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.     

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.     

Atmospheric and soil drought reduce nocturnal conductance in live oaks

Nocturnal and daytime whole-canopy transpiration rate (E) and conductance (g = E/VPD, where VPD is leaf to air vapor pressure difference) were assessed gravimetrically in drought-treated and well-watered 3-year-old saplings of live oak species (Quercus series Virentes Nixon) from the southeastern USA (Quercus virginiana Mill.) and Central America (Q. oleoides Cham. and Schlecter). Our objectives were to: (1) quantify nocturnal and daytime E and g in a controlled environment; (2) determine the impact of severe drought on nocturnal E and g; and (3) examine whether unavoidable water loss through the epidermis could account for nocturnal water loss. We calculated daytime E during peak daylight hours (between 0930 and 1330 h) and nocturnal E during complete darkness (between 2200 and 0500 h). In addition to reducing E and g during the daytime, drought-treated plants reduced nocturnal E and g on a whole-canopy basis by 62-64% and 59-61%, respectively, and on a leaf-level basis by 27-28% and 19-26%, respectively. In well-watered plants, nocturnal g declined with increasing VPD, providing evidence for stomatal regulation of nocturnal transpiration. In drought-treated plants, g was low and there was no relationship between nocturnal g and VPD, indicating that water loss could not be reduced further through stomatal regulation. Both daytime and nocturnal g declined curvilinearly with predawn water potential for all plants, but nocturnal g was unrelated to predawn water potentials below -1 MPa. The reductions in daytime and nocturnal E and g during drought were associated with decreases in whole-plant and leaf hydraulic conductances. Observed nocturnal g was within the same range as epidermal conductance for oak species determined in previous studies under a range of conditions. Nocturnal E rose from 6-8% of daytime E for well watered plants to 19-20% of daytime E for drought-treated plants. These results indicate that, during drought, saplings of live oak species reduce g to a minimum through stomatal closure, and experience unavoidable water loss through the epidermis.     

Effects of phenology,water availability and seed source on loblolly pine biomass partitioning and transpiration

One-year-old loblolly pine (Pinus taeda L.) seedlings from four seed sources (Arkansas, Georgia, Texas and Virginia) grown in 1-m-deep sand-filled pits in two water regimes (well-watered and drought) were studied, to gain insight into the process of seedling establishment. Whole-plant transpiration was measured biweekly from July to December. Whole-plant harvests were conducted at 6-week intervals from April to December. Whole-plant transpiration and transpiration per unit leaf and root area were affected by treatment, seedlot and phenology. Seedlings of the Arkansas seedlot maintained significantly higher transpiration rates per unit leaf and root area during drought than seedlings of the Virginia, Georgia or Texas seedlots, but did not accumulate greater biomass. The high transpiration rates of the Arkansas seedlings were attributed to their deep root systems. Allometric relationships indicated that, relative to the whole plant, biomass allocation to needles of drought-treated seedlings was enhanced during the summer (allometric ratio 1.09), whereas allocation to roots was enhanced in the spring and fall (allometric ratios of 1.13 and 1.09, respectively). Relative to the whole plant, biomass allocation to needles of well-watered seedlings was enhanced throughout the experiment (allometric ratio of 1.16 declining to 1.05), whereas the allometric ratio of root to total biomass was 0.89 or less throughout. Allometric relationships also indicated variation in biomass partitioning to roots in three soil layers (0-30, 30-60 and 60-100 cm), which differed among harvests in each soil layer. Root growth in both well-watered and drought-treated seedlings was concentrated in the top soil layer in the spring, shifted to the middle and bottom soil layers in the summer, and then increased in the top soil layer in the fall. Compared with well-watered seedlings, drought-treated seedlings had higher rates of root growth in the bottom soil layer in the fall, a characteristic that would confer tolerance to future periods of limited soil water availability.     

