Light environment alters ozone uptake per net photosynthetic rate in black cherry trees

Foliar ozone uptake rates of different-sized black cherry (Prunus serotina Ehrh.) trees were compared within a deciduous forest and adjacent openings in north-central Pennsylvania during one growing season. Study trees included open-grown seedlings and saplings, forest understory seedlings and saplings, and sunlit and shaded portions of mature canopy tree crowns. Instantaneous ozone uptake rates were highest in high-light environments primarily because of higher stomatal conductances. Low ozone uptake rates of seedlings and saplings in the forest understory could be attributed partially to lower average ambient ozone concentrations compared to the canopy and open environments. Among the tree size and light combinations tested, ozone uptake rates were highest in open-grown seedlings and lowest in forest-grown seedlings. Despite lower ozone uptake rates of foliage in shaded environments, ozone uptake per net photosynthesis of foliage in shaded environments was significantly higher than that of foliage in sunlit environments because of weaker coupling between net photosynthesis and stomatal conductance in shaded environments. The potential for greater ozone injury in shaded environments as a result of greater ozone uptake per net photosynthesis is consistent with previous reports of greater ozone injury in shaded foliage than in sunlit foliage.     

Ethylenediurea (EDU) effects on hybrid larch saplings exposed to ambient or elevated ozone over three growing seasons

Ground-level ozone (O3) pollution is a persis-tent environmental issue that can lead to adverse effects on trees and wood production,thus indicating a need for forestry interventions to mediate O3 effects.We treated hybrid larch (Larix gmelinii var.japonica × L.kaempferi)saplings grown in nutrient-poor soils with 0 or 400 mg L-1 water solutions of the antiozonant ethylenediurea(EDU0,EDU400) and exposed them to ambient O3 (AOZ;08:00-18:00 ≈ 30 nmol mol-1) or elevated O3 (EOZ;08:00-18:00≈ 60 nmol mo1-1) over three growing sea-sons.We found that EDU400 protected saplings against most effects of EOZ,which included extensive visible foliar injury,premature senescence,decreased photosyn-thetic pigment contents and altered balance between pig-ments,suppressed gas exchange and biomass production,and impaired leaf litter decay.While EOZ had limited effects on plant growth (suppressed stem diameter),it decreased the total number of buds per plant,an effect that was not observed in the first growing season.These results indicate that responses to EOZ might have implications to plant competitiveness,in the long term,as a result of decreased potential for vegetative growth.However,when buds were standardized per unit of branches biomass,EOZ significantly increased the number of buds per unit of biomass,suggest-ing a potentially increased investment to bud development,in an effort to enhance growth potential and competitiveness in the next growing season.EDU400 minimized most of these effects of EOZ,significantly enhancing plant health under O3-induced stress.The effect of EDU was attributed mainly to a biochemical mode of action.Therefore,hybrid larch,which is superior to its parents,can be significantly improved by EDU under long-term elevated O3 exposure,providing a perspective for enhancing afforestation practices.     

Ecophysiology of shade needles of Picea glauca saplings in relation to removal of competing hardwoods and degree of prior shading

We studied shade needles of Picea glauca (Moench) Voss saplings that were growing in 15, 45 or 90% shade from competing vegetation at northern boreal forest sites in Alberta and Saskatchewan. At each site, in late May or early June 1990, all hardwoods were removed within a 2-m radius of each of eight saplings in each shade treatment (released saplings), and eight saplings in each shade treatment were left as controls. Light-saturated net assimilation (NA), stomatal and mesophyll conductance and water use efficiency of one-year-old needles were measured four times during the 1990 growing season and in the spring of 1991. There was a trend of increased photosynthetic capacity within one week following release. By August 1990 and in the following spring, NA was higher in released trees than in controls. The increase in NA appeared to be related to increased stomatal conductance to water vapor and to increased foliar nitrogen and resulting increases in mesophyll conductance to CO(2). There was no measurable effect of degree of shading prior to release on NA following release. Foliage of the released saplings appeared capable of rapid acclimation to the open conditions.     

