Seasonal variation in the photosynthetic capacity of a willow (Salix cv. Aquatica gigantea) canopy. 2 Comparison of the structure and function of chloroplasts at different levels in the canopy

Development of leaf chloroplast ultrastructure at five levels in a willow (Salix cv. Aquatica gigantea) canopy was followed during one growing season in the field. Changes in chloroplast ultrastructure were compared with the rate of CO(2) uptake of the same leaves. The highest rates of CO(2) uptake were recorded in young leaves exposed to full available sunlight. In these leaves, the area of the grana stacks was less than 20% of the total chloroplast area and the degree of thylakoid stacking was less than 1.5. The chloroplasts of these leaves contained large amounts of starch and small amounts of plastoglobuli. As the canopy grew and the leaves in the lower parts of the canopy became shaded, the structure of the chloroplast thylakoids gradually changed. In leaves at the two lowest levels of the canopy, the degree of stacking at the end of the growing season was close to 2 and correspondingly the rate of CO(2) uptake was low. The areas of grana stacks and plastoglobuli in these chloroplasts increased and were about 30 and 10% of the chloroplast area, respectively, by the end of the growing season. The increase in the degree of thylakoid stacking was caused by increased biosynthesis of grana lamellae, which in general were thinner than the lamellae of young leaves. The length of the stroma lamellae did not change with leaf age. Morphometric measurements showed that the structure of the chloroplasts in leaves 160 cm above ground was dynamic and responsive to environmental conditions so that photosynthetic capacity remained high for 7 weeks despite an increase in leaf shading.     

Transient etiolation: protochlorophyll(ide) and chlorophyll forms in differentiating plastids of closed and breaking leaf buds of horse chestnut (Aesculus hippocastanum)

An accompanying paper reports the accumulation of photoactive protochlorophyllide (Pchlide) in the innermost leaf primordia of buds of many tree species. In this paper, we describe plastid differentiation, changes in pigment concentrations and spectral properties of bud scales and leaf primordia of horse chestnut (Aesculus hippocastanum L.) from January until the end of bud break in April. The bud scales contained plastids with grana, stroma thylakoids characteristic of chloroplasts and large dense bodies within the stroma. In January, proplastids and young chloroplasts were present in the leaf primordia, and the fluorescence spectra of the primordia were similar to those of green leaves except for a minor band at 630 nm, indicative of a protochlorophyll(ide). During bud break, the pigment concentrations of the green bud scales and the outermost leaf primordia increased, and Pchlide forms with emission maxima at 633, 644 and 655 nm accumulated in the middle and innermost leaf primordia. Depending on the position of the leaf primordia within the bud, their plastids and their pigment concentrations varied. Etio-chloroplasts with prolamellar bodies (PLBs) and prothylakoids with developing grana were observed in the innermost leaves. Besides the above-mentioned Pchlide forms, the middle and innnermost leaf primordia contained only a Chl band with an emission maximum at 686 nm. The outermost leaf primordia contained etio-chloroplasts with well-developed grana and small, narrow-type PLBs. These outermost leaves contained only chlorophyll forms like the mature green leaves. No Pchlide accumulation was observed after bud break, indicating that etiolation of the innermost and middle leaves is transient. The Pchlide forms and the plastid types of the primordia in buds grown in nature were similar to those of leaves of dark-germinated seedlings and to those of the leaf primordia of dark-forced buds. We conclude that transient etiolation occurs under natural conditions. The formation of PLBs and etio-chloroplasts and the accumulation of the light-dependent NADPH:protochlorophyllide oxidoreductase are involved in the natural greening process and ontogenesis of young leaf primordia of horse chestnut buds.     

Effects of CO2 enrichment on the photosynthetic light response of sun and shade leaves of canopy sweetgum (Liquidambar styraciflua) in a forest ecosystem

