Effects of carbon dioxide concentration and nutrition on photosynthetic functions of white birch seedlings

To investigate the interactive effects of atmospheric carbon dioxide concentration ([CO(2)]) and nutrition on photosynthesis and its acclimation to elevated [CO(2)], a two-way factorial experiment was carried out with two nutritional regimes (high- and low-nitrogen (N), phosphorus (P) and potassium (K)) and two CO(2) concentrations (360 and 720 ppm) with white birch seedlings (Betula papyrifera Marsh.) grown for four months in environment-controlled greenhouses. Elevated [CO(2)] enhanced maximal carboxylation rate (V(cmax)), photosynthetically active radiation-saturated electron transport rate (J(max)), actual photochemical efficiency of photosystem II (PSII) in the light (DeltaF/F(m)') and photosynthetic linear electron transport to carboxylation (J(c)) after 2.5 months of treatment, and it increased net photosynthetic rate (A(n)), photosynthetic water-use efficiency (WUE), photosynthetic nitrogen-use efficiency (NUE) and photosynthetic phosphorus-use efficiency (PUE) after 2.5 and 3.5 months of treatment, but it reduced stomatal conductance (g(s)), transpiration rate (E) and the fraction of total photosynthetic linear electron transport partitioned to oxygenation (J(o)/J(T)) after 2.5 and 3.5 months of treatment. Low nutrient availability decreased A(n), WUE, V(cmax), J(max), triose phosphate utilization (TPU), (/F(m)' - F)//F(m)' and J(c), but increased J(o)/J(T) and NUE. Generally, V(cmax) was more sensitive to nutrient availability than J(max). There were significant interactive effects of [CO(2)] and nutrition over time, e.g., the positive effects of high nutrition on A(n), V(cmax), J(max), DeltaF/F(m)' and J(c) were significantly greater in elevated [CO(2)] than in ambient [CO(2)]. In contrast, the interactive effect of [CO(2)] and nutrition on NUE was significant after 2.5 months of treatment, but not after 3.5 months. High nutrient availability generally increased PUE after 3.5 months of treatment. There was evidence for photosynthetic up-regulation in response to elevated [CO(2)], particularly in seedlings receiving high nutrition. Photosynthetic depression in response to low nutrient availability was attributed to biochemical limitation (or increased mesophyll resistance) rather than stomatal limitation. Elevated [CO(2)] reduced leaf N concentration, particularly in seedlings receiving low nutrition, but had no significant effect on leaf P or K concentration. High nutrient availability generally increased area-based leaf N, P and K concentrations, but had negligible effects on K after 2.5 months of treatment.     

Parameterization and testing of a biochemically based photosynthesis model for walnut (Juglans regia) trees and seedlings

The biochemically based leaf photosynthesis model proposed by Farquhar et al. (1980) and the stomatal conductance model proposed by Jarvis (1976) were parameterized for walnut. Responses of photosynthesis to CO(2) and irradiance were used to determine the key parameters of the photosynthesis model. Concurrently, stomatal conductance responses to leaf irradiance (Q), leaf temperature (T(l)), water vapor pressure deficit at the leaf surface (D), and air CO(2) concentration at the leaf surface (C(s)) were used to parameterize the stomatal conductance model. To test the generality of the model parameters, measurements were made on leaves from a 20-year-old tree growing in the field, and from sunlit and shaded greenhouse-grown seedlings. The three key parameters of the photosynthesis model (maximum carboxylation rate V(cmax), electron transport capacity J(max), and dark respiration rate R(d)) and the key parameter of the conductance model (reference stomatal conductance, g(sref)) were linearly correlated with the amount of leaf nitrogen per unit leaf area. Unique relationships could be used to describe nitrogen effects on these parameters for leaves from both the tree and the seedlings. Our data allowed separation of the effects of increasing total photosynthetic apparatus per unit leaf area from the effects of partitioning nitrogen among different pools of this apparatus for foliage acclimation to leaf irradiance. Strong correlations were found between stomatal conductance g(s) and Q, D and C(s), whereas the relationship between g(s) and T(l) was weak. Based on these parameterizations, the model adequately predicted leaf photosynthesis and stomatal conductance when tested with an independent set of data obtained for the tree and seedlings. Total light-driven electron flows derived from chlorophyll fluorescence data obtained at different leaf temperatures were consistent with values computed by the model. The model was also tested with branch bag data acquired from a three-year-old potted walnut tree. Despite a relatively large variance between observed and simulated values, the model predicted stomatal conductance and photosynthesis reasonably well at the branch scale. The results indicate that the photosynthesis-conductance model developed here is robust and can be applied to walnut trees and seedlings under various environmental conditions where water is non-limiting.     

