Stomatal control by both [ABA] in the xylem sap and leaf water status: a test of a model for draughted or ABA-fed field-grown maize

A model of maize stomatal behaviour has been developed, in which stomatal conductance is linked to the concentration of abscisic acid ([ABA]) in the xylem sap, with a sensitivity dependent upon the leaf water potential (Ψ₁). It was tested against two alternative hypotheses, namely that stomatal sensitivity to xylem [ABA] would be linked to the leaf-to-air vapour pressure difference (VPD), or to the flux of ABA into the leaf. Stomatal conductance (g_s) was studied: (1) in field-grown plants whose xylem [ABA] and Ψ₁ depended on soil water status and evaporative demand; (2) in field-grown plants fed with ABA solutions such that xylem [ABA] was artificially raised, thereby decreasing g_s and increasing Ψ₁ and leaf-to-air VPD; and (3) in ABA-fed detached leaves exposed to varying evaporative demands, but with a constant and high Ψ₁. The same relationships between g_s, xylem [ABA] and Ψ₁, showing lower stomatal sensitivity to [ABA] at high Ψ₁, applied whether variations in xylem [ABA] were due to natural increase or to feeding, and whether variations in Ψ₁, were due to changes in evaporative demand or to the increased Ψ₁ observed in ABA-fed plants. Conversely, neither the leaf-to-air VPD nor the ABA flux into the leaf accounted for the observed changes in stomatal sensitivity to xylem [ABA]. The model, using parameters calculated from previous field data and the detached-leaf data, was tested against the observations of both ABA-fed and droughted plants in the field. It accounted with reasonable accuracy for changes in g_s (r² ranging from 0.77 to 0.81). These results support the view that modelling of stomatal behaviour requires consideration of both chemical and hydraulic aspects of root-to-shoot communication.     

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.     

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.     

What information is conveyed by an ABA signal from maize roots in drying field soil?

During two seasons, ABA concentrations were monitored in roots, leaves and xylem sap of field-grown maize. The water status of soil and plant was also measured. Plants were grown on plots with compacted or non-compacted soil, which were irrigated or remained unwatered. ABA concentration in the xylem sap before dawn and in the roots increases 25-fold and five-fold, respectively, as the soil dried, with a close correlation with the soil water status, but with no clear effect of the soil structure. In contrast to the results of several laboratory experiments, no appreciable increase in xylem [ABA] and reduction in stomatal conductance were observed with dehydration of the part of the root system located in soil upper layers. These responses only occurred when the water reserve of the whole soil profile was close to depletion and the transpiration declined. Xylem [ABA] measured during the day was appreciably higher in the compacted treatment than in non-compacted treatment, unlike that measured before dawn. Since a mechanical message is unlikely to undergo such day-night alterations, we suggest that this was due to a faster decrease in root water potential and water flux in the compacted treatment, linked to the root spatial arrangement. These results raise the possibility that ABA concentration in the xylem sap could be controlled by two coexisting mechanisms: (1) the rate of ABA synthesis in the roots linked to the soil or root water status, as shown in laboratory experiments; (2) the dilution of ABA in the water flow from roots, which could be an overriding mechanism in field conditions. This second mechanism would allow the plant to sense the water flux through the root system.     

The ecosystem wilting point defines drought response and recovery of a Quercus-Carya forest

Soil and atmospheric droughts increasingly threaten plant survival and productivity around the world. Yet, conceptual gaps constrain our ability to predict ecosystem-scale drought impacts under climate change. Here, we introduce the ecosystem wilting point (Ψ_EWP), a property that integrates the drought response of an ecosystem's plant community across the soil–plant–atmosphere continuum. Specifically, Ψ_EWP defines a threshold below which the capacity of the root system to extract soil water and the ability of the leaves to maintain stomatal function are strongly diminished. We combined ecosystem flux and leaf water potential measurements to derive the Ψ_EWP of a Quercus-Carya forest from an “ecosystem pressure–volume (PV) curve,” which is analogous to the tissue-level technique. When community predawn leaf water potential (Ψ_pd) was above Ψ_EWP (=−2.0 MPa), the forest was highly responsive to environmental dynamics. When Ψ_pd fell below Ψ_EWP, the forest became insensitive to environmental variation and was a net source of carbon dioxide for nearly 2 months. Thus, Ψ_EWP is a threshold defining marked shifts in ecosystem functional state. Though there was rainfall-induced recovery of ecosystem gas exchange following soaking rains, a legacy of structural and physiological damage inhibited canopy photosynthetic capacity. Although over 16 growing seasons, only 10% of Ψ_pd observations fell below Ψ_EWP, the forest is commonly only 2–4 weeks of intense drought away from reaching Ψ_EWP, and thus highly reliant on frequent rainfall to replenish the soil water supply. We propose, based on a bottom-up analysis of root density profiles and soil moisture characteristic curves, that soil water acquisition capacity is the major determinant of Ψ_EWP, and species in an ecosystem require compatible leaf-level traits such as turgor loss point so that leaf wilting is coordinated with the inability to extract further water from the soil.     