To investigate whether sun and shade leaves respond differently to CO2 enrichment, we examined photosynthetic light response of sun and shade leaves in canopy sweetgum (Liquidambar styraciflua L.) trees growing at ambient and elevated (ambient + 200 microliters per liter) atmospheric CO2 in the Brookhaven National Laboratory/Duke University Free Air CO2 Enrichment (FACE) experiment. The sweetgum trees were naturally established in a 15-year-old forest dominated by loblolly pine (Pinus taeda L.). Measurements were made in early June and late August 1997 during the first full year of CO2 fumigation in the Duke Forest FACE experiment. Sun leaves had a 68% greater leaf mass per unit area, 63% more leaf N per unit leaf area, 27% more chlorophyll per unit leaf area and 77% greater light-saturated photosynthetic rates than shade leaves. Elevated CO2 strongly stimulated light-saturated photosynthetic rates of sun and shade leaves in June and August; however, the relative photosynthetic enhancement by elevated CO2 for sun leaves was more than double the relative enhancement of shade leaves. Elevated CO2 stimulated apparent quantum yield by 30%, but there was no interaction between CO2 and leaf position. Daytime leaf-level carbon gain extrapolated from photosynthetic light response curves indicated that sun leaves were enhanced 98% by elevated CO2, whereas shade leaves were enhanced 41%. Elevated CO2 did not significantly affect leaf N per unit area in sun or shade leaves during either measurement period. Thus, the greater CO2 enhancement of light-saturated photosynthesis in sun leaves than in shade leaves was probably a result of a greater amount of nitrogen per unit leaf area in sun leaves. A full understanding of the effects of increasing atmospheric CO2 concentrations on forest ecosystems must take account of the complex nature of the light environment through the canopy and how light interacts with CO2 to affect photosynthesis.     

Leaf respiration at different canopy positions in sweetgum (Liquidambar styraciflua) grown in ambient and elevated concentrations of carbon dioxide in the field

Trees exposed to elevated CO2 partial pressure ([CO2]) generally show increased rates of photosynthesis and growth, but effects on leaf respiration are more variable. The causes of this variable response are unresolved. We grew 12-year-old sweetgum trees (Liquidambar styraciflua L.) in a Free-Air CO2 Enrichment (FACE) facility in ambient [CO2] (37/44 Pa daytime/nighttime) and elevated [CO2] (57/65 Pa daytime/nighttime) in native soil at Oak Ridge National Environmental Research Park. Nighttime respiration (R(N)) was measured on leaves in the upper and lower canopy in the second (1999) and third (2000) growing seasons of CO2 fumigation. Leaf respiration in the light (R(L)) was estimated by the technique of Brooks and Farquhar (1985) in the upper canopy during the third growing season. There were no significant short-term effects of elevated [CO2] on R(N) or long-term effects on R(N) or R(L), when expressed on an area, mass or nitrogen (N) basis. Upper-canopy leaves had 54% higher R(N) (area basis) than lower-canopy leaves, but this relationship was unaffected by CO2 growth treatment. In August 2000, R(L) was about 40% of R(N) in the upper canopy. Elevated [CO(2)] significantly increased the number of leaf mitochondria (62%), leaf mass per unit area (LMA; 9%), and leaf starch (31%) compared with leaves in ambient [CO(2)]. Upper-canopy leaves had a significantly higher number of mitochondria (73%), N (53%), LMA (38%), sugar (117%) and starch (23%) than lower-canopy leaves. Growth in elevated [CO2] did not affect the relationships (i.e., intercept and slope) between R(N) and the measured leaf characteristics. Although no factor explained more than 45% of the variation in R(N), leaf N and LMA were the best predictors for R(N). Therefore, the response of RN to CO2 treatment and canopy position was largely dependent on the magnitude of the effect of elevated [CO2] or canopy position on these characteristics. Because elevated [CO2] had little or no effect on N or LMA, there was no effect on R(N). Canopy position had large effects on these leaf characteristics, however, such that upper-canopy leaves exhibited higher R(N) than lower-canopy leaves. We conclude that elevated [CO2] does not directly impact leaf respiration in sweetgum and that barring changes in leaf nitrogen or leaf chemical composition, long-term effects of elevated [CO2] on respiration in this species will be minimal.     

Spatial distribution of leaf morphological and physiological characteristics in relation to local radiation regime within the canopies of 3-year-old Populus clones in coppice culture

Spatial distributions of leaf characteristics relevant to photosynthesis were compared within high-density coppice canopies of Populus spp. of contrasting genetic origin. We studied three clones representative of the range in growth potential, leaf morphology, coppice and canopy structure: Clone Hoogvorst (Hoo) (Populus trichocarpa Torr. & Gray x Populus deltoides Bartr. & Marsh), Clone Fritzi Pauley (Fri) (Populus trichocarpa Torr. & Gray) and Clone Wolterson (Wol) (Populus nigra L.). Leaf area index ranged from 2.7 (Fri and Wol) to 3.8 (Hoo). The clones exhibited large vertical variation in leaf area density (0.02-1.42 m2 m-3). Leaf dry mass per unit leaf area (DM(A)) increased with increasing light in Clones Hoo and Fri, from about 56 g m-2 at the bottom of the canopy to 162 g m-2 at the top. In Clone Wol, DM(A) varied only from 65 to 100 g m-2, with no consistent relationship with respect to light. Conversely, nitrogen concentration on a mass basis was nearly constant (around 1.3-2.1%) within the canopies of Clones Hoo and Fri, but increased strongly with light in Clone Wol, from 1.4% at the bottom of the canopy to 4.1% at the top. As a result, nitrogen per unit leaf area (N(A)) increased with light in the canopies of all clones, from 0.9 g m-2 at the bottom to 2.9 g m-2 at the top. Although a single linear relationship described the dependence of maximum carboxylation rate (17-93 micromol CO2 m-2 s-1) or electron transport capacity (45-186 micromol electrons m-2 s-1) on N(A), for all clones, Clone Wol differed from Clones Hoo and Fri by exhibiting a higher dark respiration rate at low N(A) (1.8 versus 0.8 micromol CO2 m-2 s-1).     

