Chlorophyll fluorescence and CO(2) assimilation of black spruce seedlings following frost in different temperature and light conditions

Effects of artificial frosts on light-saturated photosynthesis (A(max)) and ground, maximal and variable fluorescence variables (F(o), F(m), and F(v) and F(v)/F(m)) were monitored on 1-year-old foliage of black spruce seedlings (Picea mariana (Mill.) BSP) grown at high (25 degrees C), moderate (15 degrees C) and low (5 degrees C) temperatures and moderate (240 &mgr;mol m(-2) s(-1)) and low (80 &mgr;mol m(-2) s(-1)) irradiances. Photoinhibition of 1-year-old foliage was greater in seedlings grown in moderate light than in seedlings grown in low light. Photoinhibition increased with decreasing growth chamber temperature at both irradiances. Most changes in F(v)/F(m) were caused by changes in F(v). Exposure to -4 degrees C decreased both F(v)/F(m) and A(max) compared with control values. The effect of the -4 degrees C frost treatment was greater in seedlings grown in low light than in seedlings grown in moderate light, probably because seedlings grown in moderate light were already partially photoinhibited before the frost treatment. Following -4 degrees C treatment, neither F(v)/F(m) nor A(max) recovered in seedlings grown in low light. Light-saturated photosynthesis decreased with decreasing growth chamber temperature. Light-saturated photosynthesis was more sensitive to the -3 and -4 degrees C frost treatments in seedlings grown at 25 degrees C than in seedlings grown at 15 and 5 degrees C. The A(max) of seedlings grown at 15 degrees C was sensitive only to the -4 degrees C frost treatment, whereas A(max) of seedlings grown at 5 degrees C was not sensitive to any of the frost treatments. Recovery of A(max) following frost took longer in seedlings grown at high temperatures than in seedlings grown at low temperatures. For seedlings grown at the same temperature but under different irradiances, both A(max) and F(v)/F(m) reflected damage to the photosynthetic system following a moderate frost. However, for seedlings grown at the same irradiance but different temperatures, A(max) provided a more sensitive indicator of frost damage to the photosynthetic system than F(v)/F(m) ratio.     

Regulation of photosynthesis in interior spruce during water stress: changes in gas exchange and chlorophyll fluorescence

Photosynthetic response to water stress was analyzed in 1-year-old interior spruce (Picea glauca (Moench) Voss x P. engelmanni Parry hybrid complex) seedlings and emblings produced from somatic embryogenesis. Carbon dioxide uptake, oxygen evolution and chlorophyll fluorescence at 20 degrees C were monitored as predawn shoot water potential (Psi) decreased. Concurrently with stomatal closure, carbon assimilation declined rapidly as Psi decreased to -1.0 MPa. Oxygen evolution at 10,000 micro l CO(2) l(-1) declined continuously as Psi decreased to -1.6 MPa. At photon flux densities (PFD) above 50 micro mol m(-2) s(-1), photochemical efficiency of photosystem (PS) II observed during actinic light exposure (Phi(II), calculated as DeltaF/F(m)') decreased as Psi decreased. At the same PFDs, photochemical quenching (q(P)) declined with decreasing Psi and nonphotochemical quenching (q(N)) increased steadily. At PFDs below 50 micro mol m(-2) s(-1), major decreases in q(N) were not observed until Psi decreased below -1.6 MPa. We identified three phases of photosynthetic response to progressive water stress in interior spruce: a pronounced decline in gas exchange, subsequent photoprotective changes in chlorophyll fluorescence as primary photochemistry was down-regulated, and a decline in photochemical efficiency of dark-adapted needles.     

Photosynthetic traits around budbreak in pre-existing needles of Sakhalin spruce (Picea glehnii) seedlings grown under elevated CO2 concentration assessed by chlorophyll fluorescence measurements

To assess the effects of elevated CO(2) concentration ([CO(2)]) on the photosynthetic properties around spring budbreak, we monitored the total leaf sugar and starch content, and chlorophyll fluorescence in 1-year-old needles of Sakhalin spruce (Picea glehnii Masters) seedlings in relation to the timing of budbreak, grown in a phytotron under natural daylight at two [CO(2)] levels (ambient: 360?μmol mol(-1) and elevated: 720?μmol mol(-1)). Budbreak was accelerated by elevated [CO(2)] accompanied with earlier temporal declines in the quantum yield of PSII electron transport (Φ(PSII)) and photochemical quenching (q(L)). Plants grown under elevated [CO(2)] showed pre-budbreak leaf starch content twice as high with no significant difference in Φ(PSII) from ambient-CO(2)-grown plants when compared at the same measurement [CO(2)], i.e., 360 or 720?μmol mol(-1), suggesting that the enhanced pre-budbreak leaf starch accumulation might not cause down-regulation of photosynthesis in pre-existing needles under elevated [CO(2)]. Conversely, lower excitation pressure adjusted for the efficiency of PSII photochemistry ((1?-?q(P)) F(v)'/F(m)') was observed in plants grown under elevated [CO(2)] around budbreak when compared at their growth [CO(2)] (i.e., comparing (1?-?q(P)) F(v)'/F(m)' measured at 720?μmol mol(-1) in elevated-CO(2)-grown plants with that at 360?μmol mol(-1) in ambient-CO(2)-grown plants), which suggests lower rate of photoinactivation of PSII in the elevated-CO(2)-grown plants around spring budbreak. The degree of photoinhibition, as indicated by the overnight-dark-adapted F(v)/F(m), however, showed no difference between CO(2) treatments, thereby suggesting that photoprotection during the daytime or the repair of PSII at night was sufficient to alleviate differences in the rate of photoinactivation.     

