Stomatal conductance, growth and root signaling in young oak seedlings subjected to partial soil drying

Leaf conductance, water relations, growth, and abscisic acid (ABA) concentrations in xylem sap, root apices and leaves were assessed in oak seedlings (Quercus robur L.) grown with a root system divided between two compartments and subjected to one of four treatments: (a) well watered, WW; (b) half of root system exposed to soil drying and half kept well watered, WD; (c) whole root system exposed to drought, DD; and (d) half of root system severed, RE. Sharp decreases in plant stomatal conductance, leaf water potential, hydraulic conductance and leaf growth were observed during DD treatment. No significant differences in plant leaf water potential and stomatal conductance were detected between the WW and WD treatments. Nevertheless, the WD treatment resulted in inhibition of leaf expansion and stimulation of root elongation only in the well-watered compartment. Abscisic acid concentrations did not change in leaves, root tips, or xylem sap of WD- compared to WW-treated plants. Increased concentrations of ABA were observed in xylem sap from DD-treated plant roots, but the total flux of ABA to shoots was reduced compared to that in WW-treated plants, because of decreases in transpiration flux. Similar plant responses to the WD and RE treatments indicate that the responses observed in the WD-treated plants were probably not triggered by a positive signal originating from drying roots.     

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.     

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.     

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.     

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.     

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.     

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.     

Drought tolerance,xylem sap abscisic acid and stomatal conductance during soil drying: a comparison of young plants of four temperate deciduous angiosperms

Patterns of water relations, xylem sap abscisic acid (ABA) concentration ([ABA]) and stomatal aperture were compared in drought-sensitive black walnut (Juglans nigra L.) and black willow (Salix nigra Marsh.), less drought-sensitive sugar maple (Acer saccharum Marsh.) and drought-tolerant white oak (Quercus alba L.). Strong correlations among reduction in predawn water potential, increase in xylem sap [ABA] and stomatal closure were observed in all species. Stomatal response was more highly correlated with xylem [ABA] than with ABA flux. Xylem sap pH and ion concentrations appeared not to play a major role in the stomatal response of these species. Stomata were more sensitive to relative changes in [ABA] in drought-sensitive black walnut and black willow than in sugar maple and white oak. In the early stages of drought, increased [ABA] in the xylem sap of black walnut and black willow was probably of root origin and provided a signal to the shoot of the water status of the roots. In sugar maple and white oak, leaf water potential declined with the onset of stomatal closure, so that stomatal closure also may have occurred in response to the change in leaf water potential.     

Abscisic acid in leaves and roots of willow: significance for stomatal conductance

Excised leaves and roots of willow (Salix dasyclados Wimm.) accumulated abscisic acid (ABA) in response to desiccation. The accumulation of ABA was greater in young leaves and roots than in old leaves and roots. In mature leaves, ABA accumulation was related to the severity and duration of the desiccation treatment. Water loss equal to 12% of initial fresh weight caused the ABA content of mature leaves to increase measurably within 30 min and to double in 2.5 h. The drying treatment caused significant (P = 0.05) reductions in leaf water potential and stomatal conductance. Recovery of leaf water potential to the control value occurred within 10 min of rewatering the dehydrated leaves, but recovery of stomatal conductance took an hour or longer, depending on the interval between dehydration and rewatering. The addition of ABA to the transpiration stream of well-watered excised leaves was sufficient to cause partial stomatal closure within 1 h and, depending on ABA concentration, more or less complete stomatal closure within 3 h. When the ABA solution was replaced with water, stomatal conductance increased at a rate inversely related to the concentration of the ABA solution with which the leaves had been supplied.     

Stomatal response of an anisohydric grapevine cultivar to evaporative demand, available soil moisture and abscisic acid

