Drought resistance of Ailanthus altissima: root hydraulics and water relations

Drought resistance of Ailanthus altissima (Mill.) Swingle is a major factor underlying the impressively wide expansion of this species in Europe and North America. We studied the specific mechanism used by A. altissima to withstand drought by subjecting potted seedlings to four irrigation regimes. At the end of the 13-week treatment period, soil water potential was -0.05 MPa for well-watered control seedlings (W) and -0.4, -0.8 and -1.7 MPa for drought-stressed seedlings (S) in irrigation regimes S1, S2 and S3, respectively. Root and shoot biomass production did not differ significantly among the four groups. A progressively marked stomatal closure was observed in drought-stressed seedlings, leading to homeostasis of leaf water potential, which was maintained well above the turgor loss point. Root and shoot hydraulics were measured with a high-pressure flow meter. When scaled by leaf surface area, shoot hydraulic conductance did not differ among the treated seedlings, whereas root hydraulic conductance decreased by about 20% in S1 and S2 seedlings and by about 70% in S3 seedlings, with respect to the well-watered control value. Similar differences were observed when root hydraulic conductance was scaled by root surface area, suggesting that roots had become less permeable to water. Anatomical observations of root cross sections revealed that S3 seedlings had shrunken cortical cells and a multilayer endodermal-like tissue that probably impaired soil-to-root stele water transport. We conclude that A. altissima seedlings are able to withstand drought by employing a highly effective water-saving mechanism that involves reduced water loss by leaves and reduced root hydraulic conductance. This water-saving mechanism helps explain how A. altissima successfully competes with native vegetation.     

Root growth potential as an indicator of drought stress history

Container-grown quiescent Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) seedlings were air dried to plant water potentials of -0.2, -2.2 or -3.8 MPa (unstressed, moderate, and severe stress treatments, respectively). Trees from each treatment were either placed in root mist chambers held at 10, 20, or 28 degrees C for 28 days and root growth potential (RGP) and plant water potential (PWP) measured weekly, or potted in a 1/1 mix of peat and vermiculite, watered only once, and height growth and survival recorded after 10 weeks in an unheated greenhouse. Root growth potential of unstressed trees was greater than that of moderately stressed trees at all temperatures. Root growth potential of severely stressed trees was zero. Predawn plant water potentials of unstressed and moderately stressed trees were initially high, fell to -0.5 to -0.8 MPa, and then increased. Predawn plant water potential of severely stressed trees declined continuously over the 28-day experiment. Survival and height growth of the severely stressed trees were reduced compared to the unstressed and moderately stressed trees. Among the root growth potential measurements, RGP measured after 7 days at 10 degrees C was most sensitive to drought stress history and revealed differences in vigor that were not apparent from the survival and height growth data.     

Effects of drought and rewatering on growth and transpiration in European beech seedlings late in the growing season

Drought stress is one of the most important environmental factors affecting plant growth and survival. To date, most studies aim at understanding of post-stress physiological and anatomical adaptation to drought stress; however only few studies focus on plant recovery. In the present study, transpiration, shoot water potential, and anatomical and morphological measurements were performed on 4-year-old European beech seedlings with fully developed leaves. The seedlings were exposed to three levels of soil water potential (well-watered, moderate drought stress and severe drought stress) and followed by rewatering under greenhouse conditions. Reduced transpiration rates were observed in the stressed seedlings as a response to drought stress, whereas anatomical and morphological variables remained unchanged. Three days after rewatering, transpiration rates in both moderately and severely stressed seedlings recovered to the levels of those of well-watered seedlings. Drought stress promoted leaf budding, resulting in higher shoot dry mass of stressed seedlings. Our findings indicate that anatomical and morphological adaptations of European beech seedlings to drought stress are visibly limited during late-season growth stages. These results will help us to further understand factors involved in drought adaptation potential of European beech seedlings faced with expected climate-related environmental changes. To complete our findings, further experiments on plant recovery from drought stress should be focused on different periods of growing season.     

