RESUMO
Most leaf functional trait studies in the Amazon basin do not consider ontogenetic variations (leaf age), which may influence ecosystem productivity throughout the year. When leaf age is taken into account, it is generally considered discontinuous, and leaves are classified into age categories based on qualitative observations. Here, we quantified age-dependent changes in leaf functional traits such as the maximum carboxylation rate of ribulose-1,5-biphosphate carboxylase/oxygenase (Rubisco) (Vcmax), stomatal control (Cgs%), leaf dry mass per area and leaf macronutrient concentrations for nine naturally growing Amazon tropical trees with variable phenological strategies. Leaf ages were assessed by monthly censuses of branch-level leaf demography; we also performed leaf trait measurements accounting for leaf chronological age based on days elapsed since the first inclusion in the leaf demography, not predetermined age classes. At the tree community scale, a nonlinear relationship between Vcmax and leaf age existed: young, developing leaves showed the lowest mean photosynthetic capacity, increasing to a maximum at 45 days and then decreasing gradually with age in both continuous and categorical age group analyses. Maturation times among species and phenological habits differed substantially, from 8 ± 30 to 238 ± 30 days, and the rate of decline of Vcmax varied from -0.003 to -0.065 µmol CO2 m-2 s-1 day-1. Stomatal control increased significantly in young leaves but remained constant after peaking. Mass-based phosphorus and potassium concentrations displayed negative relationships with leaf age, whereas nitrogen did not vary temporally. Differences in life strategies, leaf nutrient concentrations and phenological types, not the leaf age effect alone, may thus be important factors for understanding observed photosynthesis seasonality in Amazonian forests. Furthermore, assigning leaf age categories in diverse tree communities may not be the recommended method for studying carbon uptake seasonality in the Amazon, since the relationship between Vcmax and leaf age could not be confirmed for all trees.
Assuntos
Ecossistema , Árvores , Dióxido de Carbono , Fotossíntese , Folhas de PlantaRESUMO
Photosynthetic carbon uptake by tropical forests is of critical importance in regulating the earth's climate, but rising temperatures threaten this stabilizing influence of tropical forests. Most research on how temperature affects photosynthesis focuses on fully sun-exposed leaves, and little is known about shade leaves, even though shade leaves greatly outnumber sun leaves in lowland tropical forests. We measured temperature responses of light-saturated photosynthesis, stomatal conductance, and the biochemical parameters VCMax (maximum rate of RuBP carboxylation) and JMax (maximum rate of RuBP regeneration, or electron transport) on sun and shade leaves of mature tropical trees of three species in Panama. As expected, biochemical capacities and stomatal conductance were much lower in shade than in sun leaves, leading to lower net photosynthesis rates. However, the key temperature response traits of these parameters-the optimum temperature (TOpt) and the activation energy-did not differ systematically between sun and shade leaves. Consistency in the JMax to VCMax ratio further suggested that shade leaves are not acclimated to lower temperatures. For both sun and shade leaves, stomatal conductance had the lowest temperature optimum (~25 °C), followed by net photosynthesis (~30 °C), JMax (~34 °C) and VCMax (~38 °C). Stomatal conductance of sun leaves decreased more strongly with increasing vapor pressure deficit than that of shade leaves. Consistent with this, modeled stomatal limitation of photosynthesis increased with increasing temperature in sun but not shade leaves. Collectively, these results suggest that modeling photosynthetic carbon uptake in multi-layered canopies does not require independent parameterization of the temperature responses of the biochemical controls over photosynthesis of sun and shade leaves. Nonetheless, to improve the representation of the shade fraction of carbon uptake dynamics in tropical forests, better understanding of stomatal sensitivity of shade leaves to temperature and vapor pressure deficit will be required.
Assuntos
Fotossíntese , Árvores , Dióxido de Carbono , Panamá , Folhas de Planta , TemperaturaRESUMO
Leaf canopy carbon exchange processes, such as photosynthesis and respiration, are substantial components of the global carbon cycle. Climate models base their simulations of photosynthesis and respiration on an empirical understanding of the underlying biochemical processes, and the responses of those processes to environmental drivers. As such, data spanning large spatial scales are needed to evaluate and parameterize these models. Here, we present data on four important biochemical parameters defining leaf carbon exchange processes from 626 individuals of 98 species at 12 North and Central American sites spanning ~53° of latitude. The four parameters are the maximum rate of Rubisco carboxylation (Vcmax ), the maximum rate of electron transport for the regeneration of Ribulose-1,5,-bisphosphate (Jmax ), the maximum rate of phosphoenolpyruvate carboxylase carboxylation (Vpmax ), and leaf dark respiration (Rd ). The raw net photosynthesis by intercellular CO2 (A/Ci ) data used to calculate Vcmax , Jmax , and Vpmax rates are also presented. Data were gathered on the same leaf of each individual (one leaf per individual), allowing for the examination of each parameter relative to others. Additionally, the data set contains a number of covariates for the plants measured. Covariate data include (1) leaf-level traits (leaf mass, leaf area, leaf nitrogen and carbon content, predawn leaf water potential), (2) plant-level traits (plant height for herbaceous individuals and diameter at breast height for trees), (3) soil moisture at the time of measurement, (4) air temperature from nearby weather stations for the day of measurement and each of the 90 d prior to measurement, and (5) climate data (growing season mean temperature, precipitation, photosynthetically active radiation, vapor pressure deficit, and aridity index). We hope that the data will be useful for obtaining greater understanding of the abiotic and biotic determinants of these important biochemical parameters and for evaluating and improving large-scale models of leaf carbon exchange.
Assuntos
Carbono/metabolismo , Folhas de Planta/metabolismo , Dióxido de Carbono , América Central , Fotossíntese , Ribulose-Bifosfato Carboxilase , ÁrvoresRESUMO
The inactivation of the chloroplast ascorbate peroxidases (chlAPXs) has been thought to limit the efficiency of the water-water cycle and photo-oxidative protection under stress conditions. In this study, we have generated double knockdown rice (Oryza sativa L.) plants in both OsAPX7 (sAPX) and OsAPX8 (tAPX) genes, which encode chloroplastic APXs (chlAPXs). By employing an integrated approach involving gene expression, proteomics, biochemical and physiological analyses of photosynthesis, we have assessed the role of chlAPXs in the regulation of the protection of the photosystem II (PSII) activity and CO2 assimilation in rice plants exposed to high light (HL) and methyl violagen (MV). The chlAPX knockdown plants were affected more severely than the non-transformed (NT) plants in the activity and structure of PSII and CO2 assimilation in the presence of MV. Although MV induced significant increases in pigment content in the knockdown plants, the increases were apparently not sufficient for protection. Treatment with HL also caused generalized damage in PSII in both types of plants. The knockdown and NT plants exhibited differences in photosynthetic parameters related to efficiency of utilization of light and CO2. The knockdown plants overexpressed other antioxidant enzymes in response to the stresses and increased the GPX activity in the chloroplast-enriched fraction. Our data suggest that a partial deficiency of chlAPX expression modulate the PSII activity and integrity, reflecting the overall photosynthesis when rice plants are subjected to acute oxidative stress. However, under normal growth conditions, the knockdown plants exhibit normal phenotype, biochemical and physiological performance.