Drought tolerance,xylem sap abscisic acid and stomatal conductance during soil drying: a comparison of canopy trees of three temperate deciduous angiosperms

Patterns of water relations, xylem sap abscisic acid concentration ([ABA]) and stomatal aperture were characterized and compared in drought-sensitive black walnut (Juglans nigra L.), less drought-sensitive sugar maple (Acer saccharum Marsh.) and drought-tolerant white oak (Quercus alba L.) trees co-occurring in a second-growth forest in Missouri, USA. There were strong correlations among reduction in predawn leaf water potential, increased xylem sap [ABA] and stomatal closure in all species. Stomatal conductance was more closely correlated with xylem sap ABA concentration than with ABA flux or xylem sap pH and cation concentrations. In isohydric black walnut, increased concentrations of ABA in the xylem sap appeared to be primarily of root origin, causing stomatal closure in response to soil drying. In anisohydric sugar maple and white oak, however, there were reductions in midday leaf water potential associated with stomatal closure, making it uncertain whether drought-induced xylem sap ABA was of leaf or root origin. The role of root-originated xylem sap ABA in these species as a signal to the shoot of the water status of the roots is, therefore, less certain.     

Hydraulic patterns and safety margins, from stem to stomata, in three eastern U.S. tree species

Adequate water transport is necessary to prevent stomatal closure and allow for photosynthesis. Dysfunction in the water transport pathway can result in stomatal closure, and can be deleterious to overall plant health and survival. Although much is known about small branch hydraulics, little is known about the coordination of leaf and stem hydraulic function. Additionally, the daily variations in leaf hydraulic conductance (K(leaf)), stomatal conductance and water potential (Ψ(L)) have only been measured for a few species. The objective of the current study was to characterize stem and leaf vulnerability to hydraulic dysfunction for three eastern U.S. tree species (Acer rubrum, Liriodendron tulipifera and Pinus virginiana) and to measure in situ daily patterns of K(leaf), leaf and stem Ψ, and stomatal conductance in the field. Sap flow measurements were made on two of the three species to compare patterns of whole-plant water use with changes in K(leaf) and stomatal conductance. Overall, stems were more resistant to hydraulic dysfunction than leaves. Stem P50 (Ψ resulting in 50% loss in conductivity) ranged from -3.0 to -4.2 MPa, whereas leaf P50 ranged from -0.8 to -1.7 MPa. Field Ψ(L) declined over the course of the day, but only P. virginiana experienced reductions in K(leaf) (nearly 100% loss). Stomatal conductance was greatest overall in P. virginiana, but peaked midmorning and then declined in all three species. Midday stem Ψ in all three species remained well above the threshold for embolism formation. The daily course of sap flux in P. virginiana was bell-shaped, whereas in A. rubrum sap flux peaked early in the morning and then declined over the remainder of the day. An analysis of our data and data for 39 other species suggest that there may be at least three distinct trajectories of relationships between maximum K(leaf) and the % K(leaf) at Ψ(min). In one group of species, a trade-off between maximum K(leaf) and % K(leaf) at Ψ(min) appeared to exist, but no trade-off was evident in the other two trajectories.     

黑河流域荒漠地区水分胁迫对梭梭苗木影响的试验研究(英文)

梭梭(HaloxylonAmmodendronBge,一种C4灌木)苗种植在15升的容器中,给予不同的水分胁迫处理,研究了其水分关系和气体交换特征。结果表明:当土壤水分含量大于11%时,梭梭苗有高的蒸腾量;土壤水分含量低于6%时,苗木就不能从土壤中吸取水分;很好供水的苗木的蒸腾量与潜在蒸发量成线型相关。气体交换测定发现,随着土壤水分含量的下降,造成了不同程度的气孔导度、叶蒸腾强度和光合作用的下降。对同一苗木而言,由于这个地区有高的水气压亏缺(VPD),很好和中度供水的苗木在气孔反应方面有较宽的范围,气孔在决定光合作用方面起着较小的作用,二者没有明显的线型相关关系。虽然水分胁迫使蒸腾速率比光合速率下降的更快,提高了水分利用效率,而较高的蒸发需求增加了蒸腾量,限制了光合作用,但是总的趋势是光合作用和蒸腾强度成线型相关。图6表2参15。     

