Responses of loblolly pine seedlings to elevated CO(2) and fluctuating water supply

Osmotic adjustment of loblolly pine (Pinus taeda L.) seedlings to fluctuating water supply in elevated CO(2) was investigated. Seedlings were grown in controlled-environment chambers in either 350 or 700 micro l l(-1) CO(2) with weekly watering for four months, after which they were either watered weekly (well-watered treatment) or every two weeks (water-stress treatment) for 59 days. Osmotic adjustment was assessed by pressure-volume analysis of shoots and by analysis of soluble carbohydrates and free amino acids in roots during the last drying cycle. In well-watered seedlings, elevated CO(2) increased the concentration of soluble sugars in roots by 68%. Water stress reduced the soluble sugar concentration in roots of seedling growing in ambient CO(2) to 26% of that in roots of well-watered seedlings. Elevated CO(2) mitigated the water stress-induced decrease in the concentration of soluble sugars in roots. However, this was probably due, in part, to carbohydrate loading during the first four months when all seedlings were grown in the presence of a high water supply, rather than to osmotic adjustment to water stress. Water stress caused a doubling in the concentration of free primary amino acids in roots, whereas elevated CO(2) reduced primary amino acid and nitrogen concentrations to 32 and 74%, respectively, of those in roots of seedlings grown in ambient CO(2). There was no indication of large-scale osmotic adjustment to water stress or that elevated CO(2) enhanced osmotic adjustment in loblolly pine.     

Carry-over effects of ozone on root growth and carbohydrate concentrations of ponderosa pine seedlings

Ozone exposure decreases belowground carbon allocation and root growth of plants; however, the extent to which these effects persist and the cumulative impact of ozone stress on plant growth are poorly understood. To evaluate the potential for plant compensation, we followed the progression of ozone effects, with particular emphasis on the development of new roots. Ponderosa pine (Pinus ponderosa Dougl. ex Laws.) seedlings were exposed to ozone for 2 years. Following removal of the seedlings from ozone, root growth was assessed to characterize the carry-over effects on new root production, and carbohydrate concentrations were measured to determine if allocation strategies differed among ozone treatments. Four months after removal from ozone, dormant seedlings had significantly lower starch concentrations in stems, coarse roots and fine roots than control seedlings. Following root flushing, starch concentrations in all seedlings decreased, with ozone-treated seedlings containing significantly less starch, sucrose, fructose, glucose and total monosaccharides than control seedlings. There was some evidence that stem starch was mobilized to compensate partially for the lower concentrations of root starch in ozone-treated seedlings; however, there was significantly less new root production in seedlings previously exposed to ozone for 2 years than in control seedlings. Early senescence of older needle age classes, perhaps resulting in inadequate available photosynthate, may be responsible for the reduction in new root production during the year following exposure to ozone. Stored carbohydrate reserves, which were depleted in seedlings previously exposed to ozone, were insufficient to compensate for the ozone-induced reduction in canopy photosynthate. We conclude that there are carry-over effects of ozone exposure on ponderosa pine seedlings, including an enhanced potential for seedling susceptibility to other stresses even in respite years when ozone concentrations are low.     

水分胁迫对云锦杜鹃光合特性日变化的影响

以盆栽5年生云锦杜鹃苗木为材料,分对照、轻度胁迫、中度胁迫、重度胁迫4组,研究了土壤水分胁迫对云锦杜鹃叶片光合特性日变化的影响。结果表明:随着水分胁迫的加重,云锦杜鹃的光合日变化曲线由典型的双峰型逐渐转变为峰值很小的单峰型,峰值的降低在轻度水分胁迫下由气孔限制引起,而在中度和重度水分胁迫下则由非气孔限制引起。轻度、中度和重度水分胁迫对云锦杜鹃光合特性日变化的影响差异显著。经逐步多元回归得到了净光合速率日变化的回归方程。     

Carbon partitioning and allocation in northern red oak seedlings and mature trees in response to ozone

Northern red oak (Quercus rubra L.) seedlings and trees differ in their response to ozone. Previous work reported reductions in net photosynthesis, carboxylation efficiency and quantum yield of mature tree leaves, whereas seedling processes were unaffected by the same ozone exposure. To further characterize differences in ozone response between seedlings and mature trees, we examined carbon partitioning and allocation in 32-year-old trees and 4-year-old seedlings of northern red oak after exposure to subambient (seasonal SUM00 dose (sum of all hourly ozone exposures) = 31 ppm-h), ambient (SUM00 dose = 85 ppm-h) and twice ambient (SUM00 dose = 151 ppm-h) ozone concentrations for three growing seasons. For mature trees, ozone exposure decreased foliar starch partitioning, increased starch partitioning in branches and increased (14)C retention in leaves. In contrast, starch partitioning in leaves and branches, and foliar (14)C retention in seedlings were unaffected by ozone exposure, but soluble carbohydrate concentrations in coarse and fine roots of seedlings were reduced. Differences in carbohydrate demand between seedlings and mature trees may underlie the higher leaf ozone uptake rates and greater physiological response to ozone in mature northern red oak trees compared with seedlings.     

