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.     

Abscisic acid triggers whole-plant and fruit-specific mechanisms to increase fruit calcium uptake and prevent blossom end rot development in tomato fruit

Calcium (Ca) uptake into fruit and leaves is dependent on xylemic water movement, and hence presumably driven by transpiration and growth. High leaf transpiration is thought to restrict Ca movement to low-transpiring tomato fruit, which may increase fruit susceptibility to the Ca-deficiency disorder, blossom end rot (BER). The objective of this study was to analyse the effect of reduced leaf transpiration in abscisic acid (ABA)-treated plants on fruit and leaf Ca uptake and BER development. Tomato cultivars Ace 55 (Vf) and AB2 were grown in a greenhouse environment under Ca-deficit conditions and plants were treated weekly after pollination with water (control) or 500 mg l(-1) ABA. BER incidence was completely prevented in the ABA-treated plants and reached values of 30-45% in the water-treated controls. ABA-treated plants had higher stem water potential, lower leaf stomatal conductance, and lower whole-plant water loss than water-treated plants. ABA treatment increased total tissue and apoplastic water-soluble Ca concentrations in the fruit, and decreased Ca concentrations in leaves. In ABA-treated plants, fruit had a higher number of Safranin-O-stained xylem vessels at early stages of growth and development. ABA treatment reduced the phloem/xylem ratio of fruit sap uptake. The results indicate that ABA prevents BER development by increasing fruit Ca uptake, possibly by a combination of whole-plant and fruit-specific mechanisms.     

Chemical signals and their regulations on the plant growth and water use efficiency of cotton seedlings under partial root-zone drying and different nitrogen applications

Partial root-zone drying during irrigation (PRD) has been shown effective in enhancing plant water use efficiency (WUE), however, the roles of chemical signals from root and shoot that are involved and the possible interactions affected by nitrogen nutrition are not clear. Pot-grown cotton (Gossypium spp.) seedlings were treated with three levels of N fertilization and PRD. The concentrations of nitrate (NO₃⁻), abscisic acid (ABA) and the pH value of leaf and root xylem saps, biomass and WUE were measured. Results showed that PRD plants produced larger biomass and higher WUE than non-PRD plants, with significant changes in leaf xylem ABA, leaf and root xylem NO₃⁻ concentrations and pH values, under heterogeneous soil moisture conditions. Simultaneously, high-N treated plants displayed larger changes in leaf xylem ABA and higher root xylem NO₃⁻ concentrations, than in the medium- or low-N treated plants. However, the WUE of plants in the low-N treatment was higher than that of those in the high- and medium-N treatments. PRD and nitrogen levels respectively induced signaling responses of ABA/NO₃⁻ and pH in leaf or root xylem to affect WUE and biomass under different watering levels, although significant interactions of PRD and nitrogen levels were found when these signal molecules responded to soil drying. We conclude that these signaling chemicals are regulated by interaction of PRD and nitrogen status to regulate stomatal behavior, either directly or indirectly, and thus increase PRD plant WUE under less irrigation.     

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.     

根源信号参与调控气孔行为的机制及其农业节水意义

在土壤干旱情况下,根源信号一方面向植物地上部分的长距离传输,为地上部分提供了土壤水分获取能力的测度,另一方面调控气孔开度,抑制蒸腾作用并提高植物的水分利用效率．文中综述了根源信号参与调控植物水分利用的生理机制和理论模型,指出该模型与根系吸水模型、气孔导度模型耦合,能够更好地反映植物叶片对土壤干旱以及大气干旱的响应、评述了在根源信号参与调控植物水分关系的基础上发展的调亏灌溉(RDI)、部分根系干旱(PRD)和控制性交替灌溉(CAI)等有效灌溉手段,有助于合理配置根系层供水量,通过根土相互作用和信号物质的传输,降低蒸腾和提高水分利用效率、另外,根源信号在调控根系生长发育、延缓地上部分生长以调节根冠比例,优化资源分配以利于生殖生长等方面均有所为,为全面提高农田水分利用效率提供节水生理基础。     

