Photoperiodic regulation of the seasonal pattern of photosynthetic capacity and the implications for carbon cycling.

Although temperature is an important driver of seasonal changes in photosynthetic physiology, photoperiod also regulates leaf activity. Climate change will extend growing seasons if temperature cues predominate, but photoperiod-controlled species will show limited responsiveness to warming. We show that photoperiod explains more seasonal variation in photosynthetic activity across 23 tree species than temperature. Although leaves remain green, photosynthetic capacity peaks just after summer solstice and declines with decreasing photoperiod, before air temperatures peak. In support of these findings, saplings grown at constant temperature but exposed to an extended photoperiod maintained high photosynthetic capacity, but photosynthetic activity declined in saplings experiencing a naturally shortening photoperiod; leaves remained equally green in both treatments. Incorporating a photoperiodic correction of photosynthetic physiology into a global-scale terrestrial carbon-cycle model significantly improves predictions of seasonal atmospheric CO(2) cycling, demonstrating the benefit of such a function in coupled climate system models. Accounting for photoperiod-induced seasonality in photosynthetic parameters reduces modeled global gross primary production 2.5% (∼4 PgC y(-1)), resulting in a >3% (∼2 PgC y(-1)) decrease of net primary production. Such a correction is also needed in models estimating current carbon uptake based on remotely sensed greenness. Photoperiod-associated declines in photosynthetic capacity could limit autumn carbon gain in forests, even if warming delays leaf senescence.

Separating foliar physiology from morphology reveals the relative roles of vertically structured transpiration factors within red maple crowns and limitations of larger scale models.

A spatially explicit mechanistic model, MAESTRA, was used to separate key parameters affecting transpiration to provide insights into the most influential parameters for accurate predictions of within-crown and within-canopy transpiration. Once validated among Acer rubrum L. genotypes, model responses to different parameterization scenarios were scaled up to stand transpiration (expressed per unit leaf area) to assess how transpiration might be affected by the spatial distribution of foliage properties. For example, when physiological differences were accounted for, differences in leaf width among A. rubrum L. genotypes resulted in a 25% difference in transpiration. An in silico within-canopy sensitivity analysis was conducted over the range of genotype parameter variation observed and under different climate forcing conditions. The analysis revealed that seven of 16 leaf traits had a ≥5% impact on transpiration predictions. Under sparse foliage conditions, comparisons of the present findings with previous studies were in agreement that parameters such as the maximum Rubisco-limited rate of photosynthesis can explain ∼20% of the variability in predicted transpiration. However, the spatial analysis shows how such parameters can decrease or change in importance below the uppermost canopy layer. Alternatively, model sensitivity to leaf width and minimum stomatal conductance was continuous along a vertical canopy depth profile. Foremost, transpiration sensitivity to an observed range of morphological and physiological parameters is examined and the spatial sensitivity of transpiration model predictions to vertical variations in microclimate and foliage density is identified to reduce the uncertainty of current transpiration predictions.

Exploring the importance of within-canopy spatial temperature variation on transpiration predictions.

Models seldom consider the effect of leaf-level biochemical acclimation to temperature when scaling forest water use. Therefore, the dependence of transpiration on temperature acclimation was investigated at the within-crown scale in climatically contrasting genotypes of Acer rubrum L., cv. October Glory (OG) and Summer Red (SR). The effects of temperature acclimation on intracanopy gradients in transpiration over a range of realistic forest growth temperatures were also assessed by simulation. Physiological parameters were applied, with or without adjustment for temperature acclimation, to account for transpiration responses to growth temperature. Both types of parameterization were scaled up to stand transpiration (expressed per unit leaf area) with an individual tree model (MAESTRA) to assess how transpiration might be affected by spatial and temporal distributions of foliage properties. The MAESTRA model performed well, but its reproducibility was dependent on physiological parameters acclimated to daytime temperature. Concordance correlation coefficients between measured and predicted transpiration were higher (0.95 and 0.98 versus 0.87 and 0.96) when model parameters reflected acclimated growth temperature. In response to temperature increases, the southern genotype (SR) transpiration responded more than the northern (OG). Conditions of elevated long-term temperature acclimation further separate their transpiration differences. Results demonstrate the importance of accounting for leaf-level physiological adjustments that are sensitive to microclimate changes and the use of provenance-, ecotype-, and/or genotype-specific parameter sets, two components likely to improve the accuracy of site-level and ecosystem-level estimates of transpiration flux.

Measuring and modeling the variation in species-specific transpiration in temperate deciduous hardwoods.

