Limitation of leaf carbon gain by stomatal and photochemical processes in the top canopy of Macaranga conifera,a tropical pioneer tree

Diurnal changes in gas exchange and chlorophyll fluorescence were measured in the top canopy leaves of the tropical rainforest tree species, Macaranga conifera (Zoll.) Muell. Arg. during a drought year. Maximum values of net photosynthetic rate (P(n), 10 &mgr;mol m(-2) s(-1)) and stomatal conductance (g(s), 0.2 mol m(-2) s(-1)) were found in east-facing leaves in early morning. After 1000 h, both P(n) and g(s) decreased. Minimum daytime values of P(n), g(s), and photosystem II (PSII) quantum yield (DeltaF/F(m)') were found in horizontally fixed leaves. At a given electron transport rate through PSII (ETR), P(n) was higher in early morning than at midday, suggesting a high rate of photorespiration at midday. We tested the hypothesis that the effect of low leaf temperature (T(leaf)) on P(n) is significant in the early morning, whereas the effect of low g(s) on P(n) predominates at midday. In the early morning, when T(leaf) was increased from 32 to 38 degrees C by artificial heating, P(n) at a given ETR decreased 29%, suggesting that the low T(leaf) was associated with a high P(n). When T(leaf) at midday was decreased from 37 to 32 degrees C by artificial cooling, P(n) increased 22%, but P(n) at a given ETR was higher in early morning than at midday, even at the same low T(leaf) (32 degrees C). This suggests that the rate of photorespiration was higher at midday than in early morning because low g(s) at midday caused a reduction in leaf intercellular CO(2) concentration. We conclude that low P(n) at midday was the result of both a reduction in the photochemical process and an increase in stomatal limitation.     

Light-use properties in two sun-adapted shrubs with contrasting canopy structures

We investigated the impact of high solar irradiance and elevated temperature on carbon gain by two, co-occurring, sun-adapted, dwarf shrub species, Planchonella obovata var. dubia (Koidz.) Hatusima and Hibiscus glaber Matsumura, growing on sun-exposed ridges in the Bonin Islands, in the subtropical Pacific Ocean. Planchonella had steeply inclined, longer lived, sclerophyllous leaves, whereas Hibiscus has thinner, more horizontally oriented, and shorter lived leaves. We tested the hypothesis that leaf physiological tolerance to high light is lower in Planchonella than in Hibiscus. Under relatively high irradiances (photosynthetic photon flux density, PPFD, > 500 micromol m(-2) s(-1)), net photosynthetic rate (P(n)) was about 8.0 and 0.4 micromol m(-2) s(-1) in mature and young leaves of Planchonella, and about 12.4 and 10.3 micromol m(-2) s(-1) in mature and young leaves of Hibiscus, respectively. Both P(n) and photosystem II (PSII) quantum yield at a given PPFD were lower in Planchonella than in Hibiscus, whereas non-photochemical quenching (NPQ) at a given PPFD was higher in Planchonella. When leaf discs were exposed to high light (1900 micromol m(-2) s(-1) PPFD) at 37, 40 or 43 degrees C for 3 h, the recovery of PSII quantum yield (F(v)/F(m)) in the following 60-min dark period was slower in Planchonella than in Hibiscus, indicating that the ability of PSII to tolerate high light and high temperature was less in Planchonella than in Hibiscus. We postulate that there is a linkage between leaf display and leaf photochemical ability in sun-adapted shrub species.     

Interactive effects of leaf age and self-shading on leaf structure,photosynthetic capacity and chlorophyll fluorescence in the rain forest tree,Dryobalanops aromatica

