Physiological responses of potato (Solanum tuberosum L.) to partial root-zone drying: ABA signalling, leaf gas exchange, and water use efficiency

The physiological responses of potato (Solanum tuberosum L. cv. Folva) to partial root-zone drying (PRD) were investigated in potted plants in a greenhouse (GH) and in plants grown in the field under an automatic rain-out-shelter. In the GH, irrigation was applied daily to the whole root system (FI), or to one-half of the root system while the other half was dried, for 9 d. In the field, the plants were drip irrigated either to the whole root system near field capacity (FI) or using 70% water of FI to one side of the roots, and shifted to the other side every 5-10 d (PRD). PRD plants had a similar midday leaf water potential to that of FI, whereas in the GH their root water potential (Psi(r)) was significantly lowered after 5 d. Stomatal conductance (g(s)) was more sensitive to PRD than photosynthesis (A) particularly in the field, leading to greater intrinsic water use efficiency (WUE) (i.e. A/g(s)) in PRD than in FI plants on several days. In PRD, the xylem sap abscisic acid concentration ([ABA](xylem)) increased exponentially with decreasing Psi(r); and the relative [ABA](xylem) (PRD/FI) increased exponentially as the fraction of transpirable soil water (FTSW) in the drying side decreased. In the field, the leaf area index was slightly less in PRD than in FI treatment, while tuber biomass was similar for the two treatments. Compared with FI, PRD treatment saved 30% water and increased crop water use efficiency (WUE) by 59%. Restrictions on leaf area expansion and g(s) by PRD-induced ABA signals might have contributed to reduced water use and increased WUE.     

Water relations,photosynthesis, growth and water-use efficiency in tomato plants subjected to partial rootzone drying and regulated deficit irrigation

Abstract

Partial rootzone drying (PRD) and regulated deficit irrigation (RDI) are water-saving irrigation systems that have been developed to increase water-use efficiency (WUE) without significant yield reduction. In order to investigate whether a high-value horticultural crop such as tomato responded differently to RDI and PRD, we compared the physiological and growth responses of tomato plants using a split-root system. Plants were grown in a greenhouse under controlled conditions with their roots separated equally between two soil compartments. Three irrigation treatments were imposed: (i) Control, receiving an amount of water equivalent to 100% of plant transpiration; (ii) PRD, in which one compartment was watered with 50% of the amount of water supplied to the controls, allowing one-half of the root system to be exposed to dry soil and switching irrigation between sides weekly; and (iii) RDI, in which 50% of the amount of water given to the controls was supplied, half to each side of the root system. Leaf RWC and midday leaf Ψ decreased substantially in RDI-treated plants, while the PRD plants exhibited relatively higher Ψ and RWC values. Both PRD and RDI treatments reduced by about 30% the total plant dry mass compared with the control. However, plant transpiration was reduced by about 50% in both PRD and RDI, allowing a significant improvement in whole-plant WUE. Stomatal conductance (Gs) and leaf growth were also significantly reduced by PRD and RDI. These results may be related to a significant increase in xylem sap pH and leaf apoplastic pH. Generally, the photosynthetic apparatus of tomato leaves had a high resistance to restricted water availability. In fact, the decreased Gs had no major negative impact on carbon assimilation. However, V _cmax, i.e. Rubisco efficiency, was significantly decreased in RDI plants with respect to control ones. This may imply that, although the differences between the PRD and RDI treatments in our study were subtle, they may become more marked with a more prolonged and severe water deficit.     

Accounting for sap flow from different parts of the root system improves the prediction of xylem ABA concentration in plants grown with heterogeneous soil moisture

