A model is presented which solves simultaneously for leafâscale stomatal conductance, CO2 assimilation and the energy balance as a function of leaf position within canopies of wellâwatered vegetation. Fluxes and conductances were calculated separately for sunlit and shaded leaves. A linear dependence of photosynthetic capacity on leaf nitrogen content was assumed, while leaf nitrogen content and light intensity were assumed to decrease exponentially within canopies. Separate extinction coefficients were used for diffuse and direct beam radiation. An efficient Gaussian integration technique was used to compute fluxes and mean conductances for the canopy. The multilayer model synthesizes current knowledge of radiation penetration, leaf physiology and the physics of evaporation and provides insights into the response of whole canopies to multiple, interacting factors. The model was also used to explore sources of variation in the slopes of two simple parametric models (nitrogenâ and lightâuse efficiency), and to set bounds on the magnitudes of the parameters. For canopies low in total N, daily assimilation rates are â¼10% lower when leaf N is distributed uniformly than when the same total N is distributed according to the exponentially decreasing profile of absorbed radiation. However, gains are negligible for plants with high N concentrations. Canopy conductance, Gc should be calculated as Gc=AI(fslgsl+fshgsh), where I is leaf area index, fsi and fsh are the fractions of sunlit and shaded leaves at each level, and gsi and gsh are the corresponding stomatal conductances. Simple addition of conductances without this weighting causes errors in transpiration calculated using the âbigâleafâ version of the PenmanâMonteith equation. Partitioning of available energy between sensible and latent heat is very responsive to the parameter describing the sensitivity of stomata to the atmospheric humidity deficit. This parameter also affects canopy conductance, but has a relatively small impact on canopy assimilation. Simple parametric models are useful for extrapolating understanding from small to large scales, but the complexity of real ecosystems is thus subsumed in unexplained variations in parameter values. Simulations with the multilayer model show that both nitrogenâ and radiationâuse efficiencies depend on plant nutritional status and the diffuse component of incident radiation, causing a 2â to 3âfold variation in these efficiencies.

Leaf nitrogen, photosynthesis, conductance and transpiration : scaling from leaves to canopies

A process-based leaf gas exchange model for C3 plants was developed which specifically describes the effects observed along light gradients of shifting nitrogen investment in carboxylation and bioenergetics and modified leaf thickness due to altered stacking of photosynthetic units. The model was parametrized for the late-successional, shadetolerant deciduous species Acer saccharum Marsh. The specific activity of ribulose-1,5-bisphosphate carboxylase (Rubisco) and the maximum photosynthetic electron transport rate per unit cytochrome f (cyt f) were used as indices that vary proportionally with nitrogen investment in the capacities for carboxylation and electron transport. Rubisco and cyt f per unit leaf area are related in the model to leaf dry mass per area (MA), leaf nitrogen content per unit leaf dry mass (Nm), and partitioning coefficients for leaf nitrogen in Rubisco (PR) and in bioenergetics (PB). These partitioning coefficients are estimated from characteristic response curves of photosynthesis along with information on leaf structure and composition. While PR and PB determine the light-saturated value of photosynthesis, the fraction of leaf nitrogen in thylakoid light-harvesting components (PL) and the ratio of leaf chlorophyll to leaf nitrogen invested in light harvesting ( CB), which is dependent on thylakoid stoichiometry, determine the initial photosynthetic light utilization efficiency in the model. Carbon loss due to mitochondrial respiration, which also changes along light gradients, was considered to vary in proportion with carboxylation capacity. Key model parameters ‐ Nm, PR, PB, PL, CB and stomatal sensitivity with respect to changes in net photosynthesis ( Gf) ‐ were examined as a function of MA, which is linearly related to irradiance during growth of the leaves. The results of the analysis applied to A. saccharum indicate that PB and PR increase, and Gf, PL and CB decrease with increasing MA. As a result of these effects of irradiance on nitrogen partitioning, the slope of the light-saturated net photosynthesis rate per unit leaf dry mass (A m max ) versus Nm relationship

A model separating leaf structural and physiological effects on carbon gain along light gradients for the shade‐tolerant species Acer saccharum

