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.     

Coordination of crown structure, leaf plasticity and carbon gain within the crowns of three winter-deciduous mature trees

We examined the vertical profiles of leaf characteristics within the crowns of two late-successional (Fagus crenata Blume and Fagus japonica Maxim.) and one early-successional tree species (Betula grossa Sieb. et Zucc.) in a Japanese forest. We also assessed the contributions of the leaves in each crown layer to whole-crown instantaneous carbon gain at midday. Carbon gain was estimated from the relationship between electron transport and photosynthetic rates. We hypothesized that more irradiance can penetrate into the middle of the crown if the upper crown layers have steep leaf inclination angles. We found that such a crown has a high whole-crown carbon gain, even if leaf traits do not change greatly with decreasing crown height. Leaf area indices (LAIs) of the two Fagus trees (5.26-5.52) were higher than the LAI of the B. grossa tree (4.50) and the leaves of the F. crenata tree were more concentrated in the top crown layers than were leaves of the other trees. Whole-crown carbon gain per unit ground area (micromol m(-2) ground s(-1)) at midday on fine days in summer was 16.3 for F. crenata, 11.0 for F. japonica, and 20.4 for B. grossa. In all study trees, leaf dry mass (LMA) and leaf nitrogen content (N) per unit area decreased with decreasing height in the crown, but leaf N per unit mass increased. Variations (plasticity) between the uppermost and lowermost crown layers in LMA, leaf N, the ratio of chlorophyll to N and the ratio of chlorophyll a to b were smaller for F. japonica and B. grossa than for F. crenata. The light extinction coefficients in the crowns were lower for the F. japonica and B. grossa trees than for the F. crenata tree. The leaf carbon isotope ratio (delta(13)C) was higher for F. japonica and B. grossa than for F. crenata, especially in the mid-crown. These results suggest that, in crowns with low leaf plasticity but steep leaf inclination angles, such as those of F. japonica and B. grossa trees, irradiance can penetrate into the middle of the crowns, thereby enhancing whole-crown carbon gain.     

Photosynthetic acclimation to light changes in tropical monsoon forest woody species differing in adult stature

We studied morphological and physiological leaf and whole-plant features of seedlings of six late-successional woody species common in the Xishuangbanna lowland rain forest in southwest China. Study species differed in adult stature and shade tolerance and included the shrubs Lasianthus attenuatus Jack and Lasianthus hookeri C.B. Clarke ex Hook. f.; the sub-canopy species Barringtonia macrostachya (Jack) Kurz and Linociera insignis C.B. Clarke; the canopy tree Pometia tomentosa (Blume) Teijsm. & Binn.; and the emergent species Shorea chinensis (Wang Hsie) H. Zhu. After 1 year of growth in low light (4.5% full sun), seedlings were transferred to high light (24.5% full sun) to investigate acclimation responses of existing leaves to forest gap opening and to determine whether seedling capacity for acclimation is a limiting factor in its natural regeneration. Leaves of the shrub species are shade-adapted, as indicated by their low photosynthetic capacity, efficiency in using sunflecks, low stomatal density, low Chl a/b ratio and high spongy/palisade mesophyll ratio. The shrub species utilized sunflecks efficiently because they had a short photosynthetic induction time and low induction loss. In all species, transfer of seedlings to high light resulted in a substantial initial reduction in the dark-adapted quantum yield of photosystem II (variable chlorophyll fluorescence/maximum chlorophyll fluorescence; Fv/Fm) at midday. Predawn Fv/Fm of the taller species did not change greatly, but predawn Fv/Fm of the shrub species decreased significantly without complete recovery within 25 days of transfer to high light, indicating chronic photoinhibition and damage to the previously shade-adapted leaves. Maximum net photosynthetic rate and dark respiration of the four taller species increased considerably after transfer to high light, but not in the shrub species. Similar trends were observed for the number of newly formed leaves and relative height growth rate. We conclude that the shrubs L. hookeri and L. attenuatus have limited potential for developmental and physiological acclimation to high light, which explains their absence from forest gaps. Compared with the shrub species, the taller tree species, which are more likely to experience high light during their life span, showed a greater potential for light acclimation. Physiological differences among the four tree species were not consistent with differences in adult stature.     