Seasonal patterns of light-saturated photosynthesis and leaf conductance for mature and seedling Quercus rubra L. foliage: differential sensitivity to ozone exposure

Extrapolation of the effects of ozone on seedlings to large trees and forest stands is a common objective of current assessment activities, but few studies have examined whether seedlings are useful surrogates for understanding how mature trees respond to ozone. This two-year study utilized a replicated open-top chamber facility to test the effects of subambient, ambient and twice ambient ozone concentrations on light-saturated net photosynthesis (P(max)) and leaf conductance (g(l)) of leaves from mature trees and genetically related seedlings of northern red oak (Quercus rubra L.). Gas exchange measurements were collected four times during the 1992 and 1993 growing seasons. Both P(max) and g(l) of all foliage followed normal seasonal patterns of ontogeny, but mature tree foliage had greater P(max) and g(l) than seedling foliage at physiological maturity. At the end of the growing season, P(max) and g(l) of the mature tree foliage exposed to ambient ( approximately 80-100 ppm-h) and twice ambient ( approximately 150-190 ppm-h) exposures of ozone were reduced 25 and 50%, respectively, compared with the values for foliage in the subambient ozone treatment ( approximately 35 ppm-h). In seedling leaves, P(max) and g(l) were less affected by ozone exposure than in mature leaves. Extrapolations of the results of seedling exposure studies to foliar responses of mature forests without considering differences in foliar anatomy and stomatal response between juvenile and mature foliage may introduce large errors into projections of the response of mature trees to ozone.     

Long-term photosynthetic acclimation to increased atmospheric CO(2) concentration in young birch (Betula pendula) trees

To study the long-term response of photosynthesis to elevated atmospheric CO(2) concentration in silver birch (Betula pendula Roth.), 18 trees were grown in the field in open-top chambers supplied with 350 or 700 &mgr;mol mol(-1) CO(2) for four consecutive growing seasons. Maximum photosynthetic rates, stomatal conductance and CO(2) response curves were measured over the fourth growing season with a portable photosynthesis system. The photosynthesis model developed by Farquhar et al. (1980) was fitted to the CO(2) response curves. Chlorophyll, soluble proteins, total nonstructural carbohydrates, nitrogen and Rubisco activity were determined monthly. Elevated CO(2) concentration stimulated photosynthesis by 33% on average over the fourth growing season. However, comparison of maximum photosynthetic rates at the same CO(2) concentration (350 or 700 &mgr;mol mol(-1)) revealed that the photosynthetic capacity of trees grown in an elevated CO(2) concentration was reduced. Analysis of the response curves showed that acclimation to elevated CO(2) concentration involved decreases in carboxylation efficiency and RuBP regeneration capacity. No clear evidence for a redistribution of nitrogen within the leaf was observed. Down-regulation of photosynthesis increased as the growing season progressed and appeared to be related to the source-sink balance of the trees. Analysis of the main leaf components revealed that the reduction in photosynthetic capacity was accompanied by an accumulation of starch in leaves (100%), which was probably responsible for the reduction in Rubisco activity (27%) and to a lesser extent for reductions in other photosynthetic components: chlorophyll (10%), soluble protein (9%), and N concentrations (12%) expressed on an area basis. Despite a 21% reduction in stomatal conductance in response to the elevated CO(2) treatment, stomatal limitation was significantly less in the elevated, than in the ambient, CO(2) treatment. Thus, after four growing seasons exposed to an elevated CO(2) concentration in the field, the trees maintained increased photosynthetic rates, although their photosynthetic capacity was reduced compared with trees grown in ambient CO(2).     

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.     

Vertical profiles reveal impact of ozone and temperature on carbon assimilation of Betula pendula and Populus tremula

Rising temperature and tropospheric ozone (O(3)) concentrations are likely to affect carbon assimilation processes and thus the carbon sink strength of trees. In this study, we investigated the joint action of elevated ozone and temperature on silver birch (Betula pendula) and European aspen (Populus tremula) saplings in field conditions by combining free-air ozone exposure (1.2?×?ambient) and infrared heaters (ambient +1.2 °C). At leaf level measurements, elevated ozone decreased leaf net photosynthesis (P(n)), while the response to elevated temperature was dependent on leaf position within the foliage. This indicates that leaf position has to be taken into account when leaf level data are collected and applied. The ozone effect on P(n) was partly compensated for at elevated temperature, showing an interactive effect of the treatments. In addition, the ratio of photosynthesis to stomatal conductance (P(n)/g(s) ratio) was decreased by ozone, which suggests decreasing water use efficiency. At the plant level, the increasing leaf area at elevated temperature resulted in a considerable increase in photosynthesis and growth in both species.     