Gas exchange characteristics of a Canarian laurel forest tree species (Laurus azorica) in relation to environmental conditions and leaf canopy position

Diurnal courses of gas exchange were measured over a 1-year period in fully expanded current-year leaves in the upper (sun-exposed, 18 m above ground) and the lower (shaded, 12 m above ground) canopy of Laurus azorica (Seub.) Franco, a major canopy species of the Canarian laurel forest in Tenerife, Canary Islands, Spain. Laurus azorica exhibited high leaf plasticity in gas exchange characteristics, with a maximum carbon assimilation rate (Amax) of shade leaves about 50% that of sun leaves. This difference reflects the high leaf area index (LAI) of the stand and the correspondingly sharp light attenuation with increasing canopy depth. In sun leaves, Amax peaked at about 11 micromol m-2 s-1 and maximum transpiration (E) was about 8 mmol m-2 s-1, which corresponded with a maximum stomatal conductance (gs) of about 650 mmol m-2 s-1. Mean maximum instantaneous water-use efficiency (WUE) was 1.5 mmol mol-1 and the mean maximum A/gs was 20-35 micromol mol-1. Mean minimum internal CO2 concentration (Ci) was 225 micromol mol-1. Although high air vapor pressure deficit (VPD) caused a small decrease in gs, it remained high enough to maintain relatively high A and E. These gas exchange characteristics indicate a non-conservative use of water, which is appropriate for a species subject to droughts that are mild or of short duration. In this respect, Laurus azorica differs from its congener, L. nobilis L., of the Mediterranean region and other shrubs growing in Mediterranean-type climates in California and Chile that have to withstand more severe or more prolonged droughts.     

Do elevation of CO(2) concentration and nitrogen fertilization alter storage and remobilization of carbon and nitrogen in pedunculate oak saplings?

Soil nitrogen can alter storage and remobilization of carbon and nitrogen in forest trees and affect growth responses to elevated carbon dioxide concentration ([CO(2)]). We investigated these effects in oak saplings (Quercus robur L.) exposed for two years to ambient or twice ambient [CO(2)] in combination with low- (LN, 0.6 mmol N l(-1)) or high-nitrogen (HN, 6.1 mmol N l(-1)) fertilization. Autumn N retranslocation efficiency from senescing leaves was less in HN saplings than in LN saplings, but about 15% of sapling N was lost to the litter. During the dormant season, nonstructural carbohydrates made up 20 to 30% of the dry mass of perennial organs. Starch was stored mainly in large roots where it represented 35-46% of dry mass. Accumulation of starch increased in large roots in response to LN but was unaffected by elevated [CO(2)]. The HN treatment resulted in high concentrations of N-soluble compounds, and this effect was reduced by elevated [CO(2)], which decreased soluble protein N (-17%) and amino acid N (-37%) concentrations in the HN saplings. Carbon and N reserves were labeled with (13)C and (15)N, respectively, at the end of the first year. In the second year, about 20% of labeled C and 50% of labeled N was remobilized for spring growth in all treatments. At the end of leaf expansion, 50-60% of C in HN saplings originated from assimilation versus only 10-20% in LN saplings. In HN saplings only, N uptake occurred, and some newly assimilated N was allocated to new shoots. Through effects on the C and N content of perennial organs, elevated [CO(2)] and HN increased remobilization capacity, thereby supporting multiple shoot flushes, which increased leaf area and subsequent C acquisition in a positive feedback loop.     