Application of chlorophyll fluorescence to evaluate Mn tolerance of deciduous broad-leaved tree seedlings native to northern Japan

We used chlorophyll fluorescence to examine photosynthetic responses to excess Mn accumulation in leaves of four tree species differing in successional traits. Betula ermanii Cham. (Be) and Alnus hirsuta Turcz. (Ah) were studied as representatives of early-successional species. Ulmus davidiana Planch. var. japonica (Rehder) Nakai (Ud) was selected as a mid-successional species, and Acer mono Maxim. var. glabrum (Lév. et Van't.) Hara (Am) was chosen as a late-successional species. In Be, Ah and Am, high foliar concentrations of Mn had little effect on maximum photochemical efficiency of photosystem II (PSII), as indicated by the values of dark-adapted F(v)/F(m), whereas a significant decrease was observed in Ud. Photochemical quenching (qP) and the excitation capture efficiency of open PSII (F'(v)/F'(m)) decreased with increasing leaf Mn concentration at photosynthetic steady state after a 15-min exposure to 430 &mgr;mol m(-2) s(-1) PPFD. Compared with early-successional species, these decreases were greater in mid- and late-successional species leading to lower effective quantum efficiencies of PSII (DeltaF/F'(m) = qP x F'(v)/F'(m) = (F'(m) - F)/F'(m)). To determine the extent of photoinhibition, F(v)/F(m) of the illuminated leaves was remeasured after a 15-min dark period. Compared with the dark-adapted F(v)/F(m), we observed a significant decrease in F(v)/F(m) in Am leaves containing high concentrations of Mn. These chlorophyll fluorescence studies indicate that the early-successional species Be and Ah have a higher tolerance to excessive accumulations of Mn in leaves than the mid- and late-successional species Ud and Am.     

Photosynthetic induction responses to variable light under field conditions in three species grown in the gap and understory of a Fagus crenata forest

Photosynthetic induction responses to abrupt increases in photon flux density (PFD) to 800 and 1500 &mgr;mol m(-2) s(-1) from either darkness or 100 &mgr;mol m(-2) s(-1) were examined in situ in leaves of Fagus crenata Blume, Daphniphyllum humile Maxim., and Acer rufinerve Siebold & Zucc. growing in a gap and the understory of an F. crenata forest. Among the species studied, F. crenata exhibited the highest assimilation rate (A(100)), stomatal conductance (g(s100)) at the background PFD of 100 &mgr;mol m(-2) s(-1), and A(100)/A(max) (A(max) = maximum assimilation rate), in both the gap and the understory. Time required for full induction depended on both background PFD and maximum PFD. The induction period was 2-4-fold shorter at a background PFD of 100 &mgr;mol m(-2) s(-1) than in darkness. For the three understory species, time required to full induction was 2-3-fold longer when irradiance was increased from darkness to 800 &mgr;mol m(-2) s(-1) than when irradiance was increased from darkness to 1500 &mgr;mol m(-2) s(-1). Acer rufinerve showed higher initial stomatal conductance (g(s0)) and a shorter induction period in the understory than in the gap. Fagus crenata exhibited a similar g(s0) and induction period in both habitats. Daphniphyllum humile demonstrated lower g(s0) and a longer induction period in the understory than in the gap. These findings indicate that initial stomatal conductance is closely correlated with the photosynthetic induction response. We conclude that the photosynthetic induction response is affected by the light conditions experienced by plants before the sudden increase in irradiance and by the extent of the increase in irradiance.     