Diurnal and seasonal changes in the impact of CO(2) enrichment on assimilation, stomatal conductance and growth in a long-term study of Mangifera indica in the wet-dry tropics of Australia

We studied assimilation, stomatal conductance and growth of Mangifera indica L. saplings during long-term exposure to a CO(2)-enriched atmosphere in the seasonally wet-dry tropics of northern Australia. Grafted saplings of M. indica were planted in the ground in four air-conditioned, sunlit, plastic-covered chambers and exposed to CO(2) at the ambient or an elevated (700 micro mol mol(-1)) concentration for 28 months. Light-saturating assimilation (A(max)), stomatal conductance (g(s)), apparent quantum yield (phi), biomass and leaf area were measured periodically. After 28 months, the CO(2) treatments were changed in all four chambers from ambient to the elevated concentration or vice versa, and A(max) and g(s) were remeasured during a two-week exposure to the new regime. Throughout the 28-month period of exposure, A(max) and apparent quantum yield of leaves in the elevated CO(2) treatment were enhanced, whereas stomatal conductance and stomatal density of leaves were reduced. The relative impacts of atmospheric CO(2) enrichment on assimilation and stomatal conductance were significantly larger in the dry season than in the wet season. Total tree biomass was substantially increased in response to atmospheric CO(2) enrichment throughout the experimental period, but total canopy area did not differ between CO(2) treatments at either the first or the last harvest. During the two-week period following the change in CO(2) concentration, A(max) of plants grown in ambient air but measured in CO(2)-enriched air was significantly larger than that of trees grown and measured in CO(2)-enriched air. There was no difference in A(max) between trees grown and measured in ambient air compared to trees grown in CO(2)-enriched air but measured in ambient air. No evidence of down-regulation of assimilation in response to atmospheric CO(2) enrichment was observed when rates of assimilation were compared at a common intercellular CO(2) concentration. Reduced stomatal conductance in response to atmospheric CO(2) enrichment was attributed to a decline in both stomatal aperture and stomatal density.     

Genotypic variation in growth and physiological responses of Finnish hybrid aspen (Populus tremuloides x P. tremula) to elevated tropospheric ozone concentration

Saplings of six Finnish hybrid aspen (Populus tremuloides Michx. x P. tremula L.) clones were exposed to 0, 50, 100 and 150 ppb ozone (O3) for 32 days in a chamber experiment to determine differences in O3 sensitivity among genotypes. Based on the chamber experiment, three clones with intermediate sensitivity to O3 were selected for a free-air O3 enrichment experiment in which plants were exposed for 2 months to either ambient air (control) or air containing 1.3 x the ambient O3 concentration. We measured stem height and radial growth, number of leaves, dry mass and relative growth rate of leaves, stem and roots, visible leaf injuries, net photosynthesis and stomatal conductance of the clones. There was high clonal variation in susceptibility to O3 in the chamber experiment, indicated by foliar injuries and differential reductions in growth and net photosynthesis. In the free-air O3 enrichment experiment, ozone caused a shift in resource allocation toward stem height growth, thereby altering the shoot to root balance. In both experiments, low O3 concentrations tended to stimulate growth of most clones, whereas 100 and 150 ppb O3 in the chamber experiment impaired growth of most clones. However, growth of the most O3-tolerant clone was not significantly affected by any O3 treatment.     