Moderate shade can increase net gas exchange and reduce photoinhibition in citrus leaves

Daily variations in net gas exchange, chlorophyll a fluorescence and water relations of mature, sun-acclimated grapefruit (Citrus paradisi Macfady.) and orange (Citrus sinensis L. Osbeck) leaves were determined in tree canopies either shaded with 50% shade screens or left unshaded (sunlit). Mean daily maximum photosynthetic photon flux density (PPFD) under shade varied from 500 to 700 micromol m-2 s-1 and was sufficient to achieve maximum net CO2 assimilation rates (A CO2). Responses of grapefruit and orange leaves to shading were remarkably similar. At midday, on bright clear days, the temperatures of sunlit leaves were 2-6 degrees C above air temperature and 1-4 degrees C above the temperatures of shaded leaves. Although midday depressions of stomatal conductance (gs) and A CO2 were observed in both sunlit and shaded leaves, shaded leaves had lower leaf-to-air vapor pressure differences (D) along with higher gs, A CO2 and leaf water-use efficiency than sunlit leaves. Estimated stomatal limitation to A CO2 was generally less than 25% and did not differ between shaded and sunlit leaves. Leaf intercellular CO2 partial pressure was not altered by shade treatment and did not change substantially with increasing D. Radiation and high temperature stress-induced non-stomatal limitation to A CO2 in sunlit leaves was greater than 40%. Reversible photoinhibition of photosystem II efficiency was more pronounced in sunlit than in shaded leaves. Thus, non-stomatal factors play a major role in regulating A CO2 of citrus leaves during radiation and high temperature stress.     

Leaf nutrition and photosynthetic performance of sugar maple (Acer saccharum) in stands with contrasting health conditions

Leaf nutrition and photosynthetic performance of sugar maple (Acer saccharum Marsh.) were compared between two sugar maple stands in northwestern Vermont with contrasting health conditions as indicated by annual basal area growth, degree of crown dieback, and foliar appearance. Observations made during the diurnal cycle of both stands showed no apparent leaf water stress. In both stands, leaves had similar concentrations of major non-structural carbohydrates (starch and sucrose). Over two consecutive growing seasons (1991 and 1992), we consistently observed lower leaf Ca and Mg concentrations in the declining stand than in the healthy stand. Compared with the healthy stand, lower leaf chlorophyll concentrations and apparent leaf chlorosis were observed in the declining stand, and some trees had very low foliar Ca and Mg concentrations (0.31 +/- 0.03% and 0.09 +/- 0.01%, respectively). Trees in the declining stand had lower light-saturated net photosynthetic rates on a dry mass basis at both ambient CO(2) (P(n,amb)) and saturating CO(2) (P(n,sat)) than trees in the healthy stand. There were significant linear correlations between P(n,amb) and leaf mass per unit area (LMA) and between P(n,sat) per unit leaf area and LMA. There were also linear correlations between both P(n,amb) and P(n,sat) and leaf N when expressed on an area basis in both stands, indicating that variation in LMA may have been largely responsible for the observed photosynthesis-nitrogen relationship. The values of P(n,amb) and P(n,sat) were not significantly correlated with leaf N on a mass basis but were weakly correlated with leaf Ca and Mg on a mass basis. We conclude that low leaf Ca or Mg concentrations may limit leaf CO(2) assimilation and tree carbohydrate status in the declining stand.     

Effects of leaf,shoot and fruit development on photosynthesis of lychee trees (Litchi chinensis)

Changes in gas exchange with leaf age and fruit growth were determined in lychee trees (Litchi chinensis Sonn.) growing in subtropical Queensland (27 degrees S). Leaves expanded in a sigmoid pattern over 50 days during spring, with net CO2 assimilation (A) increasing from -4.1 +/- 0.9 to 8.3 +/- 0.5 micromol m-2 s-1 as the leaves changed from soft and red, to soft and light green, to hard and dark green. Over the same period, dark respiration (Rd) decreased from 5.0 +/- 0.8 to 2.0 +/- 0.1 micromol CO2 m-2 s-1. Net CO2 assimilation was above zero about 30 days after leaf emergence or when the leaves were half fully expanded. Chlorophyll concentrations increased from 0.7 +/- 0.2 mg g-1 in young red leaves to 10.3 +/- 0.7 mg g-1 in dark green leaves, along with stomatal conductance (gs, from 0.16 +/- 0.09 to 0.47 +/- 0.17 mol H2O m-2 s-1). Fruit growth was sigmoidal, with maximum values of fresh mass (29 g), dry mass (6 g) and fruit surface area (39 cm2) occurring 97 to 115 days after fruit set. Fruit CO2 exchange in the light (Rl) and dark (Rd) decreased from fruit set to fruit maturity, whether expressed on a surface area (10 to 3 micromol CO2 m-2 s-1 and 20 to 3 micromol CO2 m-2 s-1, respectively) or on a dry mass basis (24 to 2 nmol CO2 g-1 s-1 and 33 to 2 nmol CO2 g-1 s-1, respectively). Photosynthesis never exceeded respiration, however, the difference between Rl and Rd was greatest in young green fruit (4 to 8 micromol CO2 m-2 s-1). About 90% of the carbon required for fruit growth was accounted for in the dry matter of the fruit, with the remainder required for respiration. Fruit photosynthesis contributed about 3% of the total carbon requirement of the fruit over the season. Fruit growth was mainly dependent on CO2 assimilation in recently expanded dark green leaves.     