In the tropical canopy tree, Dryobalanops aromatica Gaertn. f., upper-canopy leaves (UL) develop under sunlit conditions but are subjected to self-shading within the crown as they age. In contrast, lower-canopy leaves (LL) are exposed to uniform dim light conditions throughout their life span. By comparing leaf morphology and physiology of UL and LL, variations in leaf characteristics were related to leaf age and self-shading. Mass-based chlorophyll (chl) concentration and the chlorophyll/nitrogen (chl/N) ratio were lower and the chl a/b ratio was higher in UL than in LL. In UL, the chl/N ratio gradually increased and the chl a/b ratio gradually decreased with leaf aging, whereas these ratios remained unchanged with leaf age in LL. The effective quantum yield of photosystem II (PSII) (DeltaF/F(m)') at a given irradiance remained unchanged with leaf age in LL, whereas DeltaF/F(m)' changed with leaf age in UL. These data indicate N reallocation within the leaves from carbon fixation components to light harvesting components and a dynamic regulation of photochemical processes of PSII in response to increased self-shading of UL. Despite the difference in light environment with leaf age between UL and LL, maximum photosynthetic rates and nitrogen-use efficiency decreased with leaf aging in both UL and LL. Constancy in the chl/N ratio with leaf age in LL indicated that the decrease in photosynthetic capacity was caused by effects other than shading, such as leaf aging. We conclude that N reallocation and acclimation of PSII to self-shading occurred even in mature leaves, whereas the change in photosynthetic capacity with leaf age was more conservative.     

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.     

Interrelationships among light,photosynthesis and nitrogen in the crown of mature Pinus contorta ssp. latifolia

Scaling leaf-level measurements to estimate carbon gain of entire leaf crowns or canopies requires an understanding of the distribution of photosynthetic capacity and corresponding light microenvironments within a crown. We have compared changes in the photosynthetic light response and nitrogen (N) content (per unit leaf area) of Pinus contorta Dougl. ssp. latifolia Engelm. (lodgepole pine) leaves in relation to their age and light microenvironment. The vertical gradient in integrated daily photosynthetic photon flux density (PPFD) from the upper to the lower crown of lodgepole pine was similar in magnitude to the horizontal gradient in daily PPFD along shoots from young to old leaves. The relationship between light-saturated net photosynthesis (A(max)) and daily PPFD was significant for both young and old leaves. However, old leaves had a lower A(max) than young leaves in a similar daily irradiance regime. For leaves of all ages from throughout the crown, A(max) was linearly related to the estimated daily net carbon gain that leaves could achieve in their natural PPFD environment (estimated A(day)) (r(2) = 0.84, P < 0.001, n = 39), indicating that estimated A(day) may be dominated by carbon fixed when leaves are light-saturated and operating at A(max). Comparison of the PPFD required to achieve A(max) and the PPFD available to the leaves showed that all of the measured leaves (n = 39), regardless of their position in the crown or age, were in light environments that could light-saturate photosynthesis for a similar proportion of the day. For all data pooled, foliar N was weakly correlated with daily PPFD. Analyzing each leaf age class separately showed that foliar N was significantly related to daily PPFD, A(max), and estimated A(day) for the youngest leaves but not for middle-aged or old leaves. Therefore, the general theory that foliar N is allocated within a crown according to total daily light availability was supported only for young (1-4 years old) leaves in this study.     

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.     

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.     

Shoot structure and growth along a vertical profile within a Populus-Tilia canopy

We investigated shoot growth patterns and their relationship to the canopy radiation environment and the distribution of leaf photosynthetic production in a 27-m-tall stand of light-demanding Populus tremula L. and shade-tolerant Tilia cordata Mill. The species formed two distinct layers in the leaf canopy and showed different responses in branch architecture to the canopy light gradient. In P. tremula, shoot bifurcation decreased rapidly with decreasing light, and leaf display allowed capture of multidirectional light. In contrast, leaf display in T. cordata was limited to efficient interception of unidirectional light, and shoot growth and branching pattern facilitated relatively rapid expansion into potentially unoccupied space even in the low light of the lower canopy. At the canopy level, T. cordata had higher photosynthetic light-use efficiency than P. tremula, whereas P. tremula had higher nitrogen-use efficiency than T. cordata. However, at the individual leaf level, both species had similar efficiencies under comparable light conditions. Production of new leaf area in the canopy followed the pattern of photosynthetic production. However, the species differed substantially in extension growth and space-filling strategy. Light-demanding P. tremula expanded into new space with a few long shoots, with shoot length strongly dependent on photosynthetic photon flux density (PPFD). Production of new leaf area and extension growth were largely uncoupled in this species because short shoots, which do not contribute to extension growth, produced many new leaves. Thus, in P. tremula, the growth pattern was strongly directed toward the top of the canopy. In contrast, in shade-tolerant T. cordata, shoot growth was weakly related to PPFD and more was invested in long shoot growth on a leaf area basis compared with P. tremula. However, this extension growth was not directed and may serve as a passive means of avoiding self-shading. This study supports the hypothesis that, for a particular species, allocation patterns and crown architecture contribute as much to shade tolerance as leaf-level photosynthetic acclimation.     

