The effect of root cooling on hormone content, leaf conductance and root hydraulic conductivity of durum wheat seedlings (Triticum durum L.)

Root cooling of 7-day-old wheat seedlings decreased root hydraulic conductivity causing a gradual loss of relative water content during 45 min (RWC). Subsequently (in 60 min), RWC became partially restored due to a decrease in transpiration linked to lower stomatal conductivity. The decrease in stomatal conductivity cannot be attributed to ABA-induced stomatal closure, since no increase in ABA content in the leaves or in the concentration in xylem sap or delivery of ABA from roots was found. However, decreased stomatal conductance was associated with a sharp decline in the content of cytokinins in shoots that was registered shortly after the start of root cooling and linked to increases in the activity of cytokinin-oxidase. This decrease in shoot cytokinin content may have been responsible for closing stomata, since this hormone is known to maintain stomatal opening when applied to plants. In support of this, pre-treatment with synthetic cytokinin benzyladenine was found to increase transpiration of wheat seedlings with cooled roots and bring about visible loss of turgor and wilting.     

Does Cytokinin Transport from Root-To-Shoot in the Xylem Sap Regulate Leaf Responses to Root Hypoxia?

We have examined the hypothesis that cytokinins transportedfrom roots to shoots affects leaf growth, stomatal conductance,and cytokinin concentration of leaves of Phaseolus and a hybridpoplar (Populus trichocarpa x Populus deltoides) with hypoxicroots. Because cytokinins may interact with other substances,potassium and calcium concentrations were determined in xylemsap of Populus plants with hypoxic and aerated roots while gibberellin(GA) concentrations were measured in shoot tissues. Root hypoxiadecreased leaf growth and closed stomata in both species. Inboth species, fluxes of cytokinins out of the roots were reduced,but no differences in bulk leaf concentrations were measuredbetween the hypoxic and aerated plants. Shoots with aeratedroots contained slightly higher concentrations of GA₁ and GA₃than shoots from hypoxic plants. There were no differences incalcium or potassium concentrations in xylem sap between aerationtreatments. Exogenously applied cytokinins did not alleviatethe growth or stomatal responses caused by root hypoxia. Informationon the site(s) and mechanism(s) of cytokinin action and theways in which cytokinins are compartmentalized within plantcells will be required to understand the physiological significanceof cytokinin transport in the transpirational stream. Key words: Cytokinins, hypoxia, Populus, Phaseolus     

Soil temperature and flooding effects on two species of citrus

Summary Rough lemon (Citrus jambhiri Lush.) and sour orange (C. aurantium L.) seedlings were grown at constant soil temperatures of 16, 24, and 33 C for 3 months. Shoot and root growth of rough lemon was greatest at 33 C while growth of sour orange was greatest at 24 C. There were no significant effects of soil temperature on shoot: root ratio, leaf water potential or stomatal conductance. The hydraulic conductivity of intact root systems of both species was highest when seedlings were grown at 16 C. Thus, acclimation through greater root conductivity at low soil temperature may have compensated for decreased root growth at 16 C and negated effects of soil temperature on plant water relations. Half the plants growing at each soil temperature were subsequently flooded. Within 1 week, the soil redox potential (Eh) dropped below zero mV, reaching a minimum Eh of –250mV after 3 weeks of flooded conditions. Flooded plants exhibited lower root conductivity, a cessation of shoot growth, lower leaf water potentials, lower stomatal conductances, and visual sloughing of fibrous roots. Decreases in root conductivity in response to flooding were large enough to account for the observed decreases in stomatal conductance.Florida Agricultural Experiment Stations Journal Series No. 4080.     

Ability of bacterium Bacillus subtilis to produce cytokinins and to influence the growth and endogenous hormone content of lettuce plants

Hormone production by micro-organisms selected as antagonists of pathogenic fungi and the effect of their introduction into soil on hormone content and growth of lettuce plants were studied. Hormones in bacterial cultural media and in plant extracts were immunopurified and assayed using specific antibodies to indolyl-3-acetic acid (IAA), abscisic acid (ABA), and different cytokinins (zeatin riboside (ZR), dihydrozeatinriboside (DHZR) and isopentenyladenosine (iPA)). ZR was shown to be the main cytokinin present in bacterial cultural media as a complex with a high molecular weight component. Inoculation of lettuce plants with bacteria increased the cytokinin content of both shoots and roots. Accumulation of zeatin and its riboside was greatest in roots shortly 2days after inoculation, when their content was 10 times higher than in control. Changes in the content of other hormones (ABA and IAA) were observed at the end of experiments only. Accumulation of cytokinins in inoculated lettuce plants was associated with an increase in plant shoot and root weight of approximately 30% over 8days.     