Photosynthetic capacity in a central Amazonian rain forest

The vertical profile in leaf photosynthetic capacity was investigated in a terra firme rain forest in central Amazonia. Measurements of photosynthesis were made on leaves at five levels in the canopy, and a model was fitted to describe photosynthetic capacity for each level. In addition, vertical profiles of photosynthetic photon flux density, leaf nitrogen concentration and specific leaf area were measured. The derived parameters for maximum rate of electron transport (J(max)) and maximum rate of carboxylation by Rubisco (V(cmax)) increased significantly with canopy height (P < 0.05). The highest J(max) for a single canopy level was measured at the penultimate canopy level (20 m) and was 103.9 &mgr;mol m(-2) s(-1) +/- 24.2 (SE). The highest V(cmax) per canopy height was recorded at the top canopy level (24 m) and was 42.8 +/- 5.9 &mgr;mol m(-2) s(-1). Values of J(max) and V(cmax) at ground level were 35.8 +/- 3.3 and 20.5 +/- 1.3 &mgr;mol m(-2) s(-1), espectively. The increase in photosynthetic capacity with increasing canopy height was strongly correlated with leaf nitrogen concentration when examined on a leaf area basis, but was only weakly correlated on a mass basis. The correlation on an area basis can be largely explained by the concomitant decrease in specific leaf area with increasing height. Apparent daytime leaf respiration, on an area basis, also increased significantly with canopy height (P < 0.05). We conclude that canopy photosynthetic capacity can be represented as an average vertical profile, perturbations of which may be explained by variations in the environmental variables driving photosynthesis.     

Within-canopy nitrogen and photosynthetic gradients are unaffected by soil fertility in field-grown Eucalyptus globulus

A significant and well-supported hypothesis is that increased growth following nitrogen (N) fertilization is a function of the relationships among photosynthesis, tissue N content and the light environment-specifically, the within-canopy allocation of N among leaves and the within-leaf allocation of N between Rubisco and chlorophyll. We tested this hypothesis in a field trial that included annual applications of N,P,K fertilizer (from planting) to a Eucalyptus globulus Labill. plantation growing on uniform leached sands. Growth of 4-year-old E. globulus increased significantly in response to fertilization. Leaf N and phosphorus concentrations were 0.1-0.5 g m(-2) and 0.4-0.5 g m(-2) higher in fertilized trees compared to unfertilized trees, respectively. Stomatal conductance (g(s)) at the maximum photosynthetic rate (A(max)) was between 0.2 and 0.4 mol m(-2) s(-1) higher in fertilized trees, but A(max) and the concentration of Rubisco (Rub(a)) were unaffected by fertilization. This seeming paradox, where there was no response of A(max) to fertilization despite increases in g(s) and leaf N concentration, was explained by reduced in vivo specific activity of Rubisco in fertilized trees. Acclimation to light, measured by redistribution of N between Rubisco and chlorophyll, was unaffected by fertilization. Distribution of leaf N followed irradiance gradients, but A(max) did not. Maximum photosynthetic rate was correlated with leaf N concentration only in unfertilized trees. These findings indicate that the relationships among photosynthesis, N and the light environment in E. globulus are affected by N,P,K fertilization.     

