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     

Water relations and abscisic acid levels of watermelon as affected by rooting volume restriction

Rooting volume restriction (RVR) reduces shoot growth of plantsprovided with sufficient water or nutrients. The effects ofRVR on water status, abscisic acid (ABA) levels in leaves, roots,or xylem sap from detopped plants of watermelon [Citrullus lanatus(Thunb.) Matsum. and Nakai ‘StarBrite’] seedlingswere evaluated with five rooting volumes (18, 26, 36, 46, or80 cm3). Shoot water potential increased with increasing rootingvolume, with no difference between plants from 18 and 26 cm³cells or between plants from 36 and 46 cm³ cells. Stomatal conductancewas not consistently affected by RVR; at 10 and 20 DAE, stomatalconductance in plants grown in 36 cm³ cells was higher thanthat of plants grown in any other cell volume. Severe RVR (18and/or 26 cm³) tended to produce plants with higher ABA levelsin roots (15 DAE only), xylem sap (all dates), and leaves (5and 10 DAE). Plants grown in 18 and 26 cm³ cells had higherroot ABA levels than those from 46 and 80 cm³ cells at 15 DAE.Plants grown in 18 cm³ cells had the highest xylem sap ABA levelat all dates, but ABA levels did not differ among plants grownin the other cell volumes. Plants grown in 18 cm³ cells at 5DAE and 18 and 26 cm³ cells at 10 DAE also had higher leaf ABAlevels than those from other rooting volumes. The results suggestthat ABA may act as a signal for reduced growth of plants underRVR conditions. Key words: Abscisic acid, ABA, root signals, root volume restriction, water relations     

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     

Concentrations of abscisic acid and other solutes in xylem sap from root systems of tomato and castor-oil plants are distorted by wounding and variable sap flow rates

We investigated if concentrations of abscisic acid (ABA) andother solutes measured in the first few droplets of xylem sapfrom detopped root systems, are good estimates of those in thetranspiration stream as it enters the shoot-base of whole plants.Xylem sap from root systems of pot-grown tomato plants (Lycopersiconesculentum Mill., cv. Ailsa Craig), at the seven-leaf stage,was obtained by placing root systems in chambers pressurizedto 0.3 MPa with air. The first sample was taken from the cut-surfaceof the hypo-cotyl stump within 30 s of removing the shoot. ABA,sucrose and other osmolytes were more concentrated in the initial100–200 mm³ of xylem sap than in subsequent samples. Thissuggested the sap was contaminated and not unchanged transpirationfluid. The effect was reproduced on the same plant, severaltimes, by recutting the hypocotyl prior to reassembling thesap collecting set-up and repressurizing. Similar results werefound with castor-oil plants (Ricinus communis L., cv. Gibsonii).However, neither release of ABA from the cut surface of thehypocotyl stump, nor the effects of pressure to the roots causedthe contamination. Instead, small radial pressures exerted bya rubber sleeve attached to the hypocotyl stump, for collectingthe sap, were responsible. The effect was reproduced by lightlysqueezing the hypocotyl by hand. The possibility was examined that reliable estimates of ABAconcentrations in transpiration stream fluid may be obtainedfrom sap samples taken immediately after rejecting the initial,contaminated 200 mm³. However, ABA concentrations in these latersamples were also unsatisfactory since they changed with rateof sap flow. The problem may be overcome by analysing sap inducedto flow through detached root systems at rates close to thoseof whole-plant transpiration. Key words: Tomato, Lycopersicon esculentum Mill., Castor-oil plant, Ricinus communis L., roots, root to shoot communication, xylem sap, abscisic acid, sucrose, transpiration stream     

Growth of Zea mays L. plants with their seminal roots only. Effects on plant development, xylem transport, mineral nutrition and the flow and distribution of abscisic acid (ABA) as a possible shoot to root signal