When soil moisture is heterogeneous, sap flow from, and ABA status of, different parts of the root system impact on leaf xylem ABA concentration ([X-ABA]leaf). The robustness of a model for predicting [X-ABA]leaf was assessed. 'Two root-one shoot' grafted sunflower (Helianthus annuus L.) plants received either deficit irrigation (DI, each root system received the same irrigation volumes) or partial rootzone drying (PRD, only one root system was watered and the other dried the soil). Irrespective of whether relative sap flow was assessed using sap flow sensors in vivo or by pressurization of de-topped roots, each root system contributed similarly to total sap flow during DI, while sap flow from roots in drying soil declined linearly with soil water potential (Psisoil) during PRD. Although Psisoil of the irrigated pot determined the threshold Psisoil at which sap flow from roots in drying soil decreased, the slope of this decrease was independent of the wet pot Psisoil. Irrespective of whether sap was collected from the wet or dry root system of PRD plants, or a DI plant, root xylem ABA concentration increased as Psisoil declined. The model, which weighted ABA contributions of each root system according to the sap flow from each, almost perfectly explained [X-ABA] immediately above the graft union. That the model overestimated measured [X-ABA]leaf may result from changes in [X-ABA] along the transport pathway or an artefact of collecting xylem sap from detached leaves. The implications of declining sap flow through partially dry roots during PRD for the control of stomatal behaviour and irrigation scheduling are discussed.     

Daily irrigation attenuates xylem abscisic acid concentration and increases leaf water potential of Pelargonium × hortorum compared with infrequent irrigation

The physiological response of plants to different irrigation frequencies may affect plant growth and water use efficiency (WUE; defined as shoot biomass/cumulative irrigation). Glasshouse‐grown, containerized Pelargonium × hortorum BullsEye plants were irrigated either daily at 100% of plant evapotranspiration (ET) (well‐watered; WW), or at 50% ET applied either daily [frequent deficit irrigation (FDI)] or cumulatively every 4 days [infrequent deficit irrigation (IDI)], for 24 days. Both FDI and IDI applied the same irrigation volume. Xylem sap was collected from the leaves, and stomatal conductance (g_s) and leaf water potential (Ψ_leaf) measured every 2 days. As soil moisture decreased, g_s decreased similarly under both FDI and IDI throughout the experiment. Ψ_leaf was maintained under IDI and increased under FDI. Leaf xylem abscisic acid (ABA) concentrations ([X‐ABA]_leaf) increased as soil moisture decreased under both IDI and FDI, and was strongly correlated with decreased g_s, but [X‐ABA]_leaf was attenuated under FDI throughout the experiment (at the same level of soil moisture as IDI plants). These physiological changes corresponded with differences in plant production. Both FDI and IDI decreased growth compared with WW plants, and by the end of the experiment, FDI plants also had a greater shoot fresh weight (18%) than IDI plants. Although both IDI and FDI had higher WUE than WW plants during the first 10 days of the experiment (when biomass did not differ between treatments), the deficit irrigation treatments had lower WUE than WW plants in the latter stages when growth was limited. Thus, ABA‐induced stomatal closure may not always translate to increased WUE (at the whole plant level) if vegetative growth shows a similar sensitivity to soil drying, and growers must adapt their irrigation scheduling according to crop requirements.     

Abscisic acid signalling when soil moisture is heterogeneous: decreased photoperiod sap flow from drying roots limits abscisic acid export to the shoots

To investigate the contribution of different parts of the root system to total sap flow and leaf xylem abscisic acid (ABA) concentration ([X-ABA]_leaf), individual sunflower ( Helianthus annuus L.) shoots were grafted onto the root systems of two plants grown in separate pots and sap flow through each hypocotyl measured below the graft union. During deficit irrigation (DI), both pots received the same irrigation volumes, while during partial root zone drying (PRD) one pot ('wet') was watered and another ('dry') was not. During PRD, once soil water content ( θ ) decreased below a threshold, the fraction of sap flow from drying roots declined. As θ declined, root xylem ABA concentration increased in both irrigation treatments, and [X-ABA]_leaf increased in DI plants, but [X-ABA]_leaf of PRD plants actually decreased within a certain θ range. A simple model that weighted ABA contributions of wet and dry root systems to [X-ABA]_leaf according to the sap flow from each, better predicted [X-ABA]_leaf of PRD plants than either [X-ABA]_dry, [X-ABA]_wet or their mean. Model simulations revealed that [X-ABA]_leaf during PRD exceeded that of DI with moderate soil drying, but continued soil drying (such that sap flow from roots in drying soil ceased) resulted in the opposite effect.     

Does engineering abscisic acid biosynthesis in Nicotiana plumbaginifolia modify stomatal response to drought?