Stomatal responsiveness to evaporative demand (air vapour pressure deficit (VPD)) ranges widely between species and cultivars, and mechanisms for stomatal control in response to VPD remain obscure. The interaction of irrigation and soil moisture with VPD on stomatal conductance is particularly difficult to predict, but nevertheless is critical to instantaneous transpiration and vulnerability to desiccation. Stomatal sensitivity to VPD and soil moisture was investigated in Semillon, an anisohydric Vitis vinifera L. variety whose leaf water potential (Ψ(l)) is frequently lower than that of other grapevine varieties grown under similar conditions in the warm grape-growing regions of Australia. A survey of Semillon vines across seven vineyards revealed that, regardless of irrigation treatment, midday Ψ(l) was dependent on not only soil moisture but VPD at the time of measurement. Predawn Ψ(l) was more closely correlated to not only soil moisture in dry vineyards but to night-time VPD in drip-irrigated vineyards, with incomplete rehydration during high night-time VPD. Daytime stomatal conductance was low only under severe plant water deficits, induced by extremes in dry soil. Stomatal response to VPD was inconsistent across irrigation regime; however, in an unirrigated vineyard, stomatal sensitivity to VPD-the magnitude of stomatal response to VPD-was heightened under dry soils. It was also found that stomatal sensitivity was proportional to the magnitude of stomatal conductance at a reference VPD of 1kPa. Exogenous abscisic acid (ABA) applied to roots of Semillon vines growing in a hydroponic system induced stomatal closure and, in field vines, petiole xylem sap ABA concentrations rose throughout the morning and were higher in vines with low Ψ(l). These data indicate that despite high stomatal conductance of this anisohydric variety when grown in medium to high soil moisture, increased concentrations of ABA as a result of very limited soil moisture may augment stomatal responsiveness to low VPD.     

Adaptations to soil drying in woody seedlings of African locust bean, (Parkia biglobosa (Jacq.) Benth.)

Stomatal conductance, transpiration and xylem pressure potential of African locust bean (Parkia biglobosa (Jacq.) Benth.) seedlings subjected from the sixth week after emergence to four weeks of continuous soil drought did not differ from those of well-watered, control plants until two-thirds of the available soil water had been used. In both well-watered and drought-treated plants, stomatal conductance was highest early in the day when vapor pressure deficits were low, but decreased sharply by midday when evaporative demand reached its highest value. There was no increase in stomatal conductance later in the day as vapor pressure deficit declined. The relationship between transpiration rate and xylem pressure potential showed non-linearity and hysteresis in both control and drought-treated plants, which seems to indicate that the plants had a substantial capacity to store water. The rate of leaf extension in African locust bean seedlings subjected to six consecutive 2-week cycles of soil drought declined relative to that of well-watered, control plants, whereas relative root extension increased. It appears that African locust bean seedlings minimized the impact of drought by: (1) restricting transpiration to the early part of the day when a high ratio of carbon gain to water loss can be achieved; (2) utilizing internally stored water during periods of rapid transpiration; (3) reducing the rate of leaf expansion and final leaf size in response to soil drought without reducing the rate of root extension, thereby reducing the ratio of transpiring leaf surface area to absorbing root surface area.     

Hydraulic responses of whole vines and individual roots of kiwifruit (Actinidia chinensis) following root severance

Whole vine (K(plant)) and individual root (K(root)) hydraulic conductances were measured in kiwifruit (Actinidia chinensis Planch. var. chinensis 'Hort16A') vines to observe hydraulic responses following partial root system excision. Heat dissipation and compensation heat pulse techniques were used to measure sap flow in trunks and individual roots, respectively. Sap flux and measurements of xylem pressure potential (Ψ) were used to calculate K(plant) and K(root) in vines with zero and ～80% of roots severed. Whole vine transpiration (E), Ψ and K(plant) were significantly reduced within 24 h of root pruning, and did not recover within 6 weeks. Sap flux in intact roots increased within 24 h of root pruning, driven by an increase in the pressure gradient between the soil and canopy and without any change in root hydraulic conductance. Photosynthesis (A) and stomatal conductance (g(s)) were reduced, without significant effects on leaf internal CO(2) concentration (c(i)). Shoot growth rates were maintained; fruit growth and dry matter content were increased following pruning. The woody roots of kiwifruit did not demonstrate a rapid dynamic response to root system damage as has been observed previously in monocot seedlings. Increased sap flux in intact roots with no change in K(root) and only a moderate decline in shoot A suggests that under normal growing conditions root hydraulic conductance greatly exceeds requirements for adequate shoot hydration.     

Growth,morphology and photosynthetic activity in flooded <Emphasis Type="Italic">Alnus japonica</Emphasis> seedlings

This study was conducted on Alnus japonica seedlings subjected to flooding for 2, 4, and 6 weeks to examine responses in growth, morphology, and photosynthesis to different periods of flooding. Seedlings subjected to flooding for 2 and 4 weeks were drained after flooding then watered daily. Increases in biomass of leaves, roots, and whole plants were less for 6-week-flooded seedlings. Rate of photosynthesis and stomatal conductance of flooded seedlings decreased within 2 weeks. For 2-week-flooded seedlings recovery from reduced stomatal conductance and recovery of photosynthetic activity occurred after drainage. For the 6-week-flooded seedlings stomatal conductance recovered by the end of the experiment. Adventitious root formation by the 4 and 6-week-flooded seedlings was observed from the third week of flooding. These results suggest that recovery of reduced function in leaves may progress with development of adventitious roots during the period of flooding.     