干旱胁迫对不同种源沙柳苗木水势和水分利用效率的影响

通过人工模拟方式对5个种源沙柳苗木进行干旱胁迫处理,研究干旱处理不同阶段苗木小枝水势日变化、叶片相对含水量、水分饱和亏缺、保水力及水分利用效率。结果表明:干旱胁迫下5个种源的沙柳苗木小枝水势和叶片相对含水量均下降,而水分利用效率在轻度干旱时呈上升趋势,随干旱胁迫的加剧而下降。干旱处理15天后,民勤种源的沙柳日平均小枝水势下降幅度最大为0.77 MPa,然后依次为达拉特旗0.57 MPa、盐池0.36 MPa、榆林0.25 MPa和乌审旗0.12 MPa,其中民勤种源的苗木在充分供水和干旱胁迫下小枝水势差异达到显著。干旱处理18天后叶片相对含水量分别下降:民勤1.68%、盐池7.03%、达拉特旗7.25%,榆林9.35%和乌审旗11.07%,水分饱和亏缺民勤<盐池<达拉特旗<榆林<乌审旗。结果表明,民勤种源的沙柳抗旱的能力较强,而盐池和达拉特旗沙柳苗木在受到水分胁迫时水分利用效率下降幅度较小,即水分胁迫对苗木干物质的积累影响较小,因此具有更高的经济价值。     

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.     

Effects of regulated deficit irrigation during the pre-harvest period on gas exchange, leaf development and crop yield of mature almond trees

We investigated the effects of regulated deficit irrigation (RDI) during the pre-harvest period (kernel-filling stage) on water relations, leaf development and crop yield in mature almond (Prunus dulcis (Mill.) D.A. Webb cv. Cartagenera) trees during a 2-year field experiment. Trees were either irrigated at full-crop evapotranspiration (ETc=100%) (well-irrigated control treatment) or subjected to an RDI treatment that consisted of full irrigation for the full season, except from early June to early August (kernel-filling stage), when 20% ETc was applied. The severity of water stress was characterized by measurements of soil water content, predawn leaf water potential (Psipd) and relative water content (RWC). Stomatal conductance (gs), net CO2 assimilation rate (A), transpiration rate (E), leaf abscission, leaf expansion rate and crop yield were also measured. In both years, Psipd and RWC of well-irrigated trees were maintained above -1.0 MPa and 92%, respectively, whereas the corresponding values for trees in the RDI treatment were -2.37 MPa and 82%. Long-term water stress led to a progressive decline in gs, A and E, with significant reductions after 21 days in the RDI treatment. At the time of maximum stress (48 days after commencement of RDI), A, gs and E were 64, 67 and 56% lower than control values, respectively. High correlations between A, E and gs were observed. Plant water status recovered within 15 days after the resumption of irrigation and was associated with recovery of soil water content. A relatively rapid and complete recovery of A and gs was also observed, although the recovery was slower than for Psipd and RWC. Severe water stress during the kernel-filling stage resulted in premature defoliation (caused by increased leaf abscission) and a reduction in leaf growth rate, which decreased tree leaf area. Although kernel yield was correlated with leaf water potential, RDI caused a nonsignificant 7% reduction in kernel yield and had no effect on kernel size. The RDI treatment also improved water-use efficiency because about 30% less irrigation water was applied in the RDI treatment than in the control treatment. We conclude that high-cropping almonds can be successfully grown in semiarid regions in an RDI regime provided that Psipd is maintained above a threshold value of -2 MPa.     

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。     

Survival and growth of drought hardened <Emphasis Type="Italic">Eucalyptus pilularis</Emphasis> Sm. seedlings and vegetative cuttings