Growth, nutrition and response to water stress of Pinus pinaster inoculated with ten dikaryotic strains of Pisolithus sp

Reconstituted dikaryons of Pisolithus sp. (Pers.) Coker & Couch from South Africa influenced growth parameters (shoot length, shoot/root ratio and leaf area), nutrition and physiological indicators (transpiration rate, stomatal conductance and xylem water potential) of maritime pine (Pinus pinaster Ait.) seedlings during drought and recovery from drought. Seedlings colonized with certain dikaryons were more sensitive to water stress and showed less mycorrhiza formation under water stress than seedlings colonized with other dikaryons. Control (uninoculated) seedlings were significantly smaller than those inoculated with dikaryons. Transpiration rate, stomatal conductance and xylem water potential varied among mycorrhizal treatments during the water stress and recovery periods. After rewatering, the controls and seedlings inoculated with dikaryon 34 x 20 had a weaker recovery in transpiration rate, stomatal conductance and xylem water potential than the other treatments and appeared to have experienced damage due to the water stress. Concentrations of various elements differed in the shoots of Pinus pinaster colonized by the various dikaryons. It is suggested that breeding of ectomycorrhizal fungi could constitute a new tool for improving reforestation success in arid and semi-arid zones.     

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。     

The response of Pinus sylvestris to drought: stomatal control of transpiration and hydraulic conductance

We investigated the impact of drought on the physiology of 41-year-old Scots pine (Pinus sylvestris L.) in central Scotland. Measurements were made of the seasonal course of transpiration, canopy stomatal conductance, needle water potential, xylem water content, soil-to-needle hydraulic resistance, and growth. Comparison was made between drought-treated plots and those receiving average precipitation. In response to drought, transpiration rate declined once volumetric water content (VWC) over the top 20 cm of soil reached a threshold value of 12%. Thereafter, transpiration was a near linear function of soil water content. As the soil water deficit developed, the hydraulic resistance between soil and needles increased by a factor of three as predawn needle water potential declined from -0.54 to -0.71 MPa. A small but significant increase in xylem embolism was detected in 1-year-old shoots. Stomatal control of transpiration prevented needle water potential from declining below -1.5 MPa. Basal area, and shoot and needle growth were significantly reduced in the drought treatment. In the year following the drought, canopy stomatal conductance and soil-to-needle hydraulic resistance recovered. Current-year needle extension recovered, but a significant reduction in basal area increment was evident one year after the drought. The results suggest that, in response to soil water deficit, mature Scots pine closes its stomata sufficiently to prevent the development of substantial xylem embolism. Reduced growth in the year after a severe soil water deficit is most likely to be the result of reduced assimilation in the year of the drought, rather than to any residual embolism carried over from one year to the next.     

Variation in drought resistance, drought acclimation and water conservation in four willow cultivars used for biomass production

Growth and water-use parameters of four willow (Salix spp.) clones grown in a moderate drought regime or with ample water supply were determined to characterize their water-use efficiency, drought resistance and capacity for drought acclimation. At the end of the 10-week, outdoor pot experiment, clonal differences were observed in: (1) water-use efficiency of aboveground biomass production (WUE); (2) resistance to xylem cavitation; and (3) stomatal conductance to leaf-specific, whole-plant hydraulic conductance ratio (g(st)/K(P); an indicator of water balance). Across clones and regimes, WUE was positively correlated with the assimilation rate to stomatal conductance ratio (A/g(st)), a measure of instantaneous water-use efficiency. Both of these water-use efficiency indicators were generally higher in drought-treated trees compared with well-watered trees. However, the between-treatment differences in (shoot-based) WUE were smaller than expected, considering the differences in A/g(st) for two of the clones, possibly because plants reallocated dry mass from shoots to roots when subject to drought. Higher root hydraulic conductance to shoot hydraulic conductance ratios (K(R)/K(S)) during drought supports this hypothesis. The same clones were also the most sensitive to xylem cavitation and, accordingly, showed the strongest reduction in g(st)/K(P) in response to drought. Drought acclimation was manifested in decreased g(st), g(st)/K(P), osmotic potential and leaf area to vessel internal cross-sectional area ratio, and increased K(R), K(P) and WUE. Increased resistance to stem xylem cavitation in response to drought was observed in only one clone. It is concluded that WUE and drought resistance traits are inter-linked and that both may be enhanced by selection and breeding.     