Seasonal photosynthetic gas exchange and leaf reflectance characteristics of male and female cottonwoods in a riparian woodland

Cottonwoods (Populus spp.) are dioecious phreatophytes of hydrological and ecological importance in riparian woodlands throughout the Northern Hemisphere. In streamside zones of southern Alberta, groundwater and soil water typically decline between May and September. To understand how narrowleaf cottonwoods (Populus angustifolia James) are adapted to this seasonal decrease in water availability, we measured photosynthetic gas exchange, leaf reflectance, chlorophyll fluorescence and stable carbon isotope composition (delta(13)C) in trees growing in the Oldman River valley of southern Alberta during the 2006 growth season. Accompanying the seasonal recession in river flow, groundwater table depth (Z(gw)) declined by 1.6 m, but neither mean daily light-saturated net photosynthetic rate (A(max)) nor stomatal conductance (g(s)) was correlated with this change. Both A(max) and g(s) followed a parabolic seasonal pattern, with July 24 maxima of 15.8 micromol m(-2) s(-1) and 559 mmol m(-2) s(-1), respectively. The early summer rise in A(max) was related to an increase in the chlorophyll pool during leaf development. Peak A(max) coincided with the maximum quantum efficiency of Photosystem II (F(v)/F(m)), chlorophyll index (CI) and scaled photochemical reflectance index (sPRI), but occurred one month after maximum volumetric soil water (theta(v)) and minimum Z(gw). In late summer, A(max) decreased by 30-40% from maximum values, in weak correlation with theta(v) (r(2) = 0.50). Groundwater availability limited late-season water stress, so that there was little variation in mean daily transpiration (E). Decreasing leaf nitrogen (% dry mass), CI, F(v)/F(m) and normalized difference vegetation index (NDVI) were also consistent with leaf aging effects. There was a strong correlation between A(max) and g(s) (r(2) = 0.89), so that photosynthetic water-use efficiency (WUE; A(max)/E) decreased logarithmically with increasing vapor pressure deficit in both males (r(2) = 0.75) and females (r(2) = 0.95). The male:female ratio was unequal (2:1, chi(2) = 16.5, P < 0.001) at the study site, but we found no significant between-sex differences in photosynthetic gas exchange, leaf reflectance or chlorophyll fluorescence that might explain the unequal ratio. Females tended to display lower NDVI than males (P = 0.07), but mean WUE did not differ significantly between males and females (2.1 +/- 0.2 versus 2.5 +/- 0.2 mmol mol(-1)), and delta(13)C remained in the -28.8 to -29.3 per thousand range throughout the growth season, in both sexes. These results demonstrate changes in photosynthetic and water-use characteristics that collectively enable vigorous growth throughout the season, despite seasonal changes in water supply and demand.     

Effect of elevated [CO(2)] and varying nutrient application rates on physiology and biomass accumulation of Sitka spruce (Picea sitchensis)

Sitka spruce (Picea sitchensis (Bong.) Carr.) seedlings were supplied with solutions containing nitrogen (N) at 0.1 x or 2 x the optimum rate (low-N and high-N supply, respectively) and grown either outside in a control plot or inside open-top chambers and exposed to ambient (355 &mgr;mol mol(-1)) or elevated (700 &mgr;mol mol(-1)) CO(2) concentration ([CO(2)]). Gas exchange measurements, chlorophyll determinations and nutrient analysis were made on current-year (< 1-year-old) shoots of the upper whorl after the seedlings had been growing in the [CO(2)] treatments for 17 months and the nutrient treatments for 6 months. Total seedling biomass and biomass allocation were assessed at the end of the experiment. Nutrient treatment had a significant effect on the light response curves, irrespective of [CO(2)] or chamber treatment; seedlings supplied with high-N rates had higher net photosynthetic rates than seedlings supplied with low-N rates. The degree of photosynthetic stimulation in response to elevated [CO(2)] was larger in seedlings receiving high-N rates than in seedlings receiving low-N rates. Light-saturated net photosynthesis of seedlings grown and measured in elevated [CO(2)] was 26% higher than that of seedlings grown and measured in ambient [CO(2)]. There was no significant effect of [CO(2)] or chamber treatment on the CO(2) response curves of seedlings receiving High-N supply rates. In contrast, analysis of the CO(2) response curves of seedlings receiving Low-N supply rates showed acclimation to elevated [CO(2)]. Both maximum rate of carboxylation (V(cmax)) and maximum electron transport capacity (J(max)) were lower and J(max)/V(cmax) higher in seedlings in the elevated [CO(2)] treatment. There was no effect of elevated [CO(2)] on stomatal conductance, although it was highly dependent on foliar [N], ranging from ~60 mmol m(-2) s(-1) at ~1.5 g N m(-2) to 200 mmol m(-2) s(-1) at ~5 g N m(-2). In the high-N and low-N treatments, foliar N concentration was 10 and 28% lower in seedlings grown in elevated [CO(2)] than in seedlings grown in ambient [CO(2)], respectively. There was no [CO(2)] effect on foliar phosphorus concentration ([P]). Chlorophyll concentration increased with increasing N supply in all treatments. There was no significant effect of elevated [CO(2)] on specific leaf area. Chlorophyll concentration expressed either on an area or dry mass basis for a given foliar [N] was higher in seedlings grown in elevated [CO(2)] than in seedings grown in ambient [CO(2)]. Elevated [CO(2)] increased total biomass accumulation by 37% in seedlings in the high-N treatment but had no effect in seedlings in the low-N treatment. There was a proportionally bigger allocation of biomass to roots of seedlings in the elevated [CO(2)] + low-N supply rate treatment compared with seedlings in other treatments. This resulted in a reduction in aboveground biomass compared with corresponding seedlings grown in ambient [CO(2)].     