Forestry requires low mortality of transplanted seedlings. Mortality shortly after planting is often associated with inadequate hydration of transplants. Seedlings can be hardened to the drought conditions they may experience after transplanting by exposing them to controlled drought conditions in the nursery. Eucalyptus pilularis Sm. seedlings were drought hardened by providing nil (severe treatment) or half (mild treatment) the daily irrigation routinely received (control treatment) for up to two non-consecutive days per week during the last 4 weeks of growth in the nursery. Drought hardening reduced stem diameter, seedling leaf area, leaf area per root biomass and seedling quality measured by the Dickson quality index, but increased root:shoot ratio. Hardened seedlings had lower stomatal conductance and leaf water potential on the days they received less irrigation that the control treatment. Hardened seedlings had greater stomatal conductance and were less water stressed than seedlings experiencing drought for the first time indicating hardened seedlings had adjusted physiologically to drought. Survival after transplanting in the controlled drought environment in a glasshouse was enhanced by the hardening treatments. Non hardened seedlings that had had their upper leaves manually removed immediately prior to transplanting to reduce leaf area (top-clipped) had similar survival to hardened seedlings. Stomatal conductance and leaf water potential after transplanting were higher in hardened and top-clipped seedlings than unhardened control seedlings or vegetative cuttings. Survival in the field trial was over 95% for all treatments, possibly as rain fell within 4 days of planting and follow-up rain occurred in the subsequent weeks. Neither the hardened or top-clipped seedlings planted in the field trial had reduced growth, increased propensity to form double leaders or worse stem form than control seedlings when measured at age 3 years.     

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.     

水分胁迫对白杨杂种无性系生理和生长的影响

为探索水分胁迫对白杨杂种无性系气体变换和生长的影响,以白杨双杂交杂种新无性系Ｂ４３０苗木为试验材料,在３种水分胁迫条件下,对各无性系生理和生长指标进行了分析。结果表明,在不同水分胁迫下,无性系苗木的Ｐｎ、Ｔｒ,Ｇｓ,Ｌａ,Ｃｉ等日变化模式基本一致,但受胁迫的苗木各生理过程受到明显抑制。各生长指标对水分胁迫的敏感性不同,苗高和单叶叶面积对水分胁迫最敏感,叶片数和生物量的敏感性较差,随着水分胁迫加强,     

Xylem sap flow and drought stress of Fraxinus excelsior saplings

We report on diurnal and seasonal variations in sap flow rate and stem water potential of Fraxinus excelsior L. saplings growing at the edge of a Fraxino-Aceretum forest in western Germany. Because of shallow soil, the trees were subjected to drought in summer. When soil water availability was not limiting, sap flow rate was related to changes in solar radiation and vapor pressure deficit. Maximum transpiration rates per leaf area were 3.5-7.4 mmol m-2 s-1, and maximum daily totals were 1.7-3.3 kg m-2 day-1. Under drought conditions, stem water potential dropped to midday minima of -2.6 to -3.5 MPa and sap flow rate was strongly related to this parameter. After the drought period, reduced apparent (whole-plant) hydraulic conductance was observed, which was attributed to a continued reduction in stomatal conductance after the drought stress had ceased. A model was developed that linked sap flow rate directly to climatic variables and stem water potential. Good correlation between measured and simulated sap flow rates allowed the model to be used for data interpretation.     

Effects of irrigation deprivation during the harvest period on leaf persistence and function in mature almond trees

In nut tree orchards in California, irrigation is typically withheld during the harvest period to reduce the likelihood of bark damage during mechanical shaking of the trees. The ensuing water stress, however, may result in premature defoliation and subsequent yield declines. Our objective was to establish and quantify the water stress resulting from irrigation deprivation and determine its impact on leaf function and persistence in mature almond trees (Prunus dulcis (Mill.) D.A. Webb cv. Nonpareil) during a 3-year field experiment. The severity of the water stress was characterized by measurements of predawn leaf (Psi(pd)) and midday stem (Psi(ms)) water potentials, stomatal conductance (gs), net CO2 assimilation rate (A) and leaf abscission. During 1995, Psi(ms) of fully irrigated (FI) trees was maintained above -1.0 MPa. In trees in the moderate- (MS) and severe-stress (SS) treatments, Psi(ms) was reduced to -1.4 to -2.0 MPa and -2.0 to -2.6 MPa, respectively. After 18 days of irrigation deprivation, A was reduced by 32 and 58% at midday and early afternoon, respectively, compared with morning values. A significant decrease in morning values of A only occurred after 30 days of irrigation deprivation. Water-use efficiency and A declined as evaporative demand increased from morning to afternoon. Assimilation also declined seasonally as leaves aged. Midday stem water potential was highly correlated with A, but less so with gs. The coefficient of determination between Psi(ms) and gs improved considerably when vapor pressure deficit and wind were multiply regressed with Psi(ms). Although A recovered rapidly when MS trees were irrigated, recovery in SS trees was slower and incomplete. Integrating the MS and SS effects for an extended period during 1995 resulted in 14 and 30% declines in A, and 6 and 20% declines in gs, respectively. The apparent Psi(ms) threshold for leaf abscission was -1.8 MPa. Daily canopy light interception declined with decreasing Psi(ms) as a result of premature defoliation (and perhaps altered leaf angles) from 67.9% in FI trees to 61.4 and 60.7% in MS and SS trees, respectively.     