Leaf-age effects on seasonal variability in photosynthetic parameters and its relationships with leaf mass per area and leaf nitrogen concentration within a Pinus densiflora crown

In the temperate zone of Japan, Pinus densiflora Sieb. et Zucc. bears needles of up to three age classes in the upper crown and up to five age classes in the lower crown. To elucidate the effects of leaf age on photosynthetic parameters and its relationships with leaf mass per unit area (LMA) and leaf nitrogen (N(l)) concentration on an area (N(a)) and mass (N(m)) basis, we measured seasonal variations in LMA, N(l), light-saturated photosynthetic rate (A(max)), stomatal conductance (g(s)), maximum rate of carboxylation (V(cmax)) and maximum rate of electron transport (J(max)) in leaves of all age classes in the upper and lower crown. Leaf mass per unit area increased by 27% with increasing leaf age in the lower crown, but LMA did not depend on age in the upper crown. Leaf age had a significant effect on N(m) but not on N(a) in both crown positions, indicating that decreases in N(m) resulted from dilution. Photosynthetic parameters decreased significantly with leaf age in the lower crown (39% for A(max) and 43% for V(cmax)), but the effect of leaf age was not as great in the upper crown, although these parameters exhibited seasonal variation in both crown positions. Regression analysis indicated a close relationship between LMA and N(a), regardless of age class or when each age class was pooled (r(2) = 0.57-0.86). Relationships between LMA and N(a) and among A(max), V(cmax) and J(max) were weak or not significant when all age classes were examined by regression analysis. However, compared with older leaves, relationships among LMA, N(a) and A(max) were stronger in younger leaves. These results indicate that changes in LMA and N(l) mainly reflect light acclimation during leaf development, but they are only slightly affected by irradiance in mature leaves. In conclusion, LMA and N(l) are useful parameters for estimating photosynthetic capacity, but age-related effects need to be taken into account, especially in evergreen conifers.     

Interpreting the decrease in leaf photosynthesis during flowering in mango

Little is known about the effect of flowering on leaf photosynthesis. To understand why net photosynthesis (A(net)) is lower in Mangifera indica L. leaves close to inflorescences than in leaves on vegetative shoots, we measured nitrogen and carbohydrate concentrations, chlorophyll a fluorescence and gas exchange in recently matured leaves on vegetative terminals and on floral terminals of 4-year-old trees. We used models to estimate photosynthetic electron fluxes and mesophyll conductance (g(m)). Lower A(net) in leaves close to developing inflorescences was attributable to substantial decreases in stomatal conductance and g(m), and also in photosynthetic capacity as indicated by the decrease in the light-saturated rate of photosynthetic electron transport (J(max)). The decrease in J(max) was the result of decreases in the amount of foliar nitrogen per unit leaf area, and may have been triggered by a decrease in sink activity as indicated by the increase in the hexose:sucrose ratio. Parameters measured on leaves close to panicles bearing set fruits were generally intermediate between those measured on leaves on vegetative shoots and on leaves close to inflorescences, suggesting that the changes in A(net) associated with flowering are reversible.     

Height-related decreases in mesophyll conductance, leaf photosynthesis and compensating adjustments associated with leaf nitrogen concentrations in Pinus densiflora

Hydraulic limitations associated with increasing tree height result in reduced foliar stomatal conductance (g(s)) and light-saturated photosynthesis (A(max)). However, it is unclear whether the decline in A(max) is attributable to height-related modifications in foliar nitrogen concentration (N), to mesophyll conductance (g(m)) or to biochemical capacity for photosynthesis (maximum rate of carboxylation, V(cmax)). Simultaneous measurements of gas exchange and chlorophyll fluorescence were made to determine g(m) and V(cmax) in four height classes of Pinus densiflora Sieb. & Zucc. trees. As the average height of growing trees increased from 3.1 to 13.7 m, g(m) decreased from 0.250 to 0.107 mol m(-2) s(-1), and the CO(2) concentration from the intercellular space (C(i)) to the site of carboxylation (C(c)) decreased by an average of 74 μmol mol(-1). Furthermore, V(cmax) estimated from C(c) increased from 68.4 to 112.0 μmol m(-2) s(-1) with the increase in height, but did not change when it was calculated based on C(i). In contrast, A(max) decreased from 14.17 to 10.73 μmol m(-2) s(-1). Leaf dry mass per unit area (LMA) increased significantly with tree height as well as N on both a dry mass and an area basis. All of these parameters were significantly correlated with tree height. In addition, g(m) was closely correlated with LMA and g(s), indicating that increased diffusive resistance for CO(2) may be the inevitable consequence of morphological adaptation. Foliar N per unit area was positively correlated with V(cmax) based on C(c) but negatively with A(max), suggesting that enhancement of photosynthetic capacity is achieved by allocating more N to foliage in order to minimize the declines in A(max). Increases in the N cost associated with carbon gain because of the limited water available to taller trees lead to a trade-off between water use efficiency and photosynthetic nitrogen use efficiency. In conclusion, the height-related decrease in photosynthetic performance appears to result mainly from diffusive resistances rather than biochemical limitations.     