Plant growth and physiology under heterogeneous salinity

Background

Soil salinity is heterogeneous, and within the root-zone of single plants the salinity of the soil solution can vary widely.

Scope

This review shows that water uptake by roots from the least saline part of the soil is the key factor driving shoot growth; plants with part of the root at low salinity (0–10?mM NaCl) had 3- to 10-fold higher shoot dry mass than plants with roots in uniformly saline (50–800?mM NaCl) media. Plants in heterogeneous salinity had shoot water potentials similar to those of plants growing in uniform low-salt media, and this was likely a result of uptake of low salinity water and reduced stomatal conductance. Under heterogeneous conditions, roots in saline media took up ions, resulting in higher shoot Na⁺ and Cl^- concentrations compared with plants growing in low-salt media.

Conclusions

Results from split-root experiments complement knowledge of plant responses to uniform salinities; the next challenge is to develop new protocols so that this understanding can be extrapolated to more complex soil- and field-based systems. More work is also required to understand the physiological mechanisms underlying changes in stomatal conductance and shoot ion regulation in plants under heterogeneous salinities and how these are linked to the saline parts of the root-zone.     

The shoot controls zeatin riboside export from pea roots. Evidence from the branching mutant rms4

The rms4 mutant of pea ( Pisum sativum L.) was used in grafting studies and cytokinin analyses of the root xylem sap to provide evidence that, at least for pea, the shoot can modify the import of cytokinins from the root. The rms4 mutation, which confers a phenotype with increased branching in the shoot, causes a very substantial decrease (down to 40-fold less) in the concentration of zeatin riboside (ZR) in the xylem sap of the roots. Results from grafts between wild-type (WT) and rms4 plants indicate that the concentration of cytokinins in the xylem sap of the roots is determined almost entirely by the genotype of the shoot. WT scions normalize the cytokinin concentration in the sap of rms4 mutant roots, whereas mutant scions cause WT roots to behave like those of self-grafted mutant plants. The mechanism whereby rms4 shoots of pea cause a down-regulation in the export of cytokinins from the roots is unknown at this time. However, our data provide evidence that the shoot transmits a signal to the roots and thereby controls processes involved in the regulation of cytokinin biosynthesis in the root.     

Role of roots in regulating the growth rate and cytokinin content in leaves

The role of roots in the enhancement of cytokinin content and leaf growth of Phaseolus vulgaris plants after decapitation and partial defoliation was investigated. Partial excision of the roots of plants which were decapitated above the primary leaf node resulted in a reduction of leaf growth and soluble proteins accumulation in the primary leaves. Roots excision was done at time of decapitation and repeated 8 days later. Endogenous cytokinins, known to be involved in enhancing shoot growth, accumulated in the leaves and stems of decapitated and partially defoliated plants. Lower levels of cytokinins were detected in the leaves of decapitated plants with only a partial root system. The level of cytokinins in the roots of decapitated plants was reduced by partial root excision. The growth and accumulation of cytokinins in leaves were, however, not totally suppressed by removing a large proportion of the roots. At the commencement of the experiment the stem had a higher cytokinin content than both the leaves and roots. This suggests that the stem could be an alternative source of cytokinins to the leaves. The cytokinin complement in the leaves of decapitated plants is not identical to that in the roots. It appears that cytokinins supplied by the roots are metabolized in the leaves, or that alternatively certain cytokinins are synthesized in the leaves themselves.     

Involvement of plant growth substances in the alteration of leaf gas exchange of flooded tomato plants

Ethylene, abscisic acid, and cytokinins were tested for their ability to either induce or prevent the changes which occur in gas exchange characteristics of tomato (Lycopersicon esculentum Mill. cv. Rheinlands Ruhm) leaves during short-term soil flooding. Ethylene, which increases in the shoots of flooded plants, had no effect on stomatal conductance or photosynthetic capacity of drained plants. Abscisic acid, which also accumulates in the shoots of flooded plants, could reproduce the stomatal behavior of flooded plants when sprayed on the leaves of drained plants. However, photosynthetic capacity of drained plants was unaffected by abscisic acid sprays. Cytokinin export from the roots to the shoots declines in flooded plants. Spray applications of benzyladenine increased stomatal conductance in both flooded and drained plants. In addition, the decline in photosynthetic capacity during flooding was largely prevented by supplementary cytokinin applications. The possible involvement of these growth substances in modifying leaf gas exchange during flooding is discussed.     