Stomatal conductance in relation to xylem sap abscisic acid concentrations in two tropical trees, Acacia confusa and Litsea glutinosa

Two tropical trees, Acacia confusa and Litsea glutinosa, were grown under controlled conditions with their roots subjected to soil drying and soil compaction treatments. In both species, a decline in stomatal conductance resulting from soil drying took place much earlier than the decline of leaf water potential. Soil compaction treatment also resulted in a substantial decrease in stomatal conductance but had little effect on leaf water potential. A rapid and substantial increase in xylem abscisic acid (ABA) concenation ([ABA]), rather than hulk leaf ABA, was closely related to soil drying and soil compaction. A significant relationship between stomatal conductance (g_s) and xylem [ABA] was observed in both species. Artificially feeding ABA solutions to excised leaves of both species showed that the relationship bet ween g_s and [ABA] was very similar to that obtained from the whole plant, i.e. the relationship between g_s and xylem [ABA]. These results suggest that xylem ABA may act as a stress signal in the control of stomatal conductance.     

Differences in osmotic adjustment,foliar abscisic acid dynamics,and stomatal regulation between an isohydric and anisohydric woody angiosperm during drought

Species are often classified along a continuum from isohydric to anisohydric, with isohydric species exhibiting tighter regulation of leaf water potential through stomatal closure in response to drought. We investigated plasticity in stomatal regulation in an isohydric (Eucalyptus camaldulensis) and an anisohydric (Acacia aptaneura) angiosperm species subject to repeated drying cycles. We also assessed foliar abscisic acid (ABA) content dynamics, aboveground/belowground biomass allocation and nonstructural carbohydrates. The anisohydric species exhibited large plasticity in the turgor loss point (Ψ_TLP), with plants subject to repeated drying exhibiting lower Ψ_TLP and correspondingly larger stomatal conductance at low water potential, compared to plants not previously exposed to drought. The anisohydric species exhibited a switch from ABA to water potential‐driven stomatal closure during drought, a response previously only reported for anisohydric gymnosperms. The isohydric species showed little osmotic adjustment, with no evidence of switching to water potential‐driven stomatal closure, but did exhibit increased root:shoot ratios. There were no differences in carbohydrate depletion between species. We conclude that a large range in Ψ_TLP and biphasic ABA dynamics are indicative of anisohydric species, and these traits are associated with exposure to low minimum foliar water potential, dense sapwood and large resistance to xylem embolism.     

周期性土壤干旱和叶片水势对气孔响应木质部ABA灵敏度的影响

研究了周期性土壤干旱期间气孔对木质部ＡＢＡ响应的灵敏度的变化以及叶片水势对灵敏度的影响。实验结果证明了木质部ＡＢＡ浓度是反映根系周围土壤水分状况的一个指标的结论。土壤周期性干旱不影响木质部ＡＢＡ浓度对土壤水分状况的依赖关系,但显著地提高了气孔对木质部ＡＢＡ 响应的灵敏度。根据对实测数据的数学模拟结果显示,引起气孔导度下降５０％ 所需的木质部ＡＢＡ浓度从第一轮土壤干旱的７５０ ｎｍｏｌ／Ｌ降至第二轮土壤干旱的５５０ ｎｍｏｌ／Ｌ。分根实验的结果表明,叶片水分亏缺显著提高了气孔对木质部ＡＢＡ 的响应的灵敏程度,全根干旱中引起气孔导度下降５０ ％ 所需的木质部ＡＢＡ 浓度比半根干旱的小２ ～４ 倍。这表明,气孔对木质部ＡＢＡ响应的灵敏度不是一个固定的特性,可随植物生长环境及许多其他因素的变化而表现出很大的差异     

Effects of Periodical Soil Drying and Leaf Water Potential on the Sensitivity of Stomatal Response to Xylem ABA

The study on the changes of stomatal sensitivity in relation to xylem ABA during periodical soil drying and the effect of leaf water status on the stomatal sensitivity has confirmed that xylem ABA concentration is a good indicator of soil water status around roots and the relation between xylem ABA concentration and predawn leaf water potential remained constant during the three consecutive soil drying cycles based on the slopes of the fitted lines. The sensitivity of stomata to xylem ABA increased substantially as the soil drying cycles progressed, and the xylem ABA concentration needed to cause a 50% decrease of stomatal conductance was as low as 550 mnoL/L in the next two soil drying cycle, as compared with the 750 nmol/L ABA in the first cycle of soil drying. The results using the split-root system showed that leaf water deficit significantly enhanced the stomatal response to xylem ABA and the xylem ABA concentration needed to cause a 50% decrease in stomatal conductance was 2 to 4 times smaller in the whole-root-drying treatment than those in the semi-root- drying treatment. These results suggested that the sensitivity of stomata to xylem ABA concentration is not a fixed characteristic.     