不同氮素水平对红厚壳幼苗生长及光合特性的影响

采用温室盆栽方法,设置0(对照)、50、100、150、200、300、400、600 mg.株-18个氮素处理,研究氮素供应对红厚壳幼苗生长及光合特性的影响。结果表明:随供氮量的增加,红厚壳幼苗高、地径、叶面积及整株生物量均呈现出先增加后降低的趋势,均在200 mg.株-1时达到峰值,分别为22.06 cm、0.65 cm、327.27 cm2和10.43 g.株-1,是对照的1.59、1.25、2.58、1.90倍;地下生物量N5显著高于N7、N8处理,但与其余处理无显著性差异,根冠比随供氮量的增加呈现单调递减趋势;红厚壳幼苗叶绿素a、b及总量随供氮量的增加而增加;红厚壳幼苗的净光合速率、气孔导度、胞间CO2浓度以及蒸腾速率随供氮量的增加呈现出先增加后减小的趋势,均在200 mg.株-1时达最大值,分别是7.29μmol.m-2.s-1、0.071 mol.m-2.s-1、220.22μmol.mol-1和1.34 mmol.m-2.s-1,分别是对照的2.26、1.92、1.37、1.79倍,不同氮素处理间红厚壳光合气体交换参数差异均显著。结果表明:施氮量200 mg.株-1左右为红厚壳幼苗温室栽培的最佳施肥量。     

Carbon dioxide exchange of larch (Larix gmelinii) cones during development

Larch (Larix gmelinii (Rupr.) Rupr.) cone scales are green, but little is known of their photosynthetic role in cone development or about how they differ in gas exchange characteristics from needle leaves. In contrast to leaf photosynthesis (Pleaf), we found that stomatal regulation of cone photosynthetic rate (Pcone) was marginal because the photosynthetic carbon came from internal recycling of respiratory carbon dioxide (CO2). Photosynthetic recycling of respired CO2 was confirmed by the finding that the intercellular CO2 concentration (Ci) in cone scales was much higher than ambient [CO2]; also, there was a positive correlation between Pcone and Ci, whereas Pleaf was almost constant as Ci varied. Low chlorophyll (Chl) concentration was a limiting factor for Pcone, but not for Pleaf, as indicated by the correlation between Pcone and chlorophyll concentration. Moreover, chlorophyll utilization efficiency (Psat/Chl a+b) for cone scales was lower than that for leaves. In both cones and leaves, nitrogen (N) was positively correlated with photosynthetic capacity (P), but the P/N value was much lower for cones than for leaves. For both organs, the ratio of respiration to N was broadly similar. Although mature cones have no photosynthetic capacity, Pcone of young cones was as high as 5.3 micromol m(-2) s(-1), about 1.26 times the value of Pleaf, and accounted for the refixation of 30-40% of the respiratory CO2 produced by cones, equivalent to the photosynthetic capacity of a bundle of short shoots near each cone. Thus, Pcone may be an important additional source of photosynthate for cones, given the weak assimilating capacity of leaves that are not fully expanded during cone development.     

Inhibitory effect of flowering and early fruit growth on leaf photosynthesis in mango