Leaf traits in relation to crown development, light interception and growth of elite families of loblolly and slash pine

Crown architecture and size influence leaf area distribution within tree crowns and have large effects on the light environment in forest canopies. The use of selected genotypes in combination with silvicultural treatments that optimize site conditions in forest plantations provide both a challenge and an opportunity to study the biological and environmental determinants of forest growth. We investigated tree growth, crown development and leaf traits of two elite families of loblolly pine (Pinus taeda L.) and one family of slash pine (P. elliottii Mill.) at canopy closure. Two contrasting silvicultural treatments -- repeated fertilization and control of competing vegetation (MI treatment), and a single fertilization and control of competing vegetation treatment (C treatment) -- were applied at two experimental sites in the West Gulf Coastal Plain in Texas and Louisiana. At a common tree size (diameter at breast height), loblolly pine trees had longer and wider crowns, and at the plot-level, intercepted a greater fraction of photosynthetic photon flux than slash pine trees. Leaf-level, light-saturated assimilation rates (A(max)) and both mass- and area-based leaf nitrogen (N) decreased, and specific leaf area (SLA) increased with increasing canopy depth. Leaf-trait gradients were steeper in crowns of loblolly pine trees than of slash pine trees for SLA and leaf N, but not for A(max). There were no species differences in A(max), except in mass-based photosynthesis in upper crowns, but the effect of silvicultural treatment on A(max) differed between sites. Across all crown positions, A(max) was correlated with leaf N, but the relationship differed between sites and treatments. Observed patterns of variation in leaf properties within crowns reflected acclimation to developing light gradients in stands with closing canopies. Tree growth was not directly related to A(max), but there was a strong correlation between tree growth and plot-level light interception in both species. Growth efficiency was unaffected by silvicultural treatment. Thus, when coupled with leaf area and light interception at the crown and canopy levels, A(max) provides insight into family and silvicultural effects on tree growth.     

Contrasting seasonal leaf habits of canopy trees between tropical dry-deciduous and evergreen forests in Thailand

We compared differences in leaf properties, leaf gas exchange and photochemical properties between drought-deciduous and evergreen trees in tropical dry forests, where soil nutrients differed but rainfall was similar. Three canopy trees (Shorea siamensis Miq., Xylia xylocarpa (Roxb.) W. Theob. and Vitex peduncularis Wall. ex Schauer) in a drought-deciduous forest and a canopy tree (Hopea ferrea Lanessan) in an evergreen forest were selected. Soil nutrient availability is lower in the evergreen forest than in the deciduous forest. Compared with the evergreen tree, the deciduous trees had shorter leaf life spans, lower leaf masses per area, higher leaf mass-based nitrogen (N) contents, higher leaf mass-based photosynthetic rates (mass-based P(n)), higher leaf N-based P(n), higher daily maximum stomatal conductance (g(s)) and wider conduits in wood xylem. Mass-based P(n) decreased from the wet to the dry season for all species. Following onset of the dry season, daily maximum g(s) and sensitivity of g(s) to leaf-to-air vapor pressure deficit remained relatively unchanged in the deciduous trees, whereas both properties decreased in the evergreen tree during the dry season. Photochemical capacity and non-photochemical quenching (NPQ) of photosystem II (PSII) also remained relatively unchanged in the deciduous trees even after the onset of the dry season. In contrast, photochemical capacity decreased and NPQ increased in the evergreen tree during the dry season, indicating that the leaves coped with prolonged drought by down-regulating PSII. Thus, the drought-avoidant deciduous species were characterized by high N allocation for leaf carbon assimilation, high water use and photoinhibition avoidance, whereas the drought-tolerant evergreen was characterized by low N allocation for leaf carbon assimilation, conservative water use and photoinhibition tolerance.     

