Exploring the optimum nitrogen partitioning to predict the acclimation of C3 leaf photosynthesis to varying growth conditions.

The distribution of leaf nitrogen among photosynthetic proteins (i.e. chlorophyll, the electron transport system, Rubisco, and other soluble proteins) responds to environmental changes. We hypothesize that this response may underlie the biochemical aspect of leaf acclimation to the growth environment, and describe an analytical method to solve optimum nitrogen partitioning for maximized photosynthesis in C3 leaves. The method predicts a high investment of nitrogen in Rubisco under conditions leading to excessive energy supply relative to metabolic demand (e.g. low temperature, high light, low nitrogen, or low CO2). Conversely, more nitrogen is invested in chlorophyll when the energy supply is limiting. Overall, our optimization results are qualitatively consistent with literature reports. Commonly reported changes in photosynthetic parameters with growth temperature were emergent properties of the optimum nitrogen partitioning. The method was used to simulate dynamic acclimation under varying environmental conditions, using first-order kinetics. Simulated diurnal patterns of leaf photosynthetic rates as a result of acclimation differed greatly from those without acclimation (Awithout). However, differences in predicted photosynthesis integrated over a day or over the growing season from Awithout depended on the value of the kinetic time constant (τ), suggesting that τ is a critical parameter determining the overall impact of nitrogen distribution on acclimated photosynthesis.

On the relative magnitudes of photosynthesis, respiration, growth and carbon storage in vegetation.

BACKGROUND AND AIMS: The carbon balance of vegetation is dominated by the two large fluxes of photosynthesis (P) and respiration (R). Mechanistic models have attempted to simulate the two fluxes separately, each with their own set of internal and external controls. This has led to model predictions where environmental change causes R to exceed P, with consequent dieback of vegetation. However, empirical evidence suggests that the R : P ratio is constrained to a narrow range of about 0.4-0.5. Physiological explanations for the narrow range are not conclusive. The aim of this work is to introduce a novel perspective by theoretical study of the quantitative relationship between the four carbon fluxes of P, R, growth and storage (or its inverse, remobilization). METHODS: Starting from the law of conservation of mass - in this case carbon - equations are derived for the relative magnitudes of all carbon fluxes, which depend on only two parameters: the R : P ratio and the relative rate of storage of carbon in remobilizable reserves. The equations are used to explain observed flux ratios and to analyse incomplete data sets of carbon fluxes. KEY RESULTS: The storage rate is shown to be a freely varying parameter, whereas R : P is narrowly constrained. This explains the constancy of the ratio reported in the literature. With the information thus gained, a data set of R and P in grassland was analysed, and flux estimates could be derived for the periods after cuts in which plant growth is dominated by remobilization before photosynthesis takes over. CONCLUSIONS: It is concluded that the relative magnitudes of photosynthesis, respiration, growth and substrate storage are indeed tightly constrained, but because of mass conservation rather than for physiological reasons. This facilitates analysis of incomplete data sets. Mechanistic models, as the embodiment of physiological mechanisms, need to show consistency with the constraints.

No evidence for substantial aerobic methane emission by terrestrial plants: a 13C-labelling approach.

* The results of a single publication stating that terrestrial plants emit methane has sparked a discussion in several scientific journals, but an independent test has not yet been performed. * Here it is shown, with the use of the stable isotope (13)C and a laser-based measuring technique, that there is no evidence for substantial aerobic methane emission by terrestrial plants, maximally 0.3% (0.4 ng g(-1) h(-1)) of the previously published values. * Data presented here indicate that the contribution of terrestrial plants to global methane emission is very small at best. * Therefore, a revision of carbon sequestration accounting practices based on the earlier reported contribution of methane from terrestrial vegetation is redundant.

Some quantitative relationships between leaf area index and canopy nitrogen content and distribution.

In a previous study (Yin et al. 2000. Annals of Botany 85: 579-585), a generic logarithmic equation for leaf area index (L) in relation to canopy nitrogen content (N) was developed: L=(1/ktn)1n(1+ktnN/nb). The equation has two parameters: the minimum leaf nitrogen required to support photosynthesis (nb), and the leaf nitrogen extinction coefficient (ktn). Relative to nb, there is less information in the literature regarding the variation of ktn. We therefore derived an equation to theoretically estimate the value of ktn. The predicted profile of leaf nitrogen in a canopy using this theoretically estimated value of ktn is slightly more uniform than the profile predicted by the optimum nitrogen distribution that maximizes canopy photosynthesis. Relative to the optimum profile, the predicted profile is somewhat closer to the observed one. Based on the L-N logarithmic equation and the theoretical ktn value, we further quantified early leaf area development of a canopy in relation to nitrogen using simulation analysis. In general, there are two types of relations between L and N, which hold for canopies at different developmental phases. For a fully developed canopy where the lowest leaves are senescing due to nitrogen shortage, the relationship between L and N is described well by the logarithmic model above. For a young, unclosed canopy (i.e. L < 1.0), the relation between L and N is nearly linear. This linearity is virtually the special case of the logarithmic model when applied to a young canopy where its total nitrogen content approaches zero and the amount of nitrogen in its lowest leaves is well above nb. The expected patterns of the L-N relationship are discussed for the phase of transition from young to fully developed canopies.