The consequences of manipulating abscisic acid (ABA) biosynthesis rates on stomatal response to drought were analysed in wild‐type, a full‐deficient mutant and four under‐producing transgenic lines of N. plumbaginifolia. The roles of ABA, xylem sap pH and leaf water potential were investigated under four experimental conditions: feeding detached leaves with varying ABA concentration; injecting exogenous ABA into well‐watered plants; and withholding irrigation on pot‐grown plants, either intact or grafted onto tobacco. Changes in ABA synthesis abilities among lines did not affect stomatal sensitivity to ABA concentration in the leaf xylem sap ([ABA]_xyl), as evidenced with exogenous ABA supplies and natural increases of [ABA]_xyl in grafted plants subjected to drought. The ABA‐deficient mutant, which is uncultivable under normal evaporative demand, was grafted onto tobacco stock and then presented the same stomatal response to [ABA]_xyl as wild‐type and other lines. This reinforces the dominant role of ABA in controlling stomatal response to drought in N. plumbaginifolia whereas roles of leaf water potential and xylem sap pH were excluded under all studied conditions. However, when plants were submitted to soil drying onto their own roots, stomatal response to [ABA]_xyl slightly differed among lines. It is suggested, consistently with all the results, that an additional root signal of soil drying modulates stomatal response to [ABA]_xyl.     

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.

     

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.     

Potential of partial rootzone drying as an alternative irrigation technique for potatoes (Solanum tuberosum)

The increasing demands on limited water supplies worldwide require the adoption of more efficient irrigation techniques for sustainable production in agriculture. Partial rootzone drying (PRD) is one of the techniques that offer potential saving of irrigation water. This technique involves alternate irrigation to two sides of a plant root system. The studies reported here investigated PRD irrigation regimes and the optimum time of starting PRD in potatoes grown in a protected environment. In the first experiment, plants of the potato cv. Estima were exposed to five different irrigation treatments and a fully watered control at tuber initiation. The treatment that performed most similar to the control was alternate PRD to field capacity (APRD₁₀₀). This treatment produced similar total leaf area, haulm fresh and dry weights, plant water status and no significant yield reduction compared with the control plants. The APRD₁₀₀ treatment utilised 29% less water and increased water use efficiency (WUE) by 19%. In the second experiment, the APRD₁₀₀ irrigation was started at 2, 4, 6, 8 and 10 weeks after plant emergence. Vegetative growth and yield increased with the delay of the APRD₁₀₀. APRD₁₀₀ started at 6 weeks after emergence did not significantly reduce fresh tuber yield but received 21% less total water with a 19% increase in WUE. The results indicate that PRD may have potential use in the potato crop for conserving irrigation water with minimal loss of yield.     

Growth response of barley and tomato to nitrogen stress and its control by abscisic acid,water relations and photosynthesis

Barley (Hordeum vulgare L.) and tomato Lycopersicon esculentum Mill.) were grown hydroponically and examined 2, 5, and 10 d after being deprived of nitrogen (N) supply. Leaf elongation rate declined in both species in response to N stress before there was any reduction in rate of dryweight accumulation. Changes in water transport to the shoot could not explain reduced leaf elongation in tomato because leaf water content and water potential were unaffected by N stress at the time leaf elongation began to decline. Tomato maintained its shoot water status in N-stressed plants, despite reduced water absorption per gram root, because the decline in root hydraulic conductance with N stress was matched by a decline in stomatal conductance. In barley the decline in leaf elongation coincided with a small (8%) decline in water content per unit area of young leaves; this decline occurred because root hydraulic conductance was reduced more strongly by N stress than was stomatal conductance. Nitrogen stress caused a rapid decline in tissue NO ₃ ^- pools and in NO ₃ ^- flux to the xylem, particularly in tomato which had smaller tissue NO ₃ ^- reserves. Even in barley, tissue NO ₃ ^- reserves were too small and were mobilized too slowly (60% in 2 d) to support maximal growth for more than a few hours. Organic N mobilized from old leaves provided an additional N source to support continued growth of N-stressed plants. Abscisic acid (ABA) levels increased in leaves of both species within 2 d in response to N stress. Addition of ABA to roots caused an increase in volume of xylem exudate but had no effect upon NO ₃ ^- flux to the xylem. After leaf-elongation rate had been reduced by N stress, photosynthesis declined in both barley and tomato. This decline was associated with increased leaf ABA content, reduced stomatal conductance and a decrease in organic N content. We suggest that N stress reduces growth by several mechanisms operating on different time scales: (1) increased leaf ABA content causing reduced cell-wall extensibility and leaf elongation and (2) a more gradual decline in photosynthesis caused by ABA-induced stomatal closure and by a decrease in leaf organic N.Abbreviation and symbols ABA abscisic acid - c_i leaf internal CO₂ concentration - L_p root hydraulic conductance     