Short-term effects of potassium fertilization on the hydraulic conductance of Laurus nobilis L

This study reports experimental evidence on the effect of short-term potassium fertilization on potassium uptake, tissue concentration and hydraulic conductance of pot-grown laurel plants. Potassium uptake and loading into the xylem of laurel seedlings increased within 24 h after fertilization. Potassium was not accumulated in roots and leaves, but the [K(+)] of xylem sap was 80% higher in fertilized plants (+K) than in potassium-starved plants (-K), as a likely result of recirculation between xylem and phloem. Increased xylem sap [K(+)] resulted in a 45% increase in transpiration rate, a 30% increase in plant hydraulic conductance (K(plant)) and a 120% increase in leaf-specific conductivity of the shoot (k(shoot)). We suggest that this increase was due to ion-mediated up-regulation of xylem hydraulics, possibly caused by the interaction of potassium ions with the pectic matrix of intervessel pits. The enhancement of hydraulic conductance following short-term potassium fertilization is a phenomenon that can be of advantage to plants for maintaining cell turgor, stomatal aperture and gas exchange rates under moderate drought stress. Our data provide additional support for the important role of potassium nutrition in agriculture and forestry.     

Water stress responses of seedlings of four Mediterranean oak species

Effects of water stress on phenology, growth, stomatal activity and water status were assessed from April to November 1996 in 2-year-old seedlings of Quercus frainetto Ten. (Quercus conferta Kit.), Quercus pubescens Willd., Quercus macrolepis Kotschy (Quercus aegilops auct.) and Quercus ilex L. growing in containers in northern Greece. All four species developed more than 50% of their total leaf area before the beginning of June--an adaptation to arid climates. Well-irrigated plants tended to develop greater individual leaf area, number of leaves per plant, total plant leaf area, height and root:shoot ratios than water-stressed plants, but the difference between treatments was not significant for any parameter in any species. Quercus macrolepis appeared to be the most drought-tolerant of the four species. It maintained the highest number of leaves of the smallest size and increased the proportion of fine roots during drought. In all species, drought caused significant decreases in stomatal conductance and predawn and midday water potentials from mid-July until the end of August, when the lowest soil water content and highest mean daily air temperatures and midday leaf temperatures occurred; however, the responses were species-specific. Among the four species, Quercus macrolepis sustained the highest stomatal conductance despite very low water potentials, thus overcoming drought by means of desiccation tolerance. Quercus ilex decreased stomatal conductance even before severe water stress occurred, thereby avoiding desication during drought. Quercus pubescens had the highest water potential despite a high stomatal conductance, indicating that its leaf water status was independent of stomatal activity. Quercus frainetto was the least drought-resistant of the four species. During drought it developed very low water potentials despite markedly reduced stomatal aperture.     

Biochemical responses to iron deficiency in kiwifruit (Actinidia deliciosa)

A comparative study of two kiwifruit genotypes (Actinidia deliciosa (A. Chev.) C.F. Liang et A.R. Ferguson var. deliciosa) with different tolerance to iron (Fe) deficiency was conducted to identify biochemical features associated with tolerance to Fe deficiency. After 14 days of growth in hydroponic culture under Fe-deficient and Fe-sufficient conditions, leaf chlorophyll concentration, activities of ferric chelate reductase (FCR), phosphoenolpyruvate carboxylase (PEPC) and citrate synthase in root extracts, concentrations of organic acids in roots, leaves and xylem sap, and xylem sap pH were measured. In response to Fe deficiency, the tolerant genotype D1 showed: (i) higher FCR activity associated with a longer lasting induction of FCR; (ii) higher PEPC activity; (iii) higher concentrations of citric acid in roots; and (iv) lower xylem sap pH compared with the susceptible genotype Hayward. These findings imply that induction of FCR and PEPC activities in roots in response to Fe deficiency are important physiological adaptations enabling Fe-efficient kiwifruit plants to tolerate Fe deficiency.     