Maize (Zea mays L.) was grown in quartz sand culture eitherwith a normal root system (controls) or with seminal roots only(‘single-rooted’). Development of adventitious rootswas prevented by using plants with an etiolated mesocotyl andthe stem base was positioned 5–8 cm above the sand. Eventhough the roots of the single-rooted plants were sufficientlysupplied with water and nutrients, the leaves experienced waterdeficits and showed decreased transpiration as trans plrationalwater flow was restricted by the constant number of xylem vesselspresent in the mesocotyl. As a consequence of this restriction,transpirational water flow velocities in the metaxylem vesselsreached mean values of 270 m h^–1 and phloem transportvelocities of 5.2 m h^–1. Despite limited xylem transportmineral nutrient concentrations in leaf tissues were not decreasedin single-rooted plants, but shoot and particularly stem developmentwas somewhat inhibited. Due to the lack of adventitious rootsthe shoot:root ratio was strongly increased in the single-rootedplants, but the seminal roots showed compensatory growth comparedto those in control plants. Consistent with decreased leaf conductance,ABA concentrations in leaves of single-rooted plants were elevatedup to 10-fold, but xylem sap ABA concentrations in these plantswere lower than in controls, in good agreement with the well-wateredconditions experienced by the seminal roots. Surprisingly, however,ABA concentrations in tissues of the seminal roots of the single-rooted plants were clearly increased compared to the controls,presumably due to increased ABA import via phloem from the water-stressedleaves. The results are discussed in relation to the role ofABA as a shoot to root signal. Key words: Zea mays, seminal roots, plant development, xylem transport, mineral nutrition, ABA, shoot-to-root signal     

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     

Salt, nutrient uptake and transport, and ABA of Populus euphratica; a hybrid in response to increasing soil NaCl

The uptake and transport of salt ions (Na⁺, Cl^-), macronutrients (K⁺, Ca²⁺, Mg²⁺) and abscisic acid (ABA) response to increasing soil salinity were examined in 2-year-old seedlings of Populus euphratica and a hybrid, P. talassica Kom 2 (P. euphratica + Salix alba L.). Leaf burn symptoms appeared in the hybrid after 8 days of exposure to salinity when soil NaCl concentration increased to 206 mM, whereas P. euphratica exhibited leaf damage after day 21 when soil NaCl exceeded 354 mM. Leaf necrosis was the result of excess salt accumulation since the injury followed an abrupt increase of endogenous salt levels. Compared with the hybrid, P. euphratica exhibited a greater capacity to exclude salt ions from leaves under increasing salinity, especially Cl^-. Salt treatment altered nutrient balance of the hybrid, leaf K⁺, Ca²⁺ and Mg²⁺ concentrations significantly declined and the same trends were observed in roots with the exception of K⁺. Although K⁺ levels decreased in salinised P. euphratica, increasing salinity did not affect the levels of Ca²⁺ and Mg²⁺ in leaves, but did increase the uptake of these nutrients when salt stress was initiated. NaCl-induced increase of ABA concentration in xylem sap [ABA] was observed in the two tested genotypes, however xylem [ABA] increased more rapidly in P. euphratica and a fivefold increase of xylem [ABA] was recorded after the first day of exposure to salt stress. Therefore, we conclude that the increase of Ca²⁺ uptake may be associated with the rise of ABA, and thus contributes to membrane integrity maintenance, which enables P. euphratica to regulate uptake and transport of salt ions under high levels of external salinity in the longer term.     

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     

Long Distance Transport of Abscisic Acid in NaCI-Treated Intact Plants of Lupinus albus

Plants of Lupinus albus were grown for 51 d under control (1.1mol m^–3 NaCl) and saline (40 mol m^–3 NaCl) conditions.Plants were harvested and changes of carbon, nitrogen and abscisicacid (ABA) contents of individual organs were determined 41d and 51 d after germination. In the period between the twoharvests xylem and phloem saps were collected and respirationand photosynthesis of individual organs were measured. Usingflows of carbon, C/ABA ratios and increments of ABA flows ofABA in phloem and xylem and rates of biosynthesis and degradationof ABA were calculated. Both under control and saline conditionsnet biosynthesis occurred in the root, the basal strata of leavesand in the inflorescence. Metabolic degradation of ABA tookplace in the stem internodes and apical leaf strata. Salt stress increased xylem transport of ABA up to 10-fold andphloem transport to the root up to 5-fold relative to that ofthe controls. A considerable amount of ABA in the xylem saporiginated from biosynthesis in the roots, i.e. 55% in salt-treatedand smaller than 28% in control plants. The remaining part ofABA in the xylem sap originated from the shoot: it was translocatedin the phloem from fully differentiated leaves towards the rootand from there it was recirculated back to the aerial partsof the plant. The data suggest that ABA may serve as a hormonalstress signal from the root system. Key words: Lupinus albus, salt stress, abscisic acid, long distance transport     

Delivery rates of abscisic acid in xylem sap of Ricinus communis L. plants subjected to part-drying of the soil