Root-to-shoot signalling when soil moisture is heterogeneous: increasing the proportion of root biomass in drying soil inhibits leaf growth and increases leaf abscisic acid concentration

To determine whether root-to-shoot signalling of soil moisture heterogeneity depended on root distribution, wild-type (WT) and abscisic acid (ABA)-deficient (Az34) barley (Hordeum vulgare) plants were grown in split pots into which different numbers of seminal roots were inserted. After establishment, all plants received the same irrigation volumes, with one pot watered (w) and the other allowed to dry the soil (d), imposing three treatments (1 d: 3 w, 2 d: 2 w, 3 d: 1 w) that differed in the number of seminal roots exposed to drying soil. Root distribution did not affect leaf water relations and had no sustained effect on plant evapotranspiration (ET). In both genotypes, leaf elongation was less and leaf ABA concentrations were higher in plants with more roots in drying soil, with leaf ABA concentrations and water potentials 30% and 0.2 MPa higher, respectively, in WT plants. Whole-pot soil drying increased xylem ABA concentrations, but maximum values obtained when leaf growth had virtually ceased (100 nm in Az34, 330 nm in WT) had minimal effects (<40% leaf growth inhibition) when xylem supplied to detached shoots. Although ABA may not regulate leaf growth in vivo, genetic variation in foliar ABA concentration in the field may indicate different root distributions between upper (drier) and lower (wetter) soil layers.     

Contrasting physiological effects of partial root zone drying in field-grown grapevine (Vitis vinifera L. cv. Monastrell) according to total soil water availability

Different spatial distributions of soil moisture were imposed on field-grown grapevines by applying the same irrigation volumes to the entire (DI; deficit irrigation) or part of the (PRD; partial root zone drying) root zone. Five treatments were applied: controls irrigated at 60% ETc (crop evapotranspiration) for the whole season (308 mm year(-1)); DI-1 and PRD-1 that received the same irrigation as controls before fruit set, 30% ETc from fruit set to harvest and 45% ETc post-harvest (192 mm year(-1)); and DI-2 and PRD-2 that were the same, except that 15% ETc was applied from fruit set to harvest (142 mm year(-1)). Compared with DI-1, PRD-1 maintained higher leaf area post-veraison and increased root water uptake, whole-plant hydraulic conductance, leaf transpiration, stomatal conductance, and photosynthesis, but decreased intrinsic gas exchange efficiency without causing differences in leaf xylem abscisic acid (ABA) concentration. Compared with DI-2, PRD-2 increased leaf xylem ABA concentration and decreased root water uptake, whole-plant hydraulic conductance, leaf transpiration, stomatal conductance, and photosynthesis, mainly at the beginning of PRD cycles. Distinctive PRD effects (e.g. greater stomatal closure) depended on the volumetric soil water content of the wet root zone, as predicted from a model of root-to-shoot ABA signalling.     