Maize stomatal conductance in the field: its relationship with soil and plant water potentials, mechanical constraints and ABA concentration in the xylem sap

Abstract. Stomatal conductance, leaf water potential, soil water potential and concentration of abscisic acid (ABA) in the xylem sap were measured on maize plants growing in the field, in two treatments with contrasting soil structures. Soil compaction affected the stomatal conductance, but this effect was no longer observed if the soil water potential was increased by irrigation. Differences in leaf water potential did not account for the differences in conductance between treatments. Conversely, the relationship between stomatal conductance and concentration of ABA in the xylem sap was consistent during the experiment. The proposed interpretation is that stomatal conductance was controlled by the root water potential via an ABA message. Control of the stomatal conductance by the leaf water potential or by an effect of mechanical stress on the roots is unlikely.     

Modelling the impact of heterogeneous rootzone water distribution on the regulation of transpiration by hormone transport and/or hydraulic pressures

Aims

A simulation model to demonstrate that soil water potential can regulate transpiration, by influencing leaf water potential and/or inducing root production of chemical signals that are transported to the leaves.

Methods

Signalling impacts on the relationship between soil water potential and transpiration were simulated by coupling a 3D model for water flow in soil, into and through roots (Javaux et al. 2008) with a model for xylem transport of chemicals (produced as a function of local root water potential). Stomatal conductance was regulated by simulated leaf water potential (H) and/or foliar chemical signal concentrations (C; H?+?C). Split-root experiments were simulated by varying transpiration demands and irrigation placement.

Results

While regulation of stomatal conductance by chemical transport was unstable and oscillatory, simulated transpiration over time and root water uptake from the two soil compartments were similar for both H and H?+?C regulation. Increased stomatal sensitivity more strongly decreased transpiration, and decreased threshold root water potential (below which a chemical signal is produced) delayed transpiration reduction.

Conclusions

Although simulations with H?+?C regulation qualitatively reproduced transpiration of plants exposed to partial rootzone drying (PRD), long-term effects seemed negligible. Moreover, most transpiration responses to PRD could be explained by hydraulic signalling alone.     

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.     

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.     

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.     

Hydraulic and Chemical Responses of Citrus Seedlings to Drought and Osmotic Stress

In this work we investigated the function of abscisic acid (ABA) as a long-distance chemical signal communicating water shortage from the root to the shoot in citrus plants. Experiments indicated that stomatal conductance, transpiration rates, and leaf water potential decline progressively with drought. ABA content in roots, leaves, and xylem sap was also increased by the drought stress treatment three- to sevenfold. The addition of norflurazon, an inhibitor of ABA biosynthesis, significantly decreased the intensity of the responses and reduced ABA content in roots and xylem fluid, but not in leaves. Polyethylene glycol (PEG)-induced osmotic stress caused similar effects and, in general, was counteracted only by norflurazon at the lowest concentration (10%). Partial defoliation was able to diminish only leaf ABA content (22.5%) at the highest PEG concentration (30%), probably through a reduction of the active sites of biosynthesis. At least under moderate drought (3–6 days without irrigation), mechanisms other than leaf ABA concentration were required to explain stomatal closure in response to limited soil water supply. Measurements of xylem sap pH revealed a progressive alkalinization through the drought condition (6.4 vs. 7.1), that was not counteracted with the addition of norflurazon. Moreover, in vitro treatment of detached leaves with buffers iso-osmotically adjusted at pH 7.1 significantly decreased stomatal conductance (more than 30%) as much as 70% when supplemented with ABA. Taken together, our results suggest that increased pH generated in drought-stressed roots is transmitted by the xylem sap to the leaves, triggering reductions in shoot water loss. The parallel rise in ABA concentration may act synergistically with pH alkalinization in xylem sap, with an initial response generated from the roots and further promotion by the stressed leaves.     