We investigated which parameters required by the MAESTRA model were most important in predicting leaf-area-based transpiration in 5-year-old trees of five deciduous hardwood species-yoshino cherry (Prunus x yedoensis Matsum.), red maple (Acer rubrum L. 'Autumn Flame'), trident maple (Acer buergeranum Miq.), Japanese flowering cherry (Prunus serrulata Lindl. 'Kwanzan') and London plane-tree (Platanus x acerifolia (Ait.) Willd.). Transpiration estimated from sap flow measured by the heat balance method in branches and trunks was compared with estimates predicted by the three-dimensional transpiration, photosynthesis and absorbed radiation model, MAESTRA. MAESTRA predicted species-specific transpiration from the interactions of leaf-level physiology and spatially explicit micro-scale weather patterns in a mixed deciduous hardwood plantation on a 15-min time step. The monthly differences between modeled mean daily transpiration estimates and measured mean daily sap flow ranged from a 35% underestimation for Acer buergeranum in June to a 25% overestimation for A. rubrum in July. The sensitivity of the modeled transpiration estimates was examined across a 30% error range for seven physiological input parameters. The minimum value of stomatal conductance as incident solar radiation tends to zero was determined to be eight times more influential than all other physiological model input parameters. This work quantified the major factors that influence modeled species-specific transpiration and confirmed the ability to scale leaf-level physiological attributes to whole-crown transpiration on a species-specific basis.

The influence of temperature on within-canopy acclimation and variation in leaf photosynthesis: spatial acclimation to microclimate gradients among climatically divergent Acer rubrum L. genotypes.

Leaf gas exchange and temperature response were measured to assess temperature acclimation within a tree canopy in climatically contrasting genotypes of Acer rubrum L. Over the course of two 50 d continuous periods, growth temperature was controlled within tree crowns and the steady-state rate of leaf gas exchange was measured. Data were then modelled to calculate the influence of genotype variation and vertical distribution of physiological activity on carbon uptake. The maximal rate of Rubisco carboxylation (V(cmax)), the maximum rate of electron transport (J(max)), leaf dark respiration rate (R(d)), maximum photosynthesis (A(max)), and the CO(2) compensation point (Gamma) increased with temperature during both (i) a constant long-term (50 d) daytime temperature or (ii) ambient daytime temperature with short-term temperature control (25-38 degrees C). In addition, within-crown variation in the temperature response of photosynthesis and R(d) was influenced by acclimation to local microclimate temperature gradients. Results indicated that carbon uptake estimates could be overestimated by 22-25% if the vertical distribution of temperature gradients is disregarded. Temperature is a major factor driving photosynthetic acclimation and within-crown gas exchange variation. Thus, this study established the importance of including spatial acclimation to temperature- and provenance-, ecotype-, and/or genotype-specific parameter sets into carbon uptake models.

Modeling intra-crown and intra-canopy interactions in red maple: assessment of light transfer on carbon dioxide and water vapor exchange.

Daily and seasonal net photosynthesis (Anet), transpiration (E), absorbed photosynthetically active radiation (Qa) and light-use efficiency (epsilonc) in a red maple container nursery were simulated with MAESTRA, a three-dimensional canopy model. Effects of canopy heterogeneity were simulated by imposing changes in crown spacing. The light transfer sub-model, a distribution model of incident, direct, diffuse and scattered radiation within MAESTRA, was validated against field measurements of light interception on an intra-crown scale. In the container nursery, we found that a fiber-optic-based method of integrating photosynthetically active radiation (Q) was more suitable for crown-layer light transfer measurements and adjustments than either orthogonal line or individual quantum sensor measurements. The model underestimated intercepted Q by 9.3, 18 and 11.1% for crown layers 1, 2 and 3, respectively; however, there were linear relationships between model estimates and observations made with each of the three measurement methods. We used the validated and parameterized light transfer model to assess intra-crown and intra-canopy light transfer on a layer, crown and canopy basis, and investigated effects of tree size ratio and tree spacing interactions on Anet, E, Qa and epsilonc in the container nursery. Heterogeneous crown and canopy photosynthesis were predicted to exceed values for a uniform canopy under space-limiting conditions. Tree size ratio had large effects on Anet, E, Qa and epsilonc when light to lower-canopy layers was limited by inadequate space between crowns. Increasing Qa at lower-crown layers had the largest impact on whole-crown and whole-canopy Anet, E, Qa and epsilonc. Increases in canopy productivity led to increased water use. Simulations of heterogeneous stands with adequate soil water indicated that light absorption is maximized under space-limiting conditions as a canopy crown moves toward heterogeneity. Nursery and plantation productivity per unit land area was optimized by tactical placement of trees of several sizes, but this was accompanied by increased canopy water use.