Leaf photosynthetic characteristics of beech (Fagus sylvatica) saplings during three years of exposure to elevated CO(2) concentration

Beech (Fagus sylvatica L.) seedlings were cultivated from seeds sown in pots or directly in the ground in outdoor chambers that were transparent to solar radiation, and provided either ambient air or CO(2)-enriched air (ambient + 350 &mgr;mol mol(-1)). The rooting volume was high in all experiments. In the short-term experiment, potted plants were assigned to a factorial CO(2) x nutrient treatment (optimal nutrient supply and severe nutrient shortage) for 1 year. In the long-term experiment, plants were grown directly in the ground and received an optimal supply of water and nutrients in both CO(2) treatments for 3 years. Nutrient stress caused carboxylation capacity (V(m)) to decrease in the potted seedlings exposed to CO(2)-enriched air during their first growing season. In the long-term experiment with optimal nutrient supply, CO(2)-enriched air did not affect V(m), but caused an upward acclimation of maximum electron transport rate (J(m)). Consequently, there was a 14% increase in the J(m)/V(m) ratio, indicating nitrogen reallocation to maintain an equilibrium between RuBP consumption and RuBP regeneration. Both V(m) and J(m) decreased during the growing season in both CO(2) treatments. Although upward acclimation of J(m) was no longer apparent at the end of the third growing season, plants in CO(2)-enriched air maintained a higher J(m)/V(m) ratio than plants in ambient air, indicating that photosynthetic acclimation always occurred. Second flush leaves appeared during each growing season. When expressed on the basis of foliar nitrogen concentration, their photosynthetic characteristics (V(m) and J(m)) were enhanced compared with other leaves. Because the number of second flush leaves was also increased in the elevated CO(2) treatment, this response should be taken into account when modeling the effects of elevated CO(2) concentration on canopy photosynthesis. Stomatal conductance decreased in response to atmospheric CO(2) enrichment; however, the stomatal response to irradiance followed a single relationship based on two stomatal conductance models.     

Leaf water relations in <Emphasis Type="Italic">Pinus densiflora</Emphasis> Sieb. et Zucc. on different soil moisture conditions

To elucidate the differences in the leaf water relations of Pinus densiflora Sieb. et Zucc. growing in different soil moisture conditions, we examined the pressure-volume curve and the diurnal changes in the stomatal conductance, the transpiration rate, and the leaf water potential. The leaf water relations were compared using field-grown 40-year-old pine trees growing on the upper and lower parts of a slope. We also compared the leaf water relations of potted 4-year-old saplings growing at pF 4.2 and pF 1.8 soil moisture levels for almost 1 year. The values of the ratio of symplasmic water at turgor loss point to symplasmic water at saturated point (V_p/V_o) and bulk modulus of elasticity () of both the adult trees on the upper part of the slope and the potted saplings growing on pF 4.2 soil moisture were higher than those values of both the adult trees on the lower part of the slope and the potted saplings growing on pF 1.8 soil moisture, respectively. The field-grown adult tree and the potted saplings growing under long-term water stress tended to reduce their stomatal conductance in response to the acute soil drying. It is suggested that P. densiflora growing under long-term water stress rapidly closed its stomata in response to soil drying and avoided losing water, and could also rapidly absorb water with reducing water loss because of the decrease in the leaf pressure potential derived from the high values.     

Leaf-level and whole-plant gas exchange characteristics of shortleaf pine exposed to ozone and simulated acid rain

Field-grown shortleaf pine (Pinus echinata Mill.) seedlings were exposed to ozone (O(3)) and simulated acid rain (SAR) in open-top chambers over three growing seasons. Ranges of O(3) and SAR spanned ambient levels found in the southern USA. Effects of O(3) on leaf-level and whole-plant gas exchange were characterized for a single measurement period immediately before the third summer of exposure. Decreased photosynthesis rates were attributed to O(3), but not SAR. Stomatal conductance decreased in response to O(3) exposure, and either increased or was unaffected by SAR. Increased internal CO(2) concentration (c(i)) in response to O(3) treatment indicated a greater effect of O(3) on photosynthetic capacity than stomatal conductance. Whole-seedling gas exchange characteristics indicated that whole-seedling carbon assimilation was more severely affected by O(3) than was evident from leaf-level gas exchange characteristics. Seedlings exposed to O(3) retained fewer flushes than seedlings grown in charcoal-filtered air.     