Quantification of the export of cytokinins from roots to shoots of Rosa hybrida and their degradation rate in the shoot

Cytokinins from the roots may be involved in regulating rose ( Rosa hybrida ) shoot growth and development. The objective of this study was to estimate the export of cytokinins from the roots and their degradation rate in the shoot, which were expected to be correlated with plant development. Hence, the total cytokinin content of the shoot, the concentration of zeatin riboside (ZR) in bleeding sap, and the transpiration rates in three stages of development were determined. The estimations performed are based on the assumption that the cytokinin concentration in bleeding sap is representative for the cytokinin concentration in xylem sap in situ. This was verified by comparing the ZR concentration in bleeding sap and in sap obtaíned after pressurizing the root system to a level equivalent to the leaf water potential; no significant differences could be found. The import of cytokinins could not be correlated with plant development, as it increased linearly with time. The estimated relative degradation rate of cytokinins in the shoot decreased as the plants matured. The half-life of cytokinins in the shoot was found to be approximately 1 day, indicating that cytokinins are rapidly metabolized in the shoot.     

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.     

Root-synthesized cytokinins improve shoot growth and fruit yield in salinized tomato (Solanum lycopersicum L.) plants

Salinity limits crop productivity, in part by decreasing shoot concentrations of the growth-promoting and senescence-delaying hormones cytokinins. Since constitutive cytokinin overproduction may have pleiotropic effects on plant development, two approaches assessed whether specific root-localized transgenic IPT (a key enzyme for cytokinin biosynthesis) gene expression could substantially improve tomato plant growth and yield under salinity: transient root IPT induction (HSP70::IPT) and grafting wild-type (WT) shoots onto a constitutive IPT-expressing rootstock (WT/35S::IPT). Transient root IPT induction increased root, xylem sap, and leaf bioactive cytokinin concentrations 2- to 3-fold without shoot IPT gene expression. Although IPT induction reduced root biomass (by 15%) in control (non-salinized) plants, in salinized plants (100?mM NaCl for 22?d), increased cytokinin concentrations delayed stomatal closure and leaf senescence and almost doubled shoot growth (compared with WT plants), with concomitant increases in the essential nutrient K(+) (20%) and decreases in the toxic ion Na(+) (by 30%) and abscisic acid (by 20-40%) concentrations in transpiring mature leaves. Similarly, WT/35S::IPT plants (scion/rootstock) grown with 75?mM NaCl for 90?d had higher fruit trans-zeatin concentrations (1.5- to 2-fold) and yielded 30% more than WT/non-transformed plants. Enhancing root cytokinin synthesis modified both shoot hormonal and ionic status, thus ameliorating salinity-induced decreases in growth and yield.     

The role of adenine in the synthesis of cytokinins in tomato plants and in cell-free root extracts

The incorporation of labelled adenine into cytokinin-like compounds was investigated in intact tomato plants, decapitated tomato roots and cell-free root extracts. In all three cases evidence was found for the incorporation of adenine into endogenous cytokinins. In intact plants and decapitated root systems no evidence was found for the incorporation into cytokinin nucleotides. Cytokinin bases and nucleosides were however, labelled. In the cell-free root extract there was some evidence for the incorporation of labelled adenine into cytokinin nucleotides. This suggests that the biosynthetic process may be strictly compartmentalized. The present results provide no evidence for the relative importance of cytokinin nucleotides in the biosynthetic process. What is clear is that the rate of adenine incorporation into cytokinins is extremely low and that only a small proportion of the cytokinins which became labelled were exported to the shoot via the root exudate.The financial support of the CSIR/Israel Collaborative Programme is gratefully acknowledged.     

Comparative effects of NaCl and polyethylene glycol on root distribution,growth, and stomatal conductance of sour orange seedlings

An experiment was conducted to study sour orange (Citrus aurantium L.) seedling root density, distribution, and morphological development under NaCl and polyethylene glycol (PEG) stresses in relation to shoot growth and stomatal conductance. Plants were treated with 2 stress levels (– 0.12 and – 0.24 MPa) of NaCl and PEG 4000 for 7 months. Root observation chambers were used to monitor root growth and distribution under stressed and non-stressed conditions. Seedlings receiving NaCl or PEG treatments produced fewer roots and shallower root systems with 46 to 65% of the roots occurring in the top portion of the soil. Fibrous root weight per unit length was increased by 24 to 30% by PEG but was not significantly increased by NaCl.Root growth rate usually alternated with shoot growth in a 2-month cycle. This alternating pattern was not shifted by NaCl and PEG stresses. In all NaCl and PEG treatments, growth was depressed and stomatal conductance was reduced. Compared to controls, plants that received NaCl or PEG had smaller shoot and root dry weights, fewer leaves, shorter height, and fewer roots. Sodium chloride usually caused less damage than PEG to sour orange seedlings suggesting that NaCl and PEG acted through different mechanisms.Florida Agricultural Experiment Station Journal Series No. 9941.     