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.     

Ecophysiological acclimatization to cyclic water stress in Eucalyptus

Drought is considered the main environmental factor limiting productivity in eucalyptus plantations in Brazil. However, recent studies have reported that exposure to water deficit conditions enables plants to respond to subsequent stresses. Thus, this study investigates the ecophysiological acclimatization of eucalyptus clones submitted to recurrent water deficit cycles. Eucalyptus seedlings were submitted to three recurrent water deficit cycles and anatomical, morphological and physiological changes were analyzed. The results were:(1) Eucalyptus seedlings responded to water deficits by directing carbohydrates to root and stem growth;(2) Size and number of stomata were reduced;(3) Stomatal conductance decreased which allowed the plants to reduce water losses through transpiration,increasing instantaneous water use efficiency;(4) The relationship between gas exchanges and available water contents allowed the seedlings to uptake the retained soil water athigher tensions;and,(5) Physiological recovery from subsequent water deficits became faster. As a result of these changes, the eucalyptus seedlings recovered from the same degree of water stress more rapidly.     

Tolerance of Japanese larch to drought is modified by nitrogen and water regimes during cultivation of container seedlings

Improving drought tolerance of container seedlings of Japanese larch is of high importance to afforestation. We hypothesized that adequate nitrogen (N) and limited water supply would increase the tolerance of container seedlings to water-deficit stress, circumventing photoinhibition, by means of (i) enhanced photosynthetic capacity with higher leaf N and (ii) decreased water loss from leaves with lower biomass allocation into aboveground parts. Container seedlings of Japanese larch were grown under the treatment combinations of adequate (+?N: 300 mg N container^?1) or limited (??N: 150 mg N container^?1) N and adequate (+?W: daily irrigation) or limited (??W: twice-a-week irrigation) water. Then, seedlings were subjected to a progressive drought treatment. Higher leaf N was observed in container seedlings grown under?+?N and???W. During progressive drought, lower stomatal conductance and net photosynthetic rate were observed in leaves with higher leaf N at a given predawn leaf water potential. Furthermore, the maximum efficiency of PSII photochemistry (F_v/F_m) was lower in leaves with higher leaf N, suggesting that higher leaf N might impair intrinsic tolerance to drought at the leaf level contrary to expectations. Conversely,???N and???W seedlings with lower shoot biomass delayed soil drying as a whole-plant response via a reduction in leaf transpiration, leading to delayed photoinhibition as indicated by a decline in F_v/F_m. To circumvent stress at the initial stage of water deficit, lower leaf N via limited N regime and smaller shoot biomass driven by limited N and water regimes would be important.

     

Decline in stomatal response to leaf water deficit in Corylus maxima cuttings

Many woody species can be propagated from leafy cuttings. However, following rooting, cuttings of Corylus maxima Mill. cv. Purpurea do not always survive the transition from a highly supportive rooting environment (e.g., fog) to a more natural environment where evaporative demand is higher. We found that it is not the supply of water to leaves, but stomatal dysfunction that leads to severe water deficits in the rooted cuttings. Two hours after well-rooted cuttings were transferred from the rooting environment, we were able to relate visible signs of leaf water deficit to high stomatal conductance (g(s)) and low relative water content (R). Small expanding leaves (L3) had unusually high g(s) and lower R than fully expanded leaves (L1). Although high cuticular conductances (g(c)) were occasionally observed in L3, SEM confirmed that increased total leaf conductance (g) was mainly a result of abnormally wide stomatal opening. We measured changes in the ability of stomata to control water loss during rooting by determining stomatal responsiveness to leaf water deficit in detached L1 and L3 harvested from cuttings during the first 75 days after severance from stock plants. Reduced stomatal responsiveness was observed within 7 days of severance, prior to adventitious root formation, and was more pronounced in L3 than in L1. A period of acclimatization after rooting (no leaf wetting, but a vapor pressure deficit of 0.20 kPa) reduced g(s) by 50% in L3 but not in L1, and partially restored stomatal responsiveness in L1 but not in L3. After rooting, the original leaves on the cutting retained substantial capacity for photosynthesis (e.g., in L1, 8 micromol m(-2) s(-1) at a photosynthetic photon flux density of 400 micromol m(-2) s(-1)). The implications of the results for post-rooting acclimatization procedures are discussed.     