Diurnal changes in leaf gas exchange characteristics in the uppermost canopy of a rain forest tree, Dryobalanops aromatica Gaertn. f

Dryobalanops aromatica Gaertn. f. is a major tropical canopy species in lowland tropical rain forests in Peninsular Malaysia. Diurnal changes in net photosynthetic rate (A) and stomatal conductance to water vapor (g(s)) were measured in fully expanded young and old leaves in the uppermost canopy (35 m above ground). Maximum A was 12 and 10 micro mol m(-2) s(-1) in young and old leaves, respectively; however, because of large variation in A among leaves, mean maximum A in young and old leaves was only 6.6 and 5.5 micro mol m(-2) s(-1), respectively. Both g(s) and A declined in young leaves when T(leaf) exceeded 34 degrees C and leaf-to-air vapor pressure deficit (DeltaW) exceeded 0.025, whereas in old leaves, g(s) and A did not start to decline until T(leaf) and DeltaW exceeded 36 degrees C and 0.035, respectively. Under saturating light conditions, A was linearly related to g(s). The coefficient of variation (CV) for the difference between the CO(2) concentrations of ambient air and the leaf intercellular air space (C(a) - C(i)) was smaller than the CV for A or g(s), suggesting that maximum g(s) was mainly controlled by mesophyll assimilation (A/C(i)). Minimum C(i)/C(a) ratios were relatively high (0.72-0.73), indicating a small drought-induced stomatal limitation to A and non-conservative water use in the uppermost canopy leaves.     

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.     

Vertical gradients in photosynthetic light response within an old-growth Douglas-fir and western hemlock canopy

We examined needle-level light response of photosynthesis across a vertical light gradient within 45-55-m-tall western hemlock (Tsuga heterophylla (Raf.) Sarg.) and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees growing in a 400-500-year-old mixed species stand. We determined: (1) whether light-saturated photosynthetic rates, light compensation points, and respiration rates varied from the upper to the lower canopy, and (2) if light-saturated photosynthetic rates, light compensation points, and respiration rates varied between Douglas-fir and western hemlock. Over a 25-m gradient from the canopy top to the lower canopy, mean light-saturated photosynthetic rates, light compensation points, and respiration rates declined in overstory Douglas-fir and western hemlock needles, paralleling a 65% decline in the mean daily photosynthetic photon flux density (PPFD). At the canopy top, increasing light-saturated photosynthetic rates relative to lower canopy needles increased carbon uptake at high PPFD. In the lower canopy, reduced respiration rates relative to upper canopy needles increased carbon uptake at low PPFD by reducing the light compensation point. At all canopy positions, western hemlock had lower mean light-saturated photosynthetic rates, light compensation points and respiration rates than Douglas-fir. As a result, western hemlock had higher net photosynthetic rates at low PPFD, but lower net photosynthetic rates at high PPFD compared with Douglas-fir.     