Influence of foliar N on foliar soluble sugars and starch of red spruce saplings exposed to ambient and elevated ozone

Red spruce (Picea rubens Sarg.) trees growing at high elevation in the northeastern United States have experienced decline in recent years but seedlings have proved to be relatively tolerant of a wide range of environmental stresses in controlled studies. One possible reason for the wide tolerance to stress in seedlings is their inherently large pool of carbohydrate reserves, which is available for maintenance during and regrowth after periods of stress. We tested for the effects of foliar N and exposure to ozone on foliar carbohydrate reserves of 20-year-old naturally regenerated saplings. The trees were maintained in native soil in 360-l containers for 5 years before the experiment. The year before the experiment, trees were fertilized with N,P,K to provide a population of trees from N deficient to N sufficient. As foliar N decreased below 0.9%, length of current-year shoots and specific needle area of current-year needles declined. Foliar N concentration was correlated with foliar sugar and starch concentrations, but relationships varied with time of year. Before bud break, foliar carbohydrates and N, in general, were positively correlated, and date of bud break was delayed in N-deficient trees. During active growth, foliar soluble sugars and N were positively correlated, but starch concentrations were negatively correlated with N. By late September, neither starch nor sugar concentration was correlated with N concentration. Ozone and foliar N concentrations did not interact to change foliar carbohydrate concentrations or shoot and needle growth in this relatively short-term study.     

Seasonal air and soil temperature effects on photosynthesis in red spruce (Picea rubens) saplings

Net photosynthesis and stomatal conductance were measured in ten red spruce (Picea rubens Sarg.) saplings, growing near Ithaca, New York, throughout the early spring and late-fall growing periods. Gas exchange and daily minimum and maximum soil and air temperatures were also measured. Linear regression analysis showed that rates of net photosynthesis were positively correlated with both minimum daily soil and air temperatures but that minimum soil temperature was a better predictor of net photosynthesis. Moreover, net photosynthesis was more sensitive to changes in soil temperature than to changes in air temperature, and photosynthesis was approximately twice as sensitive to temperature changes during the fall than during the spring.     

Effects of foliar nitrogen concentration on photosynthesis and water use efficiency in Douglas-fir

Leaf-level physiological processes were studied in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) to determine whether apparent increases in stand-level water use efficiency (WUE) observed in response to nitrogen (N) fertilization were attributable to foliar N effects on carbon fixation rates or on stomatal control of water loss. Photosynthesis and transpiration were measured at different light intensities and ambient CO(2) molar fractions and comparisons were made between current-year shoots with average foliar N concentrations of 1.58% (High-N) and 1.25% (Low-N). Photosynthetic rates and foliar N concentrations were positively correlated. In response to light, photosynthesis and stomatal conductance were closely coupled and a similar coupling was observed in response to different ambient CO(2) concentrations. Partitioning the photosynthetic responses into mesophyll and stomatal components indicated that foliar N altered mesophyll conductance but not stomatal control of water loss. High-N shoots had significantly greater rates of photosynthesis and transpiration than Low-N shoots and, as a result, instantaneous WUE did not differ significantly between High-N and Low-N shoots.     

Testing the unifying theory of ozone sensitivity with mature trees of Fagus sylvatica and Picea abies

The broad range in plant responses to chronic O(3) exposure compels a search for integrative, underlying principles. One such approach is the unifying theory proposed by Reich (1987), which combines the O(3) response of contrasting physiognomic classes of plants on the basis of their intrinsic leaf diffusive conductance and, hence, capacity for O(3) uptake. Physiognomic classes differ in the proportional decline in photosynthesis and growth when compared on the basis of cumulative O(3) exposure per unit time, but converge when compared on the basis of O(3) uptake per unit time or cumulative O(3) uptake over the entire lifetime of the leaf. The theory is based on observations on a large number of contrasting plant species, relying primarily on studies of juvenile trees subjected to short-term O(3) exposure. To test the applicability of the unifying theory to mature trees, broadleaf deciduous European beech (Fagus sylvatica L.) and the evergreen conifer Norway spruce (Picea abies (L.) Karst.) in a mature mixed stand were exposed to either ambient air (control) or air with twice the ambient O(3) concentration delivered into the canopy by means of a free-air fumigation system. We accounted for differences in growing season length, leaf longevity and O(3)-related effects on leaf diffusive conductance in determining total O(3) uptake over the lifetime of the leaf. On this basis, Norway spruce needles required 5 years to take up as much O(3) as did beech leaves in one growing season. The core of the unifying theory on O(3) sensitivity was substantiated in relation to O(3) exposure and uptake. However, contrary to the unifying theory, which was formulated on the basis of results with juvenile trees, the O(3) response of mature trees in a natural stand was more complex. The increased complexity was attributed to additional environmental stressors, stress compensation at the whole-tree level, and differential O(3) sensitivities of leaves according to age class and position within the canopy. Contrary to the theory, photosynthesis was no less sensitive to O(3) in Norway spruce than that of beech, and was reduced in the twice-ambient O(3) regime in the first year of exposure.     