Effect of elevated carbon dioxide concentration and root restriction on net photosynthesis, water relations and foliar carbohydrate status of loblolly pine seedlings

To determine the effects of CO(2)-enriched air and root restriction on photosynthetic capacity, we measured net photosynthetic rates of 1-year-old loblolly pine seedlings grown in 0.6-, 3.8- or 18.9-liter pots in ambient (360 micro mol mol(-1)) or 2x ambient CO(2) (720 micro mol mol(-1)) concentration for 23 weeks. We also measured needle carbohydrate concentration and water relations to determine whether feedback inhibition or water stress was responsible for any decreases in net photosynthesis. Across all treatments, carbon dioxide enrichment increased net photosynthesis by approximately 60 to 70%. Net photosynthetic rates of seedlings in the smallest pots decreased over time with the reduction occurring first in the ambient CO(2) treatment and then in the 2x ambient CO(2) treatment. Needle starch concentrations of seedlings grown in the smallest pots were two to three times greater in the 2x ambient CO(2) treatment than in the ambient CO(2) treatment, but decreased net photosynthesis was not associated with increased starch or sugar concentrations. The reduction in net photosynthesis of seedlings in small pots was correlated with decreased needle water potentials, indicating that seedlings in the small pots had restricted root systems and were unable to supply sufficient water to the shoots. We conclude that the decrease in net photosynthesis of seedlings in small pots was not the result of CO(2) enrichment or an accumulation of carbohydrates causing feedback inhibition, but was caused by water stress.     

Assimilation and stomatal conductance responses of red spruce to midwinter frosts and the constituent ions of acid mist

Red spruce (Picea rubens Sarg.) seedlings growing outside in open-top chambers were sprayed twice weekly with artificial mists at either pH 2.5 or 5.6, for five months during the 1988 growing season. The mists contained one of the following: water, pH 5.6 (control); (NH(4))(2)SO(4), pH 5.6; NH(4)NO(3), pH 5.6; HNO(3), pH 2.5; H(2)SO(4), pH 2.5; or (NH(4))(2)SO(4) + NH(4)NO(3), pH 2.5. During January 1989, the light responses of assimilation and stomatal conductance were assessed in the laboratory following a 4-day equilibration at 12 degrees C. The aerial portions of the intact trees were then subjected to a mild (-10 degrees C) frost for three hours during the night and the rate of recovery of light-saturated assimilation (A(max)) was determined the following day using the same branches as were used for the assimilation studies before the frost treatment. The same trees were then subjected to a second frost of -18 degrees C for three hours during the following night and the recovery of A(max) of the same branches was measured the next day. All of the acid mist treatments increased A(max) and apparent quantum yield relative to the control treatment when measured before the frost treatments. Frosts of -10 and -18 degrees C resulted in a significant decline in A(max) of seedlings in all treatments except the control. Stomatal conductance increased with increasing irradiance in seedlings in the acid mist treatments that did not contain SO(4) (2-) ion. Stomatal conductance of seedlings in acid mist treatments containing SO(4) (2-) ion was insensitive to changes in irradiance over the range 50-1500 micro mol m(-2) s(-1). It is concluded that acid precipitation increased the sensitivity of the assimilation response to midwinter frosts that follow a brief warm period. The SO(4) (2-) ion appears to be significant in causing increased sensitivity to frost and in causing stomatal insensitivity to light flux density.     

Physiological adjustment of two full-sib families of ponderosa pine to elevated CO(2)