Carbohydrate and nitrogen contents, chlorophyll fluorescence and gas exchange were measured in leaves from both vegetative and reproductive terminal shoots of 12-year-old flowering mango trees. Reproductive shoot leaves were close to swelling floral buds, inflorescences or panicles bearing set fruits. Leaves close to inflorescences had lower rates of mitochondrial respiration (Rd) and net photosynthesis (Anet), and lower stomatal conductance (gs) and quantum efficiency of photosystem II under actinic light than vegetative shoot leaves. Leaf nitrogen concentration, which decreased from the beginning until the end of flowering, was lower in leaves close to inflorescences than in vegetative shoot leaves. However, these differences and changes were counterbalanced by an increase in leaf mass-to-area ratio so that leaf nitrogen per unit leaf area (Na) remained nearly constant during the whole flowering period, except in leaves close to panicles bearing set fruits. Net CO2 assimilation rate simulated by a biochemical model of leaf photosynthesis (Urban et al. 2003) was much higher than Anet measured at an ambient CO2 partial pressure (Ca) of either 36 or 70 Pa. The overestimation of Anet was more pronounced in leaves close to inflorescences, to panicles bearing set fruits and to reversing inflorescences (characterized by the appearance of leaves in terminal positions on inflorescences) than in vegetative shoot leaves. It is concluded that low Anet in leaves close to inflorescences was probably due neither to changes in Na nor to a decrease in Rubisco activity induced by low gs, but rather to a decrease in electron flow in photosystem II. This decrease was not directly associated with higher starch or soluble sugar contents.     

Internal leaf anatomy and photosynthetic resource-use efficiency: interspecific and intraspecific comparisons

Leaf mass per unit area (LMA) and internal leaf anatomy often affect net gas exchange because of their effects on internal CO2 conductance to the site of carboxylation, internal shading, competition for CO2 among carboxylation sites, nitrogen concentration and its partitioning. To evaluate effects of LMA and leaf anatomy on CO2 assimilation, water-use efficiency (WUE) and nitrogen-use efficiency (NUE), we measured LMA, leaf thickness, the thickness of mesophyll components, and gas exchange rates at ambient CO2 concentration in leaves of six woody deciduous and evergreen species with different leaf life spans. In two species, CO2 assimilation was also estimated at saturating CO2 concentrations. There were interspecific differences in all morphological variables studied. Long-lived leaves had higher LMA and were thicker than short-lived leaves. Species with high LMA had low assimilation rates and NUE, both in ambient and saturating CO2 concentrations. Thus, in species with high LMA, assimilation was reduced by non-stomatal limitations, possibly because of a lower allocation of N to the photosynthetic machinery than in species with low LMA. Within a species, thicker leaves tended to have a lower tissue density. In intraspecific comparisons under field conditions, increasing internal air volume had positive effects on WUE, probably because of enhanced internal CO2 conductance to the site of carboxylation. We conclude that, in interspecific comparisons, different patterns of N partitioning strongly influence NUE, whereas in intraspecific comparisons, internal leaf anatomy is a key factor regulating resource-use efficiency.     

Influence of canopy light environment and nitrogen availability on leaf photosynthetic characteristics and photosynthetic nitrogen-use efficiency of field-grown nectarine trees

Relationships between CO(2) assimilation at light saturation (A(max)), nitrogen (N) content and weight per unit area (W(A)) were studied in leaves grown with contrasting irradiances (outer canopy versus inner canopy) and N supply rates in field-grown nectarine trees Prunus persica L. Batsch. cv. Fantasia. Both A(max) and N content per unit leaf area (N(A)) were linearly correlated to W(A), but leaves in the high-N treatment had higher N(A) and A(max) for the same value of W(A) than leaves in the low-N treatment. The curvilinear relationship between photosynthesis and total leaf N was independent of treatments, both when expressed per unit leaf area A(maxA) and N(A)) and per unit leaf weight (A(maxW) and N(W)), but the relationship was stronger when data were expressed on a leaf area basis. Both A(maxA) and N(A) were higher for outer canopy leaves than for inner canopy leaves and A(maxW) and N(W) were higher for leaves in the high-N treatment than for leaves in the low-N treatment. The relationship between A(max) and N resulted in a similar photosynthetic nitrogen-use efficiency at light saturation (A(max)NUE) for both N and light treatments. Photosynthetic nitrogen-use efficiency was similar among treatments throughout the whole light response curve of photosynthesis. Leaves developed in shade conditions did not show higher N-use efficiency at low irradiance. At any intercellular CO(2) partial pressure (C(i)), photosynthetic CO(2) response curves were higher for outer canopy leaves and, within each light treatment, were higher for the high-N treatments than for the low-N treatments. Consequently, most of the differences among treatments disappeared when photosynthesis was expressed per unit N. However, slightly higher assimilation rates per unit N were found for outer canopy leaves compared with inner canopy leaves, in both N treatments. Because higher daily irradiance within the canopies of the low-N trees more than compensated for the lower photosynthetic performances of these leaves compared to the leaves of high-N trees, daily carbon gain (and N-use efficiency on a daily assimilation basis) per leaf was higher for the low-N treatment than for the high-N treatment in both outer and inner canopy leaves.     