Increased photosynthesis following partial defoliation of field-grown Eucalyptus globulus seedlings is not caused by increased leaf nitrogen

Increased photosynthetic rates following partial defoliation may arise from changes in leaf biochemistry, water relations or nutrient status. Twelve-month-old field-grown Eucalyptus globulus Labill. seedlings were pruned from below to reduce the green crown depth by 50 (D50) or 70% (D70). Photosynthetic responses to light and CO2 concentration were examined before and one, three and five weeks after partial defoliation. One week after defoliation, photosynthetic rates were greater in seedlings in the D50 (21 micromol m(-2) s(-1)) and D70 (23 micromol m(-2) s(-1)) treatments than in control seedlings (15 micromol m(-2) s(-1)); however, there was little difference in photosynthetic rates between partially defoliated seedlings and control seedlings after 5 weeks. An analysis of the sensitivity of photosynthesis to biochemical parameters revealed that the transient increase in photosynthetic rate in response to partial defoliation was largely a function of the maximum rate of carboxylation (85-87%) and the maximum rate of RuBP regeneration (55-60%) rather than stomatal conductance (12-13%). Nitrogen increased in leaves following partial defoliation (increases of 0.6 and 1.2 g m(-2) for D50 and D70, respectively), but was accumulated in a non-photosynthetic form (i.e., there was no increase in nitrogen concentration of Rubisco or chlorophyll). Increased photosynthetic rates immediately following partial defoliation were primarily a result of increased activity rather than amount of photosynthetic machinery. There was no evidence that phosphorus was responsible for the increase in photosynthetic rates after partial defoliation.     

Seasonal fluctuations and temperature dependence in photosynthetic parameters and stomatal conductance at the leaf scale of Populus euphratica Oliv

A combined model to simulate CO? and H?O gas exchange at the leaf scale was parameterized using data obtained from in situ leaf-scale observations of diurnal and seasonal changes in CO? and H?O gas exchange. The Farquhar et al.-type model of photosynthesis was parameterized by using the Bayesian approach and the Ball et al.-type stomatal conductance model was optimized using the linear least-squares procedure. The results show that the seasonal physiological changes in photosynthetic parameters (e.g., V(cmax25), J(max25), R(d25) and g(m25)) in the biochemical model of photosynthesis and m in the stomatal conductance model should be counted in estimating long-term CO? and H?O gas exchange. Overall, the coupled model successfully reproduced the observed response in net assimilation and transpiration rates.     

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)].     

Photosynthetic responses of cottonwood seedlings grown in glacial through future atmospheric [CO2] vary with phosphorus supply

Plants often exhibit proportionately larger photosynthetic responses to the transition from glacial to modern [CO(2)] than from modern to future [CO(2)]. Although this pattern may reflect increased nutrient demand with increasing [CO(2)], few studies have examined the role of nutrient supply in regulating responses to the range of [CO(2)] from glacial to future [CO(2)]. In this study, we examined the effects of P supply (0.004-0.5 mM) on photosynthetic responses of Populus deltoides (cottonwood) seedlings to glacial (200 micromol mol(-1)), modern (350 μmol mol(-1)) and future (700 micromol mol(-1)) [CO(2)]. The A(sat) (light-saturated net photosynthetic rates at the growth [CO(2)]) response to future [CO(2)] decreased with decreasing P supply such that there was no response at the lowest P supply. However, P supply did not affect A(sat) responses to an increase from glacial to modern [CO(2)]. Photosynthetic capacity [e.g., final rubisco activity, apparent, maximal Rubisco-limited rate of photosynthesis (V(cmax)), apparent, maximal electron transport-limited rate of photosynthesis (J(max))], stomatal conductance (g(s)) and leaf P generally increased with increasing P supply but decreased with increasing [CO(2)]. Measures of carbohydrate sink capacity (e.g., leaf mass per unit leaf area, leaf starch) increased with both increasing P supply and increasing [CO(2)]. Changes in V(cmax) and g(s) together accounted for 78% of the variation in A(sat) among [CO(2)] and P treatments, suggesting significant biochemical and stomatal controls on photosynthesis. However, A(sat) responses to increasing [CO(2)] did not reflect the changes in the carbohydrate sink capacity. These results have important implications because low P already constrains responses to increasing [CO(2)] in many ecosystems, and our results suggest that the P demand will increasingly affect A(sat) in cottonwood as [CO(2)] continues to increase.     

