Abstract. The empirical Ball–Berry stomatal conductance model is commonly used in Earth system models to simulate biotic regulation of evapotranspiration. However, the dependence of stomatal conductance (gs) on vapor pressure deficit (Ds) and soil moisture must both be empirically parameterized. We evaluated the Ball–Berry model used in the Community Land Model version 4.5 (CLM4.5) and an alternative stomatal conductance model that links leaf gas exchange, plant hydraulic constraints, and the soil–plant–atmosphere continuum (SPA) to numerically optimize photosynthetic carbon gain per unit water loss while preventing leaf water potential dropping below a critical minimum level. We evaluated two alternative optimization algorithms: intrinsic water-use efficiency (Δ An/Δ gs, the marginal carbon gain of stomatal opening) and water-use efficiency (Δ An/Δ El, the marginal carbon gain of water loss). We implemented the stomatal models in a multi-layer plant canopy model, to resolve profiles of gas exchange, leaf water potential, and plant hydraulics within the canopy, and evaluated the simulations using: (1) leaf analyses; (2) canopy net radiation, sensible heat flux, latent heat flux, and gross primary production at six AmeriFlux sites spanning 51 site–years; and (3) parameter sensitivity analyses. Without soil moisture stress, the performance of the SPA stomatal conductance model was generally comparable to or somewhat better than the Ball–Berry model in flux tower simulations, but was significantly better than the Ball–Berry model when there was soil moisture stress. Functional dependence of gs on soil moisture emerged from the physiological theory linking leaf water-use efficiency and water flow to and from the leaf along the soil-to-leaf pathway rather than being imposed a priori, as in the Ball–Berry model. Similar functional dependence of gs on Ds emerged from the water-use efficiency optimization. Sensitivity analyses showed that two parameters (stomatal efficiency and root hydraulic conductivity) minimized errors with the SPA stomatal conductance model. The critical stomatal efficiency for optimization (ι) was estimated from leaf trait datasets and is related to the slope parameter (g1) of the Ball–Berry model. The optimized parameter value was consistent with this estimate. Optimized root hydraulic conductivity was consistent with estimates from literature surveys. The two central concepts embodied in the stomatal model, that plants account for both water-use efficiency and for hydraulic safety in regulating stomatal conductance, imply a notion of optimal plant strategies and provide testable model hypotheses, rather than empirical descriptions of plant behavior.