Light availability and photosynthesis of Pseudotsuga menziesii seedlings grown in the open and in the forest understory

The light environment, photosynthetic dynamics and steady-state net photosynthetic rates of lateral branch shoots of Pseudotsuga menziesii var. glauca (Beissn.) Franco seedlings growing in the open and in the forest understory were investigated in situ. Mean incident photosynthetic photon flux density (PPFD) was 702.5 micro mol m(-2) s(-1) on open-grown branches and 52.0 micro mol m(-2) s(-1) on understory-grown branches. Mean daily durations of PPFD greater than 500, 200, and 50 micro mol m(-2) s(-1) were 8.5, 31.5, and 270.3 min, respectively, on understory-grown branches, and 559.1, 700.7, and 803.3 min, respectively, on open-grown branches. Sunflecks accounted for 32.4% of total daily photosynthetically active radiation incident on understory branches. Following 10 min at a PPFD of 50 micro mol m(-2) s(-1), the induction time required for net photosysnthesis to reach 50 and 90% of steady-state rates was shorter at a PPFD of 200 than at a PPFD of 500 micro mol m(-2) s(-1) and shorter in understory-grown branches than in open-grown branches. On a leaf area basis, dark respiration rates of understory-grown branches were lower and net photosynthetic rates were higher than those of open-grown branches exposed to low PPFD. However, at high PPFDs, understory-grown branches had lower photosynthetic rates than open-grown branches. When measurements were expressed on a leaf dry mass basis, there was no difference in dark respiration rates between understory branches and open-grown branches, but net photosynthetic rates of understory branches were equal to or higher than those of open-grown branches at all PPFDs.     

Fruit load and branch ring-barking affect carbon allocation and photosynthesis of leaf and fruit of Coffea arabica in the field

Increasing fruit load (from no berries present to 25, 50 and 100% of the initial fruit load) significantly decreased branch growth on 5-year-old coffee (Coffea arabica L.) trees of the dwarf cultivar 'Costa Rica 95', during their third production cycle. Ring-barking the branches further reduced their growth. Berry dry mass at harvest was significantly reduced by increasing fruit load. Dry matter allocation to berries was four times that allocated to branch growth during the cycle. Branch dieback and berry drop were significantly higher at greater fruit loads. This illustrates the importance of berry sink strength and indicates that there is competition for carbohydrates between berries and shoots and also among berries. Leaf net photosynthesis (P(n)) increased with increasing fruit load. Furthermore, leaves of non-isolated branches bearing full fruit load achieved three times higher P(n) than leaves of isolated (ring-barked) branches without berries, indicating strong relief of leaf P(n) inhibition by carbohydrate demand from berries and other parts of the coffee tree when excess photoassimilates could be exported. Leaf P(n) was significantly higher in the morning than later during the day. This reduction in leaf P(n) is generally attributed to stomatal closure in response to high irradiance, temperature and vapor pressure deficit in the middle of the day; however, it could also be a feedback effect of reserves accumulating during the morning when climatic conditions for leaf P(n) were optimal, because increased leaf mass ratio was observed in leaves of ring-barked branches with low or no fruit loads. Rates of CO(2) emission by berries decreased and calculated photosynthetic rates of berries increased with increasing photosynthetic photon flux (PPF) especially at low PPFs (0 to 100 micromol m(-2) s(-1)). The photosynthetic contribution of berries at the bean-filling stage was estimated to be about 30% of their daily respiration costs and 12% of their total carbon requirements at PPF values commonly experienced in the field (200 to 500 micromol m(-2) s(-1)).     

Three-dimensional lamina architecture alters light-harvesting efficiency in Fagus: a leaf-scale analysis

Modification of foliage exposition and morphology by seasonal average integrated quantum flux density (Qint) was investigated in the canopies of the shade-tolerant late-successional deciduous tree species Fagus orientalis Lipsky and Fagus sylvatica L. Because the leaves were not entirely flat anywhere in the canopy, the leaf lamina was considered to be three-dimensional and characterized by the cross-sectional angle between the leaf halves (theta). Both branch and lamina inclination angles with respect to the horizontal scaled positively with irradiance in the canopy, allowing light to penetrate to deeper canopy horizons. Lamina cross-sectional angle varied from 170 degrees in the most shaded leaves to 90-100 degrees in leaves in the top of the canopy. Thus, the degree of leaf rolling increased with increasing Qint, further reducing the light-interception efficiency of the upper-canopy leaves. Simulations of the dependence of foliage light-interception efficiency on theta demonstrated that decreases in theta primarily reduce the interception efficiency of direct irradiance, but that diffuse irradiance was equally efficiently intercepted over the entire range of theta values in our study. Despite strong alteration in foliage light-harvesting capacity within the canopy and greater transmittance of the upper crown compared with the lower canopy, mean incident irradiances varied more than 20-fold within the canopy, indicating inherent limitations in light partitioning within the canopy. This extensive canopy light gradient was paralleled by plastic changes in foliar structure and chemistry. Leaf dry mass per unit area varied 3-4-fold between the canopy top and bottom, providing an important means of scaling foliage nitrogen contents and photosynthetic capacity per unit area with Qint. Although leaf structure versus light relationships were qualitatively similar in all cases, there were important tree-to-tree and species-to-species variations, as well as evidence of differences in investments in structural compounds within the leaf lamina, possibly in response to contrasting leaf water availability in different trees.     

