A biochemical model of photosynthesis for mango leaves: evidence for the effect of fruit on photosynthetic capacity of nearby leaves

Variations in leaf nitrogen concentration per unit mass (Nm) and per unit area (Na), mass-to-area ratio (Ma), total nonstructural carbohydrates (Ta), and photosynthetic capacity (maximum carboxylation rate, electron transport capacity, rate of phosphate release in triose phosphate utilization and dark respiration rate) were studied within the digitized crowns of two 3-year-old mango trees (Mangifera indica L.) on La Réunion Island. Additional measurements of Nm, Na, Ma, Ta and photosynthetic capacities were performed on young, fully expanded leaves of 11-year-old mango trees. Leaves of similar gap fractions were taken far from and close to developing fruits. Unlike Nm, both Na and Ta were linearly correlated to gap fraction. Similar relationships were found for all leaves whatever their age and origin, except for Ta, for which we found a significant tree effect. Photosynthetic capacity was nonlinearly correlated to Na, and a unique relationship was obtained for all types of leaves. Photosynthetic acclimation to light was mainly driven by changes in Ma, but allocation of total leaf N between the different photosynthetic functions also played a substantial role in acclimation to the lowest irradiances. Leaves close to developing fruits exhibited a higher photosynthetic capacity than other leaves, but similar Ta. Our data suggest that Ta does not control photosynthetic capacity in mango leaves. We used the data to parameterize a biochemically based model of photosynthesis and an empirical stomatal conductance model, allowing accurate predictions of net photosynthesis of leaves in field-grown mango trees.     

Effects of crown development on leaf irradiance,leaf morphology and photosynthetic capacity in a peach tree

The three-dimensional (3-D) architecture of a peach tree (Prunus persica L. Batsch) growing in an orchard near Avignon, France, was digitized in April 1999 and again four weeks later in May 1999 to quantify increases in leaf area and crown volume as shoots developed. A 3-D model of radiation transfer was used to determine effects of changes in leaf area density and canopy volume on the spatial distribution of absorbed quantum irradiance (PAR(a)). Effects of changes in PAR(a) on leaf morphological and physiological properties were determined. Leaf mass per unit area (M(a)) and leaf nitrogen concentration per unit leaf area (N(a)) were both nonlinearly related to PAR(a), and there was a weak linear relationship between leaf nitrogen concentration per unit leaf mass (N(m)) and PAR(a). Photosynthetic capacity, defined as maximal rates of ribulose-1,5-bisphosphate carboxylase (Rubisco) carboxylation (V(cmax)) and electron transport (J(max)), was measured on leaf samples representing sunlit and shaded micro-environments at the same time that the tree crown was digitized. Both V(cmax) and J(max) were linearly related to N(a) during May, but not in April when the range of N(a) was low. Photosynthetic capacity per unit N(a) appeared to decline between April and May. Variability in leaf nitrogen partitioning between Rubisco carboxylation and electron transport was small, and the partitioning coefficients were unrelated to N(a). Spatial variability in photosynthetic capacity resulted from acclimation to varying PAR(a) as the crown developed, and acclimation was driven principally by changes in M(a) rather than the amount or partitioning of leaf nitrogen.     

Changes in photosynthesis and leaf characteristics with tree height in five dipterocarp species in a tropical rain forest

Variations in leaf photosynthetic, morphological and biochemical properties with increasing plant height from seedlings to emergent trees were investigated in five dipterocarp species in a Malaysian tropical rain forest. Canopy openness increased significantly with tree height. Photosynthetic properties, such as photosynthetic capacity at light saturation, light compensation point, maximum rate of carboxylation and maximum rate of photosynthetic electron transport, all increased significantly with tree height. Leaf morphological and biochemical traits, such as leaf mass per area, palisade layer thickness, nitrogen concentration per unit area, chlorophyll concentration per unit dry mass and chlorophyll to nitrogen ratio, also changed significantly with tree height. Leaf properties had simple and significant relationships with tree height, with few intra- and interspecies differences. Our results therefore suggest that the photosynthetic capacity of dipterocarp trees depends on tree height, and that the trees adapt to the light environment by adjusting their leaf morphological and biochemical properties. These results should aid in developing models that can accurately estimate carbon dioxide flux and biomass production in tropical rain forests.     