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.     

Mechanisms of drought response in Thuja occidentalis L. II. Post-conditioning water stress and stress relief

We examined the extent of osmotic adjustment and the changes in relative water content (RWC) and transpiration rate (i.e., relative stomatal function) that occur in water-deficit-conditioned 6-year-old Thuja occidentalis L. (eastern white cedar) trees in response to a severe drought. Trees conditioned by successive cycles of mild or moderate nonlethal water stress (conditioning) and nonconditioned trees were exposed to drought (i.e., -2.0 MPa predawn water potential) to determine if water deficit conditioning enhanced tolerance to further drought stress. Following drought, all trees were well watered for 11 days to evaluate how quickly osmotic potential, RWC and transpiration rate returned to preconditioning values. Both nonconditioned trees and mildly conditioned trees exhibited similar responses to drought, whereas moderately conditioned trees maintained higher water potentials and transpiration rates were 38% lower. Both conditioned and nonconditioned trees exhibited a similar degree of osmotic adjustment (-0.39 MPa) in response to drought relative to the well-watered control trees. The well-watered control trees, nonconditioned trees and mildly conditioned trees had similar leaf RWCs that were about 3% lower than those of the moderately conditioned trees. Following the 11-day stress relief, there were no significant differences in osmotic potential between the well-watered control trees and any of the drought-treated trees. Daily transpiration rates and water potential integrals (WPI) of all drought-treated trees approached those of the well-watered control trees during the stress relief period. However, the relationship between cumulative transpiration and WPI showed that previous exposure to drought stress reduced transpiration rates. Leaf RWC of the moderately conditioned trees remained slightly higher than that of the nonconditioned and mildly conditioned trees.     

Competition for soil water between perennial bunch-grass (Elymus glaucus B.B.) and blue oak seedlings (Quercus douglasii H. & A.)

The competition effects of the perennial bunch-grass (Elymus glaucus B.B.) on the growth and survival of the oak seedlings (Quercus douglasii H. & A.) were investigated. There were four levels of Elymus competition, replicated three times. The three densities ofElymus employed were zero (control), 50 (Low — ‘L’ -), 116 (Medium — ‘M’ -) and 199 (High — ‘H’ -) plants m^?2. Rates of soil water depletion, stomatal conductance, transpiration, shoot elongation and leaf expansion rates were measured between 23 March and 26 May 1988. Rates of soil water depletion, stomatal conductance and transpiration differed amongst the treatments and were higher in the control for the duration of the experiment. Shoot elongation rate (SER) and leaf expansion rate (LER) of blue oak seedling were directly related to soil water potentials. Zero values of LER rates for all treatments were observed at soil water potentials lower than?1.91 MPa, and concurrent reductions of stomatal conductance indicated stomatal closure due to the soil water deficit. In the control treatment, transpiration alone was not high enough to deplete soil moisture and to reduce LER of the oak seedlings. Leaf dessication occurred first in the H and M treatments (53% of seedlings dessicated) and two weeks later in the L treatment (37% dessicated) when the soil water potential was approximately ?4.0 MPa. The number of reproductive tillers and seed dry weight indicated thatElymus plants were under water stress from April 25 and concluded on May 25 with an early summer dormancy in all treatments. Data indicated that light intensity of 50% of ambient did not limit the development of oak seedlings. The results suggested that density of the perennial bunch-grassElymus glaucus lower than 50 plants m^?2 could allow survival and successful establishment of blue oak in understories.     