Stomatal sensitivity to vapor pressure deficit and its relationship to hydraulic conductance in Pinus palustris

We studied the response of stomatal conductance at leaf (gS) and canopy (GS) scales to increasing vapor pressure deficit (D) in mature Pinus palustris Mill. (longleaf pine) growing in a sandhill habitat in the coastal plain of the southeastern USA. Specifically, we determined if variation in the stomatal response to D was related to variation in hydraulic conductance along the soil-to-leaf pathway (KL) over the course of a growing season. Reductions in KL were associated with a severe growing season drought that significantly reduced soil water content (theta) in the upper 90-cm soil profile. Although KL recovered partially following the drought, it never reached pre-drought values. Stomatal sensitivity to D was well correlated with maximum gS at low D at both leaf and canopy scales, and KL appeared to influence this response by controlling maximum gS. Our results are consistent with the hypothesis that stomatal response to D occurs to regulate minimum leaf water potential, and that the sensitivity of this response is related to changes in whole-plant hydraulics.     

Genotypic variation in drought tolerance of poplar in relation to abscisic acid

We investigated effects of water stress and external abscisic acid (ABA) supply on shoot growth, stomatal conductance and water status in 1-year-old cuttings of a drought-sensitive poplar genotype Populus x euramericana cv. I-214 (Italica) and a drought-tolerant genotype P. 'popularis 35-44' (popularis). Populus popularis was more productive and maintained higher leaf water potentials throughout the drought treatment than cv. Italica. Supply of ABA to the xylem sap caused a greater decline in growth and more leaf abscission in shoots of cv. Italica than in shoots of P. popularis. Immediately after initiation of the drought treatment in P. popularis, the ABA concentration ([ABA]) of the xylem increased rapidly and stomatal conductance declined; however, stomatal conductance had returned to control values by the third day of the drought treatment, coincident with a gradual decline in xylem [ABA]. In contrast, xylem [ABA] of cv. Italica initially increased more slowly than that of P. popularis in response to the drought treatment, but the increase continued for 3 days at which time a tenfold increase in xylem [ABA] was observed that was followed by abscission of more than 40% of the leaves. We conclude that sensitivity of poplar roots to variation in soil water content varies by clone and that a rapid short-term accumulation of ABA in shoots in response to water stress may contribute to drought tolerance.     

Changes in whole-tree water relations during ontogeny of Pinus flexilis and Pinus ponderosa in a high-elevation meadow

We measured sap flux in Pinus ponderosa Laws. and Pinus flexilis James trees in a high-elevation meadow in northern Arizona that has been invaded by conifers over the last 150 years. Sap flux and environmental data were collected from July 1 to September 1, 2000, and used to estimate leaf specific transpiration rate (El), canopy conductance (Gc) and whole-plant hydraulic conductance (Kh). Leaf area to sapwood area ratio (LA/SA) increased with increasing tree size in P. flexilis, but decreased with increasing tree size in P. ponderosa. Both Gc and Kh decreased with increasing tree size in P. flexilis, and showed no clear trends with tree size in P. ponderosa. For both species, Gc was lower in the summer dry season than in the summer rainy season, but El did not change between wet and dry summer seasons. Midday water potential (Psi(mid)) did not change across seasons for either species, whereas predawn water potential (Psi(pre)) tracked variation in soil water content across seasons. Pinus flexilis showed greater stomatal response to vapor pressure deficit (VPD) and maintained higher Psi(mid) than P. ponderosa. Both species showed greater sensitivity to VPD at high photosynthetically active radiation (PAR; > 2500 micromol m-2 s-1) than at low PAR (< 2500 micromol m-2 s-1). We conclude that the direction of change in Gc and Kh with increasing tree size differed between co-occurring Pinus species, and was influenced by changes in LA/SA. Whole-tree water use and El were similar between wet and dry summer seasons, possibly because of tight stomatal control over water loss.     