Abscisic acid (ABA) moving from roots to shoots in the transpirationstream is a potential hormonal message integrating perceptionof a root stress with adaptive changes in the shoot. A twinroot system was used to study ways of estimating effects ofdroughting the upper roots of Ricinus communis L. on abscisicacid (ABA) transport to the shoot in the transpiration stream.Droughted plants transpired more slowly than controls. Droughtingalso increased concentrations of ABA up to I I-fold in sap inducedto flow from the roots of freshly decapitated plants at ratesof whole plant transpiration. However, because of dilution effectsarising from the different sap flows in control and droughtedplants, these changes in ABA concentration in the xylem sapdid not accurately reflect amounts of ABA transported. To overcomethis problem, delivery rates were calculated by multiplyingconcentration with sap flow rate to generate ABA delivery interms of µmol s–1 per plant. Droughting for 24 hor more increased ABA delivery from roots to shoots by 5-fold.Since droughting can alter the relative sizes of the roots andshoots and also the root:shoot ratio these delivery rates wererefined in several ways to reflect both the amount of root generatingthe ABA message and the size of the recipient shoot system. Key words: Abscisic acid, Ricinus communis L., soil drying, xylem sap     

The Effects of Mechanical Impedance to Growth on the Levels of ABA and IAA in Root Tips of Zea mays L.

Extraction and analytical methods have been refined and newones devised to allow precise determinations by GC-EC of thelevels of abscisic acid (ABA) and indol-3ylacetic acid (IAA)in samples of maize root tips as small as 1.0 g fr. wt. Seminalroots of 5-d-old maize seedlings grown in normal (bulk density1200 kg m^–3) and compacted (bulk density 1600 kg m^–3)sand/garden loam mixtures have been examined. Seminal rootsfrom compacted soil had an average length of about 40% of thatof control roots and were much thicker. The ABA levels in 10mm tips of impeded roots (c. 25–35 ng g^–1 fr.wt.)did not differ significantly from those of normal root tipson both a fresh and dry weight basis. The levels in 0–1mm tips were approximately double those in the remaining 1–10mm zones. IAA levels were increased by about 3 times in impededroots (176.3 as compared with 52.4 ng g^–1 fr.wt) and itis concluded that this response is likely to be the main causeof the morphological and growth changes brought about by soilcompaction.     

Carbon Flow from Nodulated Roots to the Shoots of Soybean (Glycine max L. Merr.) Plants: An Estimation of the Contribution of Current Photosynthate to Ureides in the Xylem Stream

Kouchi, H. and Higuchi, T. 1988. Carbon flow from nodulatedroots to the shoots of soybean {Glycine max L. Merr.) plants:An estimation of the contribution of current photosynthate toureides in the xylem stream.–J. exp. Bot. 39: 1015–1023. Well-nodulated, water-cultured soybean plants were allowed toassimilate ¹³CO₂ at a constant specific activity for 10 h andthe ¹³C-labelling of total carbon and ureides in xylem sap wasinvestigated. Labelled carbon appeared very rapidly in the xylem stream. Percentageof labelled carbon (relative specific activity, RSA) in xylemsap was 18% at 2 h after the start of ¹³CO₂ assimilation andreached 53% at the end of the 10 h assimilation. The amountof labelled carbon exported from nodulated roots to the shootsvia the xylem during the 10 h labelling period accounted for33% of total labelled carbon imported into the nodulated roots.Ureides (allantoin and allantoic acid) in xylem sap were stronglydependent on currently assimilated carbon. The RSA of ureidesin xylem sap had reached 83% at the end of the assimilationperiod. Labelled carbon in ureides accounted for 51% of totallabelled carbon returned from nodulated roots to the shootsvia the xylem during the 10 h assimilation period. A treatmentwith 20 mol m^–3 nitrate in the culture medium for 2 ddecreased the ureide concentration in the xylem sap slightly,but greatly decreased the RSA of ureides. By comparing the data with the results of analysis of the xylemsap of nodule-detached plants, it was concluded that the majorityof labelled carbon exported to the xylem stream from noduleswas in ureide form. A considerable amount of carbon was alsoreturned from roots to shoots via the xylem stream but it wasmore dependent on (non-labelled) carbon reserved in the roottissues. Key words: Soybean(Glycine max L.), root nodule, carbon partitoning, ¹³CO₂ assimilation, xylem     

Inhibition of Ion Transport in Excised Barley Roots by Abscisic Acid; Relation to Water Permeability of the Roots

Previous papers have shown that abscisic acid can inhibit transportof ions across the root to the xylem vessels, resulting in reducedexudation from excised roots or inhibiting guttation from intactplants. However, it has not been established whether the inhibitionwas due to a reduction in salt transport (J_s) or in permeabilityof the roots to water (L_p). This paper investigates the effectof ABA on L_p and J_s separately. It is shown that L_p increasedin ABA and then fell, but was about the same as in control rootswhen transport was inhibited. The effect of ABA on exudationtherefore appeared to be mainly due to reduction in J_s. Inhibitionof J_s was also present in intact, transpiring plants and sowas not due to reduced water flow. The inhibition of ion releaseto the xylem affected Na⁺, Mg²⁺, Ca²⁺, and phosphate as wellas the major ion in the exudate, K⁺. It is concluded that ABAinhibits salt transport to the shoot by acting on ion transportinto the xylem, and not by reducing water flow coupled withsalt transport.     