Photosynthetic capacity in relation to nitrogen in the canopy of a Quercus robur, Fraxinus angustifolia and Tilia cordata flood plain forest

We measured gas exchange and various leaf parameters of ash (Fraxinus angustifolia Vahl.) and oak (Quercus robur L.) in the high canopy and of lime (Tilia cordata Mill.) in the lower canopy of a planted, 120-year-old floodplain forest in southern Moravia, Czech Republic. The high-canopy leaves of F. angustifolia and Q. robur had nitrogen concentrations on a leaf area basis (N(area)) that were twice those of low-canopy leaves of T. cordata. Upper-canopy leaves of F. angustifolia had a photosynthetic rate at light saturation (A(max)) of about 16 micromol CO2 m(-2) s(-1), whereas A(max) of the upper-canopy foliage of Q. robur achieved only about two thirds of this value. Contrary to previous investigations of photosynthetic performance in monospecific stands, leaves of the uppermost branches of T. cordata at 15-m height had the highest A(max) and transpiration rate among the species studied. Water-use efficiency (WUE) was low in T. cordata at 15-m canopy height, whereas WUE was significantly higher for Q. robur leaves at 27-m height than for the other species. Leaves of T. cordata at 15-m height showed the strongest relationship between A(max) and N(area) (R2 = 0.90) followed by F. angustifolia (R2 = 0.69). The strong correlation between photosynthesis and nitrogen concentration in T. cordata at 15 m, together with the steep regression slope for the A(max):N(area) relationship, indicated that nitrogen allocation to the photosynthetic apparatus resulted in high nitrogen-use efficiency of light-saturated photosynthesis (PNUE). Despite differences in PNUE among species, PNUE was fairly constant for leaves sampled from the same canopy position, suggesting that single-leaf parameters are matched to optimize PNUE for prevailing light conditions. High PNUE in T. cordata at 15 m partially compensated for the species' subordinate position in the canopy, and may be an important mechanism for its coexistence in highly structured vegetation.     

Seasonal changes in photosynthesis of trees in the flooded forest of the Mapire River

We studied the flood tolerance of five tree species growing in the flooded forest adjacent to the Mapire river, in SW Venezuela. Mean photosynthetic rate and leaf conductance were 11 &mgr;mol m(-2) s(-1) and 700 mmol m(-2) s(-1), respectively. Xylem water potential ranged from -0.08 to -1.15 MPa. Based on leaf gas exchange as a criterion of tolerance to flooding, two response patterns were identified: (1) decreasing photosynthetic rate with increasing flooding and leaf conductance (Psidium ovatifolium Berg. ex Desc., Campsiandra laurifolia Benth., Symmeria paniculata Benth. and Acosmium nitens (Vog.) Benth); and (2) independence of photosynthesis and leaf conductance from flooding (Eschweilera tenuifolia (Berg.) Miers.). In the first response pattern, declining photosynthetic rate with flooding may be interpreted as a sign of reduced flood tolerance, whereas the second response pattern may indicate increased flood tolerance. An increase in xylem water potential with depth of water column was found for all species (with the possible exception of P. ovatifolium), indicating that flooding does not cause water stress in these trees. Submerged leaves that had been under water for between four days and four months generally had photosynthetic rates and leaf conductances similar to those of aerial leaves, indicating maintenance of photosynthetic capacity under water. Daily positive oscillations in glucan content in submerged leaves of P. ovatifolium and C. laurifolia suggest that submerged leaves do not represent a sink for photosynthates produced by aerial leaves.     