Modelling functional trait acclimation for trees of different height in a forest light gradient: emergent patterns driven by carbon gain maximization

Forest trees show large changes in functional traits as they develop from a sapling in the shaded understorey to an adult in the light-exposed canopy. The adaptive function of such changes remains poorly understood. The carbon gain hypothesis suggests that these changes should be adaptive (acclimation) and that they serve to maximize net vegetative or reproductive growth. We explore the carbon gain hypothesis using a mechanistic model that combines an above-ground plant structure, a biochemical photosynthesis model and a biophysical stomatal conductance model. Our simulations show how forest trees that maximize their carbon gain increase their total leaf area, sapwood area and leaf photosynthetic capacity with tree height and light intensity. In turn, they show how forest trees increased crown stomatal conductance and transpiration, and how the carbon budget was affected. These responses in functional traits to tree height (and light availability) largely differed from the responses exhibited by exposed trees. Forest and exposed trees nevertheless shared a number of emergent patterns: they showed a similar decrease in the average leaf water potential and intercellular CO(2) concentration with tree height, and kept almost constant values for the ratio of light absorption to electron transport capacity, the ratio of photosynthetic capacity to water supply capacity, and nitrogen partitioning between electron transport and carboxylation. While most of the predicted qualitative responses in individual traits are consistent with field or lab observations, the empirical support for capacity balances is scarce. We conclude that modelling functional trait optimization and carbon gain maximization from underlying physiological processes and trade-offs generates a set of predictions for functional trait acclimation and maintenance of capacity balances of trees of different height in a forest light gradient, but actual tests of the predicted patterns are still scarce.     

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.     

Distribution of leaf mass per unit area and leaf nitrogen concentration determine partitioning of leaf nitrogen within tree canopies

Distribution of leaf nitrogen with respect to leaf mass per unit area (M(a)), nitrogen per unit mass (N(m)) and nitrogen per unit area (N(a)) within peach (Prunus persica L.) tree canopies was studied in two field experiments. In one experiment, leaf light exposure and M(a) were measured on leaves from different canopy positions of peach trees subjected to five nitrogen (N) fertilization treatments. Leaf light exposure and M(a) were linearly related and the relationship was independent of N fertilization. In a subsequent experiment, N fertilizer was applied to previously unfertilized trees in midsummer, after shoot growth had terminated. Application of N fertilizer did not affect mean canopy M(a). Fertilization increased N(m) of all leaves throughout the canopy compared with non-fertilized trees. No significant relationship between N(m) and M(a) was found in either fertilized or control trees. There was a linear relationship between N(a) and M(a) and the slope of the relationship was increased by N fertilizer application. We conclude that distribution of N(a) in peach tree canopies is primarily a function of M(a) partitioning with light and N(m), which is related to soil N availability.     

Photosynthetic capacity and foliar nitrogen distribution in Eucalyptus nitens is altered by high-intensity thinning

The changes in photosynthetic rates, light environment and foliar nutrient concentrations following thinning were examined in an 8-year-old Eucalyptus nitens (Deane and Maiden) Maiden plantation. The objectives of the study were to: (1) determine the extent to which maximum photosynthetic rates (Amax) of E. nitens are affected by stand thinning; (2) relate the spatial pattern of Amax within the crown to the changes in light environment caused by thinning; and (3) establish if the responses of Amax to thinning are driven by changes in area-based foliar nitrogen (Na) or phosphorus (Pa) concentrations. Photosynthetic rates measured under saturating light availability increased throughout the crown after thinning. The greatest increases were observed in the lower and middle crown zones. Photosynthetic rate was positively related to foliar N concentration. Thinning increased Na and Pa because of a significant decrease in specific leaf area (SLA) after thinning. Changes in photosynthetic rates, SLA and foliar nutrient distributions with thinning were well correlated with changes in incident solar irradiance throughout the tree crown.     

Spatial and temporal variations in leaf area index, specific leaf area and leaf nitrogen of two co-occurring savanna tree species