Relationship between photosynthesis and leaf nitrogen concentration in ambient and elevated [CO2] in white birch seedlings

To study the effects of elevated CO2 concentration ([CO2]) on relationships between nitrogen (N) nutrition and foliar gas exchange parameters, white birch (Betula papyrifera Marsh.) seedlings were exposed to one of five N-supply regimes (10, 80, 150, 220, 290 mg N l(-1)) in either ambient [CO2] (360 micromol mol(-1)) or elevated [CO2] (720 micromol mol(-1)) in environment-controlled greenhouses. Foliar gas exchange and chlorophyll fluorescence were measured after 60 and 80 days of treatment. Photosynthesis showed a substantial down-regulation (up to 57%) in response to elevated [CO2] and the magnitude of the down-regulation generally decreased exponentially with increasing leaf N concentration. When measured at the growth [CO2], elevated [CO2] increased the overall rate of photosynthesis (P(n)) and instantaneous water-use efficiency (IWUE) by up to 69 and 236%, respectively, but decreased transpiration (E) and stomatal conductance (g(s)) in all N treatments. However, the degree of stimulation of photosynthesis by elevated [CO2] decreased as photosynthetic down-regulation increased from 60 days to 80 days of treatment. Elevated [CO2] significantly increased total photosynthetic electron transport in all N treatments at 60 days of treatment, but the effect was insignificant after 80 days of treatment. Both P(n) and IWUE generally increased with increasing leaf N concentration except at very high leaf N concentrations, where both P(n) and IWUE declined. The relationships of P(n) and IWUE with leaf N concentration were modeled with both a linear regression and a second-order polynomial function. Elevated [CO2] significantly and substantially increased the slope of the linear regression for IWUE, but had no significant effect on the slope for P(n). The optimal leaf N concentration for P(n) and IWUE derived from the polynomial function did not differ between the CO2 treatments when leaf N was expressed on a leaf area basis. However, the mass-based optimal leaf N concentration for P(n) was much lower in seedlings in elevated [CO2] than in ambient [CO2] (31.88 versus 37.00 mg g(-1)). Elevated [CO2] generally decreased mass-based leaf N concentration but had no significant effect on area-based leaf N concentration; however, maximum N concentration per unit leaf area was greater in elevated [CO2] than in ambient [CO2] (1.913 versus 1.547 g N m(-2)).     

Fruit load and canopy shading affect leaf characteristics and net gas exchange of 'Spring' navel orange trees

Five-year-old 'Spring' navel (Citrus sinensis (L.) Osbeck) orange trees were completely defruited, 50% defruited or left fully laden to study effects of fruit load on concentrations of nitrogen (N) and carbohydrate, net assimilation of CO2 (Ac) and stomatal conductance (gs) of mature leaves on clear winter days just before fruit harvest. Leaves on defruited trees were larger, had higher starch concentrations and greater leaf dry mass per area (LDMa) than leaves on fruited trees. Both Ac and gs were more than 40% lower in sunlit leaves on defruited trees than in sunlit leaves on trees with fruit. Leaves immediately adjacent to fruit were smaller, had lower leaf nitrogen and carbohydrate concentrations, lower LDMa and lower Ac than leaves on non-fruiting branches of the same trees. Removing half the crop increased individual fruit mass, but reduced fruit color development. Half the trees were shaded with 50% shade cloth for 4 months before harvest to determine the effects of lower leaf temperature (Tl) and leaf-to-air vapor pressure difference on leaf responses. On relatively warm days when sunlit Tl > 25 degrees C, shade increased Ac and gs, but had no effect on the ratio of internal to ambient CO2 (Ci/Ca) concentration in leaves, implying that high mesophyll temperatures in sunlit leaves were more important than gs in limiting Ac. Sunlit leaves were more photoinhibited than shaded leaves on cooler days when Tl < 25 degrees C. Shade decreased total soluble sugar concentrations in leaves, but had no effect on leaf starch concentrations. Shading had no effects on canopy volume, yield or fruit size, but shaded fruit developed better external color than sun-exposed fruit. Overall, the presence of a normal fruit crop resulted in lower foliar carbohydrate concentrations and higher Ac compared with defruited trees, except on warm days when Ac was reduced by high leaf temperatures.     