Characteristics of water relations in seedling of Machilus yunnanensis and Cinnamomum camphora under soil drought condition

Introduction Soil drought not only influences agriculture and forestry production, but also brings much trouble for cultivating plants of gardening in the urban. Although the mean annual precipitation of Kunming City, China is 1079 mm, its distribution is…     

Physiological and morphological adaptations of the fruit tree Ziziphus rotundifolia in response to progressive drought stress

The physiological basis of drought resistance in Ziziphus rotundifolia Lamk., which is an important, multipurpose fruit tree of the northwest Indian arid zone, was investigated in a greenhouse experiment. Three irrigation regimes were imposed over a 34-day period: an irrigation treatment, a gradual drought stress treatment (50% of water supplied in the irrigation treatment) and a rapid drought stress treatment (no irrigation). Changes in gas exchange, water relations, carbon isotope composition and solute concentrations of leaves, stems and roots were determined. The differential rate of stress development in the two drought treatments did not result in markedly different physiological responses, but merely affected the time at which they were expressed. The initial response to decreasing soil water content was reduced stomatal conductance, effectively maintaining predawn leaf water potential (Psi(leaf)), controlling water loss and increasing intrinsic water-use efficiency, while optimizing carbon gain during drought. Carbon isotope composition (delta13C) of leaf tissue sap provided a more sensitive indicator of changes in short-term water-use efficiency than delta13C of bulk leaf tissue. As drought developed, osmotic potential at full turgor decreased and total solute concentrations increased in leaves, indicating osmotic adjustment. Decreases in leaf starch concentrations and concomitant increases in hexose sugars and sucrose suggested a shift in carbon partitioning in favor of soluble carbohydrates. In severely drought-stressed leaves, high leaf nitrate reductase activities were paralleled by increases in proline concentration, suggesting an osmoprotective role for proline. As water deficit increased, carbon was remobilized from leaves and preferentially redistributed to stems and roots, and leaves were shed, resulting in reduced whole-plant transpiration and enforced dormancy. Thus, Z. rotundifolia showed a range of responses to different drought intensities indicating a high degree of plasticity in response to water deficits.     

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.     

Leaf growth, photosynthesis and tissue water relations of greenhouse-grown Eucalyptus marginata seedlings in response to water deficits

Leaf growth, rate of leaf photosynthesis and tissue water relations of shoots of Eucalyptus marginata Donn ex Sm. (jarrah) seedlings were studied during a soil drying and rewatering cycle in a greenhouse experiment. Rates of leaf growth and photosynthesis were sensitive to water deficits. The rate of leaf growth decreased linearly with predawn leaf water potential to reach zero at -1.5 MPa. Rate of leaf growth did not recover completely within the first three days after rewatering. Midday photosynthetic rates declined to 40% of those of well-watered seedlings at a predawn leaf water potential of -1.0 MPa and reached zero at -2.2 MPa. Photosynthetic rate recovered rapidly following rewatering and almost fully recovered by the second day after rewatering. All tissue water relations parameters, except the bulk modulus of elasticity, changed significantly as the soil dried and recovered completely by the third day after rewatering. Changes in osmotic pressure at full turgor of 0.4 MPa indicated considerable capacity for osmotic adjustment. However, because there was little osmotic adjustment until predawn leaf water potential fell below -1.5 MPa, this capacity would not have enhanced seedling growth, although it may have increased seedling survival. The sensitivity of photosynthesis and relative water content to water deficits suggests that greenhouse-grown E. marginata seedlings behave like mesophytic plants, even though E. marginata seedlings naturally grow in a drought-prone environment.     