Functional characterisation of a cytokinin oxidase gene DSCKX1 in Dendrobium orchid

Cytokinin oxidase plays an important role in the cytokinin regulatory processes. We have cloned a novel putative cytokinin oxidase, DSCKX1 (D endrobium Sonia cytokinin oxidase), by mRNA differential display from shoot apices of Dendrobium Sonia cultured in the presence of BA. The DSCKX1 gene appears to have three alternative splicing forms and its expression of DSCKX1 was induced in a tissue-specific manner by cytokinins. In transgenic orchid plants overexpressing DSCKX1, the elevated level of cytokinin oxidase activity was accompanied by a reduction of cytokinin content. These plants exhibited slow shoot growth with numerous and long roots in vitro. Their calli also showed decreased capability of shoot formation. Conversly, antisense transgenic plants showed rapid proliferation of shoots and inhibition of root growth combined with a higher endogenous cytokinin content than wild-type plants. Thus DSCKX1 appears to play an important role on cytokinin metabolism and the related developmental programmes in orchid.     

Exogenous zeatin accumulation in wheat root cells shows its role in regulation of cytokinin transport

Here we have shown that 24 hours after addition of zeatin to the nutrient solution the cytokinin content in xylem sap of wheat plants appears to be about 10 times lower that in the nutrient solution. Cytokinins accumulated mostly in roots and not in shoots of treated plants. These data demonstrate the existence of some barrier on cytokinin pathway from the nutrient solution to the plant shoot. With the help of Sudan III an increase in lignin and suberin deposition in the endodermis could be detected, being stronger with the increase in the distance from the root tip. The increase in deposition of suberin and lignin coincided with the decrease in cytokinin immunolabeling in root cells revealed with the help of monoclonal cytokinin antibodies and the second gold-labelled antibodies. Simultaneously exogenous cytokinins accumulated in root stele cells showing that the Casparian band was not only barrier on cytokinin pathway to plant shoot. It is concluded that high cytokinin immunolabe ling in the stele parenchyma cells around the stele vessels demonstrated accumulation of cytokinins by these cells, which could be important in regulation of cytokinin loading to the xylem vessels during there transport to the shoot. The role of cytokinin transporters is discussed.     

A Negative Hydraulic Message from Oxygen-Deficient Roots of Tomato Plants? (Influence of Soil Flooding on Leaf Water Potential,Leaf Expansion,and Synchrony between Stomatal Conductance and Root Hydraulic Conductivity)

Four to 10 h of soil flooding delayed and suppressed the normal daily increase in root hydraulic conductance (Lp) in tomato (Lycopersicon esculentum Mill. cv Ailsa Craig) plants. The resulting short-term loss of synchrony between Lp and stomatal conductance decreased leaf water potential ([psi]L) relative to well-drained plants within 2 h. A decrease in [psi]L persisted for 8 h and was mirrored by decreased leaf thickness measured using linear displacement transducers. After 10 h of flooding, further closing of stomata and re-convergence of Lp in flooded and well-drained roots returned [psi]L to control values. In the second photoperiod, Lp in flooded plants exceeded that in well-drained plants in association with much increased Lp and decreased stomatal conductance. Pneumatic balancing pressure applied to roots of intact flooded plants to prevent temporary loss of [psi]L in the 1st d did not modify the patterns of stomatal closure or leaf expansion. Thus, the magnitude of the early negative hydraulic message was neither sufficient nor necessary to promote stomatal closure and inhibit leaf growth in flooded tomato plants. Chemical messages are presumed to be responsible for these early responses to soil flooding.     

A model of water flow through plants incorporating shoot/root 'message' control of stomatal conductance