Water relations and gas exchange of Acer saccharum seedlings in contrasting natural light and water regimes

Field measurements were made of leaf photosynthesis (A), stomatal conductance (g) and leaf water relations for sugar maple (Acer saccharum Marsh.) seedlings growing in a forest understory, small gap or large clearing habitat in southwestern Wisconsin, USA. Predawn water status, leaf gas exchange and plasticity in field and laboratory water relations characteristics were compared among contrasting light environments in a wet year (1987) and a dry year (1988) to evaluate possible interactions between light and water availability in these habitats. Leaf water potentials (Psi(leaf)) at predawn and midday were lower for clearing than gap or understory seedlings. Acclimation of tissue osmotic potentials to light environment was observed among habitats but did not occur within any of the habitats in response to prolonged drought. During a summer drought in 1988, decreases in daily maximum g (g(max)) and maximum A (A(max)) in clearing seedlings were correlated with predawn Psi(leaf), which reached a seasonal minimum of -2.0 MPa. Under well-watered conditions, diurnal fluctuations in Psi(leaf) of up to 2.0 MPa in clearing seedlings occurred along with large midday depressions of A and g. In a wet year, strong stomatal responses to leaf-to-air vapor pressure difference (VPD) in sunny habitats were observed over nine diurnal courses of gas exchange measurements on seedlings in a gap and a clearing. Increasing stomatal limitations to photosynthesis appeared to be responsible for the reduction in A at high VPD for clearing seedlings. In understory seedlings, however, low water-use efficiency and development of leaf water deficits in sunflecks was related to reduced stomatal limitations to photosynthesis relative to seedlings in sunny habitats. Predawn Psi(leaf) and VPD appear to be important factors limiting carbon assimilation in sugar maple seedlings in light-saturating irradiances, primarily through stomatal closure. The overall results are consistent with the idea that sugar maple seedlings exhibit "conservative" water use patterns and have low drought tolerance. Leaf water relations and patterns of water use should be considered in studies of acclimation and species photosynthetic performance in contrasting light environments.     

Seasonal changes in photosynthesis and growth of Zizyphus attopensis seedlings in three contrasting microhabitats in a tropical seasonal rain forest

We hypothesized that photosynthesis and growth of tropical vegetation at its most northern distribution in Asia (Xishuangbanna, SW China) is adversely affected by seasonal drought and chilling temperatures. To test this hypothesis, we measured photosynthetic and growth characteristics of Zizyphus attopensis Pierre seedlings grown in three contrasting forest microhabitats: the understory, a small gap and a large gap. Photosynthetic capacity (light-saturated photosynthetic rate (A(max)), maximum rate of carboxylation and electron transport rate) and partitioning of leaf nitrogen (N) into carboxylation and electron transport differed significantly among seasons and microhabitats. Specific leaf area (SLA) did not change seasonally, but differed significantly among microhabitats and showed a negative linear relationship with daily integrated photon flux (PPF(i)). In contrast, leaf N concentration per unit area (N(a)) changed seasonally but did not differ among microhabitats. Measurements of maximum PSII photochemical efficiency (F(v)/F(m)) indicated that chronic photoinhibition did not occur in seedlings in any of the microhabitats during the study. Photosynthetic capacity was greatest in the wet season and lowest in the cool season. During the cool and dry seasons, the reduction in A(max) was greater in seedlings grown in the large gap than in in the understory and the small gap. Close logarithmic relationships were detected between PPF(i), leaf N(a) and photosynthetic capacity. Stem mass ratio decreased, and root mass ratio increased, in the dry season. We conclude that seasonal acclimation in growth and photosynthesis of the seedlings was associated with changes in biochemical features (particularly N(a) and partitioning of total leaf N between the different photosynthetic pools) and biomass allocation, rather than with changes in leaf morphological features (such as SLA). Local irradiance is the main factor driving seasonal variations in growth and photosynthesis in the study area, where the presence of heavy fog during the cool and dry seasons limits irradiance, but supplies water to the soil surface layers.     