Cycling of Amino-Nitrogen and other Nutrients between Shoots and Roots in Cereals--A Possible Mechanism Integrating Shoot and Root in the Regulation of Nutrient Uptake

A split root system was used to investigate the cycling of nitrogenbetween shoots and roots in young wheat and rye plants. ¹⁵N-nitratewas supplied to one part of the root system for various periods,at the end of which these roots were excised. Xylem sap wasthen collected from the other roots which had not been supplieddirectly with ¹⁵N-nitrate. ¹⁵N detected in the xylem sap indicatedcycling of nitrogen between shoots and roots. Calculations showedthat over 60% of the amino-N flux in the xylem was cycling.Thus nitrate assimilation in the root could account for onlya minor part of amino-N in the xylem sap. The specific activity of ¹⁵N in the total N of xylem sap washigher than in the total N of roots and shoots through whichit had cycled. This is because exchange between amino-N in thetransport pools and bulk tissue N is limited. It is proposedthat there is, in effect, a single regulatory pool of amino-N,common to shoots and roots, and that this pool may be a keyelement in the control of N uptake at the level of the wholeplant. The likely energy costs of cycling and implications for thepartitioning of N between shoots and roots are discussed. Infurther investigations the cycling of ⁴²K-potassium and ³²S-sulphurwas demonstrated. Key words: Potassium, sulphur, transport, xylem     

Characterization of Cd translocation and identification of the Cd form in xylem sap of the Cd-hyperaccumulator Arabidopsis halleri

Arabidopsis halleri is a Cd hyperaccumulator; however, the mechanismsinvolved in the root to shoot translocation of Cd are not wellunderstood. In this study, we characterized Cd transfer fromthe root medium to xylem in this species. Arabidopsis halleriaccumulated 1,500 mg kg^–1 Cd in the shoot without growthinhibition. A time-course experiment showed that the releaseof Cd into the xylem was very rapid; by 2 h exposure to Cd,Cd concentration in the xylem sap was 5-fold higher than thatin the external solution. The concentration of Cd in the xylemsap increased linearly with increasing Cd concentration in theexternal solution. Cd transfer to the xylem was completely inhibitedby the metabolic inhibitor carbonyl cyanide 3-chlorophenylhydrazone(CCCP). Cd concentration in the xylem sap was decreased by increasingthe concentration of external Zn, but enhanced by Fe deficiencytreatment. Analysis with ¹¹³Cd-nuclear magnetic resonance (NMR)showed that the chemical shift of ¹¹³Cd in the xylem sap wasthe same as that of Cd(NO₃)₂. Metal speciation with Geochem-PCalso showed that Cd occurred mainly in the free ionic form inthe xylem sap. These results suggest that Cd transfer from theroot medium to the xylem in A. halleri is an energy-dependentprocess that is partly shared with Zn and/or Fe transport. Furthermore,Cd is translocated from roots to shoots in inorganic forms.     

Normalizing Development of Cultured Triticum aestivum L. Embryos: I. LOW OXYGEN TENSIONS AND EXOGENOUS ABA

Wheat (Triticum aestivum L.) embryos form in dynamically-regulatedovular environments. Our objectives were to improve developmentof cultured immature wheat embryos by simulating, in vitro,abscisic acid (ABA) levels and O₂ tensions as found in wheatovules during zygotic embryogenesis. We characterized from intactwheat kernels embryo respiration, embryo morphology and embryoand endosperm + ABA levels at 13, 19 and 25 d post-anthesis(DPA). Young (13 DPA) embryos were then excised and culturedin vitro, where they were exposed to 0·2 or 2·Ommol m^–3 ±ABA and 2.·1, 2·5 or 7·4mol m^–3 (6, 7 and 21%, respectively) gaseous O₂. At 6and 12 d in culture, + ABA levels, embryo respiration and embryomorphology were characterized by treatment. Thirteen-day-oldembryos from two different plant populations differed by 17-foldin initial ABA content. However, this difference did not affectprecocious germination in vitro, nor did it affect the amountof exogenous ABA required to reduce precocious germination by40%. In this respect, embryos from both populations were equallysensitive to exogenous ABA. Cavity sap O₂ levels (2·1to 2·5 mol m^–3) were much more effective in preventingprecocious germination of cultured embryos than were cavitysap levels of ABA (0·2 to 2·0 mmol m^–3).The combination of physiological levels of both ABA and O₂ largelynormalized DW accumulation and embryo morphology without alteringendogenous + ABA levels. Residual respiration of cultured embryoswas higher than that of embryos grown in situ, and was not influencedby the exogenous O₂ and ABA treatments Key words: Abscisic acid, embryo development, oxygen tensions, respiration, wheat     