Seeds from two full-sib families of ponderosa pine (Pinus ponderosa) with known differences in growth rates were germinated and grown in an ambient (350 micro l l(-1)) or elevated (700 micro l l(-1)) CO(2) concentration. Gas exchange at both ambient and elevated CO(2) concentrations was measured 1, 6, 39, and 112 days after the seed coat was shed. Initial stimulation of CO(2) exchange rate (CER) by elevated CO(2) was large (> 100%). On Day 1, CER of seedlings grown in elevated CO(2) and measured at ambient CO(2) was significantly lower than the CER of seedlings grown and measured at ambient CO(2), indicating physiological adjustment of the seedlings exposed to elevated CO(2). Physiological acclimation to elevated CO(2) was complete by Day 39 when there was no significant difference in CER between seedlings grown and measured at ambient CO(2) and seedlings grown and measured at elevated CO(2). After 4 months, the light response of seedlings in the two treatments was determined at both ambient and elevated CO(2). Light compensation point, CER at light saturation, and apparent quantum efficiency of seedlings grown and measured at ambient CO(2) were not significantly different from those of seedlings grown and measured at elevated CO(2). With a short-term increase in CO(2), CER at light saturation (5.16 +/- 0.52 versus 3.13 +/- 0.30 micro mol CO(2) m(-2) s(-1)) and apparent quantum efficiency (0.082 +/- 0.011 versus 0.045 +/- 0.003 micro mol CO(2) micro mol(-1) quanta) were significantly increased. Leaf C/N ratio was significantly increased in the elevated CO(2) treatment. There were few significant differences between families for any response to elevated CO(2). Under the experimental conditions, high growth rate was not correlated with a greater response to elevated CO(2).     

Analysis of the relationships among O(3) uptake, conductance, and photosynthesis in needles of Pinus ponderosa

We studied the effects of O(3) uptake on conductance (g(wv)) and photosynthesis (A) in needles of ponderosa pine (Pinus ponderosa Laws.) seedlings exposed for 70 days to one of three O(3) regimes-Low-O(3) (0.1 micro mol mol(-1) daily peak), High-O(3) (0.2 micro mol mol(-1) daily peak), and Low/High-O(3) (alternating 2 days Low-O(3) and 2 days High-O(3)). Seedlings exposed to charcoal-filtered air served as controls. Total O(3) exposures, expressed as ppm-h (the sum of the average hourly concentration in ppm ( micro mol mol(-1)) over the exposure period), were 77, 135, 105 and 4 for the Low-O(3), High-O(3), Low/High-O(3) and control treatments, respectively. Conductance (g(wv)) declined to about 60% of the value in control seedlings by Day 6 in seedlings in the High-O(3) treatment and by Day 37 in seedlings in the Low/High-O(3) treatment, but g(wv) did not decline at all in seedlings in the Low-O(3) treatment. At the end of the 70-day experiment, cumulative O(3) uptake, calculated from measured g(wv) values and assuming an internal O(3) concentration of zero, was 12.2, 13.5, and 14.7 mmol m(-2) for seedlings in the Low-O(3), Low/High-O(3), and High-O(3) treatments, respectively; however, O(3) uptake was reduced by 0, 24, and 36%, respectively, from that expected if there had been no decline in g(wv). With increasing total O(3) exposure, A declined, but the reduction was not strictly cumulative, i.e., A measured on Days 49 and 70 was similar for a given treatment even though both total O(3) exposure and uptake had increased. At the end of the experiment, A at near saturating CO(2) (1000 micro mol mol(-1)) and saturating photosynthetic photon flux density was reduced by about 25, 40 and 50% in seedlings in the Low-O(3), Low/High-O(3) and High-O(3) treatments, respectively, compared to the control seedlings. The ratio of internal to external CO(2) concentrations, an indicator of relative change in stomatal limitation of A, did not change over time and did not differ among treatments, suggesting that A and g(wv) decreased in parallel. After 40-60 days without O(3), A of seedlings in all O(3) treatments was not significantly different. Our data indicate that O(3)-induced stomatal closure was a result of reduced A and that decreased g(wv) reduced O(3) uptake to a rate that needles of ponderosa pine could tolerate without exhibiting further reductions in gas exchange capacity.     

Nutrient contents and concentrations in relation to growth of Picea abies and Fagus sylvatica along a European transect