Photosynthetic responses of Scots pine to elevated CO(2) and nitrogen supply: results of a branch-in-bag experiment

Naturally seeded Scots pine (Pinus sylvestris L.) trees, age 25-30 years, were subjected to two soil-nitrogen-supply regimes and to elevated atmospheric CO(2) concentrations by the branch-in-bag method from April 15 to September 15 for two or three years. Gas exchange in detached shoots was measured in a diffuse radiation field. Seven parameters associated with photosynthetic performance and two describing stomatal conductance were determined to assess the effects of treatments on photosynthetic components. An elevated concentration of CO(2) did not lead to a significant downward regulation in maximum carboxylation rate (V(cmax)) or maximum electron transport rate (J(max)), but it significantly decreased light-saturated stomatal conductance (g(sat)) and increased minimum stomatal conductance (g(min)). Light-saturated rates of CO(2) assimilation were higher (24-31%) in shoots grown and measured at elevated CO(2) concentration than in shoots grown and measured at ambient CO(2) concentration, regardless of treatment time or nitrogen-supply regime. High soil-nitrogen supply significantly increased photosynthetic capacity, corresponding to significant increases in V(cmax) and J(max). However, the combined elevated CO(2) + high nitrogen-supply treatment did not enhance the photosynthetic response above that observed in the elevated CO(2) treatment alone.     

Effects of water stress on irradiance acclimation of leaf traits in almond trees

Photosynthetic acclimation to highly variable local irradiance within the tree crown plays a primary role in determining tree carbon uptake. This study explores the plasticity of leaf structural and physiological traits in response to the interactive effects of ontogeny, water stress and irradiance in adult almond trees that have been subjected to three water regimes (full irrigation, deficit irrigation and rain-fed) for a 3-year period (2006-08) in a semiarid climate. Leaf structural (dry mass per unit area, N and chlorophyll content) and photosynthetic (maximum net CO(2) assimilation, A(max), maximum stomatal conductance, g(s,max), and mesophyll conductance, g(m)) traits and stem-to-leaf hydraulic conductance (K(s-l)) were determined throughout the 2008 growing season in leaves of outer south-facing (S-leaves) and inner northwest-facing (NW-leaves) shoots. Leaf plasticity was quantified by means of an exposure adjustment coefficient (ε=1-X(NW)/X(S)) for each trait (X) of S- and NW-leaves. Photosynthetic traits and K(s-l) exhibited higher irradiance-elicited plasticity (higher ε) than structural traits in all treatments, with the highest and lowest plasticity being observed in the fully irrigated and rain-fed trees, respectively. Our results suggest that water stress modulates the irradiance-elicited plasticity of almond leaves through changes in crown architecture. Such changes lead to a more even distribution of within-crown irradiance, and hence of the photosynthetic capacity, as water stress intensifies. Ontogeny drove seasonal changes only in the ε of area- and mass-based N content and mass-based chlorophyll content, while no leaf age-dependent effect was observed on ε as regards the physiological traits. Our results also indicate that the irradiance-elicited plasticity of A(max) is mainly driven by changes in leaf dry mass per unit area, in g(m) and, most likely, in the partitioning of the leaf N content.     