Stomatal responses to drought at a Mediterranean site: a comparative study of co-occurring woody species differing in leaf longevity

We studied stomatal responses to decreasing predawn water potential (Psipd) and increasing leaf-to-air water vapor pressure difference (VPD) of co-occurring woody Mediterranean species with contrasting leaf habits and growth form. The species included two evergreen oaks (Quercus ilex subsp. ballota (Desf.) Samp. and Q. suber L.), two deciduous oaks (Q. faginea Lam. and Q. pyrenaica Willd.) and two deciduous shrubs (Pyrus bourgaeana Decne. and Crataegus monogyna Jacq.). Our main objective was to determine if stomatal sensitivity is related to differences in leaf life span and leaf habit. The deciduous shrubs had the least conservative water-use characteristics, with relatively high stomatal conductance and low stomatal sensitivity to soil and atmospheric drought. As a result, Psipd decreased greatly in both species during the growing season, resulting in early leaf abscission in the summer. The deciduous oaks showed intermediate water-use characteristics, having maximum stomatal conductances and CO2 assimilation rates similar to or even higher than those of the deciduous shrubs. However, they had greater stomatal sensitivity to soil drying and showed less negative Psipd values than the deciduous shrubs. The evergreen oaks, and especially the species with the greatest leaf longevity, Q. ilex, exhibited the most conservative water-use behavior, having lower maximum stomatal conductances and greater sensitivity to VPD than the deciduous species. As a result, Psipd decreased less during the growing season in the evergreens than in the deciduous species, which may contribute to greater leaf longevity by avoiding irreversible damage during the summer drought. However, the combination of low maximum CO2 assimilation rates and high stomatal sensitivity to drought must have a negative impact on the final carbon budget of leaves with a long life span.     

Stomatal conductance, transpiration and sap flow of tropical montane rain forest trees in the southern Ecuadorian Andes

We investigated tree water relations in a lower tropical montane rain forest at 1950-1975 m a.s.l. in southern Ecuador. During two field campaigns, sap flow measurements (Granier-type) were carried out on 16 trees (14 species) differing in size and position within the forest stand. Stomatal conductance (g(s)) and leaf transpiration (E(l)) were measured on five canopy trees and 10 understory plants. Atmospheric coupling of stomatal transpiration was good (decoupling coefficient Omega = 0.25-0.43), but the response of g(s) and E(l) to the atmospheric environment appeared to be weak as a result of the offsetting effects of vapor pressure deficit (VPD) and photosynthetic photon flux (PPF) on g(s). In contrast, sap flow (F) followed these atmospheric parameters more precisely. Daily F depended chiefly on PPF sums, whereas on short time scales, VPD impeded transpiration when it exceeded a value of 1-1.2 kPa. This indicates an upper limit to transpiration in the investigated trees, even when soil water supply was not limiting. Mean g(s) was 165 mmol m(-2) s(-1) for the canopy trees and about 90 mmol m(-2) s(-1) for the understory species, but leaf-to-leaf as well as tree-to-tree variation was large. Considering whole-plant water use, variation in the daily course of F was more pronounced among trees differing in size and crown status than among species. Daily F increased sharply with stem diameter and tree height, and ranged between 80 and 120 kg day(-1) for dominant canopy trees, but was typically well below 10 kg day(-1) for intermediate and suppressed trees of the forest interior.     

Stomatal conductance of Acer rubrum ecotypes under varying soil and atmospheric water conditions: predicting stomatal responses with an abscisic acid-based model

A multiplicative model of stomatal conductance was developed and tested in two functionally distinct ecotypes of Acer rubrum L. (red maple). The model overcomes the main limitation of the commonly used Ball-Berry model (Ball et al. 1987) by accounting for stomatal behavior under soil drying conditions. We combined the Ball-Berry model with an integrated expression of abscisic acid (ABA)-based stomatal response to ABA concentration ([ABA]) in bulk leaf tissue (gfac), which coupled physiological changes at the leaf level with those in the root. The factor gfac = exp(-beta[ABA]L) incorporated the stomatal response to [ABA] into the Ball-Berry model by down regulating stomatal conductance (gs) in response to physiological changes in the root. The down regulation of gs is pertinent under conditions where soil drying may modify the delivery of chemical signals to leaf stomata. Model testing indicated that the multiplicative model was capable of predicting gs in red maple under wide ranges of soil and atmospheric conditions. Concordance correlation coefficients were high (between 0.59 and 0.94) for the tested ecotypes under three environmental conditions (atmospheric, rhizospheric and minimal stress). The study supported the use of gfac as a gas exchange function that controls water stress effects on gs and aids in the prediction of gs responses.     