Reduction, assimilation and transport of N in normal and gibberellin-deficient tomato plants

A fast-growing normal and a slow-growing gibberellin-deficient mutant of Lycopersicon esculentum (L.) Mill. cv. Moneymaker were used to test the hypothesis that slow-growing plants reduce NO₃^? in the root to a greater extent than do fast-growing plants. Plants that reduce NO₃^? in the root may grow more slowly due to the higher energetic and carbon costs associated with root-based NO₃^? reduction compared to photosynthetically driven shoot NO₃^? reduction. The plants were grown hydroponically with a complete nutrient solution containing 10 mM NO₃^? and the biomass production, gas exchange characteristics, root respiratory O₂ consumption, nitrate reductase activity and translocation of N in the xylem were measured. The gibberellin-deficient mutants accumulated more total N unit^?1 dry weight than did the faster-growing normal plants. There were no significant differences between the genotypes in the rates of photosynthesis expressed on a leaf dry weight basis. The plants differed in the proportion of photosynthetic carbon available to growth due to a greater proportion of daily photo-synthate production being consumed by respiration in the slow-growing genotype. This difference in allocation of carbon was associated with differences in the specific leaf area and specific root length. In addition, a greater leaf weight ratio in the fast-growing than in the slow-growing plants indicates a greater investment of carbon into biomass supporting photosynthetic production in the former. We did not find differences in the activity or distribution of nitrate reductase or in the N composition of the xylem sap between the genotypes. We thus conclude that the growth rate was determined by the efficiency of carbon partitioning and that the site of NO₃^? reduction and assimilation was not related to the growth rate of these plants.     

Can stomatal closure caused by xylem ABA explain the inhibition of leaf photosynthesis under soil drying?

Effects of leaf water deficit and increase in endogenous ABA on photosynthesis of two tropical trees, t Acacia confusa and t Leucaena leucocephala, were investigated with two soil-drying methods, i.e. half or whole root system was subjected to soil drying. Half-root drying was achieved by allowing upper layer of soil column to dry and lower layer of soil column to remain watered. Half-root drying had little effect on leaf water potential, but when compared to the well-watered control, both methods of soil drying substantially increased the ABA concentration in xylem and reduced leaf conductance in both species. There was a significant relationship between leaf conductance and xylem ABA concentrations in both species, which was comparable to the same relationship that was generated by feeding ABA to excised twigs. The rate of photosynthesis was inhibited substantially in both soil-drying treatments and in both species, but photochchemical efficiency, measured as a ratio of variable fluorescence to a peak fluorescence emission of a dark-adapted leaf (F_v/F_m), was not reduced except in the whole root-dried t L. leucocephala plants where leaf water potential was reduced to –2.5 MPa. In all the cases where photosynthesis was inhibited, there was a concomitant reduction in both leaf conductance and calculated internal CO₂ concentration. After two days of rewatering, leaf water potential and xylem ABA concentration rapidly returned to pre-treatment levels, but leaf conductance and photosynthesis of both whole-root and half root dried t L. leucocephala remained inhibited substantially. Rewatering led to a full recovery of both stomatal conductance and photosynthesis in soil-dried t A. confusa, although its photosynthesis of whole-root dried plants did not recover fully but such difference was not significant statistically. These results suggest that drought-induced decline of photosynthesis was mainly a result of the stomatal factor caused by the increase of ABA concentration in the xylem sap. Non-stomatal factors, e.g. reduced photochemical activity and/or carbon metabolic activity, were species-specific and were brought about only at very low water potential.     

Inhibition of tomato shoot growth by over‐irrigation is linked to nitrogen deficiency and ethylene

Although physiological effects of acute flooding have been well studied, chronic effects of suboptimal soil aeration caused by over‐irrigation of containerized plants have not, despite its likely commercial significance. By automatically scheduling irrigation according to soil moisture thresholds, effects of over‐irrigation on soil properties (oxygen concentration, temperature and moisture), leaf growth, gas exchange, phytohormone [abscisic acid (ABA) and ethylene] relations and nutrient status of tomato (Solanum lycopersicum Mill. cv. Ailsa Craig) were studied. Over‐irrigation slowly increased soil moisture and decreased soil oxygen concentration by 4%. Soil temperature was approximately 1°C lower in the over‐irrigated substrate. Over‐irrigating tomato plants for 2 weeks significantly reduced shoot height (by 25%) and fresh weight and total leaf area (by 60–70%) compared with well‐drained plants. Over‐irrigation did not alter stomatal conductance, leaf water potential or foliar ABA concentrations, suggesting that growth inhibition was not hydraulically regulated or dependent on stomatal closure or changes in ABA. However, over‐irrigation significantly increased foliar ethylene emission. Ethylene seemed to inhibit growth, as the partially ethylene‐insensitive genotype Never ripe (Nr) was much less sensitive to over‐irrigation than the wild type. Over‐irrigation induced significant foliar nitrogen deficiency and daily supplementation of small volumes of 10 mM Ca(NO₃)₂ to over‐irrigated soil restored foliar nitrogen concentrations, ethylene emission and shoot fresh weight of over‐irrigated plants to control levels. Thus reduced nitrogen uptake plays an important role in inhibiting growth of over‐irrigated plants, in part by stimulating foliar ethylene emission.     