Mineral nutrition of Norway spruce (Picea abies (L.) Karst.) and beech (Fagus sylvatica L.) was investigated along a transect extending from northern Sweden to central Italy. Nitrogen (N) concentrations of needles and leaves in stands growing on acid soils did not differ significantly between central Italy and southern Sweden (1.0 +/- 0.1 mmol N g(-1) for needles and 1.9 +/- 0.14 mmol N g(-1) for leaves). In both species, foliar N concentrations were highest in Germany (1.2 mmol N g(-1) for needles and 2.0 mmol N g(-1) for leaves) and decreased by 50% toward northern Sweden (0.5 mmol N g(-1)). Both species showed constant S/N and P/N ratios along the transect. Calcium, K and Mg concentrations generally reflected local soil conditions; however, Mg concentrations reached deficiency values in Germany. Leaf area per unit dry weight varied significantly along the transect with lowest values for Norway spruce recorded in northern Sweden and Italy (3.4 m(2) kg(-1)) and a maximum in central Europe (4.7 m(2) kg(-1)). A similar pattern was observed for beech. Despite the low variation in foliar N concentrations on the large geographic scale, local and regional variations in N concentrations equalled or exceeded the variation along the entire continental transect. Furthermore, nutrient contents (i.e., nutrient concentration x dry weight per needle or leaf) showed a greater variation than nutrient concentrations along the transect. Nitrogen contents of Norway spruce needles reached minimum values in northern Sweden (2.4 micro mol N needle(-1)) and maximum values in Denmark (5.0 micro mol N needle(-1)). The N content of beech leaves was highest in Denmark (242 micro mol N leaf(-1)). At the German site, foliar N content rather than N concentration reflected the seasonal dynamics of foliar growth and N storage of the two species. During foliage expansion, there was an initial rapid increase in N content and a decrease in N concentration. This pattern lasted for about 2 weeks after bud break and was followed by 6 weeks during which dry weight and N content of the foliage increased, resulting in a further decrease in N concentration. During summer, dry weight and N content of mature needles of Norway spruce increased further to reach a maximum in autumn, whereas N concentration remained constant. In spring, reallocation of N from 1- and 2-year-old needles was 1.5 and 1.0 micro mol N needle(-1), respectively. This remobilized N was a major source of N for the development of new needles, which had an N content of 1.5 micro mol N needle(-1) after bud break. The seasonal remobilization of N from old foliage decreased with increasing needle age. Needle N content and dry weight decreased progressively with age (1 micro mol N needle(-1) between age classes 2 and 5), whereas N concentrations remained constant. For Norway spruce, annual stemwood production was correlated with needle N content but not with foliar N concentration or with the total amount of N in the canopy. Interspecific and geographical differences in plant nutrition are discussed on the basis of competitive demands for C and N between growth of foliage and wood.     

Leaf physiology, production, water use, and nitrogen dynamics of the grassland invader Acacia smallii at elevated CO(2) concentrations

Invasion by woody legumes can alter hydrology, nutrient accumulation and cycling, and carbon sequestration on grasslands. The rate and magnitude of these changes are likely to be sensitive to the effects of atmospheric CO(2) enrichment on growth and water and nitrogen dynamics of leguminous shrubs. To assess potential effects of increased atmospheric CO(2) concentrations on plant growth and acquisition and utilization of water and nitrogen, seedlings of Acacia smallii Isely (huisache) were grown for 13 months at CO(2) concentrations of 385 (ambient), 690, and 980 micro mol mol(-1). Seedlings grown at elevated CO(2) concentrations exhibited parallel declines in leaf N concentration and photosynthetic capacity; however, at the highest CO(2) concentration, biomass production increased more than 2.5-fold as a result of increased leaf photosynthetic rates, leaf area, and N(2) fixation. Measurements of leaf gas exchange and aboveground biomass production and soil water balance indicated that water use efficiency increased in proportion to the increase in atmospheric CO(2) concentration. The effects on transpiration of an accompanying decline in leaf conductance were offset by an increase in leaf area, and total water loss was similar across CO(2) treatments. Plants grown at elevated CO(2) fixed three to four times as much N as plants grown at ambient CO(2) concentration. The increase in N(2) fixation resulted from an increase in fixation per unit of nodule mass in the 690 micro mol mol(-1) CO(2) treatment and from a large increase in the number and mass of nodules in plants in the 980 micro mol mol(-1) CO(2) treatment. Increased symbiotic N(2) fixation by woody invaders in response to CO(2) enrichment may result in increased N deposition in litterfall, and thus increased productivity on many grasslands.     

Selection of white spruce families in the context of climate change: heat tolerance

To assess the responses and plasticity of white spruce seedlings (Picea glauca (Moench) Voss) to high temperatures, 12 open-pollinated families differing in growth performance were exposed to a 30-min heat treatment of 42, 44, 46, 48, or 50 degrees C with or without heat preconditioning at 38 degrees C for 5 h. Damage was evaluated based on chlorophyll fluorescence parameters after heat preconditioning, after the heat treatments and during a 7-day recovery period. Visible needle damage was also evaluated after the heat treatments and 14 days later. Chlorophyll fluorescence parameters indicated that seedlings subjected to a heat treatment of 42-43 degrees C lost the ability to phosphorylate and donate water to photosystem II (PSII). A heat treatment of 44-46 degrees C severely limited the ability of the seedlings to use NADPH and ATP in the Calvin cycle. Based on visible needle damage, families with superior height-growth performance were more sensitive to heat stress than families with intermediate or inferior height-growth performance. Moreover, families with superior height-growth performance had low photochemical efficiencies in the light (DeltaF/F(m)') after heat treatment. Heat preconditioning increased the thermotolerance of the seedlings. However, the data suggest that white spruce seedlings exhibiting fast-growing characteristics under present conditions may not grow as well at higher temperatures.     