Mean canopy stomatal conductance responses to water and nutrient availabilities in Picea abies and Pinus taeda

We compared sap-flux-scaled, mean, canopy stomatal conductance (GS) between Picea abies (L.) Karst. in Sweden and Pinus taeda (L.) in North Carolina, both growing on nutritionally poor soils. Stomatal conductance of Picea abies was approximately half that of Pinus taeda and the sensitivity of GS in Picea abies to vapor pressure deficit (D) was lower than in Pinus taeda. Optimal fertilization increased leaf area index (L) two- and threefold in Pinus taeda and Picea abies, respectively, regardless of whether irrigation was increased. Although it increased L, fertilization did not increase GS in Picea abies unless irrigation was also provided. In Pinus taeda growing on coarse, sandy soils, the doubling of L in response to fertilization reduced GS sharply unless irrigation was also provided. The reduction in GS with fertilization in the absence of irrigation resulted from the production of fine roots with low saturated hydraulic conductivity. When Pinus taeda received both fertilization and irrigation, the increase in L was accompanied by a large increase in GS. In Pinus taeda, a reference GS (defined as GS at D = 1 kPa; GSR) decreased in all treatments with decreasing volumetric soil water content (theta). In Picea abies, theta varied little within a treatment, but overall, GSR declined with theta, reaching lowest values when drought was imposed by the interception of precipitation. Despite the large difference in GS both between Picea abies and Pinus taeda and among treatments, stem growth was related to absorbed radiation, and stem growth response to treatment reflected mostly the changes in L.     

Environmental control of ozone uptake in Larix decidua Mill.: a comparison between different altitudes

We studied ozone (O(3)) uptake by European larch (Larix decidua Mill.) needles under ambient conditions at both a low and a high elevation site. At a given ambient O(3) concentration, the rate of O(3) uptake by needles was effectively controlled by stomatal conductance and, hence, by factors such as light, humidity and water status, which control stomatal conductance. At both study sites, atmospheric water vapor pressure deficit (VPD) was the climatic factor most closely correlated with ambient O(3) concentration. Thus, when ambient O(3) concentrations were highest, O(3) flux into the needles tended to be restricted by narrowing of the stomata. Mitigation of potential O(3) stress by stomatal closure was most marked at the low elevation site where both soil water stress and atmospheric VPD were greater than at the high elevation site.     

A comparison of three approaches to modeling leaf gas exchange in annually drought-stressed ponderosa pine forests

We tested, compared and modified three models of stomatal conductance at the leaf level in a forest ecosystem where drought stress is a major factor controlling forest productivity. The models were tested against 2 years (1999 and 2000) of leaf-level measurements on ponderosa pine (Pinus ponderosa Dougl. ex Laws.) growing in the Mediterranean climate of California, USA. The Ball, Woodrow and Berry (1987) (BWB) model was modified to account for soil water stress. Among the models, results of the modified BWB model were in the closest agreement with observations (r2 = 0.71). The Jarvis (1976) model showed systematic simulation errors related to vapor pressure deficit (r2 = 0.65). Results of the Williams, Rastetter, Fernandes et al. (1996) (SPA) model showed the poorest correlation with empirical data, but this model has only one calibration parameter (r2 = 0.60). Sensitivity analyses showed that, in all three models, predictions of stomatal conductance were most responsive to photosynthetically active radiation and soil water content. Stomatal conductance showed little sensitivity to vapor pressure deficit in the Jarvis model, whereas in both the BWB and SPA models, vapor pressure deficit (or relative humidity) was the third most important variable. Parameterization of the SPA model was in accordance with the parameterization of the modified BWB model, although the two models differ greatly. Measured and modeled results indicate that stomatal behavior is not water conservative during spring; however, during summer, when soil water content is low and vapor pressure deficit is high, stomatal conductance decreases and, according to the models, intrinsic water- use efficiency increases.     