亚低温与干旱胁迫对番茄植株水分传输和形态解剖结构的影响

为探讨亚低温和干旱对植株水分传输的影响机制,以番茄幼苗为试材,利用人工气候室设置常温(昼25 ℃/夜18 ℃)和亚低温(昼15 ℃/夜8 ℃)环境,采用盆栽进行正常灌水(75%～85%田间持水量)和干旱处理(55%～65%田间持水量),分析了温度和土壤水分对番茄植株水分传输、气孔和木质部导管形态解剖结构的影响。结果表明: 与常温正常灌水处理相比,干旱处理使番茄叶水势、蒸腾速率、气孔导度、水力导度、茎流速率、气孔长度和叶、茎、根导管直径显著减小,而使叶、茎、根导管细胞壁厚度和抗栓塞能力增强;亚低温处理下番茄叶水势、蒸腾速率、气孔导度、水力导度和叶、茎、根导管直径显著降低,但气孔变大,叶、根导管细胞壁厚度和叶、茎、根抗栓塞能力显著升高。亚低温条件下土壤水分状况对番茄叶水势、蒸腾速率、气孔导度、水力导度、气孔形态、叶、根导管结构均无显著影响。总之,干旱处理下番茄通过协同调控叶、茎、根结构使植株水分关系重新达到稳态;亚低温处理下番茄植株水分关系的调控主要通过改变叶和根导管结构实现,且受土壤水分状况的影响较小。     

The efficiency of water use in water stressed plants is increased due to ABA induced stomatal closure

Gas exchange and abscisic acid content of Digitalis lanata EHRH. have been examined at different levels of plant water stress. Net photosynthesis, transpiration and conductance of attached leaves declined rapidly at first, then more slowly following the withholding of irrigation. The intercellular partial pressure of CO₂ decreased slightly. The concentration of 2-cis(S)ABA increased about eight-fold in the leaves of non-irrigated plants as compared with well-watered controls. A close linear correlation was found between the ABA content of the leaves and their conductance on a leaf area basis. In contrast, the plot of net assimilation versus ABA concentration was curvilinear, leading to an increased efficiency of water use during stress. After rewatering, photosynthesis reached control values earlier than transpiration, leaf conductance and ABA content. From these data it is concluded that transpiration through the stomata is directly controlled by the ABA content, whereas net photosynthesis is influenced additionally by other factors.Possible reasons for the responses of photosynthesis and water use efficiency to different stress and ABA levels are discussed.Abbreviations A net CO₂ assimilation - ABA abscisic acid - C_i intercellular CO₂ concentration - g stomatal conductance - T transpiration - WUE water use efficiency     

土壤干旱条件下氮素营养对玉米内源激素含量影响

在田间持水量分别保持于35％、55％和75％±5％的土壤水分条件下,利用盆栽实验研究了土壤干旱和氮素营养对玉米内源激素和气孔导度的影响．结果表明,土壤干旱下氮素营养明显降低了玉米根系木质部汁液ABA浓度,而正常供水下施氮处理间则无显著差异(施氮处理仍较低),同时测定的叶片ABA浓度则呈相反的变化趋势,表现为干旱下施氮处理要高于不施氮处理;施氮处理木质部汁液中ZRs浓度应低于相应的不施氮处理,在调控气孔行为方面并未表现拮抗ABA作用;3种土壤水分条件下,施氮玉米叶片的气孔导度均高于不施氮处理,与木质部汁液ABA浓度呈负相关,说明施氮处理较低的根源ABA浓度是导致其气孔导度较大的主要原因．     

Sequential response of whole plant water relations to prolonged soil drying and the involvement of xylem sap ABA in the regulation of stomatal behaviour of sunflower plants