Does canopy mean nitrogen concentration explain variation in canopy light use efficiency across 14 contrasting forest sites?

The maximum light use efficiency (LUE?=?gross primary production (GPP)/absorbed photosynthetic photon flux density (aPPFD)) of plant canopies has been reported to vary spatially and some of this variation has previously been attributed to plant species differences. The canopy nitrogen concentration [N] can potentially explain some of this spatial variation. However, the current paradigm of the N-effect on photosynthesis is largely based on the relationship between photosynthetic capacity (A(max)) and [N], i.e., the effects of [N] on photosynthesis rates appear under high PPFD. A maximum LUE-[N] relationship, if it existed, would influence photosynthesis in the whole range of PPFD. We estimated maximum LUE for 14 eddy-covariance forest sites, examined its [N] dependency and investigated how the [N]-maximum LUE dependency could be incorporated into a GPP model. In the model, maximum LUE corresponds to LUE under optimal environmental conditions before light saturation takes place (the slope of GPP vs. PPFD under low PPFD). Maximum LUE was higher in deciduous/mixed than in coniferous sites, and correlated significantly with canopy mean [N]. Correlations between maximum LUE and canopy [N] existed regardless of daily PPFD, although we expected the correlation to disappear under low PPFD when LUE was also highest. Despite these correlations, including [N] in the model of GPP only marginally decreased the root mean squared error. Our results suggest that maximum LUE correlates linearly with canopy [N], but that a larger body of data is required before we can include this relationship into a GPP model. Gross primary production will therefore positively correlate with [N] already at low PPFD, and not only at high PPFD as is suggested by the prevailing paradigm of leaf-level A(max)-[N] relationships. This finding has consequences for modelling GPP driven by temporal changes or spatial variation in canopy [N].     

Effects of light availability on leaf gas exchange and expansion in lychee (Litchi chinensis)

Effects of photosynthetic photon flux density (PPFD) on leaf gas exchange of lychee (Litchi chinensis Sonn.) were studied in field-grown "Kwai May Pink" and "Salathiel" orchard trees and young potted "Kwai May Pink" plants during summer in subtropical Queensland (27 degrees S). Variations in PPFD were achieved by shading the trees or plants 1 h before measurement at 0800 h. In a second experiment, potted seedlings of "Kwai May Pink" were grown in a heated greenhouse in 20% of full sun (equivalent to maximum noon PPFD of 200 micromol m(-2)xs(-1)) and their growth over three flush cycles was compared with seedlings grown in full sun (1080 micromol m(-2)xs(-1)). Young potted plants of "Kwai May Pink" were also grown outdoors in artificial shade that provided 20, 40, 70 or 100% of full sun (equivalent to maximum PPFDs of 500, 900, 1400 and 2000 micromol m(-2)xs(-1)) and measured for shoot extension and leaf area development over one flush cycle. Net CO2 assimilation increased asymptotically in response to increasing PPFD in both orchard trees and young potted plants. Maximum rates of CO2 assimilation (11.9 +/- 0.5 versus 6.3 +/- 0.2 micromol CO2 m(-2) s(-1)), dark respiration (1.7 +/- 0.3 versus 0.6 +/- 0.2 micromol CO2 m(-2) s(-1)), quantum yield (0.042 +/- 0.005 versus 0.027 +/- 0.003 mol CO2 mol(-1)) and light saturation point (1155 versus 959 micromol m(-2) s(-1)) were higher in orchard trees than in young potted plants. In potted seedlings grown in a heated greenhouse, shoots and leaves exposed to full sun expanded in a sigmoidal pattern to 69 +/- 12 mm and 497 +/- 105 cm(2) for each flush, compared with 27 +/- 7 mm and 189 +/- 88 cm(2) in shaded seedlings. Shaded seedlings were smaller and had higher shoot:root ratios (3.7 versus 3.1) than seedlings grown in full sun. In the potted plants grown outdoors in 20, 40, 70 or 100% of full sun, final leaf area per shoot was 44 +/- 1, 143 +/- 3, 251 +/- 7 and 362 +/- 8 cm(2), respectively. Shoots were also shorter in plants grown in shade than in plants grown in full sun (66 +/- 5 mm versus 101 +/- 2 mm). Photosynthesis in individual leaves of lychee appeared to be saturated at about half full sun, whereas maximum leaf expansion occurred at higher PPFDs. We conclude that lychee plants can persist as seedlings on the forest floor, but require high PPFDs for optimum growth.     