Photosynthetic responses to needle water potentials in Scots pine after a four-year exposure to elevated CO(2) and temperature

Effects of needle water potential (Psi(l)) on gas exchange of Scots pine (Pinus sylvestris L.) grown for 4 years in open-top chambers with elevated temperature (ET), elevated CO(2) (EC) or a combination of elevated temperature and CO(2) (EC + ET) were examined at a high photon flux density (PPFD), saturated leaf to air water vapor pressure deficit (VPD) and optimal temperature (T). We used the Farquhar model of photosynthesis to estimate the separate effects of Psi(l) and the treatments on maximum carboxylation efficiency (V(c,max)), ribulose-1,5-bisphosphate regeneration capacity (J), rate of respiration in the light (R(d)), intercellular partial pressure of CO(2) (C(i)) and stomatal conductance (G(s)). Depression of CO(2) assimilation rate at low Psi(l) was the result of both stomatal and non-stomatal limitations on photosynthetic processes; however, stomatal limitations dominated during short-term water stress (Psi(l) < -1.2 MPa), whereas non-stomatal limitations dominated during severe water stress. Among the nonstomatal components, the decrease in J contributed more to the decline in photosynthesis than the decrease in V(c,max). Long-term elevation of CO(2) and temperature led to differences in the maximum values of the parameters, the threshold values of Psi(l) and the sensitivity of the parameters to decreasing Psi(l). The CO(2) treatment decreased the maximum values of V(c,max), J and R(d) but significantly increased the sensitivity of V(c,max), J and R(d) to decreasing Psi(l) (P < 0.05). The effects of the ET and EC + ET treatments on V(c,max), J and R(d) were opposite to the effects of the EC treatment on these parameters. The values of G(s), which were measured simultaneously with maximum net rate of assimilation (A(max)), declined in a curvilinear fashion as Psi(l) decreased. Both the EC + ET and ET treatments significantly decreased the sensitivity of G(s) to decreasing Psi(l). We conclude that, in the future, acclimation to increased atmospheric CO(2) and temperature could increase the tolerance of Scots pine to water stress.     

Elevated CO(2) concentration affects leaf photosynthesis-nitrogen relationships in Pinus taeda over nine years in FACE

To investigate whether long-term elevated carbon dioxide concentration ([CO(2)]) causes declines in photosynthetic enhancement and leaf nitrogen (N) owing to limited soil fertility, we measured photosynthesis, carboxylation capacity and area-based leaf nitrogen concentration (N(a)) in Pinus taeda L. growing in a long-term free-air CO(2) enrichment (FACE) facility at an N-limited site. We also determined how maximum rates of carboxylation (V(cmax)) and electron transport (J(max)) varied with N(a) under elevated [CO(2)]. In trees exposed to elevated [CO(2)] for 5 to 9 years, the slope of the relationship between leaf photosynthetic capacity (A(net-Ca)) and N(a) was significantly reduced by 37% in 1-year-old needles, whereas it was unaffected in current-year needles. The slope of the relationships of both V(cmax) and J(max) with N(a) decreased in 1-year-old needles after up to 9 years of growth in elevated [CO(2)], which was accompanied by a 15% reduction in N allocation to the carboxylating enzyme. Nitrogen fertilization (110 kg N ha(-1)) in the ninth year of exposure to elevated [CO(2)] restored the slopes of the relationships of V(cmax) and J(max) with N(a) to those of control trees (i.e., in ambient [CO(2)]). The J(max):V(cmax) ratio was unaffected by either [CO(2)] or N fertilization. Changes in the apparent allocation of N to photosynthetic components may be an important adjustment in pines exposed to elevated [CO(2)] on low-fertility sites. We conclude that fundamental relationships between photosynthesis or its component processes with N(a) may be altered in aging pine needles after more than 5 years of exposure to elevated atmospheric [CO(2)].     