Effects of water stress on Haloxylon ammodendron seedlings in the desert region of Heihe inland river watershed, Gansu Province, China

The water relation and leaf gas exchange of saxoul (Haloxylon Ammodendron Bge, a C4 shrub) seedlings were studied under water stress in 2001. Saxoul seedlings maintained high transpiration when the soil moisture was above 11%. The seedlings were able to take up water from soil with above 6 % soil water content, which was the threshold level of soil moisture for seedlings. The relationship between transpiration and potential evaporation was linear for well-watered seedlings. The decrease of soil water availability led to different degrees of down-regulation of stomatal conductance, leaf transpiration and net CO2 assimilation rate. The stomata played a relatively small part in determining the net CO2 assimilation rate for the same seedling. The relationship between net CO2 assimilation rate and transpiration was linear diurnally, and reduction scale of leaf transpiration was much bigger than that of net CO2 assimilation rate by waters tress treatments, therefore intrinsic water-use-efficiency increased. High evaporative demand increased the leaf transpiration but inhibited net CO2 assimilation rate.Because of the effect of VPD on transpiration in this region, the transpiration of well-watered and mild water stress seedlings becomes responsive to change in stomatal conductance over a wider range.     

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.     

Regulation of transpirational water loss in Quercus suber trees in a Mediterranean-type ecosystem

Sap flux density in branches, leaf transpiration, stomatal conductance and leaf water potentials were measured in 16-year-old Quercus suber L. trees growing in a plantation in southern Portugal to understand how evergreen Mediterranean trees regulate water loss during summer drought. Leaf specific hydraulic conductance and leaf gas exchange were monitored during the progressive summer drought to establish how changes along the hydraulic pathway influence shoot responses. As soil water became limiting, leaf water potential, stomatal conductance and leaf transpiration declined significantly. Predawn leaf water potential reflected soil water potential measured at 1-m depth in the rhizospheres of most trees. The lowest predawn leaf water potential recorded during this period was -1.8 MPa. Mean maximum stomatal conductance declined from 300 to 50 mmol m(-2) s(-1), reducing transpiration from 6 to 2 mmol m(-2) s(-1). Changes in leaf gas exchange were attributed to reduced soil water availability, increased resistances along the hydraulic pathway and, hence, reduced leaf water supply. There was a strong coupling between changes in soil water content and stomatal conductance as well as between stomatal conductance and leaf specific hydraulic conductance. Despite significant seasonal differences among trees in predawn leaf water potential, stomatal conductance, leaf transpiration and leaf specific hydraulic conductance, there were no differences in midday leaf water potentials. The strong regulation of changes in leaf water potential in Q. suber both diurnally and seasonally is achieved through stomatal closure, which is sensitive to changes in both liquid and vapor phase conductance. This sensitivity allows for optimization of carbon and water resource use without compromising the root-shoot hydraulic link.     

Stomatal and leaf growth responses to partial drying of root tips in willow

Root tips of intact willow (Salix dasyclados Wimm., Clone 81-090) plants were partially dried by exposure to ambient greenhouse air and then kept in water-vapor-saturated air for up to 3 days. The drying treatment increased abscisic acid (ABA) concentrations in both the root tips subjected to drying and in the xylem sap, while it reduced leaf stomatal conductance and leaf extension rate. Despite the decrease in stomatal conductance, leaf water potentials were unaffected by the root drying treatment, indicating that the treatment reduced hydraulic conductivity between roots and foliage. After roots subjected to drying were returned to a nutrient solution or excised, ABA concentrations in the remaining roots and in the xylem sap, stomatal conductance of mature leaves and extension rate of unfolding leaves all returned to values observed in control plants. The 4-fold increase in xylem sap ABA concentration following the root drying treatment was not solely the result of reduced sap flow, and thus may be considered a potential cause, not merely a consequence, of the observed reduction in stomatal conductance.     