Age-related effects on leaf area/sapwood area relationships,canopy transpiration and carbon gain of Norway spruce stands (Picea abies) in the Fichtelgebirge,Germany

Stand age is an important structural determinant of canopy transpiration (E(c)) and carbon gain. Another more functional parameter of forest structure is the leaf area/sapwood area relationship, A(L)/A(S), which changes with site conditions and has been used to estimate leaf area index of forest canopies. The interpretation of age-related changes in A(L)/A(S) and the question of how A(L)/A(S) is related to forest functions are of current interest because they may help to explain forest canopy fluxes and growth. We conducted studies in mature stands of Picea abies (L.) Karst. varying in age from 40 to 140 years, in tree density from 1680 to 320 trees ha(-1), and in tree height from 15 to 30 m. Structural parameters were measured by biomass harvests of individual trees and stand biometry. We estimated E(c) from scaled-up xylem sap flux of trees, and canopy-level fluxes were predicted by a three-dimensional microclimate and gas exchange model (STANDFLUX). In contrast to pine species, A(L)/A(S) of P. abies increased with stand age from 0.26 to 0.48 m(2) cm(-2). Agreement between E(c) derived from scaled-up sap flux and modeled canopy transpiration was obtained with the same parameterization of needle physiology independent of stand age. Reduced light interception per leaf area and, as a consequence, reductions in net canopy photosynthesis (A(c)), canopy conductance (g(c)) and E(c) were predicted by the model in the older stands. Seasonal water-use efficiency (WUE = A(c)/E(c)), derived from scaled-up sap flux and stem growth as well as from model simulation, declined with increasing A(L)/A(S) and stand age. Based on the different behavior of age-related A(L)/A(S) in Norway spruce stands compared with other tree species, we conclude that WUE rather than A(L)/A(S) could represent a common age-related property of all species. We also conclude that, in addition to hydraulic limitations reducing carbon gain in old stands, a functional change in A(L)/A(S) that is related to reduced light interception per leaf area provides another potential explanation for reduced carbon gain in old stands of P. abies, even when hydraulic constraints increase in response to changes in canopy architecture and aging.     

Leaf-trait variation explained by the hypothesis that plants maximize their canopy carbon export over the lifespan of leaves

Measured values of four key leaf traits (leaf area per unit mass, nitrogen concentration, photosynthetic capacity, leaf lifespan) co-vary consistently within and among diverse biomes, suggesting convergent evolution across species. The same leaf traits co-vary consistently with the environmental conditions (light intensity, carbon-dioxide concentration, nitrogen supply) prevailing during leaf development. No existing theory satisfactorily explains all of these trends. Here, using a simple model of the carbon-nitrogen economy of trees, we show that global leaf-trait relationships and leaf responses to environmental conditions can be explained by the optimization hypothesis (MAXX) that plants maximize the total amount of carbon exported from their canopies over the lifespan of leaves. Incorporating MAXX into larger-scale vegetation models may improve their consistency with global leaf-trait relationships, and enhance their ability to predict how global terrestrial productivity and carbon sequestration respond to environmental change.     

Ontogenetic changes in leaf phenology of Ulmus davidiana var. japonica, a deciduous broad-leaved tree

To determine how plants control leaf phenology to maximize annual carbon gain, I examined ontogenetic changes in leaf phenology of Japanese elm, Ulmus davidiana var. japonica Nakai plants of different ages growing in contrasting light environments. Leaf emergence occurred earlier in 1- and 2-year-old seedlings than in current-year seedlings. Although leaf emergence was not affected by light conditions at the sites, it was influenced by plant height. The delay in leaf emergence increased with increasing plant height. These traits indicate that seedlings that received the least light during the summer intercepted light for a long period during the spring; however, the advantage of earlier leaf emergence decreased with increasing plant height. At each site, 1-year-old seedlings had a longer duration of leaf emergence than adults, because of a longer period of favorable light conditions even in the forest understory. Duration of leaf emergence, leaf duration and leaf longevity were usually longer in sun than in shade for both seedlings and adults; however, flexibility in the response to light was greater in seedlings than in adults. The plastic response in leaf phenology during the juvenile stages may contribute to the optimization of light acquisition in habitats with differing light conditions, thereby enhancing seedling survival.     