Abscisic acid accumulation in leaves of two contrasting hybrid poplar clones affected by nitrogen fertilization plus cyclic flooding and soil drying

Cuttings of hybrid Populus clones Tristis and Eugenei growing in pots in a greenhouse were treated with nitrogen fertilizer at two rates and subjected to repeated soil flooding or drying. Periodically, gas exchange measurements and radioimmunoassays, to determine abscisic acid (ABA) concentrations, were made on recently mature leaves.In both clones, photosynthesis and stomatal conductance were depressed five days after flooding, but leaf ABA concentrations remained relatively constant. In contrast, an initial, 9-day period of soil drying resulted in substantial ABA accumulation in leaves, which closely correlated with declines in photosynthesis and conductance. A second soil drying cycle of up to 9 days was less effective in modifying gas exchange and leaf ABA concentrations. High-N supply stimulated leaf ABA production as the soil dried. On the resumption of watering, gas exchange in Tristis recovered fully and rapidly and leaf ABA concentrations quickly returned to control values, whereas gas exchange in Eugenei recovered slowly and leaf ABA concentrations remained high for longer.Gas exchange in Eugenei was unaffected by soil drying until leaf ABA concentrations exceeded 100 ng g(dw) (-1), whereas Tristis showed a reduction in stomatal conductance and photosynthesis at leaf ABA concentrations of only 10 ng g(dw) (-1). A rise in internal CO(2) concentrations was associated with increased leaf ABA concentrations in Tristis, but not in Eugenei. Clonal differences in the relationship between gas exchange and leaf ABA concentration suggest contrasting physiological strategies for survival under prolonged drying conditions.     

Stomatal conductance,growth and root signaling in Betula pendula seedlings subjected to partial soil drying

Seedlings of Betula pendula Roth were grown with their root systems separated between two soil compartments. Four treatments were imposed: (i) adequate irrigation in both compartments (WW, controls); (ii) adequate irrigation in one compartment and drought in the other compartment (WD); (iii) drought in both compartments (DD); and (iv) half of the root system severed and the remainder kept well-watered (root excision, RE). Predawn leaf water potential, stomatal conductance, soil-to-leaf specific hydraulic conductance, and root and leaf growth decreased in DD-treated seedlings, which also displayed severe leaf shedding (30% loss in leaf area). The DD treatment also resulted in increased concentrations of abscisic acid (ABA) and its glucose ester in the xylem sap of roots and shoots compared to concentrations in control seedlings (about 200 versus 20 nM). Despite the difference in xylem sap concentrations, total ABA flux to the shoots was similar in the two treatments (1-2 pmol ABA m(-2) leaf area s(-1)) as a result of reduced transpiration in the DD-treated seedlings. Compared with root growth in control plants, root growth increased in the RE-treated plants and decreased in the drying compartment of the WD treatment; however, the RE and WD treatments only slightly reduced leaf expansion, and had no detectable effects on shoot water relations or ABA concentrations of the root and shoot xylem sap. We conclude that short-term soil water depletion affecting only 50% of the root system does not cause a measurable stress response in birch shoots, despite root growth cessation in the fraction of drying soil.     

Ecophysiology of Acacia species in wet–dry tropical plantations

Selected tropical Acacia species are used extensively for short-rotation plantation forestry in many parts of Asia and, to a limited degree, in Australia. We explored leaf-level photosynthetic activity and leaf water potential (Ψleaf) of three field-grown Acacia tree species (aged between 7 and 18 months) in contrasting wet–dry tropical plantations in southern Vietnam and northern Australia. Light-saturated photosynthetic rate (A1500) declined throughout the morning and early afternoon in the dry season; in the wet season, levels remained high and relatively constant throughout most of the day. Maximum daily A1500 at 09:00 ranged from 22.2 μmol?m?2?s?1 in the wet to 10.4 μmol?m?2?s?1 in the dry season. At both locations, trees were able to extract soil water such that pre-dawn leaf water potential (Ψpd) remained>?1.5?MPa even at the end of the dry season. Stomatal conductance to water vapour (gs) did not respond to decreasing Ψleaf during the wet season but was sensitive to changes in Ψleaf in the dry season. Species comparisons of the relationships between A1500 and Ψleaf revealed different strategies to balance carbon uptake and water loss in a wet–dry environment. Acacia crassicarpa and A. mangium regulated Ψleaf to a greater extent than the A. mangium×A. auriculiformis hybrid such that ?Ψleaf (determined as Ψpd?midday Ψleaf) was unaffected by season. This result suggests that the hydraulic regulation of tree water status varies amongst young tropical Acacia species. From a management perspective, for Acacia species that tend to strongly regulate water loss in environments with an extended dry season, overall productivity at the end of a rotation may be less than for species that prioritise carbon gain.     