Geographical variation in water relations, hydraulic architecture and terpene composition of Aleppo pine seedlings from Italian provinces

Ecotypic variations in leaf conductance, soil-to-leaf hydraulic conductance, components of tissue water potential, hydraulic architecture parameters and xylem embolism were examined in greenhouse-grown two-year-old Aleppo pine (Pinus halepensis Mill.) seedlings from six origins representing the geographic range of the species in Italy. Cortical resin composition of the seedlings was also determined. Measurements were made on well-watered seedlings and on seedlings subjected to recurring severe drought. Drought-stressed seedlings had lower mean leaf conductances, transpiration rates and soil-to-leaf hydraulic conductances than well-watered seedlings. They also exhibited more negative osmotic potentials, higher relative water deficit at incipient plasmolysis, but a similar maximum modulus of elasticity. Drought-stressed seedlings showed a higher degree of xylem embolism, a lower Huber value, lower leaf specific conductivity and lower specific conductivity than well-watered seedlings. Drought-stressed seedlings of provenances from more xeric habitats (Tremiti, Porto Pino and Mottola) had greater leaf conductances, transpiration rates and soil-to-leaf hydraulic conductances than drought-stressed seedlings of provenances from more mesic habitats (Imperia, Otricoli and Vico del Gargano). They also showed higher osmotic adjustment and a lower degree of xylem embolism. Among provenances, there were no significant differences in hydraulic architecture parameters in response to the drought treatment; however, Tremiti and Porto Pino seedlings displayed smaller drought-induced reductions in specific conductivity and leaf specific conductivity, respectively, than seedlings from other provenances. These differences suggest that seedlings from xeric provenances, especially Tremiti, have greater resistance to desiccation than seedlings from mesic provenances. No clear association was found between terpene variability and the other traits investigated, although terpene composition was related to the geographical distribution of the provenances. We conclude that the drought-tolerance responses of Tremiti make it a more suitable provenance than the others for establishment on sites prone to severe soil water deficits.     

干旱胁迫对白蜡苗生理指标的影响

采用聚乙二醇（PEG-6000）模拟干旱胁迫处理白蜡幼苗,对过氧化氢酶（CAT）活性、丙二醛（MDA）含量和游离脯氨酸（PRO）含量等生理指标进行测定分析。结果表明,随着PEG-6000浓度的增大,MDA含量和PRO含量也增大,CAT活性先增大后降低,表现出较强的抵御不良环境胁迫的能力。白蜡幼苗能承受PEG-6000浓度为20％的中度胁迫（相当于-0．6MPa水势）,而不能承受30％的重度胁迫（相当于-1．2MPa水势）,确定白蜡为中度耐旱植物。     

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.     

Water relations characteristics of competing singleleaf pinyon seedlings and sagebrush nurse plants

Encroachment of singleleaf pinyon (Pinus monophylla) into adjacent low sagebrush (Artemisia arbuscula) and basin big sagebrush (Artemisia tridentata ssp. tridentata) communities may be enhanced by the efficient use of limited water resources by tree seedlings. Seedlings and sagebrush nurse plants were monitored over two growing seasons to determine water-use patterns. Predawn xylem water potential of low sagebrush declined rapidly, reaching −3.5 to −5.5 MPa by late summer. Big sagebrush values dropped to −2.0 to −3.0 MPa during summer drought. The drop in sagebrush xylem water potential was related to the decline in soil water potential (r=0.68 and 0.82). The change in pinyon predawn xylem water potential was moderate, declining to values of −1.5 to −2.5 MPa. An apparent diurnal threshold xylem water potential (−2.3 to −3.0 MPa) that results in stomatal closure enables pinyon seedlings to maintain a seasonally stable xylem water potential. Water use by pinyon seedlings declined by 50% from May to August in association with a reduction in stomatal conductance. Despite reduced stomatal conductance, sagebrush water use continued to increase during summer and reached levels up to five times greater (per unit leaf area) than associated pinyon. Pinyon seedlings appear to have greater drought avoidance than sagebrush nurse plants.