterthe start of soil drying, with a maximum of 113±12nmoll^–1 during the third day. The ABA rise in the leaves oftreated plants was less significant. Indol-3yl-acetic acid (IAA) concentration in internal rootsof treated plants reached a maximum of 22.54±3.34 nmolg^–1 DW on the third day, then decreased dramatically.The protruding root system of control plants showed a maximumvalue of 16.05±1.77 nmol g^–1 DW on the sixth day. Little difference in cytokinin content of xylem sap was notedbetween control and treated plants. Hormonal variations in different parts of the plant are discussedin relation to drought stress. Key words: Soil drying, roots, ABA, IAA, cytokinins     

Genotypic variation in 'early warning signals' from roots in drying soil: intrinsic differences in ABA synthesising capacity rather than root density determines total ABA 'message' in cowpea (Vigna unguiculata L.)

In a comparison of six cowpea cultivars, we determined the variation in abscisic acid (ABA) production as an ‘early warning signal’ produced in response to drought stress. By imposing drought only to the upper 20 cm rooting zone, we compared the rates of ABA synthesis relative to (i) total root mass and (ii) inherent variation per unit root mass. We were able to relate the intensity of the stress response to these two factors, and determine which is quantitatively more important as the primary signal indicating responsiveness to drought stress. Plants were grown in 1.2 m long columns and a soil drying treatment imposed in such a way that that upper roots were in dry soil and deep roots in soil at field capacity. Relative water contents (RWC) of stressed plants were similar and not significantly different from those of well watered controls. However, roots accumulated ABA in the dehydrated zone, where root water content ranged from 10–12 g g^?1 DW. The soil moisture contents and root ‐water contents in the dry zone were similar for each of the different varieties. However, the ABA contents were significantly different in drought‐stressed (upper) roots and ranged from 7.82 nmol g^?1 DW in cv. APC 689 to 16.02 nmol g^?1 DW in cv. APC 370, such that for varieties with similar overall root weights (e.g. APC 580 and APC 540) the different ABA contents were related to the capacity for ABA synthesis. The relationship between stomatal conductance and total root ABA was assessed, with a negative relation (r= 0.90, n= 24, P= 0.05) suggesting that the intrinsic capacity of cowpea varieties for ABA synthesis could play an important role in regulating stomatal conductance in a drying soil and provide useful selection criteria for tolerance to drought stress.     

Root and bacterial secretions regulate the interaction between plants and PGPR leading to distinct plant growth promotion effects

Background and aims

Long distance signals in xylem from roots to leaves are important in plant response to drought stress. Abscisic acid (ABA) plays a key role in drought signaling in plants but apoplastic pH may modulate its effect by distributing ABA into various compartments in leaves. We aimed to reveal the dynamics of changes in sap pH and its relationships with the transport of inorganic and organic ions in eight herbaceous plant species under continuously declining soil water content. We tested several hypotheses related to the mechanism of pH changes in xylem.

Methods

We used a pressure chamber to collect xylem sap and to measure of leaf/stem water potential at various stages of soil drying. We measured pH and concentrations of the most abundant inorganic (NO₃ ^?, SO₄ ^2?, PO₄ ^3? and Cl^?) and organic (malate and citrate) anions in xylem sap.

Results

Species differed considerably in the dynamics of pH changes in xylem in drying soil. Changes in xylem sap pH during drying did not relate to the nitrogen assimilation strategy but may be affected by sap flow rate. Simultaneous changes in the concentrations of inorganic and organic anions were highly species-specific.

Conclusions

High variability among species in the observed relationships in response to drought indicates that comparisons among different studies and the generalization of results should be made with caution.

     

Is coordination of leaf and root growth mediated by abscisic acid? Opinion

Leaf growth is more inhibited than root growth when the soil is nitrogen-deficient, dry, saline, compacted, or of restricted volume. Similar differential responses in leaf and root growth occur when ABA is applied to plants in well-watered and well-fertilised conditions, and opposite responses are often found in ABA-deficient mutants. ABA levels increase in plants in dry or saline soils, suggesting a regulating role in leaf and root growth in soils of low water potential. In nitrogen-deficient or compacted soils, or soils of restricted volume, ABA only sometimes increases, and in these situations its accumulation may be of secondary importance. Use of ABA-deficient mutants has so far indicated that ABA influences leaf and root growth in unstressed plants, and plants in dry soils, but not in soils that are compacted, of restricted volume, or are nitrogen-deficient.For ABA to determine the relationship between the rate of leaf growth and the rate of root growth, there must be long-distance transport of either ABA itself or a compound that controls ABA synthesis in the growing cells of leaves and roots. ABA invariably increases in xylem sap as the soil becomes dry or saline, and sometimes when it becomes nitrogen-deficient or compacted, however the ABA is of too low a concentration to affect leaf growth. There may be a compound in xylem sap that controls the synthesis of ABA in the leaf, but no such compound has been identified. ABA accumulates in phloem sap of plants in dry or saline soil, but its function in controlling root or leaf growth is unknown.We conclude that ABA affects the ratio of root growth to leaf growth via its independent effects on root and leaf growth, and may regulate the ratio of root to leaf growth via feedforward signals in xylem or phloem, but there is no satisfactory explanation of its mechanism of control.     