Water relations of Eucalyptus nitens × Eucalyptus grandis: is there interclonal variation in response to experimentally imposed water stress?

This study tested the hypothesis that water stress increases the hydraulic efficiency of Eucalyptus nitens × E. grandis saplings as a result of osmotic and elastic adjustments. Eucalyptus nitens × E. grandis clones (NH00, NH58, NH69 and NH70) were potted in coarse river sand supplemented with a slow-release fertiliser, drip-irrigated four times daily and exposed to full sunlight for eight months. Thereafter, irrigation was withheld twice for seven consecutive days from half of the saplings of each clone, with a seven-day recovery period (regular irrigation) in-between. Relative soil moisture content did not correlate with stomatal conductance (g_s) at pre-dawn and at midday. Leaves of plants subjected to the water-stress treatment wilted in 7 d, and the reduction in g_s was significant at midday with no significant differences between clones. Stomatal conductance and all traits derived from pressure-volume graphs (e.g. osmotic potential at full turgor) were constant in the control treatment. There were no clear patterns in osmotic and elastic adjustments in both treatments. Root hydraulic conductance was constant between treatments and clones. However, water stress reduced shoot hydraulic conductance and stem hydraulic conductivity with significant interclonal effects. Plant biomass, leaf area and leaf weight ratio were significantly lower in the water-stressed plants, but there were no differences between the clones. In conclusion, the water-stress treatment did not introduce significant differences in stomatal conductance and tissue-water relations of Eucalyptus nitens × E. grandis clones. Interclonal variation in water-stress response was found in shoot hydraulic traits, and clone NH58 may be more suitable for planting across sites prone to moderate water stress.     

Canopy stomatal conductance and xylem sap abscisic acid (ABA) in mature Scots pine during a gradually imposed drought

We investigated the effect of drought on canopy stomatal conductance (g(c)), and examined the hypothesis that g(c) is controlled by the chemical messenger abscisic acid (ABA) produced in roots. Beginning in November 1994, we subjected a mature stand of Scots pine (Pinus sylvestris L.) to an imposed 11-month drought. Control plots were maintained at average-season soil water content. Xylem sap was extracted from shoots at regular intervals from April to November 1995. Soil water, sap flow and leaf water potentials (predawn to dusk) were recorded at the same time. Canopy stomatal conductance was calculated from sap flow data and xylem sap ABA concentrations ([ABA(xyl)]) were measured by radioimmunoassay. Mean [ABA(xyl)] in control trees was 250 micromol m(-3). No diurnal variation in [ABA(xyl)] was detected. With soil drying, [ABA(xyl)] increased to a maximum in summer (600 micromol m(-3)), but decreased again toward autumn; however, no significant increase in ABA flux to the leaves occurred. A decline in g(c) was detected when volumetric soil water content declined below 0.12. The decline in g(c) could not have been mediated by increasing [ABA(xyl)] because stomatal closure appeared to precede any increase in [ABA(xyl)]. Peak sap flow velocity data were used to estimate delivery times for root-to-shoot signals in 15-m tall trees. Under normal field conditions, a signal would take 12 days to travel from the site of production (roots) to the presumed site of action (shoots). However, under drought conditions it may take a chemical signal in excess of 6 weeks. We conclude that a feedforward model of short-term stomatal response to soil drying, based solely on the action of a chemical messenger from the roots, is not applicable in mature conifer trees because signal transmission is too slow.     