Sunflower plants ( Helianihus animus cv. Tall Single Yellow} were grown in the greenhouse in drain pipes (100 mm inside diameter and 1 m long) rilled with John Innes No. 2 compost. When the fifth leaf had emerged, half of the plants were left unwatered for 6 days, rewatered for 2 days and then not watered for another 12 days. Measurements of water relations and abaxial stomatal conductance were made at each leaf position at regular intervals during the experimental period. Estimates were also made of soil water potentials along the soil profile and of ABA concentrations in xylem sap and leaves.
Soil drying led to some reduction in stomatal conductance alter only 3 days but leaf turgors were not reduced until day 13 (6 days after rewatering). When the water relations of leaves did change, older leases became substantially dehydrated while high turgors were recorded in younger leaves. Leaf ABA content measured on the third youngest leaf hardly changed over the first 13 days of the experiment, despite substantial soil drying, while xylem ABA concentrations changed very significantly and dynamically as soil water status varied, even when there was no effect of soil drying on leaf water relations. We argue that the highest ABA concentrations in the xylem, found as a result of substantial soil drying, arise from synthesis in both the roots and the older leaves, and act to delay the development of water deficit in younger leases.
In other experiments ABA solutions were watered on to the root systems of sunflower plants to increase ABA concentrations in xylem sap. The stomatal response to applied ABA was quantitatively very similar to that to ABA generated as a result of soil drying. There was a log-linear relationship between the reduction of leaf conductance and the increase of ABA concentration m xylem sap.     

干旱胁迫下冠菌素对玉米幼苗光合参数和内源激素含量的影响

采用沙培称重法研究冠菌素对玉米幼苗光合参数及内源激素系统影响的结果表明,干旱胁迫下,0.01μmol·L-1冠菌素显著增加玉米幼苗茎基部粗、根长、干鲜重、光合速率、蒸腾速率和气孔导度,降低胞间二氧化碳浓度,提高幼苗的ABA和1-氨基环丙烷-1-羧酸(ACC)含量。叶片与根中ABA/GA3大时耐旱性增大。     

Xylem transport and shoot accumulation of lumichrome, a newly recognized rhizobial signal, alters root respiration, stomatal conductance, leaf transpiration and photosynthetic rates in legumes and cereals

* Root respiration, stomatal conductance, leaf transpiration and photosynthetic rates were measured in phytotron and field-grown plants following the application of 5 or 10 nM lumichrome, 10 nM ABA (abscisic acid) and 10 ml of 0.2 OD600 infective rhizobial cells. * Providing soybean and cowpea roots with their respective homologous rhizobia and/or purified lumichrome increased the concentration of this molecule in xylem sap and leaf extracts. Relative to control, rhizobial inoculation and lumichrome application significantly increased root respiration in maize, decreased it in lupin, but had no effect on the other test species. * Applying either lumichrome (10 nM), infective rhizobial cells or ABA to roots of plants for 44 h in growth chambers altered leaf stomatal conductance and transpiration in cowpea, lupin, soybean, Bambara groundnut and maize, but not in pea or sorghum. Where stomatal conductance was increased by lumichrome application or rhizobial inoculation, it resulted in increased leaf transpiration relative to control plants. Treating roots of field plants of cowpea with this metabolite up to 63 d after planting showed decreased stomatal conductance, which affected CO2 intake and reduction by Rubisco. * The effect of rhizobial inoculation closely mirrored that of lumichrome application to roots, indicating that rhizobial effects on these physiological activities were most likely due to lumichrome released into the rhizosphere.     

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     

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.     

Do increases in xylem sap pH and/or ABA concentration mediate stomatal closure following nitrate deprivation?

Stomatal conductance (g(s)) of pepper (Capsicum annuum L.) plants decreased during the second photoperiod (day 2) after withholding nitrate (N). Stomatal closure of N-deprived plants was not associated with a decreased shoot water potential (Psi(shoot)); conversely Psi(shoot) was lower in N-supplied plants. N deprivation transiently (days 2 and 3) alkalized (0.2-0.3 pH units) xylem sap exuded from de-topped root systems under root pressure, and xylem sap expressed from excised shoots by pressurization. The ABA concentration of expressed sap increased 3-4-fold when measured on days 2 and 4. On day 2, leaves detached from N-deprived and N-supplied plants showed decreased transpiration rates when fed an alkaline (pH 7) artificial xylem (AX) solution, independent of the ABA concentration (10-100 nM) supplied. Thus changes in xylem sap composition following N deprivation can potentially close stomata. However, the lower transpiration rate of detached N-deprived leaves relative to N-supplied leaves shows that factors residing within N-deprived leaves also mediate stomatal closure, and that these factors assume greater importance as the duration of N deprivation increases.