Characterization of nitrogenous solutes in tissues and xylem sap of Leucaena leucocephala

Amino acid profiles of leaf, stem, and root tissues from nodulated and nonnodulated Leucaena leucocephala (Lam.) de Wit plants were determined by gas chromatography-mass spectrometry. High concentrations of mimosine and several other potentially toxic nonprotein amino acids, including pipecolic acid and two isomers of hydroxypipecolic acid, were identified in the tissues. Five metabolites remain unidentified. Of the foliar free amino acid nitrogen, 57-66% was associated with the potentially toxic amino acids. Major constituents in the leaf tissues of nonnodulated plants were mimosine and hydroxypipecolic acid (isomer 1). Mimosine was recovered in both the neutral plus basic and acidic amino acid fractions. Major differences between amino acid profiles of nodulated and nonnodulated roots were the low percentages of asparagine + aspartate (3.6% of the total pool compared to 33% in nodulated plants) and pipecolic acid in nonnodulated roots (1% of the total compared to 12.5% in nodulated plants). A novel plant betaine (dihydroxypipecolic acid betaine) was identified by fast-atom-bombardment mass spectrometry in leaf tissues, albeit at relatively low concentrations (< 1 micro mol per gram fresh weight). Analyses of the xylem sap collected from nodulated plants confirmed that Leucaena is an asparagine transporter, as suggested by the high concentrations of asparagine and the low concentrations of ureides in its root nodules. Amino acid profiles of xylem sap from nonnodulated plants showed extremely low concentrations of asparagine + aspartate (0.12 micro mol ml(-1)), whereas asparagine + aspartate was the major constituent (4.38 micro mol ml(-1)) in the xylem sap of nodulated plants. Two nonprotein amino acids, pipecolic acid and hydroxypipecolic acid, were major constituents of the xylem sap of nodulated and nonnodulated plants, respectively. Three unidentified compounds detected in xylem sap samples from both nodulated and nonnodulated plants did not correspond with any of the peaks characterized from tissue samples.     

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.     

Patterns of photosynthesis and starch allocation in seedlings of four bottomland hardwood tree species subjected to flooding

Effects of short-term (32 days) flooding on photosynthesis, stomatal conductance, relative growth rate and tissue starch concentrations of flood-intolerant Quercus alba L. (white oak), bottomland Quercus nigra L. (water oak), bottomland Fraxinus pennsylvanica Marshall. (green ash) and flood-tolerant Nyssa aquatica L. (water tupelo) seedlings were studied under controlled conditions. Net photosynthetic rates of flooded N. aquatica seedlings were reduced by 25% throughout the 32-day flooding period. Net photosynthetic rates of flooded Q. alba seedlings fell rapidly to 25% of those of the control seedlings by Day 4 of the flooding treatment and to 5% by Day 16. In F. pennsylvanica and Q. nigra, net photosynthetic rates were reduced to 50% of control values by Day 8 but remained at approximately 30 and 23%, respectively, of control values by Day 32. Leaves of flooded Q. alba seedlings accumulated approximately twice as much starch as leaves of non-flooded control plants, whereas root starch concentrations decreased to 67% of those of control plants by the end of the 32-day flooding treatment. In contrast, flooding caused only a small increase in leaf starch concentrations of N. aquatica plants, but it increased root starch concentrations to 119% of those of the control plants by the end of the experiment. The co-occurring bottomland species, Fraxinus pennsylvanica and Q. nigra, differed from each other in their patterns of stomatal conductance and root starch concentrations. We conclude that the maintenance of low leaf starch concentrations, and high pre-flood root tissue starch concentrations are important characteristics allowing flood-tolerant species to survive in flooded soils.     

Competition for water between walnut seedlings (Juglans regia) and rye grass (Lolium perenne) assessed by carbon isotope discrimination and delta18O enrichment

Container-grown walnut seedlings (Juglans regia L.) were subjected to competition with rye grass (Lolium perenne L.) and to a 2-week soil drying cycle. One and 2 weeks after the beginning of the drought treatment, H2 18O (delta approximately equals +100%) was added to the bottom layer of soil in the plant containers to create a vertical H2 18O gradient. Rye grass competition reduced aboveground and belowground biomass of the walnut seedlings by 60%, whereas drought had no effect. The presence of rye grass reduced the dry weight of walnut roots in the upper soil layer and caused a 50% reduction in lateral root length. Rye grass competition combined with the drought treatment reduced walnut leaf CO2 assimilation rate (A) and leaf conductance (gw) by 20 and 39%, respectively. Transpiration rates in rye grass, both at the leaf level and at the plant or tiller level, were higher than in walnut seedlings. Leaf intrinsic water-use efficiency (A/gw) of walnut seedlings increased in response to drought and no differences were observed between the single-species and mixed-species treatments, as confirmed by leaf carbon isotope discrimination measurements. Measurement of delta18O in soil and in plant xylem sap indicated that the presence of rye grass did not affect the vertical profile of soil water uptake by walnut seedlings. Walnut seedlings and rye grass withdrew water from the top and middle soil layers in well-watered conditions, whereas during the drought treatment, walnut seedlings obtained water from all soil layers, but rye grass took up water from the bottom soil layer only.