Physiological responses of black spruce layers and planted seedlings to nutrient addition

We investigated effects of nutrient addition on several physiological characteristics of 60-cm-tall black spruce (Picea mariana Mill. B.S.P.) layers (i.e., rooted branches of overstory trees) and 20-cm-tall planted seedlings on a clear-cut, N-limited boreal site. After two growing seasons, current-year and one-year-old needles of fertilized trees (layers and seedlings combined) had higher net photosynthetic rates (A(n)) and maximum capacity of Rubisco for CO(2) fixation (V(max)) than unfertilized trees. One-year-old needles of fertilized trees had higher stomatal conductance (g(s)), higher water-use efficiency, and lower intercellular to ambient CO(2) ratio than unfertilized trees. Additionally, fertilized trees had higher predawn and midday shoot water potentials than unfertilized trees. Stomatal conductance of 1-year-old needles was 23% higher in seedlings than in layers, but there were no significant differences in g(s) of current-year needles between the regeneration types. For both needle age-classes, A(n) and V(max) of layers were 25 and 40% higher, respectively, than the corresponding values for seedlings. The higher values of A(n), V(max) and foliar N concentration of layers compared with seedlings after two growing seasons may be associated with the larger root systems of the layers compared with the transplanted seedlings.     

Partititioning concurrent influences of nitrogen and phosphorus supply on photosynthetic model parameters of Pinus radiata

Responses of photosynthesis (A) to intercellular CO(2) concentration (C(i)) were measured in a fast- and a slow-growing clone of Pinus radiata D. Don cultivated in a greenhouse with a factorial combination of nitrogen and phosphorus supply. Stomatal limitations scaled with nitrogen and phosphorus supply as a fixed proportion of the light-saturated photosynthetic rate (18.5%) independent of clone. Photosynthetic rates at ambient CO(2) concentration were mainly in the V(cmax)-limited portion of the CO(2) response curve at low-nitrogen supply and at the transition between V(cmax) and J(max) at high-nitrogen supply. Nutrient limitations to photosynthesis were partitioned based on the ratio of foliage nitrogen to phosphorus expressed on a leaf area basis (N(a)/P(a)), by minimizing the mean square error of segmented linear models relating photosynthetic parameters (V(cmax), J(max), T(p)) to foliar nitrogen and phosphorus concentrations. A value of N(a)/P(a) equal to 23 (mole basis) was identified as the threshold separating nitrogen (N(a)/P(a) < or = 23) from phosphorus (N(a)/P(a) > 23) limitations independent of clones. On an area basis, there were significant positive linear relationships between the parameters, V(cmax), J(max), T(p) and N(a) and P(a), but only the relationships between T(p) and N(a) and P(a) differed significantly between clones. These findings suggest that, in genotypes with contrasting growth, the responses of V(cmax) and J(max) to nutrient limitation are equivalent. The relationships between the parameters V(cmax), J(max), T(p) and foliage nutrient concentration on a mass basis were unaffected by clone, because the slow-growing clone had a significantly greater leaf area to mass ratio than the fast-growing clone. These results may be useful in discriminating nitrogen-limited photosynthesis from phosphorus-limited photosynthesis.     

Ecophysiological characteristics of Taiwan alder (Alnus formosana) seedlings adapted to the subtropical region

We investigated several ecophysiological characteristics of seedlings of a low-elevation (100-200 m) and a high-elevation (2000-2400 m) population of Taiwan alder (Alnus formosana Makino) from subtropical Taiwan. Both populations had a wide optimal temperature range for photosynthesis, and there was little difference in the optimal temperature range for photosynthesis between populations. Photosynthetic rate (P(N)) was near maximal from 20 to 35 degrees C when seedlings of both the low-elevation and the high-elevation populations were acclimated at a day/night temperature of 30/23 degrees C. When seedlings were acclimated at a day/night temperature of 20/10 degrees C, P(N) was near maximal over the range 15-35 degrees C in the low-elevation population and 15-30 degrees C in the high-elevation population. Compared with nine other tree species native to Taiwan, Taiwan alder had a high P(N) and stomatal conductance (g(s)) under well-watered conditions. Reflecting its higher transpiration rate, Taiwan alder had a significantly greater leaf-air temperature difference than camphor (Cinnamomum camphora (L.) J. Presl), a co-occurring lowland tree species with leaves similar in shape and size to those of Taiwan alder. Despite higher g(s), high root and shoot hydraulic conductances enabled Taiwan alder to maintain higher leaf water potentials than camphor under well-watered conditions. We conclude that both photosynthetic characteristics and water relations are important factors enabling Taiwan alder to adapt to a wide temperature range, thereby ensuring its success at both high and low elevations in subtropical Taiwan.