Modulation of Root Signals in Relation to Stomatal Sensitivity to Root-sourced Abscisic Acid in Drought-affected Plants

Stomatal sensitivity to root signals induced by soil drying may vary between environments and plant species. This is likely to be a result of the interactions and modulations ámong root signals. As a stress signal, abscisic acid （ABA） plays a central role in root to shoot signaling, pH and hydraulic signals may interact with ABA signals and thus, jointly regulate stomatal responses to changed soil water status, pH itself can be modified by several factors, among which the chemical compositions in the xylem stream and the live cells surrounding the vessels play crucial roles. In addition to the xylem pH, more attention should be paid to the direct modulation of leaf apoplastic pH, because many chemical compositions might strongly modify the leaf apoplastic pH while having no significant effect on the xylem pH. The direct modulation of the ABA signal intensity may be more important for the regulation of stomatal responses to soil drying than the ABA signal per se. The ABA signal is also regulated by the ABA catabolism and the supply of precursors to the roots if a sustained root to shoot communication of soil drying operates at the whole plant level. More importantly, ABA catabolism could play crucial roles in the determination of the fate of the ABA signal and thereby control the stomatal behavior of the root-sourced ABA signal.     

Modulation of Root Signals in Relation to Stomatal Sensitivity to Root-sourced Abscisic Acid in Drought-affected Plants

Stomatal sensitivity to root signals induced by soil drying may vary between environments and plant species. This is likely central role in root to shoot signaling. pH and hydraulic signals may interact with ABA signals and thus, jointly regulate stomatal responses to changed soil water status. pH itself can be modified by several factors, among which the chemical compositions In the xylem stream and the live cells surrounding the vessels play crucial roles. In addition to the xylem pH,more attention should be paid to the direct modulation of leaf apoplastic pH, because many chemical compositions might strongly modify the leaf apoplastlc pH while having no significant effect on the xylem pH. The direct modulation of the ABA signal intensity may be more important for the regulation of stomatal responses to soil drying than the ABA signal per se.The ABA signal is also regulated by the ABA catabolism and the supply of precursors to the roots If a sustained root to shoot communication of soil drying operates at the whole plant level. More importantly, ABA catabolism could play crucial roles In the determination of the fate of the ABA signal and thereby control the stomatal behavior of the root-sourced ABA signal.     

Stomatal control by fed or endogenous xylem ABA in sunflower: interpretation of correlations between leaf water potential and stomatal conductance in anisohydric species

The stomatal conductance of several anisohydric plant species, including field-grown sunflower, frequently correlates with leaf water potential (φ₁), suggesting that chemical messages travelling from roots to shoots may not play an important role in stomatal control. We have performed a series of experiments in which evaporative demand, soil water status and ABA origin (endogenous or artificial) were varied in order to analyse stomatal control. Sunflower plants were subjected to a range of soil water potentials under contrasting air vapour pressure deficits (VPD, from 0.5 to 2.5 kPa) in the field, in the glasshouse or in a humid chamber. Sunflower plants were also fed through the xylem with varying concentrations of artificial ABA, in the glasshouse and in the field. Finally, detached leaves were fed directly with varying concentrations of ABA under three contrasting VPDs. A unique relationship between stomatal conductance (g_s) and the concentration of ABA in the xylem sap (xylem [ABA]) was observed in all cases. In contrast, the relationship between φ₁ and g_s varied substantially among experiments. Its slope was positive for droughted plants and negative for ABA-fed whole plants or detached leaves, and also varied appreciably with air VPD. All observed relationships could be modelled on the basis of the assumption that φ₁ had no controlling effect on g_s. We conclude that stomatal control depended only on the concentration of ABA in the xylem sap, and that φ₁ was controlled by water flux through the plant (itself controlled by stomatal conductance). The possibility is also raised that differences in stomatal ‘strategy’ between isohydric plants (such as maize, where daytime φ₁ does not vary appreciably with soil water status) and anisohydric plants (such as sunflower) may be accounted for by the degree of influence of φ₁ on stomatal control, for a given level of xylem [ABA]. We propose that statistical relationships between φ₁ and g_s are only observed when φ₁ has no controlling action on stomatal behaviour.     

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.     