PnET is a simple, lumped-parameter, monthlytime-step model of carbon and water balances of forests built on two principal relationships: 1) maximum photosynthetic rate is a function of foliar nitrogen concentration, and 2) stomatal conductance is a function of realized photosynthetic rate. Monthyly leaf area display and carbon and water balances are predicted by combining these with standard equations describing light attenuation in canopies and photosynthetic response to diminishing radiation intensity, along with effects of soil water stress and vapor pressure deficit (VPD). PnET has been validated against field data from 10 well-studied temperate and boreal forest ecosystems, supporting our central hypothesis that aggregation of climatic data to the monthly scale and biological data such as foliar characteristics to the ecosystem level does not cause a significant loss of information relative to long-term, mean ecosystem responses. Sensitivity analyses reveal a diversity of responses among systems to identical alterations in climatic drivers. This suggests that great care should be used in developing generalizations as to how forests will respond to a changing climate. Also critical is the degree to which the temperature responses of photosynthesis and respiration might acclimate to changes in mean temperatures at decadal time scales. An extreme climate change simulation (+3° C maximum temperature, −25% precipitation with no change in minimum temperature or radiation, direct effects of increased atmospheric CO2 ignored) suggests that major increases in water stress, and reductions in biomass production (net carbon gain) and water yield would follow such a change.

A generalized, lumped-parameter model of photosynthesis, evapotranspiration and net primary production in temperate and boreal forest ecosystems.

Stomata play a key role in plant adaptation to changing environmental conditions as they control both water losses and CO(2) uptake. Particularly, in the context of global change, simulations of the consequences of drought on crop plants are needed to design more efficient and water-saving cropping systems. However, most of the models of stomatal conductance (g(s)) developed at the leaf level link g(s) to environmental factors or net photosynthesis (A(net)), but do not include satisfactorily the effects of drought, impairing our capacity to simulate plant functioning in conditions of limited water supply. The objective of this review was to draw an up-to-date picture of the g(s) models, from the empirical to the process-based ones, along with their mechanistic or deterministic bases. It focuses on models capable to account for multiple environmental influences with emphasis on drought conditions. We examine how models that have been proposed for well-watered conditions can be combined with those specifically designed to deal with drought conditions. Ideas for future improvements of g(s) models are discussed: the issue of co-regulation of g(s) and A(net); the roles of CO(2), absissic acid and H(2)O(2); and finally, how to better address the new challenges arising from the issue of global change.

https://www.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1365-3040.2010.02181.x

An overview of models of stomatal conductance at the leaf level

A model of crop primary production, which was originally developed to relate the amount of absorbed photosynthetically active radiation (APAR) to net production in field studies, is discussed in the context of coarse resolution regional remote sensing of primary production. The model depends on an approximately linear relationship between APAR and the normalized difference vegetation index. A more comprehensive form of the conventional model is shown to be necessary when different physiological types of plants or heterogeneous vegetation types occur within the study area. The predicted variable in the new model is total assimilation (net production plus respiration) rather than net production alone or harvest yield.

A model of regional primary production for use with coarse resolution satellite data

Mediterranean sclerophyll shrubs respond to seasonal drought by adjusting the amount of leaf area exposed and by reducing gas exchange via stomatal closure mechanisms. The degree to which each of these modifications can influence plant carbon and water balances under typical mediterranean-type climate conditions is examined. Leaf area changes are assessed in the context of a canopy structure and light microclimate model. Shifts in physiological response are examined with a mechanistically-based model of C3 leaf gas exchange that simulates progressive reduction of maximum photosynthesis and transpiration rates and increasingly strong midday stomatal closure over the course of drought. The results demonstrate that midday stomatal closure may effectively contribute to drought avoidance, increase water use efficiency, and strongly alter physiological efficiency in the conversion of intercepted light energy to photoproducts. Physiological adjustments lead to larger reductions in water use than occur when comparing leaf area index 3.5 to 1.5, extremes found for natural stands of sclerophyll shrubs in the California chaparral. Reductions in leaf area have the strongest effect on resource capture and use during non-water-stressed periods and the least effect under extreme drought conditions, while shifts in physiological response lead to large savings of water and efficient water use under extreme stress. An important model parameter termed GFAC (proportionality factor expressing the relation of conductance [g] to net photosynthesis rate) is utilized, which changes in response to the integrated water stress experimence of shrubs and alters the degree to which stomata may open for a given rate of carbon fixation. We attempt to interpret this parameter in terms of physiological mechanisms known to modify control of leaf gas exchange during drought. The analysis helps visualize means by which canopy gas exchange behavior may be coupled to physiological changes occurring in the root environment during soil drying.

A. Sala Serra

Papers

Factors influencing carbon fixation and water use by Mediterranean sclerophyll shrubs during summer drought