For over 40 y the dominant theory of stomatal behavior has been that plants should open stomates until the carbon gained by an infinitesimal additional opening balances the additional water lost times a water price that is constant at least over short periods. This theory has persisted because of its remarkable success in explaining strongly supported simple empirical models of stomatal conductance, even though we have also known for over 40 y that the theory is not consistent with competition among plants for water. We develop an alternative theory in which plants maximize carbon gain without pricing water loss and also add two features to both this and the classical theory, which are strongly supported by empirical evidence: (i) water flow through xylem that is progressively impaired as xylem water potential drops and (ii) fitness or carbon costs associated with low water potentials caused by a variety of mechanisms, including xylem damage repair. We show that our alternative carbon-maximization optimization is consistent with plant competition because it yields an evolutionary stable strategy (ESS)—species with the ESS stomatal behavior that will outcompete all others. We further show that, like the classical theory, the alternative theory also explains the functional forms of empirical stomatal models. We derive ways to test between the alternative optimization criteria by introducing a metric—the marginal xylem tension efficiency, which quantifies the amount of photosynthesis a plant will forego from opening stomatal an infinitesimal amount more to avoid a drop in water potential.
Earth system models are incorporating plant trait diversity into their land components to better predict vegetation dynamics in a changing climate. However, extant plant trait distributions will not allow extrapolations to novel community assemblages in future climates; that will require a mechanistic understanding of the trade-offs that determine trait diversity. In this study, we show how physiological trade-offs involving leaf mass per unit area (LMA), leaf lifespan, leaf nitrogen, and leaf respiration may explain the distribution patterns of evergreen and deciduous trees in the temperate and boreal zones based on (1) an evolutionary analysis of a simple mathematical model and (2) simulation experiments of an individual-based dynamic vegetation model (i.e. LM3-PPA). The evolutionary analysis shows that these leaf traits set up a trade-off between carbon and nitrogen use efficiency at the scale of individual trees and therefore determine competitively dominant leaf strategies. As soil nitrogen availability increases, the dominant leaf strategy switches from one that is high in nitrogen use efficiency to one that is high in carbon use efficiency or, equivalently, from high LMA/long-lived leaves (i.e. evergreen) to low LMA/ short-lived leaves (i.e. deciduous). In a region of intermediate soil nitrogen availability, the dominant leaf strategy may be either deciduous or evergreen depending on the initial conditions of plant trait abundance (i.e. founder controlled) due to feedbacks of leaf traits on soil nitrogen mineralization through litter quality. Simulated successional patterns predicted by LM3-PPA from the leaf physiological trade-offs are consistent with observed successional dynamics of evergreen and deciduous forests at three sites spanning the temperate to boreal zones.
Tropical tree size distributions are remarkably consistent despite differences in the environments that support them. With data analysis and theory, we found a simple and biologically intuitive hypothesis to explain this property, which is the foundation of forest dynamics modeling and carbon storage estimates. After a disturbance, new individuals in the forest gap grow quickly in full sun until they begin to overtop one another. The two-dimensional space-filling of the growing crowns of the tallest individuals relegates a group of losing, slow-growing individuals to the understory. Those left in the understory follow a power-law size distribution, the scaling of which depends on only the crown area–to–diameter allometry exponent: a well-conserved value across tropical forests.