Xylem ABA controls the stomatal conductance of field-grown maize subjected to soil compaction or soil drying

Stomatal conductance of individual leaves was measured in a maize field, together with leaf water potential, leaf turgor, xylem ABA concentration and leaf ABA concentration in the same leaves. Stomatal conductance showed a tight relationship with xylem ABA, but not with the current leaf water status or with the concentration of ABA in the bulk leaf. The relationship between stomatal conductance and xylem [ABA] was common for variations in xylem [ABA] linked to the decline with time of the soil water reserve, to simultaneous differences between plants grown on compacted, non-compacted and irrigated soil, and to plant-to-plant variability. Therefore, this relationship is unlikely to be fortuitous or due to synchronous variations. These results suggest that increased concentration of ABA in the xylem sap in response to stress can control the gas exchange of plants under field conditions.     

Relations between stomatal closure, leaf turgor and xylem vulnerability in eight tropical dry forest trees

This study examined the linkage between xylem vulnerability, stomatal response to leaf water potential (Ψ_L), and loss of leaf turgor in eight species of seasonally dry tropical forest trees. In order to maximize the potential variation in these traits species that exhibit a range of leaf habits and phenologies were selected. It was found that in all species stomatal conductance was responsive to Ψ_L over a narrow range of water potentials, and that Ψ_L inducing 50% stomatal closure was correlated with both the Ψ_L inducing a 20% loss of xylem hydraulic conductivity and leaf water potential at turgor loss in all species. In contrast, there was no correlation between the water potential causing a 50% loss of conductivity in the stem xylem, and the water potential at stomatal closure (Ψ_SC) amongst species. It was concluded that although both leaf and xylem characters are correlated with the response of stomata to Ψ_L, there is considerable flexibility in this linkage. The range of responses is discussed in terms of the differing leaf‐loss strategies exhibited by these species.     

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.     

Influence of leaf water status on stomatal response to humidity,hydraulic conductance,and soil drought in Betula occidentalis

Whole-canopy measurements of water flux were used to calculate stomatal conductance (g _s ) and transpiration (E) for seedlings of western water birch (Betula occidentalis Hook.) under various soil-plant hydraulic conductances (k), evaporative driving forces (ΔN; difference in leaf-to-air molar fraction of water vapor), and soil water potentials (Ψ_s). As expected, g _s dropped in response to decreased k or Ψ_S, or increased ΔN(> 0.025). Field data showed a decrease in mid-day g _s with decreasing k from soil-to-petiole, with sapling and adult plants having lower values of both parameters than juveniles. Stomatal closure prevented E and Ψ from inducing xylem cavitation except during extreme soil drought when cavitation occurred in the main stem and probably roots as well. Although all decreases in g _s were associated with approximately constant bulk leaf water potential (ψ_l), this does not logically exclude a feedback response between Ψ_L and g _s . To test the influence of leaf versus root water status on g _s , we manipulated water status of the leaf independently of the root by using a pressure chamber enclosing the seedling root system; pressurizing the chamber alters cell turgor and volume only in the shoot cells outside the chamber. Stomatal closure in response to increased ΔN, decreased k, and decreased Ψ_S was fully or partially reversed within 5 min of pressurizing the soil. Bulk Ψ_L remained constant before and after soil pressurizing because of the increase in E associated with stomatal opening. When ΔN was low (i.e., < 0.025), pressurizing the soil either had no effect on g _s , or caused it to decline; and bulk Ψ_L increased. Increased Ψ_l may have caused stomatal closure via increased backpressure on the stomatal apparatus from elevated epidermal turgor. The stomatal response to soil pressurizing indicated a central role of leaf cells in sensing water stress caused by high ΔN, low k, and low Ψ_S. Invoking a prominent role for feedforward signalling in short-term stomatal control may be premature.     

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.     

Correlations of stomatal conductance with hydraulic and chemical factors in several deciduous tree species in a natural habitat

Recent research in whole-plant stomatal physiology, conducted largely with potted plants in controlled environments, suggests that stomatal conductance ( g _s) might be more closely linked to plant chemical variables than to hydraulic variables. To test this in a field situation, seasonal g _s was examined in relation to a number of plant and environmental variables in 11 temperate, deciduous forest tree species. Stomatal conductance was generally better correlated with environmental variables (air temperature, vapor pressure deficit, PPFD) than with plant variables, and slightly better correlated with plant hydraulic variables (shoot water and osmotic potentials) than with plant chemical variables (xylem sap ABA concentration, xylem sap pH). We examined a model, developed previously for maize, which describes regulation of g _s by xylem sap ABA concentration with leaf water status acting to modify stomatal sensitivity to the ABA signal. This model explained slightly more variation in seasonal g _s in the forest trees than did single plant variables but not more variation than most single environmental variables. Response surface models, especially those incorporating environmental variables, were more consistently successful at explaining g _s across species.     

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