Concentrations and Transport of Solutes in Xylem and Phloem along the Leaf Axis of NaCl-treated Hordeum vulgare

Hordeum vulgare cv. California Mariout was established in sandculture at two different NaCl concentrations (0.5 mol m^–3‘control’ and 100 mol m^–3) in the presenceof 6.5 mol m^–3 K ⁺. Between 16 and 31 d after germination,before stem elongation started, xylem sap was collected by useof a pressure chamber. Collections were made at three differentsites on leaves 1 and 3: at the base of the sheath, at the baseof the blade, i.e. above the ligule, and at the tip of the blade.Phloem sap was collected from leaf 3 at similar sites throughaphid stylets. The concentrations of K ⁺, Na⁺, Mg2⁺ and Ca²⁺were measured. Ion concentrations in xylem sap collected at the base of leaves1 and 3 were identical, indicating there was no preferentialdelivery of specific ions to older leaves. All ion concentrationsin the xylem decreased from the base of the leaf towards thetip; these gradients were remarkably steep for young leaves,indicating high rates of ion uptake from the xylem. The gradientsdecreased with leaf age, but did not disappear completely. In phloem sap, concentrations of K⁺ and total osmolality declinedslightly from the tip to the base of leaves of both controland salt-treated plants. By contrast, Na⁺ concentrations inphloem sap collected from salt-treated plants decreased drasticallyfrom 21 mol m^–3 at the tip to 7.5 mol m^–3 at thebase. Data of K/Na ratios in xylem and phloem sap were used to constructan empirical model of Na⁺ and K⁺ flows within xylem and phloemduring the life cycle of a leaf, indicating recirculation ofNa⁺ within the leaf. Key words: Hordeum vulgare, xylem transport, phloem transport, NaCl-stress     

Unidirectional Ca2+ Fluxes in Roots of Rye (Secale cereale L). A Comparison of Excised Roots with Roots of Intact Plants

The unidirectional Ca²⁺ fluxes across the plasma membrane andtonoplast were determined in both excised roots and roots ofintact seedlings of rye (Secale cereale L. cv. Rheidol). Theunidirectional Ca²⁺ fluxes across the plasma membrane and tonoplastmeasured in excised roots were of a similar order of magnitudeto those determined in roots of intact plants. Influx and effluxof Ca²⁺ across the root plasma membrane were similar (estimatedto be between 0·7 and 3·4 µmol g     

REVIEW ARTICLE: Compartmental redistribution and long-distance transport of abscisic acid (ABA) in plants as influenced by environmental changes in the rhizosphere --a biomathematical model

Based on experimental data obtained in earlier studies on membranepermeabilities of abscisic acid (ABA) for cortex and stele cellsof roots and on measured com-partmental pH shifts after onsetor release of different types of soil-borne stresses, a biomathematicalmodel was developed which permits computer analysis of the dynamicsof compartmental ABA distribution within different root tissues(cortex, stele) and their compartments (apoplast, cytosol, vacuole),and in the xylem sap of the root stele. Metabolism and conjugationof ABA and its export from roots via the xylem and its importinto roots via phloem sap flow are also taken into consideration.We want to know which soil-borne stresses can biophysicallyprovoke a root-to-shoot signal of ABA. In this communicationwe describe the biomathematical structure of the root modeland present all necessary morphological (volumes, surfaces etc.)and physiological (pH, membrane conductances etc.) parametersof unstressed roots. This root model and an available leaf modelare integrated to a plant model (rosette plant). Simulationsreveal the fundamental role of the stele tissues, the rhizosphericABA concentration and the ABA synthesis in roots (root-to-shootcommunication). The shoot-to-root communication strongly dependson ABA synthesis in leaves, but hardly on ABA redistributioneffects after stress-induced compartmental pH-shifts in leaves. Key words: Abscisic acid, compartmental redistribution, computer model, pH shifts, root-to-shoot communication, shoot-to-root communication     

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