Variability of stomatal conductance in a small and isolated population of silver fir (Abies alba Mill.)

We analyzed the response to drought of 420 individuals from eight half-sib families from a small and isolated population of silver fir (Abies alba Mill.): 105 of them were kept in well-watered conditions as control while the remaining 315 were exposed to drought for 27 days. A model describing stomatal behavior derived from Monteith and developed in beech by Leonardi et al. was fitted to experimental transpiration data obtained simply from the difference between two daily pot weighings. The estimated parameters were maximum stomatal conductance, maximum transpiration in well-watered conditions and sensitivity to soil water deficit. The model worked well: convergence for all but four individuals and concordance between experimental and fitted data were good (R(2)=0.86). Inter-individual variability for all three estimated parameters was high and two of them (maximum stomatal conductance and sensitivity to soil water deficit) were significantly different among families, suggesting genetic control. Our results validate the simplified method used to evaluate individual stomatal parameters. We also show that in the small and isolated population of our study substantial adaptive variability remains, a crucial prerequisite to endure environmental conditions determined by climatic change foreseen for the next decades.     

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.     

Influence of soil water on the physiological and morphological components of plant water balance in Populus trichocarpa, Populus deltoides and their F(1) hybrids

Patterns of leaf growth, transpiration and whole-plant water balance in Populus trichocarpa, P. deltoides and their F(1) hybrids were studied during a soil drying cycle. Plant responses were analyzed during three distinct stages of dehydration. In stage I, the transpiration rate of drought-stressed plants remained constant and equal to that of well-watered plants even though soil water content declined by more than 40%. Stage II began as soil and plant water deficits induced stomatal closure. When soil water was expressed as a fraction of transpirable soil water, the transition from stage I to stage II occurred at soil water fractions of 0.35, 0.45 and 0.60 for P. trichocarpa, P. deltoides and their F(1) hybrids, respectively. Reductions in leaf growth coincided with the shift from stage I to stage II. As soil water declined further, decreases in relative transpiration and whole-plant leaf area were significantly greater in parental species than in F(1) hybrids. Inherent feedbacks controlling stomatal water loss and the maintenance and growth of leaf tissue appeared to differ between F(1) and parental genotypes in a pattern characteristic of an overdominant mode of inheritance.Stage III began once the ability of stomata to compensate for water loss had been exhausted. Substantial differences were found in plant survival during stage III, with F(1) hybrids surviving longer than parental species. Survival was more strongly correlated with the hydraulic conductivity of xylem tissues than with the dehydration tolerance of leaf tissues. Collectively, these responses suggest that F(1) hybrids were more drought resistant than either parental species and highlight the importance of whole-plant studies of functional relationships between plant growth, water balance and hydraulic conductivity.     

Element concentrations in the xylem sap of Picea abies (L.) Karst. seedlings extracted by various methods under different environmental conditions

We used a Scholander pressure chamber to assess the effects of various extraction methods under different environmental conditions on element concentrations in xylem sap of 3-year-old Picea abies (L.) Karst. seedlings. Sap from excised shoots contained higher element concentrations when extracted at low than at high over-pressures. When comparing plants differing in water status, we found that a high extraction over-pressure introduced a systematic error into the data. For example, in well-watered non-transpiring plants relative to unwatered transpiring plants, potassium concentrations were 70% higher in sap extracted at 0.1 MPa over-pressure, but only 10% higher in sap extracted at 1.0 MPa over-pressure. Moreover, treatment effects depended on the time of day when the sap was extracted. Increased water flux in transpiring plants relative to non-transpiring plants resulted in reduced xylem sap element concentrations when samples were collected after 9 h of transpiration, but not after 4 to 6 h of transpiration. Drought had little effect on xylem sap element concentrations, indicating that rates of element release into xylem conduits, element depletion by growing tissues, and water flow maintained a balance that may prevent nutrient stress during short-term drought.     

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.