Benefits of alternate partial root-zone irrigation on growth, water and nitrogen use efficiencies modified by fertilization and soil water status in maize

Alternate partial root-zone irrigation (APRI) is a new water-saving technique and may improve crop water use efficiency without much yield reduction. We investigated if the benefits of APRI on biomass accumulation, water and nitrogen use efficiencies could be modified by different soil fertilization and watering levels in pot-grown maize (Zea mays L. cv. super-sweet No 28, a local variety). Three irrigation methods, i.e. conventional irrigation (CI), alternate partial root-zone irrigation (APRI, alternate watering on both sides of the pot) and fixed partial root-zone irrigation (FPRI, fixed watering on one side of the pot), two watering levels, i.e. water deficit (W₁, 45–55% of field capacity) and well-watered (W₂, 70–80% of field capacity), and two N fertilization levels, i.e. no fertilization and fertilization, were designed. Results showed that APRI and FPRI methods led to more reduction in transpiration than in photosynthesis, and thus increased leaf water use efficiency (leaf WUE, i.e. the ratio of leaf net photosynthetic rate to transpiration rate). Compared to the CI treatment, APRI and FPRI increased leaf WUE by 7.7% and 8.1% before the jointing stage and 3.6% and 4.2% during the jointing stage, respectively. Under the fertilization and well-watered conditions, APRI treatment saved irrigation water by 38.4% and reduced shoot and total dry masses by 5.9% and 6.7%, respectively if compared to the CI treatment. APRI also enhanced canopy WUE (defined as the amount of total biomass per unit water used) and nitrogen (N) apparent recovery fraction (Nr, defined as the ratio of the increased N uptake to N applied) by 24.3% and 16.4%, respectively, indicating that effect of APRI can be better materialized under appropriate fertilization and water supply. Responsible Editor: Rana E. Munns     

The relations of stomatal closure and reopening to xylem ABA concentration and leaf water potential during soil drying and rewatering

Two tropical tree species, Acacia confusa and Leucaena leucocephala, were used to study the relationships among stomatal conductance, xylem ABA concentration and leaf water potential during a soil drying and rewatering cycle. Stomatal conductance of both A. confusa and L. leucocephala steadily decreased with the decreases in soil water content and pre-dawn leaf water potential. Upon rewatering, soil water content and pre-dawn leaf water potential rapidly returned to the control levels, whereas the reopening of stomata showed an obvious lag time. The length of this lag time was highly dependent not only upon the degree of water stress but also on plant species. The more severe the water stress, the longer the lag time. When A. confusa and L. leucocephala plants were exposed to the same degree of water stress (around –2.0 MPa in pre-dawn leaf water potential), the stomata of A. confusa reopened to the control level 6 days after rewatering. However, it took L. leucocephala about 14 days to reopen fully. A very similar response of leaf photosynthesis to soil water deficit was also observed for both species. Soil drying resulted in a significant increase in leaf and xylem ABA concentrations in both species. The more severe the water stress, the higher the leaf and xylem ABA concentrations. Both leaf ABA and xylem ABA returned to the control level following relief from water deficit and preceded the full recovery of stomata, suggesting that the lag phase of stomatal reopening was not controlled by leaf and/or xylem ABA. In contrast to drying the whole root system, drying half of the root system did not change the leaf water relations, but caused a significant increase in xylem ABA concentration, which could fully explain the decrease of stomatal conductance. After rewatering, the stomatal conductance of plants in which half of the roots were dried recovered more rapidly than those of whole-root dried plants, indicating that the leaf water deficit that occurred during the drying period was related to the post-stress stomatal inhibition. These results indicated that the decrease in stomatal conductance caused by water deficit was closely related to the increase in xylem ABA, but xylem ABA could not fully explain the reopening of stomata after relief of water stress, neither did the leaf ABA. Some unknown physiological and/or morphological processes in the guard cells may be related to the recovery process.     