Effects of soil water deficit on gas exchange characteristics and water relations of orchard lychee (Litchi chinensis Sonn.) trees

Eight-year-old lychee (Litchi chinensis Sonn.) trees, cv. 'Bengal,' growing in krasnozem soil were subjected to soil water deficit from one month before flowering until harvest by covering the ground with polyethylene sheeting and withholding irrigation. The ratio of daytime stomatal conductance of unirrigated to irrigated trees decreased 20% during the three months of increasing water deficit. Predawn leaf water potentials of irrigated trees averaged about -0.3 MPa throughout the period, whereas they declined progressively to -0.9 MPa in unirrigated trees. Minimum daytime leaf water potential in the unirrigated trees decreased from -1.0 to -1.1 MPa at the beginning of the drought period to -2.2 to -2.4 MPa after three months, and calculated whole-plant conductance did not change with decreasing availability of water. The calculated soil-root water potential declined to less than -1.0 MPa in unirrigated trees. Capacitance effects on the relationship between leaf water potential and transpiration were significant only at low transpiration rates. Although unirrigated trees reduced soil water content at 0-30 cm depths to an equivalent water potential of -1.0 MPa, fruit shedding was significantly less than in irrigated trees. Water deficit had no effect on the fresh weight of pericarp, but caused increased seed size and decreased fresh weight of flesh, resulting in fruit from unirrigated trees being 16% lower in total fresh weight per fruit than fruit from irrigated trees.     

Carbon dioxide enrichment improves growth, water relations and survival of droughted honey mesquite (Prosopis glandulosa) seedlings

Low water availability reduces the establishment of the invasive shrub Prosopis on some grasslands. Water deficit survival and traits that may contribute to the postponement or tolerance of plant dehydration were measured on seedlings of P. glandulosa Torr. var. glandulosa (honey mesquite) grown at CO(2) concentrations of 370 (ambient), 710, and 1050 micro mol mol(-1). Because elevated CO(2) decreases stomatal conductance, the number of seedlings per container in the elevated CO(2) treatments was increased to ensure that soil water content was depleted at similar rates in all treatments. Seedlings grown at elevated CO(2) had a greater root biomass and a higher ratio of lateral root to total root biomass than those grown at ambient CO(2) concentration; however, these seedlings also shed more leaves and retained smaller leaves. These changes, together with a reduced transpiration/leaf area ratio at elevated CO(2), may have contributed to a slight increase in xylem pressure potentials of seedlings in the 1050 micro mol mol(-1) CO(2) treatment during the first 37 days of growth (0.26 to 0.40 MPa). Osmotic potential was not affected by CO(2) treatment. Increasing the CO(2) concentration to 710 and 1050 micro mol mol(-1) more than doubled the percentage survival of seedlings from which water was withheld for 65 days. Carbon dioxide enrichment significantly increased survival from 0% to about 40% among seedlings that experienced the lowest soil water content. By increasing seedling survival of drought, rising atmospheric CO(2) concentration may increase abundance of P. glandulosa on grasslands where low water availability limits its establishment.     

Response of gas exchange rates in <Emphasis Type="Italic">Abies firma</Emphasis> seedlings to various additional stresses under chronic acid fog stress

In order to clarify the chronic influence of acid fog on the gas exchange rates of momi fir (Abies firma Sieb. et Zucc.) trees, we exposed them to simulated acid fog (pH 3) for 3 years. The composition of the acid fog was similar to that observed in a region where momi fir trees have been declining, and it contained organic acids. We then treated the firs with various additional stresses, such as drought, low temperature, fine root cutting, ozone exposure, soil acidification, nitrogen load, and rhizosphere aluminum stress. Under chronic exposure to acid fog, the momi fir seedlings exhibited a pattern of stomatal behavior whereby they excessively opened in summer and closed in winter. Furthermore, the stomata of these seedlings tended to open during drought stress, and their needles were visibly injured after ozone exposure. The net photosynthesis rates of the seedlings exposed to acid fog were regulated by their stomatal aperture, rather than directly by acid fog. These results suggest that acid fog exposure disturbs the control of stomatal function in the momi fir seedlings. In addition, we found that chronic acid fog exposure suppressed the decrease in net photosynthesis rate, due to its nitrogen load.     