Photosynthesis patterns during the establishment year within two Populus clones with contrasting morphology and phenology

Diurnal and seasonal photosynthesis patterns were studied in poplar clones Populus tristis Fisch. x P. balsamfera L. cv. Tristis #1 (NC 5260) and Populus x euramericana (Dode) Guiner cv. Eugenei (NC 5326, Carolina poplar) during their first season in the field in a short rotation, intensive culture plantation. Photosynthetic rates were low in immature leaves; increased basipetally on the shoot and peaked in leaves that had recently reached full expansion; and thereafter declined in lower-crown leaves in both clones. Photosynthesis was associated with leaf age and stomatal conductance in immature leaves; adaxial photosynthetic photon flux density (PPFD) and leaf temperature in recently mature leaves; and leaf age and adaxial PPFD in lower-crown leaves. Diurnal photosynthesis patterns within trees were highly variable due to differential light interception among leaves. Results of clonal comparisons of photosynthetic rates were dependent on which leaves were pooled for comparison and how photosynthesis was expressed. Compared to Eugenei, Tristis produced smaller leaves which had higher unit-area photosynthesis rates. The more indeterminate Eugenei outgrew Tristis principally because it more fully utilized the growing season for leaf area production. Photosynthetic production integrated over the growing season was closely related to dry matter production in both clones.     

Leaf morphology and photosynthetic adjustments among deciduous broad-leaved trees within the vertical canopy profile

Photosynthetic acclimation of deciduous broad-leaved tree species was studied along a vertical gradient within the canopy of a multi-species deciduous forest in northern Japan. We investigated variations in (1) local light regime and CO2 concentration ([CO2]), and (2) morphological (area, thickness and area per mass), biochemical (nitrogen and chlorophyll concentrations) and physiological (light-saturated photosynthetic rate) attributes of leaves of seven major species on three occasions (June, August and October). We studied early successional species, alder (Alnus hirsuta (Spach) Rupr.) and birch (Betula platyphylla var. japonica (Miq.) Hara); gap phase species, walnut (Juglans ailanthifolia Carrière) and ash (Fraxinus mandshurica var. japonica Rupr.); mid-successional species, basswood (Tilia japonica (Miq.) Simonk.) and elm (Ulmus davidiana var. japonica (Rehd.) Nakai); and the late-successional species, maple (Acer mono Bunge). All but maple initiated leaf unfolding from the lower part of the crown. The [CO2] within the vertical profile ranged from 320-350 ppm in the upper canopy to 405-560 ppm near the ground. The lowest and highest ambient [CO2] occurred during the day and during the night, respectively. This trend was observed consistently during the summer, but not when trees were leafless. Chlorophyll concentration was positively related to maximum photosynthetic rate within, but not among, species. Leaf senescence started from the inner part of the crown in alder and birch, but started either in the outer or top portion of the canopy of ash, basswood and maple. Chlorophyll (Chl) to nitrogen ratio in leaves increased with decreasing photon flux density. However, Chl b concentration in all species remained stable until the beginning of leaf senescence. Maximum photosynthetic rates observed in sun leaves of early successional species, gap phase or mid-successional species, and late successional species were 12.5-14.8 micromol m(-2) s(-1), 4.1-7.8 micromol m(-2) s(-1) and 3.1 micromol m(-2) s(-1), respectively.