Leaf gas exchange in the frankincense tree (Boswellia papyrifera) of African dry woodlands

A conceptual model was tested for explaining environmental and physiological effects on leaf gas exchange in the deciduous dry tropical woodland tree Boswellia papyrifera (Del.) Hochst. For this species we aimed at (i) understanding diurnal patterns in leaf gas exchange, (ii) exploring cause-effect relationships among external environment, internal physiology and leaf gas exchange, and (iii) exploring site differences in leaf gas exchange in response to environmental variables. Diurnal courses in gas exchange, underlying physiological traits and environmental variables were measured for 90 trees on consecutive days at two contrasting areas, one at high and the other at low altitude. Assimilation was highest in the morning and slightly decreased during the day. In contrast, transpiration increased from early morning to midday, mainly in response to an increasing vapor pressure deficit (VPD) and gradual stomatal closure. The leaf water potential varied relatively little and did not influence gas exchange during the measurement period. Our results suggest that the same cause-effect relationships function at contrasting areas. However, leaves at the higher altitude had higher photosynthetic capacity, reflecting acclimation to higher light levels. Trees at both areas nevertheless achieved similar leaf assimilation rates since assimilation was down-regulated by stomatal closure due to the higher VPD at the higher altitude, while it became more light limited at the lower altitude. Gas exchange was thus limited by a high VPD or low light levels during the wet season, despite the ability of the species to acclimate to different conditions.     

How should leaf area, sapwood area and stomatal conductance vary with tree height to maximize growth?

Conventional wisdom holds that the ratio of leaf area to sapwood area (L/S) should decline during height (H) growth to maintain hydraulic homeostasis and prevent stomatal conductance (g(s)) from declining. We contend that L/S should increase with H based on a numerical simulation, a mathematical analysis and a conceptual argument: (1) numerical simulation--a tree growth model, DESPOT (Deducing Emergent Structure and Physiology Of Trees), in which carbon (C) allocation is regulated to maximize C gain, predicts L/S should increase during most of H growth; (2) mathematical analysis--the formal criterion for optimal C allocation, applied to a simplified analytical model of whole tree carbon-water balance, predicts L/S should increase with H if leaf-level gas exchange parameters including g(s) are conserved; and (3) conceptual argument--photosynthesis is limited by several substitutable resources (chiefly nitrogen (N), water and light) and H growth increases the C cost of water transport but not necessarily of N and light capture, so if the goal is to maximize C gain or growth, allocation should shift in favor of increasing photosynthetic capacity and irradiance, rather than sustaining g(s). Although many data are consistent with the prediction that L/S should decline with H, many others are not, and we discuss possible reasons for these discrepancies.     

Sunflecks in trees and forests: from photosynthetic physiology to global change biology

Sunflecks are brief, intermittent periods of high photon flux density (PFD) that can significantly improve carbon gain in shaded forest understories and lower canopies of trees. In this review, we discuss the physiological basis of leaf-level responses to sunflecks and the mechanisms plants use to tolerate sudden changes in PFD and leaf temperature induced by sunflecks. We also examine the potential effects of climate change stresses (including elevated temperatures, rising CO(2) concentrations and drought) on the ability of tree species to use sunflecks, and advocate more research to improve our predictions of seedling and tree carbon gain in future climates. Lastly, while we have the ability to model realistic responses of photosynthesis to fluctuating PFD, dynamic responses of photosynthesis to sunflecks are not accounted for in current models of canopy carbon uptake, which can lead to substantial overestimates of forest carbon fixation. Since sunflecks are a critical component of seasonal carbon gain for shaded leaves, sunfleck regimes and physiological responses to sunflecks should be incorporated into models to more accurately capture forest carbon dynamics.     