Abstract. A model of water flow from the soil into the plant, and from the plant to the atmosphere is described. There are three state variables in the model: the soil, root and shoot water contents. The flow rate of water from the soil to the root is calculated by dividing the gradient in water potential by a resistance, comprising the resistance from the bulk soil to the root surface, and that from the root surface to the root interior. The resistance in the soil depends on the soil hydraulic conductivity, which in turn depends on the soil water potential. The flow rate from the root to the shoot is given by the gradient in water potential divided by a resistance, which depends on the structural dry mass of the plant. Transpiration is described by the Penman-Monteith equation. The plant water characteristics can be modified to take account of osmotic and cell wall rigidity parameters. The model incorporates the concept of shoot/root ‘messages’ of water stress, which influence stomatal conductance. The message works through the generation of a hormone as the pressure potential in the shoot (mesophyll) or root falls. This hormone induces a shift of osmoticum from the guard cells to the surrounding mesophyll cells, which causes an increase (i.e. closer to zero) in the osmotic potential in these cells. This, in turn, causes a decrease in their pressure potential, and so reduces stomatal conductance. The model is used as a framework to address some of the issues that have recently been raised concerning the role of water potential in describing water flow through plants. We conclude that, with the hormone present, there is unlikely to be a unique relationship between stomatal conductance and shoot total water potential, since stomatal conductance depends on the pressure potential in the guard cells, which may differ from that in other cells. Nevertheless, this does not imply that water potential is not an important, and indeed fundamental, component for describing water flow through plants. Other aspects of water flow through plants are also considered, such as diurnal patterns of shoot, root and soil water potential components. It is seen that these may differ from the commonly held view that, as the soil dries down, they all attain the same values during the dark period, and which, as we show, is largely unsubstantiated either theoretically or experimentally.     

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     

Effects of manipulation of water and nitrogen regime on the water relations of the desert shrub Larrea tridentata

Summary Water and nitrogen regimes of Larrea tridentata shrubs growing in the field were manipulated during an annual cycle. Patterns of leaf water status, leaf water relations characteristics, and stomatal behavior were followed concurrently. Large variations in leaf water status in both irrigated and nonirrigated individuals were observed. Predawn and midday leaf water potentials of nonirrigated shrubs were lowest except when measurements had been preceded by significant rainfall. Despite the large seasonal variation in leaf water status, reasonably constant, high levels of turgor were maintained. Pressure-volume curve analysis suggested that changes in the bulk leaf osmotic potential at full turgor were small and that nearly all of the turgor adjustment was due to tissue elastic adjustment. The increase in tissue elasticity with increasing water deficit manifested itself as a decrease in the relative water content at zero turgor and as a decrease in the tissue bulk elastic modulus. Because of large hydration-induced displacement in the osmotic potential and relative water content at zero turgor, it was necessary to use shoots in their natural state of hydration for pressure-volume curve determinations. Large diurnal and seasonal differences in maximum stomatal conductance were observed, but could not easily be attributed to variations in leaf water potential or leaf water relations characteristics such as the turgor loss point. The single factor which seemed to account for most of the diurnal and seasonal differences in maximum stomatal conductance between individual shrubs was an index of soil/root/ shoot hydraulic resistance. Daily maximum stomatal conductance was found to decrease with increasing soil/root/ shoot hydraulic resistance. This pattern was most consistent if the hydraulic resistance calculation was based on an estimate of total canopy transpiration rather than the more commonly used transpiration per unit leaf area. The reasons for this are discussed. It is suggested that while stomatal aperture necessarily represents a major physical resistance controlling transpiration, plant hydraulic resistance may represent the functional resistance through its effects on stomatal aperture.     

Hormonal changes induced by partial rootzone drying of irrigated grapevine

Partial rootzone drying (PRD) is a new irrigation technique which improves the water use efficiency (by up to 50%) of wine grape production without significant crop reduction. The technique was developed on the basis of knowledge of the mechanisms controlling transpiration and requires that approximately half of the root system is always maintained in a dry or drying state while the remainder of the root system is irrigated. The wetted and dried sides of the root system are alternated on a 10-14 d cycle. Abscisic acid (ABA) concentration in the drying roots increases 10-fold, but ABA concentration in leaves of grapevines under PRD only increased by 60% compared with a fully irrigated control. Stomatal conductance of vines under PRD irrigation was significantly reduced when compared with vines receiving water to the entire root system. Grapevines from which water was withheld from the entire root system, on the other hand, show a similar reduction in stomatal conductance, but leaf ABA increased 5-fold compared with the fully irrigated control. PRD results in increased xylem sap ABA concentration and increased xylem sap pH, both of which are likely to result in a reduction in stomatal conductance. In addition, there was a reduction in zeatin and zeatin-riboside concentrations in roots, shoot tips and buds of 60, 50 and 70%, respectively, and this may contribute to the reduction in shoot growth and intensified apical dominance of vines under PRD irrigation. There is a nocturnal net flux of water from wetter roots to the roots in dry soil and this may assist in the distribution of chemical signals necessary to sustain the PRD effect. It was concluded that a major effect of PRD is the production of chemical signals in drying roots that are transported to the leaves where they bring about a reduction in stomatal conductance.