Foliage growth, mass- and area-based leaf nitrogen concentrations (Nm and N a) and specific leaf area (SLA) were surveyed during a complete vegetation cycle for two co-occurring savanna tree species: Crossopteryx febrifuga (Afzel. ex G. Don) Benth. and Cussonia arborea A. Rich. The study was conducted in the natural reserve of Lamto, Ivory Coast, on isolated and clumped trees. Leaf flush occurred before the beginning of the rainy season. Maximum leaf area index (LAI), computed on a projected canopy basis for individual trees, was similar (mean of about 4) for both species. Seasonal courses of the ratio of actual to maximum LAI were similar for individuals of the same species, but differed between species. For C. febrifuga, clumped trees reached their maximum LAI before isolated trees. The LAI of C. arborea trees did not differ between clumped and isolated individuals, but maximum LAI was reached about 2 months later than for C. febrifuga. Leaf fall was associated with decreasing soil water content for C. arborea. For C. febrifuga, leaf fall started before the end of the rainy period and was independent of changes in soil water content. These features lead to a partial niche separation in time for light resource acquisition between the two species. Although Nm, N a and SLA decreased with time, SLA and N a decreased later in the vegetation cycle for C. arborea than for C. febrifuga. For both species, N a decreased and SLA increased with decreasing leaf irradiance within the canopy, although effects of light on leaf characteristics did not differ between isolated and clumped trees. Given relationships between N a and photosynthetic capacities previously reported for these species, our results show that C. arborea exhibits higher photosynthetic capacity than C. febrifuga during most of the vegetation cycle and at all irradiances.     

Foliar morphological and physiological plasticity in Picea abies and Abies alba saplings along a natural light gradient

The role of morphological versus physiological foliar plasticity in the capacity for, and mechanisms of, photosynthetic acclimation was assessed in Picea abies (L.) Karst. and Abies alba Mill. saplings in a forest gap-understory light gradient (relative irradiance, RI, ranging from 0.02 to 0.32). The species investigated showed a similar foliar morphological plasticity along the light gradient, at both the needle level (through alteration in leaf dry mass per area) and the shoot level (through alteration in the silhouette area ratio, e.g., shoot silhouette to projected needle area ratio). In both species chlorophyll (Chl) concentration on a mass basis decreased at increasing RI, but was independent of RI when expressed on an area basis. In contrast, leaf N concentration on a mass basis was independent of RI, but was positively influenced by RI when expressed on an area basis. The parameters describing photosynthetic performance at low light (dark respiration rate, apparent quantum yield and light compensation point) suggest that Abies alba was better suited to maintain a positive carbon balance in shaded conditions. By contrast, parameters describing biochemical capacity at high light (maximum electron transport rate, Jmax and maximum ribulose-1,5-biphosphate carboxylation capacity, Vcmax) indicate that only Picea abies was capable of acclimating physiologically to high photosynthetic photon flux densities (PPFDs) by increasing nitrogen partitioning to Rubisco and Vcmax/mass by increasing RI. These results support the hypothesis that interspecific differences in nitrogen partitioning within the photosynthetic apparatus may provide a mechanistic basis for species separation along a light gradient. The differences in photosynthetic plasticity observed are likely to influence regeneration patterns and habitat breadth of the species investigated. The limited ability of Abies alba saplings to exploit high-light conditions may be a competitive disadvantage in large canopy gaps and thus limit recruitment of this species to small gaps.     

Defoliation and nitrogen effects on photosynthesis and growth of Eucalyptus globulus

Plant responses to defoliation are complex. We established a field experiment in a nine-month-old Eucalyptus globulus Labill. plantation to examine the effects of pattern (upper crown versus lower crown removal), frequency (single, double or triple defoliation within a 12-month period) and severity (25 versus 38% of leaf area removed) of defoliation and the effect of soil nitrogen (N) on photosynthetic processes and stem growth. The photosynthetic responses observed following defoliation could be attributed to changes in source:sink ratios. Light-saturated CO(2) uptake (A(max)) increased with increasing severity and frequency of defoliation irrespective of defoliation pattern. Seedlings defoliated in autumn did not exhibit increases in A(max) until the following spring, whereas there was no such delay in photosynthetic responses associated with spring defoliation. Application of N before defoliation allowed trees to compensate for the effect of defoliation on stem diameter growth, which could not be explained simply in terms of increases in A(max). The observed increases in stem diameter increment following N fertilization of defoliated trees suggested increases in leaf area development, and there were changes in the leaf area:leaf dry mass ratio that may have increased light absorption by the crown. Nitrogen fertilization also increased partitioning of dry mass to branches at the expense of main stems, suggesting that N supply was important in rebuilding crowns following a defoliation event.     

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.     