Water relations and growth of original barley plants and its ABA-deficient mutants at increased air temperature

Data on the effects of air temperature increase by 4°C on leaf growth and water relation parameters in barley (Hordeum vulgare L.) plants in original cv. Steptoe and its ABA-deficient mutant (AZ24) are presented. An increase in temperature firstly resulted in the cessation of leaf elongation in both genotypes; however, later in cv. Steptoe plants, as distinct from mutants, the rate of leaf length increment was completely restored. Before air warming, transpiration was more intense in mutant plants; at increased temperature, transpiration was activated in both genotypes. After growth resumption, the water potential in cv. Steptoe plants somewhat increased as compared with initial level (before warming). In AZ34 leaves, in contrast, the water potential, which was initially below that in cv. Steptoe leaves, reduced after temperature increase. The calculation of total hydraulic conductivity of the plants and osmotic hydraulic conductivity in the roots showed that these parameters increased in cv. Steptoe and were not changed in AZ34 mutants. At temperature increase, the level of ABA was not changed in AZ34 mutants, whereas in Steptoe plants it increased in the roots and decreased in the shoots. It was concluded that a capability of ABA synthesis is required for the control of total hydraulic conductivity under changing environmental conditions.     

Phloem transport of abscisic acid in Ricinus communis L. seedlings

Sieve tube sap exuded from the cut hypocotyl of castor bean seedlings (Ricinus communis L.) was found to contain 0.2–0.5 mmol m^?3abscisic acid (ABA). The ABA concentration in the sieve tube sap always exceeded that in root pressure exudate under a wide range of water supply. Exudation of sieve tube sap from the cut hypocotyls caused water loss, and this induced ‘water shortage’ in the cotyledons which resulted in the ABA concentration in the cotyledons increasing by 3-fold and that in the sieve tube sap increasing by up to 50-fold within 7h. The wounded surface of the cut hypocotyl was not responsible for the ABA increase. Incubation of the cotyledons of endosperm-free seedlings in various ABA concentrations (up to 100 mmol m^?3) increased the ABA concentration in sieve tube sap. The concomitant increase in ABA, both in cotyledons and in sieve tube sap, had no effect on the phloem loading of sucrose, K⁺ and Mg²⁺ within the experimental period, i.e. up to 10h. It can be concluded that (i) the phloem is an important transport path for ABA, (ii) water stress at the phloem loading sites elevates phloem-mobile ABA, which may then serve as a water stress signal for sinks, for example stem and roots (not only for stomata), and (iii) the ABA concentration of cells next to or in the phloem is more important than the average ABA content in the whole cotyledon for determining the ABA concentration in sieve tube sap.     

Does Xylem Sap ABA Control the Stomatal Behaviour of Water-Stressed Sycamore (Acer pseudoplatanus L.) Seedlings?

Sycamore seedlings were grown with their root systems dividedequally between two containers. Water was withheld from onecontainer while the other container was kept well-watered. Effectsof soil drying on stomatal behaviour, shoot water status, andabscisic acid (ABA) concentration in roots, xylem sap and leaveswere evaluated. At 3 d, root ABA in the drying container increased significantly,while the root ABA in the unstressed container of the same plantsdid not differ from that of the control. The increase in rootABA was associated with the increase in xylem sap ABA and withthe decrease in stomatal conductance without any significantperturbation in shoot water status. At 7 d, despite the continuous increase in root ABA concentration,xylem sap ABA showed a marked decline when soil water contentwas depleted below 013 g g^–1. This reduction in xylemsap ABA coincided with a partial recovery of stomatal conductance.The results indicate that xylem sap ABA is a function of rootABA as well as the flow rate of water from roots to shoots,and that this ABA can be a sensitive indicator to the shootof the effect of soil drying. Key words: Acer pseudoplatanus L., soil drying, stomatal behaviour, xylem sap ABA