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。     

Water stress and crop load effects on fruit fresh and dry weights in peach (Prunus persica)

Effects of water stress on fruit fresh and dry weights were investigated in peach trees, Prunus persica (L.) Batsch., with varying crop loads: light, moderate and heavy. In well-watered controls, tree water status was independent of crop load. In trees receiving reduced irrigation, the degree of water stress increased with increasing crop load. Water stress induced fruit fresh weight reductions at all crop loads. Fruit dry weight was not reduced by water stress in trees having light to moderate crop loads, indicating that the degree of water stress imposed did not affect the dry weight sink strength of fruit. Water-stressed trees with heavy crop loads had significantly reduced fruit dry weights, which were likely due to carbohydrate source limitations resulting from large crop carbon demands and water stress limitations on photosynthesis.     

Some ecophysiological characteristics of artà (Calligonum comosum Ľ Hérit) in response to drought stress

Water deficit is an important environmental factor restricting plant growth and photosynthesis. The effect of water deprivation on leaf water status, photosynthetic gas exchange, chlorophyll content and fluorescence parameters of artà (Calligonum comosum) was studied. Five-month-old artà seedlings, grown in pots in the open air, were subjected to one of four drought treatments (i.e., mild, moderate, severe and extreme drought stress) and compared to control seedlings (normal watering regime). Results show that leaf water potential, net photosynthesis, stomatal conductance, transpiration, photosynthetic pigment content (chlorophyll a and b) decreased with increasing levels of drought stress. Inactivation of the photosynthetic apparatus was accompanied by changes in the fluorescence characteristics, providing evidence that reduction of photosynthetic rate could be attributed to non-stomatal limitations. Alterations imply changes in photochemical conversion efficiency of photosystem II by which plants could reduce water transpiration or protect their photosynthetic apparatus from destruction. These adaptations are discussed in relation to the strategies developed to grow drought-resistant artà seedlings in desert environments.     

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

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

马尾松苗木叶绿素荧光参数及相对水分亏缺测定

对广东信宜、浙江江山、湖南汝城、重庆黔江和湖北大悟等5个种源的马尾松苗木进行干旱胁迫处理,并测定其叶绿素荧光参数Fv/Fm值及相对水分亏缺。结果表明:大部分种源马尾松苗木叶绿素荧光参数Fv/Fm值在中度或重度干旱胁迫下低于对照,相对水分亏缺在中度或重度干旱胁迫下高于对照,均表现出积极的抗旱反应。     

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.     

Periodic chilling exposure during nursery culture

Five week old white spruce [Picea glauca (Moench) Voss] seedlings were placed within controlled-temperature growth chambers, and subjected to one of four periodic, chilling treatments for approximately 16 weeks. The treatments differed in the frequency of 24 hour exposure to 5 °C the seedlings received, and ranged from no chilling (control), to once every two, three or four days. After 25 weeks of growth, the control seedlings were significantly taller than all chilling-exposed seedlings, but stem diameters were similar. Chilling did not affect either needle length or needle density compared to control seedlings. During a water deficit test, frequently chilling-exposed seedlings maintained significantly higher mid-day shoot water potentials under extreme water stress. Re-watering after the drought resulted in all chilling-treated seedlings recovering faster to pre-stress shoot water potential levels than the control seedlings. Measurements of new root growth after 21 days at 10 °C indicated that previous chilling exposure delayed or inhibited new root production.     

Effects of short-term ozone exposure and soil water availability on the carbon economy of juvenile Douglas-fir

Effects of ozone and soil water availability on partitioning and translocation of assimilates were studied in three-year-old Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) seedlings exposed, in separate experiments, to 0 and 106 or 0 and 514 micro g m(-3) ozone for 8 h day(-1) for 9 days. The dynamics of carbon from assimilated (14)CO(2) were followed. No interactions between ozone and soil water content were observed. Total net uptake of carbon was reduced by low soil water content, but was unaffected by ozone. Both ozone and low soil water content increased the amount of (14)C-photosynthates retained in the current-year needles. Total starch content in old and current-year needles was unaffected by ozone, but was reduced by low water availability. Translocation of carbon to the root-soil compartment was additively affected by ozone and low soil water content. The results suggest that dry periods in summer combined with high ozone concentrations cause the greatest reduction in the supply of carbon compounds to the root-soil compartment.     

水分胁迫对栓皮栎幼苗生理特性及生长的影响

通过盆栽实验，研究了在不同的水分条件下栓皮栎幼苗生理特性的变化和苗木生长量的变化。研究表明，在水分胁迫初期，随土壤含水量的降低，栓皮栎叶片的净光合速率、蒸腾速率下降，而水分利用效率提高。在水分胁迫末期，由于长期的干旱胁迫，水分条件最差的一组苗木已干枯死亡，土壤含水量从38．123％下降到20．323％时，净光合速率和蒸腾速率均下降，水分利用效率也下降。随着土壤含水量的降低，苗高、地径和生物量均有下降的趋势，其中生物量下降的幅度最大。