Influence of nutrient supply on shade-sun acclimation of Picea abies seedlings: effects on foliar morphology,photosynthetic performance and growth

Norway spruce seedlings (Picea abies Karst.) were grown in low light for one year, under conditions of adequate and limiting nutrition, then transferred to high light. Three months after transfer we measured photosynthesis, leaf nitrogen concentration, leaf chlorophyll concentration and leaf mass per area (LMA) of current-year and 1-year-old shoots; silhouette area ratio (SAR, the ratio of shoot silhouette area to projected needle area) was also measured in current-year shoots. At the foliage level, the effects of light and nutrient treatments differed markedly. Light availability during foliage expansion primarily affected LMA and SAR (morphological acclimation at the needle and shoot level, respectively). By contrast, nutrient supply in high light affected photosynthetic capacity per unit of leaf tissue (physiological acclimation at the cellular level) but did not affect LMA and SAR. The capacity for shade-sun acclimation in foliage formed before transfer to high light differed greatly from that of foliage formed following the transfer. The morphological inflexibility of mature needles (measured by LMA) limited their shade-sun acclimation potential. In contrast, at high nutrient supply, shoots that developed just after the change in photosynthetic photon flux density largely acclimated, both morphologically and physiologically, to the new light environment. The acclimation response of both current- and 1-year-old shoots was prevented by nutrient limitation. Analysis of growth at the whole-plant level largely confirmed the conclusions drawn at the shoot level. We conclude that nutrient shortage subsequent to the opening of a canopy gap may strongly limit the acclimation response of Norway spruce seedlings. Successful acclimation was largely related to the plant's ability to produce sun foliage and adjust whole-plant biomass allocation rapidly.     

Photosynthetic nitrogen-use efficiency in evergreen broad-leaved woody species coexisting in a warm-temperate forest

Photosynthetic nitrogen-use efficiency (PNUE, photosynthetic capacity per unit leaf nitrogen) varies among species from different habitats and correlates with several ecological characteristics such as leaf life span and leaf mass per area. We investigated eight evergreen broad-leaved woody species with different leaf life spans that coexist in a warm-temperate forest. We determined photosynthetic capacity at ambient CO(2) concentration in saturated light, nitrogen concentration, and the concentration of ribulose-1,5-bisphosphate carboxylase (RuBPCase), a key enzyme of photosynthesis and the largest sink of nitrogen in leaves. Each species showed a strong correlation between photosynthetic capacity and RuBPCase concentration, and between RuBPCase concentration and nitrogen concentration. Photosynthetic capacity of leaves decreased with increasing leaf life span, whereas PNUE did not correlate significantly with leaf life span. There was a twofold variation in PNUE among species. This relatively small variation in PNUE is consistent with the argument that species that coexist in a single habitat maintain a similar PNUE. The two components of PNUE-photosynthetic rate per unit RuBPCase and RuBPCase per unit leaf nitrogen-were not significantly correlated with other leaf characteristics such as leaf life span and leaf mass per area. We conclude that differences in PNUE are relatively small among coexisting species and that differences in absolute amounts of photosynthetic proteins lead to differences in photosynthetic productivity among species.     

Interspecific variation of photosynthesis and leaf characteristics in canopy trees of five species of Dipterocarpaceae in a tropical rain forest

Photosynthetic rate, nitrogen concentration and morphological properties of canopy leaves were studied in 18 trees, comprising five dipterocarp species, in a tropical rain forest in Sarawak, Malaysia. Photosynthetic rate at light saturation (Pmax) differed significantly across species, varying from 7 to 18 micro mol m(-2) s(-1). Leaf nitrogen concentration and morphological properties, such as leaf blade and palisade layer thickness, leaf mass per area (LMA) and surface area of mesophyll cells per unit leaf area (Ames/A), also varied significantly across species. Among the relationships with leaf characteristics, Pmax had the strongest correlation with leaf mesophyll parameters, such as palisade cell layer thickness (r2 = 0.76, P < 0.001) and Ames/A (r2 = 0.73, P < 0.001). Leaf nitrogen concentration and Pmax per unit area also had a significant but weaker correlation (r2 = 0.46, P < 0.01), whereas Pmax had no correlation, or only weakly significant correlations, with leaf blade thickness and LMA. Shorea beccariana Burck, which had the highest P(max) of the species studied, also had the thickest palisade layer, with up to five or more layers. We conclude that interspecific variation in photosynthetic capacity in tropical rain forest canopies is influenced more by leaf mesophyll structure than by leaf thickness, LMA or leaf nitrogen concentration.     