Photosynthetic light acclimation in peach leaves: importance of changes in mass:area ratio, nitrogen concentration, and leaf nitrogen partitioning

Photosynthetic light acclimation of leaves can result from (i) changes in mass-based leaf nitrogen concentration, Nm, (ii) changes in leaf mass:area ratio, Ma, and (iii) partitioning of total leaf nitrogen among different pools of the photosynthetic machinery. We studied variations in Nm and Ma within the crowns of two peach (Prunus persica L. Batsch) trees grown in an orchard in Portugal, and one peach tree grown in an orchard in France. Each crown was digitized and a 3-D radiation transfer model was used to quantify the intra-crown variations in time-integrated leaf irradiance, . Nitrogen concentration, leaf mass:area ratio, chlorophyll concentration, and photosynthetic capacity were also measured on leaves sampled on five additional peach trees in the orchard in Portugal. The data were used to compute the coefficients of leaf nitrogen partitioning among carboxylation, bioenergetics, and light harvesting pools. Leaf mass:area ratio and area-based leaf nitrogen concentration, Na, were nonlinearly related to , and photosynthetic capacity was linearly related to Na. Photosynthetic light acclimation resulted mainly from changes in Ma and leaf nitrogen partitioning, and to a lesser extent from changes in Nm. This behavior contrasts with photosynthetic light acclimation observed in other tree species like walnut (Juglans regia L.) in which acclimation results primarily from changes in Ma.     

Tree size and light availability increase photochemical instead of non-photochemical capacities of Nothofagus nitida trees growing in an evergreen temperate rain forest

Nothofagus nitida (Phil.) Krasser (Nothofagaceae) regenerates under the canopy in microsites protected from high light. Nonetheless, it is common to find older saplings in clear areas and adults as emergent trees of the Chilean evergreen forest. We hypothesized that this shade to sun transition in N. nitida is supported by an increase in photochemical and non-photochemical energy dissipation capacities of both photosystems in parallel with the increase in plant size and light availability. To dissect the relative contribution of light environment and plant developmental stage to these physiological responses, the photosynthetic performance of both photosystems was studied from the morpho-anatomical to the biochemical level in current-year leaves of N. nitida plants of different heights (ranging from 0.1 to 7 m) growing under contrasting light environments (integrated quantum flux (IQF) 5-40 mol m(-2). Tree height (TH) and light environment (IQF) independently increased the saturated electron transport rates of both photosystems, as well as leaf and palisade thickness, but non-photochemical energy flux, photoinhibition susceptibility, state transition capacity, and the contents of D1 and PsbS proteins were not affected by IQF and TH. Spongy mesophyll thickness and palisade cell diameter decreased with IQF and TH. A(max), light compensation and saturation points, Rubisco and nitrogen content (area basis) only increased with light environment (IQF), whereas dark respiration (R(d)) decreased slightly and relative chlorophyll content was higher in taller trees. Overall, the independent effects of more illuminated environment and tree height mainly increased the photochemical instead of the non-photochemical energy flux. Regardless of the photochemical increase with TH, carbon assimilation only significantly improved with higher IQF. Therefore it seems that mainly acclimation to the light environment supports the phenotypic transition of N. nitida from shade to sun.     

A 10-year evaluation of the functional basis for regeneration habitat preference of trees in an African evergreen forest

The spatial distribution of tree juveniles in relation to light environments may reflect species differences in growth, survival, and functional traits and will shape the nature of forest regeneration. Long-term field experiments are important to evaluate this issue because of the potentially very long juvenile period in trees. Here, we combine a 10-year seedling survival–growth data with the results of community ordination and multivariate analyses of functional traits to ask how observed juvenile light guilds are related to species functional traits and seedling performance. We transplanted seedlings at a standardized height of 11 cm into the shaded understory and quantified their growth and survival for 10-years. Using the community-wide stem distribution data, we categorized 33 species including the focal 11 species to understory vs. gap/edge guilds. Then, we determined differences between the two guilds in seedling survival, growth, as well as seed size, adult height, and a series of leaf traits, including toughness and chemical traits (fiber, protein, phenolics, tannins, alkaloids, saponins). Among the 11 non-pioneer species whose seedlings were planted into the understory, there was no significant difference in 10-year survival between light guilds, but species in gap/edge guild tended to achieve greater height than species in the understory guild. The leaf chemical traits of 33 species did not differ between the two juvenile light guilds, but gap/edge species had smaller seeds, taller adults, and tougher leaves than understory species. We used logistic regression as a complementary approach to assess the extent to which plant traits varied between light guilds and the most parsimonious model based on AIC_c ranking included only leaf toughness and had an Akaike weight of 0.52. In addition, across the 11 species planted as seedlings, these traits were not significantly related to survivorship or growth over 10 years. A Principle Components Analysis illustrated associations among traits. We conclude that light guilds in terms of juvenile stem distribution could not be explained by long-term field performance of post-establishment seedlings alone. Earlier seedling stage or later sapling stage may be more important in differentiation of light guilds. For the species examined difference in growth rates could be linked to seed size and adult stature, but not to the adult leaf chemical traits considered. These results suggest the importance of examining ontogenetic shifts and relationships among functional traits for a better understanding of regeneration strategies of tropical trees.     