Seasonal variation in photosystem II efficiency and photochemical reflectance index of evergreen trees and perennial grasses growing at low and high elevations in subtropical Taiwan

Three pines species, three evergreen broadleaf trees, one C(3) and two C(4) perennial grasses of subtropical Taiwan were studied to elucidate the correlation between photosystem II (PSII) efficiency and photochemical reflectance index (PRI = (R(531) - R(570))/(R(531) + R(570))). Measurements were made at two sites differing in altitude (800 and 2600 m) over several growing seasons. At high elevation, potential PSII efficiency, measured by chlorophyll fluorescence (the ratio of variable to maximal fluorescence; F(v)/F(m)) at predawn, decreased with decreasing air temperature and varied greatly among species. At the lowest air temperature (-3 degrees C) studied, variation in F(v)/F(m) among species ranged from 0.33 to 0.72. In contrast, at low elevation where air temperature was moderate, seasonal variation in F(v)/F(m) was small in all of the study species. When species, elevation and season data were pooled, despite the high variation in F(v)/F(m) among species, a good correlation between F(v)/F(m) and PRI was observed. When compared at the same value of PRI, F(v)/F(m) of evergreen trees was higher than that of perennial grasses; however, when the minimum temperature on the measurement day was below 0 degrees C, F(v)/F(m) was underestimated relative to PRI. We conclude that PRI could be used as a remote indicator of photosynthetic function when air temperature is above 0 degrees C.     

The effects of ultraviolet-B radiation on photosynthesis of different aged needles in field-grown loblolly pine

We examined the effect of supplemental UV-B radiation (290-320 nm) on photosynthetic characteristics of different aged needles of 3-year-old, field-grown loblolly pine (Pinus taeda L.). Needles in four age classes were examined: I, most recently fully expanded, year 3; II, first flush, year 3; III, final flush, year 2; and IV, oldest needles still present, year 2. Enhanced UV-B radiation caused a statistically significant decrease (6%) in the ratio of variable to maximum fluorescence (F(v)/F(m)) following dark adaptation only in needles from the youngest age class, suggesting transient damage to photosynthesis. However, no effects of enhanced UV-B radiation on other instantaneous measures of photosynthesis, including maximum photosynthesis, apparent quantum yield and dark respiration, were seen for needles of any age. Foliar nitrogen concentration was unaffected by UV-B treatment. However, the (13)C/(12)C carbon isotope ratios (delta(13)C-a time integrated measure of photosynthetic function) of needles in age classes II and IV were 3% (P < 0.01) and 2% (P < 0.05) more negative, respectively, in treated plants than in control plants. Exposure to enhanced UV-B radiation caused a 20% decrease in total biomass and a 4% (P < 0.05), 25% (P < 0.01), and 9% (P < 0.01) decrease in needle length of needles in age classes I, II, and IV, respectively. The observed decreases in delta(13)C, and F(v)/F(m) of the needles in the youngest needle age class suggest subtle damage to photosynthesis, although overall growth reductions were probably a result of decreased total leaf surface rather than decreased photosynthetic capacity. Needles of age class IV had lower light- and CO(2)-saturated maximum photosynthetic rates (39%), lower dark respiration (34%), lower light saturation points (37%), lower foliar nitrogen concentration (28%), and lower delta(13)C (14%) values than needles of age class I. Apparent quantum yield and F(v)/F(m) did not change with needle age. The observed changes in photosynthesis and foliage chemical composition with needle age are consistent with previous studies of coniferous trees and may represent adaptations of older needles to shaded conditions within the canopy.     

Influence of irradiance on water relations and carbon flux during rooting of Shorea leprosula leafy stem cuttings

Single-node leafy stem cuttings of Shorea leprosula Miq. were subjected to a high, intermediate or low irradiance treatment for 16 weeks in an enclosed mist propagation system. Before rooting, maximum photosynthesis of the cuttings occurred at an irradiance of 400 micro mol m(-2) s(-1). Although none of the irradiance treatments affected the number of roots produced per cutting, the numbers of cuttings that formed roots were 50 and 30% in the high irradiance (diurnal range of 0-658 micro mol m(-2) s(-1)) and low irradiance (diurnal range of 0-98 micro mol m(-2) s(-1)) treatments, respectively, compared with 62% in the intermediate irradiance treatment (diurnal range of 0-360 micro mol m(-2) s(-1)). Low rooting frequency of cuttings in the high irradiance treatment was associated with water deficits (maximum leaf-to-air vapor pressure deficit (VPD) = 3.6 kPa), whereas cuttings in the low irradiance treatment had a low rooting frequency because they were below the light compensation point most of the time. In the intermediate irradiance treatment, cuttings withstood a daily maximum VPD of 1-2 kPa and recovered overnight from the previous day's deficit, as indicated by higher relative water content (RWC) and stomatal conductance (g(s)) in the morning than in the previous afternoon and evening. Higher RWC and g(s) of cuttings in all treatments on Days 14 and 21 compared with Day 8 probably indicated recovery from water deficit following severance and insertion of the cuttings in rooting medium. There were negative relationships between stem volume of cuttings and both number of cuttings that rooted and number of roots per cutting.     