Seasonal and yearly variations in light use and nitrogen use by seedlings of four deciduous broad-leaved tree species invading larch plantations

Several deciduous broad-leaved tree species, differing in leaf phenology, invade larch (Larix kaempferii (Lamb.) Carrière) plantations in Japan. The understory light environment of larch forests changes drastically between the leafy and leafless periods. To determine how the invading seedlings exploit the changing light environment, and if phenological differences reflect the light- and nitrogen-use traits of the seedlings, we measured leaf phenology, seasonal changes in light-saturated photosynthetic rate (P(sat)), leaf nitrogen (N) content (N(area)), chlorophyll/nitrogen ratio (Chl/N), specific leaf area (SLA) and N remobilization rate (NRMR) over 3 years. The mid-successional or gap-phase species, Magnolia hypoleuca Siebold & Zucc., had a short leafy period and high P(sat) and NRMR. In contrast, two late-successional tree species, Prunus ssiori Friedr. Schmidt, which undergoes leaf flush before larch, and Carpinus cordata Blume, which maintains green leaves until frost, both had low P(sat) and NRMR but exploited the opportunity for growth during the period when the larch canopy trees were leafless. Quercus mongolica Fisch. ex Ledeb. var. crispula (Blume) Ohashi, a mid-late-successional species that underwent leaf flush at the same time as the overstory larch, had values of photosynthetic parameters between those of the gap-phase and late-successional species. Among species, M. hypoleuca and Q. mongolica had higher photosynthetic rates and photosynthetic N-use efficiencies. In all species, the relationship between N(area) and P(sat) showed species-specific yearly fluctuations; however, there was no yearly fluctuation in the relationship between N(area) and P(sat) at CO2 saturation. Yearly fluctuations in the N(area)-P(sat) relationship appeared to be induced by changes in SLA and N-use characteristics, which in turn are affected by climatic variations.     

Photosynthesis-nitrogen relationships: interpretation of different patterns between Pseudotsuga menziesii and Populus x euroamericana in a mini-stand experiment

We compared photosynthesis-nitrogen relationships of one broad-leaved (poplar; Populus x euroamericana (Dole) Guinier) and one conifer (Douglas-fir; Pseudotsuga menziesii (Mirb.) Franco) species. Plants were grown in large pots to allow free root development and were kept well watered. We determined effects of low, intermediate and high nitrogen supply rates on area-based leaf nitrogen (Na) and chlorophyll concentrations, leaf mass per area (LMA), light-saturated photosynthesis (Amax), maximum carboxylation (Vcmax) and electron transport rate (Jmax), photosynthetic nitrogen-use efficiency (PNUE), and proportions of leaf N in active Rubisco (PR), bioenergetic pools (PB) and the light-harvesting complex (PLH). Nitrogen supply significantly affected leaf Na. Leaf mass per area did not differ between species and was unaffected by the N treatments. In both species, there was a positive correlation between leaf Na and chlorophyll concentration, and between leaf Na and the photosynthetic parameters Amax, Jmax and Vcmax. At comparable leaf Na, however, poplar showed twofold higher PNUE and a threefold steeper slope of the Amax- nitrogen relationship than Douglas-fir. Leaf Na was negatively correlated with PNUE in Douglas-fir but not in poplar. Leaf Na was also negatively correlated with PR, PB and PLH in Douglas-fir, whereas in poplar, a negative correlation was found only for PLH. Parameter PR was significantly higher in poplar than in Douglas-fir. The ratio of CO2 concentration in the intercellular space to that in ambient air was higher in poplar than in Douglas-fir. Overall, our data suggest that differences in the photosynthesis-nitrogen relationship and PNUE between Douglas-fir and poplar primarily reflect a different investment of N to active Rubisco, and possibly a different constraint to CO2 diffusion.     

Ontogenetic changes in stomatal and biochemical limitations to photosynthesis of two co-occurring Mediterranean oaks differing in leaf life span

A quantitative analysis was applied to the stomatal and biochemical limitations to light-saturated net photosynthesis under optimal field conditions in mature trees and seedlings of the co-occurring evergreen oak, Quercus ilex L., and the deciduous oak, Q. faginea Lam. Stomatal limitation to photosynthesis, maximal Rubisco activity and electron transport rate were determined from assimilation versus intercellular leaf carbon dioxide concentration response curves of leaves that were subsequently analyzed for nitrogen (N) concentration, mass per unit area, thickness and percent internal air space. In both species, seedlings had a lower leaf mass per unit area, thickness and leaf N concentration than mature trees. The root system of seedlings during their third year after planting was dominated by a taproot. A lower leaf N concentration of seedlings was associated with lower maximal Rubisco activity and electron transport rate and with assimilation rates similar to or lower than those of mature trees, despite the higher stomatal conductances and potential photosynthetic nitrogen-use efficiencies of seedlings. Consequently, stomatal limitation to photosynthesis increased with tree age in both species. In both seedlings and mature trees, a lower assimilation rate in Q. ilex than in Q. faginea was associated with lower stomatal conductance, N allocation to photosynthetic functions, maximal Rubisco activity and electron transport rate, and potential photosynthetic nitrogen-use efficiency but greater leaf thickness and leaf mass per unit area. Tree-age-related changes differed quantitatively between species, and the characteristics of the two species were more similar in seedlings than in mature trees. Despite higher stomatal conductances, seedlings are more N limited than adult trees, which contributes to lower biochemical efficiency.     