Effects of carbon dioxide, fertilization, and irrigation on photosynthetic capacity of loblolly pine trees

Branches of nine-year-old loblolly pine trees grown in a 2 x 2 factorial combination of fertilization and irrigation were exposed for 11 months to ambient, ambient + 175, or ambient + 350 micro mol mol(-1) CO(2). Rates of light-saturated net photosynthesis (A(max)), maximum stomatal conductance to water vapor (g(max)), and foliar nitrogen concentration (% dry mass) were assessed monthly from April 1993 until September 1993 on 1992 foliage (one-year-old) and from July 1993 to March 1994 on 1993 foliage (current-year). Rates of A(max) of foliage in the ambient + 175 CO(2) treatment and ambient + 350 were 32-47 and 83-91% greater, respectively, than that of foliage in the ambient CO(2) treatment. There was a statistically significant interaction between CO(2) treatment and fertilization or irrigation treatment on A(max) on only one measurement date for each age class of foliage. Light-saturated stomatal conductance to water vapor (g(max)) was significantly affected by CO(2) treatment on only four measurement dates. Light-saturated g(max) in winter was only 42% of summer g(max) even though soil water during winter was near field capacity and evaporative demand was low. Fertilization increased foliar N concentration by 30% over the study period when averaged across CO(2) treatments. During the study period, the ambient + 350 CO(2) treatment decreased average foliar N concentration of one-year-old foliage in the control, irrigated, fertilized and irrigated + fertilized plots by 5, 6.4, 9.6 and 11%, respectively, compared with one-year-old foliage in the corresponding ambient CO(2) treatments. The percent increase in A(max) due to CO(2) enrichment was similar in all irrigation and fertilization treatments and the effect persisted throughout the 11-month study period for both one-year-old and current-year foliage.     

Gas exchange characteristics of Populus trichocarpa, Populus deltoides and Populus trichocarpa x P. deltoides clones

Responses of net photosynthesis, dark respiration, photorespiration, transpiration, and stomatal conductance to irradiance, temperature, leaf-to-air vapor density difference (VDD), and plant water stress were examined in two Populus trichocarpa clones (one from a moist, coastal climate in western Washington and one from a dry, continental climate in eastern Washington), one P. deltoides clone, and two P. trichocarpa x P. deltoides clones. Light saturation of photosynthesis in greenhouse-grown trees occurred at about 800 micromol m(-2) s(-1) for P. deltoides, P. trichocarpa x P. deltoides, and the eastern Washington ecotype of P. trichocarpa, but at about 600 micromol m(-2) s(-1) for the western Washington ecotype of P. trichocarpa. Average net photosynthesis (at saturating irradiance and the optimum temperature of 25 degrees C) was 20.7, 18.8, 18.2 and 13.4 micromol CO(2) m(-2) s(-1) for P. deltoides, P. trichocarpa x P. deltoides, and the eastern and western Washington clones of P. trichocarpa, respectively. In all clones, net photosynthesis decreased about 14% as VDD increased from 3 to 18 g H(2)O m(-3). Stomatal conductance decreased sharply with decreasing xylem pressure potential (XPP) in all clones except the western Washington clone of P. trichocarpa. Stomata in this clone were insensitive to changes in XPP and did not control water loss. Complete stomatal closure (stomatal conductance < 0.05 cm s(-1)) occurred at about -2.0 MPa in the eastern Washington clone of P. trichocarpa and around -1.25 MPa in the P. deltoides and P. trichocarpa x P. deltoides clones. Transpiration rates were highest in the P. trichocarpa x P. deltoides clone and lowest in the western Washington clone of P. trichocarpa. The P. deltoides clone and eastern Washington clone of P. trichocarpa had the highest water use efficiency (WUE) and the western Washington clone of P. trichocarpa had the lowest WUE. The hybrids were intermediate. It was concluded that: (1) gas exchange characteristics of eastern and western Washington clones of P. trichocarpa reflected adaptation to their native environment; (2) crossing the western Washington clone of P. trichocarpa with the more drought resistant P. deltoides clone produced plants better adapted to the interior Pacific Northwest climate, although the stomatal response to soil water deficits in the hybrid was conservative compared with that of the eastern Washington clone of P. trichocarpa; and (3) introducing eastern Washington clones of black cottonwood into breeding programs is likely to yield lines with favorable growth characteristics combined with enhanced WUE and adaptation to soil water deficits.     