Influence of photosynthetic photon flux density on growth and transpiration in seedlings of Fagus sylvatica

Beech seedlings (Fagus sylvatica L.) were grown in various combinations of three photosynthetic photon flux densities (PPFD, 0.7, 7.3 or 14.5 mol m(-2) day(-1)) for two years in a controlled environmental chamber. Dry mass of leaves, stem and roots, leaf area and number of leaves, and unit leaf rate were affected by both previous-year and current-year PPFD. Number of shoots and length of the main shoot were affected by previous-year PPFD but not by current-year PPFD. Number of leaves per shoot did not change with PPFD, whereas leaf dry mass/leaf area ratio was mainly affected by current-year PPFD. During the first 10 days that newly emerged seedlings were grown at a PPFD of 0.7 or 14.5 mol m(-2) day(-1), transpiration rate per unit leaf area declined. Thereafter, transpiration increased to a constant new rate. Transpiration rate per seedling was closely related to leaf area but the relationship changed with time. In two-year-old seedlings grown at various PPFD combinations of 0.7, 7.3 and 14.5 mol m(-2) day(-1) during Years 1 and 2, leaf area and transpiration rate per seedling were closely correlated at Weeks 7 and 11 after bud burst. Weak correlations were found between root dry mass and transpiration rate per seedling. During Year 2, transpiration rate per leaf area was higher at a particular PPFD in seedlings grown at a previous-year PPFD of 0.7 mol m(-2) day(-1) than in seedlings grown at a previous-year PPFD of 14.5 mol m(-2) day(-1). After transfer of two-year-old seedlings at the end of the experiment to a new PPFD (7.3 or 14.5 mol m(-2) day(-1)) for one day, transpiration rates per leaf area, measured at the new PPFD, were correlated with leaf area and root dry mass, irrespective of former PPFD treatment.     

Effects of ozone on growth and gas exchange of Eucalyptus globulus seedlings

To study the effects of a low concentration of ozone on growth and gas exchange in Eucalyptus globulus Labill. seedlings, ozone was applied for 37 days at a concentration of 50 ppb for 7 h daily under conditions of low light (250 micro mol m(-2) s(-1) (PAR)) and controlled temperature (20 degrees C). The seedlings exhibited extreme sensitivity to ozone. The ozone treatment reduced total plant biomass but had no effect on the partitioning of assimilate. Photosynthesis, stomatal conductance and internal CO(2) concentration were all reduced by ozone. The decline in photosynthesis was partly the result of direct effects of ozone on the stomata.     

Leaf CO(2) exchange of Erythrina poeppigiana (Leguminosae: Phaseolae) in humid tropical field conditions

An idealized model was developed to describe leaf CO(2) exchange in the leguminous tree Erythrina poeppigiana (Walpers) O.F. Cook under well-watered field conditions. Photosynthetic rate in mature leaves (p) was modeled as a rectangular hyperbolic function of photon flux density (q) and ambient CO(2) concentration (c(a)), relative photosynthetic capacity (pi) was modeled as a logistic s-function of leaf age (l(a)), metabolic dark respiration rate (r(m)) was modeled as an exponential function of leaf temperature (T(l)), and photorespiration rate (r(p)) was modeled as a hyperbolic function of c(a). Assimilation rate (a(c)) was modeled as the difference between the product of p and pi and the sum of r(m) and r(p): a(c) = p(q,c(a))pi(l(a)) - [r(m)(T(l)) + r(p)(c(a))]. The model parameters were estimated separately for five sources of E. poeppigiana (Clones 2660, 2662, 2687 and 2693 and half-sib Family 2431) from field data measured with a portable closed-loop gas exchange system at a humid tropical site in Costa Rica. The between-source differences in leaf CO(2) exchange characteristics were small, but statistically significant. Aboveground biomass production was highest in sources that maintained high relative photosynthetic capacity throughout the leaf life span. Quantum yield varied between 0.046 and 0.067, and light-saturated assimilation rate (q = 2000 micro mol m(-2) s(-1) and T(l) = 28 degrees C) at natural atmospheric c(a) (350 micro mol mol(-1)) was 16.8-19.9 micro mol m(-2) s(-1). Increasing c(a) to 1000 micro mol mol(-1) resulted in an approximate doubling of the light-saturated assimilation rate. Foliole nitrogen concentration, which was 45.3-51.2 mg g(-1) in mature leaves, was positively correlated with relative photosynthetic capacity. Foliole nitrogen concentration, quantum yield and maximum assimilation rate of E. poeppigiana are among the highest values observed in tropical woody legumes.