The successional status of tropical rainforest tree species is associated with differences in leaf carbon isotope discrimination and functional traits

We characterised the among species variability in leaf gas exchange and morphological traits under controlled conditions of seedlings of 22 tropical rainforest canopy species to understand the origin of the variability in leaf carbon isotope discrimination (Δ) among species with different growth and dynamic characteristics (successional gradient). Our results first suggest that these species pursue a consistent strategy in terms of Δ throughout their ontogeny (juveniles grown here versus canopy adult trees from the natural forest). Second, leaf Δ was negatively correlated with WUE and N, and positively correlated with g_s, but among species differences in Δ were mainly explained by differences in WUE. Finally, species belonging to different successional groups display distinct leaf functional and morphological traits. We confirmed that fast growing early successional species maximise carbon assimilation with high stomatal conductance. In contrast, fast and slow growing late successional species are both characterised by low carbon assimilation values, but by distinct stomatal conductance and leaf morphological features. Along the successional gradient, these differences result in much lower Δ for the intermediate species (i.e. fast growing late successional) as compared to the two other groups.     

To what extent is altitudinal variation of functional traits driven by genetic adaptation in European oak and beech?

The phenotypic responses of functional traits in natural populations are driven by genetic diversity and phenotypic plasticity. These two mechanisms enable trees to cope with rapid climate change. We studied two European temperate tree species (sessile oak and European beech), focusing on (i) in situ variations of leaf functional traits (morphological and physiological) along two altitudinal gradients and (ii) the extent to which these variations were under environmental and/or genetic control using a common garden experiment. For all traits, altitudinal trends tended to be highly consistent between species and transects. For both species, leaf mass per area displayed a positive linear correlation with altitude, whereas leaf size was negatively correlated with altitude. We also observed a significant increase in leaf physiological performance with increasing altitude: populations at high altitudes had higher maximum rates of assimilation, stomatal conductance and leaf nitrogen content than those at low altitudes. In the common garden experiment, genetic differentiation between populations accounted for 0-28% of total phenotypic variation. However, only two traits (leaf mass per area and nitrogen content) exhibited a significant cline. The combination of in situ and common garden experiments used here made it possible to demonstrate, for both species, a weaker effect of genetic variation than of variations in natural conditions, suggesting a strong effect of the environment on leaf functional traits. Finally, we demonstrated that intrapopulation variability was systematically higher than interpopulation variability, whatever the functional trait considered, indicating a high potential capacity to adapt to climate change.     

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.     

Changes in shoot allometry with increasing tree height in a tropical canopy species,Elateriospermum tapos

Allometry of shoot extension units (hereafter termed "current shoots") was analyzed in a Malaysian canopy species, Elateriospermum tapos Bl. (Euphorbiaceae). Changes in current shoot allometry with increasing tree height were related to growth and maintenance of tree crowns. Total biomass, biomass allocation ratio of non-photosynthetic to photosynthetic organs, and wood density of current shoots were unrelated to tree height. However, shoot structure changed with tree height. Compared with short trees, tall trees produced current shoots of the same mass but with thicker and shorter stems. Current shoots with thin and long stems enhanced height growth in short trees, whereas in tall trees, thick and short current shoots may reduce mechanical and hydraulic stresses. Furthermore, compared with short trees, tall trees produced current shoots with more leaves of lower dry mass, smaller area, and smaller specific leaf area (SLA). Short trees adapted to low light flux density by reducing mutual shading with large leaves having a large SLA. In contrast, tall trees reduced mutual shading within a shoot by producing more small leaves in distal than in proximal parts of the shoot stem. The production of a large number of small leaves promoted light penetration into the dense crowns of tall trees. All of these characteristics suggest that the change in current shoot structure with increasing tree height is adaptive in E. tapos, enabling short trees to maximize height growth and tall trees to maximize light capture.     