Carbon gain and bud physiology in Populus tremuloides and Betula papyrifera grown under long-term exposure to elevated concentrations of CO2 and O3

Paper birch (Betula papyrifera Marsh.) and three trembling aspen clones (Populus tremuloides Michx.) were studied to determine if alterations in carbon gain in response to an elevated concentration of CO(2) ([CO(2)]) or O(3) ([O(3)]) or a combination of both affected bud size and carbohydrate composition in autumn, and early leaf development in the following spring. The trees were measured for gas exchange, leaf size, date of leaf abscission, size and biochemical characteristics of the overwintering buds and early leaf development during the 8th-9th year of free-air CO(2) and O(3) exposure at the Aspen FACE site located near Rhinelander, WI. Net photosynthesis was enhanced 49-73% by elevated [CO(2)], and decreased 13-30% by elevated [O(3)]. Elevated [CO(2)] delayed, and elevated [O(3)] tended to accelerate, leaf abscission in autumn. Elevated [CO(2)] increased the ratio of monosaccharides to di- and oligosaccharides in aspen buds, which may indicate a lag in cold acclimation. The total carbon concentration in overwintering buds was unaffected by the treatments, although elevated [O(3)] decreased the amount of starch by 16% in birch buds, and reduced the size of aspen buds, which may be related to the delayed leaf development in aspen during the spring. Elevated [CO(2)] generally ameliorated the effects of elevated [O(3)]. Our results show that both elevated [CO(2)] and elevated [O(3)] have the potential to alter carbon metabolism of overwintering buds. These changes may cause carry-over effects during the next growing season.     

Effects of leaf age and tree size on stomatal and mesophyll limitations to photosynthesis in mountain beech (Nothofagus solandrii var. cliffortiodes)

Mesophyll conductance, g(m), was estimated from measurements of stomatal conductance to carbon dioxide transfer, g(s), photosynthesis, A, and chlorophyll fluorescence for Year 0 (current-year) and Year 1 (1-year-old) fully sunlit leaves from short (2 m tall, 10-year-old) and tall (15 m tall, 120-year-old) Nothofagus solandrii var. cliffortiodes trees growing in adjacent stands. Rates of photosynthesis at saturating irradiance and ambient CO(2) partial pressure, A(satQ), were 25% lower and maximum rates of carboxylation, V(cmax), were 44% lower in Year 1 leaves compared with Year 0 leaves across both tree sizes. Although g(s) and g(m) were not significantly different between Year 0 and Year 1 leaves and g(s) was not significantly different between tree heights, g(m) was significantly (19%) lower for leaves on tall trees compared with leaves on short trees. Overall, V(cmax) was 60% higher when expressed on the basis of CO(2) partial pressure at the chloroplasts, C(c), compared with V(cmax) on the basis of intercellular CO(2) partial pressure, C(i), but this varied with leaf age and tree size. To interpret the relative stomatal and mesophyll limitations to photosynthesis, we used a model of carbon isotopic composition for whole leaves incorporating g(m) effects to generate a surface of 'operating values' of A over the growing season for all leaf classes. Our analysis showed that A was slightly higher for leaves on short compared with tall trees, but lower g(m) apparently reduced actual A substantially compared with A(satQ). Our findings showed that lower rates of photosynthesis in Year 1 leaves compared with Year 0 leaves were attributable more to increased biochemical limitation to photosynthesis in Year 1 leaves than differences in g(m). However, lower A in leaves on tall trees compared with those on short trees could be attributed in part to lower g(m) and higher stomatal, L(s), and mesophyll, L(m), limitations to photosynthesis, consistent with steeper hydraulic gradients in tall trees.