Changes in antioxidant activities and phenol content in tomato plants subjected to partial root drying and regulated deficit irrigation

Abstract

Partial rootzone drying (PRD) and regulated deficit irrigation (RDI) are water saving irrigation systems that have been developed to increase water use efficiency (WUE) without significant yield reduction. To examine whether tomato responded differently to RDI and PRD, we compared the changes in antioxidative defenses in tomato plants using a split-root system. Tomato plants were grown for 21 days under controlled conditions with their roots separated equally between two soil compartments. Three irrigation treatments were imposed: Control, receiving an amount of water equivalent to 100% of plant transpiration; PRD in which one compartment was watered with 50% of the amount of water supplied to the controls, allowing one-half of the root system to be exposed to dry soil, and switching irrigation between sides weekly; RDI in which 50% of the amount of water given to the controls was supplied, half to each side of the root system. Relative water content (RWC), midday leaf Ψ and chlorophyll content decreased largely in RDI-treated plants, whereas the PRD plants exhibited relatively higher Ψ and RWC values. An enhanced level of lipid peroxidation in both roots and leaves indicated that PRD and RDI caused oxidative stress in tomato plants. In leaves, superoxide dismutase (SOD), soluble peroxidase (POX) and polyphenol oxidase (PPO) activities showed an increase in the early phase of water deficit, and then decreased in the remaining phase of the drying cycle. However, the increase was more pronounced under RDI. Catalase (CAT) activity declined continuously from the onset of PRD and RDI treatments to below the control level, and the reduction was less under PRD than RDI. POX cell-wall associated activities exceeded the control level by 450% and 230%, respectively, under RDI and PRD. At the root level, while CAT activity also decreased under both PRD and RDI, the activities of SOD, POX and PPO significantly increased and their activities showed an alternating increase/decrease paralleling the alternating irrigation in PRD-treated roots. As a result of the difference in POX and PPO activities between the two water treatments applied, PRD-treated plants accumulated more soluble and cell-wall bound phenolic compounds.     

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.     

Plasticity in stomatal size and density of potato leaves under different irrigation and phosphorus regimes

The morphological features of stomata including their size and density could be modulated by environmental cues; however, the underlying mechanisms remain largely elusive. Here, the effect of different irrigation and phosphorus (P) regimes on stomatal size (SS) and stomatal density (SD) of potato leaves was investigated. The plants were grown in split-root pots under two P fertilization rates (viz., 0 and 100 mg kg⁻¹ soil, denoted as P0 and P1, respectively) and subjected to full (FI), deficit (DI), and partial root-zone drying (PRD) irrigation regimes. Results showed that SS and SD were unresponsive to P but significantly affected by the irrigation treatment. FI plants had the largest SS, followed by DI, and PRD the smallest; and the reverse was the case for SD. Compared to FI and DI, PRD plants had significantly lower values of specific leaf area (SLA) and leaf carbon isotope discrimination (Δ¹³C) under P0. Midday leaf water potential (Ψ_leaf) and stomatal conductance (g_s) was similar for DI and PRD, which was significantly lower than that of FI. Leaf contents of C, N, K, Ca and Mg were higher in PRD than in DI plants, particularly under P0. When analyzed across the three irrigation regimes, it was found that the P1 plants had significantly higher leaf contents of P and Mg, but significantly lower leaf K content compared to the P0 plants. Linear correlation analyses revealed that SS was positively correlated with Ψ_leaf and Δ¹³C; whereas SD was negatively correlated with Ψ_leaf, Δ¹³C and SLA, and positively correlated with leaf C, N and Ca contents. And g_s was positively correlated with SS but negatively correlated with SD. Collectively, under low P level, the smaller and denser stomata in PRD plants may bring about a more efficient stomatal control over gas exchange, hereby potentially enhance water-use efficiency as exemplified by the lowered leaf Δ¹³C under fluctuating soil moisture conditions.