Effects of elevated CO(2) concentration on leaf characteristics and photosynthetic capacity of beech (Fagus sylvatica) during the growing season

Two-year-old beech (Fagus sylvatica L.) saplings were planted directly in the ground at high density (100 per m(2)), in an experimental design that realistically mimicked field conditions, and grown for two years in air containing CO(2) at either ambient or an elevated (ambient + 350 ppm) concentration. Plant dry mass and leaf area were increased by a two-year exposure to elevated CO(2). The saplings produced physiologically distinct types of sun leaves associated with the first and second growth flushes. Leaves of the second flush had a higher leaf mass per unit area and less chlorophyll per unit area, per unit dry mass and per unit nitrogen than leaves of the first flush. Chlorophyll content expressed per unit nitrogen decreased over time in plants grown in elevated CO(2), which suggests that, in elevated CO(2), less nitrogen was invested in machinery of the photosynthetic light reactions. In early summer, the photosynthetic capacity measured at saturating irradiance and CO(2) was slightly but not significantly higher in saplings grown in elevated CO(2) than in saplings grown in ambient CO(2). However, a decrease in photosynthetic capacity was observed after July in leaves of saplings grown in CO(2)-enriched air. The results demonstrate that photosynthetic acclimation to elevated CO(2) can occur in field-grown saplings in late summer, at the time of growth cessation.     

Long-term effects of elevated carbon dioxide concentration and provenance on four clones of Sitka spruce (Picea sitchensis). II. Photosynthetic capacity and nitrogen use efficiency

Four clones of Sitka spruce (Picea sitchensis (Bong.) Carr.) from two provenances, at 53.2 degrees N (Skidegate a and Skidegate b) and at 41.3 degrees N (North Bend a and North Bend b), were grown for three growing seasons in ambient (~350 micromol per mol) and elevated (~700 micromol per mol) CO2 concentrations. The clones were grown in stress-free conditions (adequate nutrition and water) to assess the effect of elevated [CO2] on tree physiology. Growth in elevated [CO2] significantly increased instantaneous photosynthetic rates of the clonal Sitka spruce saplings by about 62%. Downward acclimation of photosynthesis (A) was found in all four clones grown in elevated [CO2]. Rubisco activity and total chlorophyll concentration were also significantly reduced in elevated [CO2]. Provenance did not influence photosynthetic capacity. Best-fit estimates of Jmax (maximum rate of electron transport), Vcmax (RuBP-saturated rate of Rubisco) and Amax (maximum rate of assimilation) were derived from responses of A to intercellular [CO2] by using the model of Farquhar et al. (1980). At any leaf N concentration, the photosynthetic parameters were reduced by growth in elevated [CO2]. However, the ratio between Jmax and Vcmax was unaffected by CO2 growth concentration, indicating a tight coordination in the allocation of N between thylakoid and soluble proteins. In elevated [CO2], the more southerly clones had a higher initial N use efficiency (more carbon assimilated per unit of leaf N) than the more northerly clones, so that they had more N available for those processes or organs that were most limiting to growth at a particular time. This may explain the initial higher growth stimulation by elevated [CO2] in the North Bend clones than in the Skidegate clones.     

Leaf senescence and late-season net photosynthesis of sun and shade leaves of overstory sweetgum (Liquidambar styraciflua) grown in elevated and ambient carbon dioxide concentrations

We examined the effects of elevated CO2 concentration ([CO2]) on leaf demography, late-season photosynthesis and leaf N resorption of overstory sweetgum (Liquidambar styraciflua L.) trees in the Duke Forest Free Air CO2 Enrichment (FACE) experiment. Sun and shade leaves were subdivided into early leaves (formed in the overwintering bud) and late leaves (formed during the growing season). Overall, we found that leaf-level net photosynthetic rates were enhanced by atmospheric CO2 enrichment throughout the season until early November; however, sun leaves showed a greater response to atmospheric CO2 enrichment than shade leaves. Elevated [CO2] did not affect leaf longevity, emergence date or abscission date of sun leaves or shade leaves. Leaf number and leaf area per shoot were unaffected by CO2 treatment. A simple shoot photosynthesis model indicated that elevated [CO2] stimulated photosynthesis by 60% in sun shoots, but by only 3% in shade shoots. Whole-shoot photosynthetic rate was more than 12 times greater in sun shoots than in shade shoots. In senescent leaves, elevated [CO2] did not affect residual leaf nitrogen, and nitrogen resorption was largely unaffected by atmospheric CO2 enrichment, except for a small decrease in shade leaves. Overall, elevated [CO2] had little effect on the number of leaves per shoot at any time during the season and, therefore, did not change seasonal carbon gain by extending or shortening the growing season. Stimulation of carbon gain by atmospheric CO2 enrichment in sweetgum trees growing in the Duke Forest FACE experiment was the result of a strong stimulation of photosynthesis throughout the growing season.