A comprehensive theory for the coupling between terrestrial carbon and water cycles, supported by stable carbon isotope measurements from leaves
Abstract:Stomata actively regulate the CO2 concentration inside plant leaves, which co-determines the biochemical rate of photosynthesis. Stomatal behaviour thus controls leaf-level water-use efficiency and the ‘exchange rate’ between the terrestrial carbon and hydrological cycles. Least-cost theory (based on the hypothesis that plants minimize the combined unit costs of maintaining the capacities for water transport and carbon uptake) predicts that (a) long-term mean values of the ci/ca ratio, i.e. the ratio of leaf-internal to ambient CO2 concentration, should be independent of both photon flux density and ca; and (b) these values should vary systematically with growing-season vapour pressure deficit, growth temperature, and atmospheric pressure.
Stable carbon isotope (δ13C) measurements provide an integrated measure of the ci/ca in C3 plants. A number of previous studies have focused on the aridity dependence of δ13C. The temperature dependence seems to have been overlooked, but the elevation dependence has been known for a long time: plants at high elevations have systematically lowered ci/ca, and correspondingly increased photosynthetic capacity (Vcmax). Why this should be is a long-standing puzzle: there are various speculative explanations in the literature, and a certain amount of controversy. By contrast, least-cost theory provides quantitative predictions of all three environmental effects. We have analysed a large (3652) set of δ13C measurements from C3 plants, spanning all latitudes and biomes, and shown that these predictions are quantitatively consistent with environmental dependences that can be shown in the measurements using a generalized linear model.
This analysis implies the ability to predict ci/ca ratios for large-scale terrestrial ecosystem modelling. Combined with the long-standing ‘co-ordination hypothesis’ for the control of photosynthetic capacity, least-cost theory provides a basis for a remarkably simple global model for gross primary production.