滴灌在林业上应用的研究与进展*

综述了滴灌在林业上应用的研究与进展，指出滴灌在林业上主要应用于经济林培育中，在用材林、苗木培育上的应用较少。滴灌在多种树木上应用的研究表明，其可促进树木的生长和提高产量，同时可大幅度提高水分利用效率。而其增产和节水的原因在于对树叶水势、光合速率、气孔导度等生理指标以及土壤水分分布及运动特性的影响。此外，还对目前节水灌溉制度探索的研究进行了概括性的总结。最后，建议对滴灌的研究应进一步深入到对优化灌溉制度的探索中，而对优化灌溉制度的研究应依据植物或树木的生长节律、生理特性与土壤的水分运动特征来确定灌溉的开始时间、灌溉频率及灌水定额，从而科学地制定灌溉计划。     

Impact of variable [CO2] and temperature on water transport structure-function relationships in Eucalyptus

Nearly 30 years ago, Whitehead and Jarvis and Whitehead et al. postulated an elegant mechanistic explanation for the observed relationship between tree hydraulic structure and function, hypothesizing that structural adjustments promote physiological homeostasis. To date, this framework has been nearly completely overlooked with regard to varying atmospheric carbon dioxide ([CO(2)]). Here, we evaluated Whitehead's hypothesis of leaf water potential (Ψ(l)) homeostasis in faster-growing (Eucalyptus saligna) and slower-growing (Eucalyptus sideroxylon) tree saplings grown under three [CO(2)] (pre-industrial, current and future) and two temperature (ambient and ambient + 4°C) treatments. We tested for relationships between physiological (stomatal conductance and Ψ(l)) and structural (leaf and sapwood areas (A(l), A(s)), height (h), xylem conductivity (k(s))) plant variables as a function of the [CO(2)] and temperature treatments to assess whether structural variables adjusted to maintain physiological homeostasis. Structural components (A(l), A(s), h) generally increased with [CO(2)] or temperature, while g(s) was negatively correlated with [CO(2)]. Contrary to Whitehead's hypothesis, Ψ(l) did not exhibit homeostasis in either species; elevated temperatures were associated with more negative Ψ(l) in faster-growing E. saligna, and less negative Ψ(l) in slower-growing E. sideroxylon. Moreover, individual structural variables were generally uncorrelated with Ψ(l). However, across both species, the integrated hydraulic property of leaf specific hydraulic conductance (K(l)) was positively correlated with an independent calculation of K(l) determined exclusively from leaf physiological variables. These results suggest that physiological homeostasis may not apply to saplings exposed to global change drivers including [CO(2)] and temperature. Nevertheless, Whitehead et al.'s formulation identified K(l) as a sensitive measure of plant structural-physiological co-variation across species.     

Modulation of leaf conductance by root to shoot signaling under water stress in Arabidopsis

Signal communication between root and shoot plays a crucial role in plant resistance to water stress. While many studies on root to shoot signals have been carried out in many plant species, no information is available for the model plant, Arabidopsis, whose adoption has great significance for further probing the molecular aspects of long distance stress signals. Here, we introduced the establishment of techniques for investigations of root to shoot signals in Arabidopsis. Stomatal movements in relation to root signals were probed by using these techniques. The results show that Arabidopsis is a suitable plant species for partial roots drying (PRD) experiments. In the PRD system, while no significant differences were found in leaf water potential between well-watered and stressed plants, water stress led to a decrease in leaf conductance, which suggests a regulation of stomatal movements by root to shoot signals. While water stress caused a significant increase in the concentration of sap abscisic acid (ABA) of xylem, no increase in xylem sap pH was observed. Moreover, the increase in the ABA content of xylem coincided with the decrease in leaf conductance, which suggests a possible role of ABA in the regulation of stomatal movements. Infrared temperature images showed that leaf temperatures of PRD plant were higher compared with those of well-watered plants, which further indicates that stomatal movements can be modulated by root signals. The confirmation of root to shoot signaling in Arabidopsis has established a basis for further investigation into the molecular mechanisms of the root to shoot signaling under water stress.