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.     

Effects of elevated carbon dioxide concentration on growth and nitrogen fixation in Alnus glutinosa in a long-term field experiment

Nitrogen-fixing plant species may respond more positively to elevated atmospheric carbon dioxide concentrations ([CO2]) than other species because of their ability to maintain a high internal nutrient supply. A key factor in the growth response of trees to elevated [CO2] is the availability of nitrogen, although how elevated [CO2] influences the rate of N2-fixation of nodulated trees growing under field conditions is unclear. To elucidate this relationship, we measured total biomass, relative growth rate, net assimilation rate (NAR), leaf area and net photosynthetic rate of N2-fixing Alnus glutinosa (L.) Gaertn. (common alder) trees grown for 3 years in open-top chambers in the presence of either ambient or elevated atmospheric [CO2] and two soil N regimes: full nutrient solution or no fertilizer. Nitrogen fixation by Frankia spp. in the root nodules of unfertilized trees was assessed by the acetylene reduction method. We hypothesized that unfertilized trees would show similar positive growth and physiological responses to elevated [CO2] as the fertilized trees. Growth in elevated [CO2] stimulated (relative) net photosynthesis and (absolute) total biomass accumulation. Relative total biomass increased, and leaf nitrogen remained stable, only during the first year of the experiment. Toward the end of the experiment, signs of photosynthetic acclimation occurred, i.e., down-regulation of the photosynthetic apparatus. Relative growth rate was not significantly affected by elevated [CO2] because although NAR was increased, the effect on relative growth rate was negated by a reduction in leaf area ratio. Neither leaf area nor leaf P concentration was affected by growth in elevated [CO2]. Nodule mass increased on roots of unfertilized trees exposed to elevated [CO2] compared with fertilized trees exposed to ambient [CO2]. There was also a biologically significant, although not statistically significant, stimulation of nitrogenase activity in nodules exposed to elevated [CO2]. Root nodules of trees exposed to elevated [CO2] were smaller and more evenly spaced than root nodules of trees exposed to ambient [CO2]. The lack of an interaction between nutrient and [CO2] effects on growth, biomass and photosynthesis indicates that the unfertilized trees maintained similar CO2-induced growth and photosynthetic enhancements as the fertilized trees. This implies that alder trees growing in natural conditions, which are often limited by soil N availability, should nevertheless benefit from increasing atmospheric [CO2].     

Ecophysiology of Cecropia schreberiana saplings in two wind regimes in an elfin cloud forest: growth,gas exchange,architecture and stem biomechanics

The hypothesis that the short, stunted development of elfin cloud forest trees is influenced by strong winds was tested by studying growth, gas exchange and biomechanics of potted saplings of Cecropia schreberiana Miq. ex. C. peltata L. (Cecropiaceae) exposed to two natural wind regimes (exposed and protected) at high elevation sites in the Luquillo Experimental Forest, Puerto Rico. The wind-exposure treatment produced several thigmomorphogenetic responses, including reductions in plant stature and crown area, changes in allocation patterns, and increased root to shoot ratio, leaf abrasion and leaf epinasty. Wind-exposure decreased maximum photosynthetic rate and respiration on an area basis, but not on a leaf-mass basis. Wind-exposed plants had lower apparent quantum yields, and higher light compensation points than wind-protected plants. Photosynthetic nitrogen-use efficiency was lower in wind-exposed plants, but such plants had higher leaf nitrogen concentration than wind-protected plants. There were no effects of treatments on stomatal conductance, transpiration rate and water-use efficiency. Stems of wind-exposed plants had lower second moment of area, apparent modulus of elasticity, flexural stiffness and stem density, but higher water content than wind-protected plants. Tissue-density-specific stiffness and the calculated critical height were not affected by the treatments. Wind-exposed plants were biomechanically less predisposed to bending and failing under their own weight than wind-protected plants because their safety factors were smaller, indicating that maintenance of an ontogenetically less developed structure enables plants to cope with wind loading. Windward trees showed a lower scaling component of the allometric relationship between diameter and height than leeward trees.