Responses of foliar photosynthetic electron transport, pigment stoichiometry, and stomatal conductance to interacting environmental factors in a mixed species forest canopy

We studied limitations caused by variations in leaf temperature and soil water availability on photosynthetic electron transport rates calculated from foliar chlorophyll fluorescence analysis (U) in a natural deciduous forest canopy composed of shade-intolerant Populus tremula L. and shade-tolerant Tilia cordata Mill. In both species, there was a positive linear relationship between light-saturated U (Umax) per unit leaf area and mean seasonal integrated daily quantum flux density (Ss, mol per square m per day). Acclimation of leaf dry mass per area and nitrogen per area to growth irradiance largely accounted for this positive scaling. However, the slopes of the Umax versus Ss relationships were greater on days when leaf temperature was high than on days when leaf temperature was low. Overall, Umax varied 2.5-fold across a temperature range of 20-30 degrees C. Maximum stomatal conductance (Gmax) also scaled positively with Ss. Although Gmax observed during daily time courses, and stomatal conductances during Umax measurements declined in response to seasonally decreasing soil water contents, was insensitive to prolonged water stress, and was not strongly correlated with stomatal conductances during its estimation. These results suggest that photorespiration was an important electron sink when intercellular CO2 concentration was low because of closed stomata. Given that xanthophyll cycle pool size (VAZ, sum of violaxanthin, antheraxanthin, and zeaxanthin) may play an important role in dissipation of excess excitation energy, the response of VAZ to fluctuating light and temperature provided another possible explanation for the stable Umax. Xanthophyll cycle carotenoids per total leaf chlorophyll (VAZ/Chl) scaled positively with integrated light and negatively with daily minimum air temperature, whereas the correlation between VAZ/Chl and irradiance was best with integrated light averaged over 3 days preceding foliar sampling. We conclude that the potential capacity for electron transport is determined by long-term acclimation of U to certain canopy light conditions, and that the rapid adjustment of the capacity for excitation energy dissipation plays a significant part in the stabilization of this potential capacity. Sustained high capacity of photosynthetic electron transport during stress periods provides an explanation for the instantaneous response of U to short-term weather fluctuations, but also indicates that U restricts potential carbon gain under conditions of water limitation less than does stomatal conductance.     

Why does leaf nitrogen decline within tree canopies less rapidly than light? An explanation from optimization subject to a lower bound on leaf mass per area

A long-established theoretical result states that, for a given total canopy nitrogen (N) content, canopy photosynthesis is maximized when the within-canopy gradient in leaf N per unit area (N(a)) is equal to the light gradient. However, it is widely observed that N(a) declines less rapidly than light in real plant canopies. Here we show that this general observation can be explained by optimal leaf acclimation to light subject to a lower-bound constraint on the leaf mass per area (m(a)). Using a simple model of the carbon-nitrogen (C-N) balance of trees with a steady-state canopy, we implement this constraint within the framework of the MAXX optimization hypothesis that maximizes net canopy C export. Virtually all canopy traits predicted by MAXX (leaf N gradient, leaf N concentration, leaf photosynthetic capacity, canopy N content, leaf-area index) are in close agreement with the values observed in a mature stand of Norway spruce trees (Picea abies L. Karst.). An alternative upper-bound constraint on leaf photosynthetic capacity (A(sat)) does not reproduce the canopy traits of this stand. MAXX subject to a lower bound on m(a) is also qualitatively consistent with co-variations in leaf N gradient, m(a) and A(sat) observed across a range of temperate and tropical tree species. Our study highlights the key role of constraints in optimization models of plant function.