Decoding leaf hydraulics with a spatially explicit model: principles of venation architecture and implications for its evolution.

Leaf venation architecture is tremendously diverse across plant species. Understanding the hydraulic functions of given venation traits can clarify the organization of the vascular system and its adaptation to environment. Using a spatially explicit model (the program K_leaf), we subjected realistic simulated leaves to modifications and calculated the impacts on xylem and leaf hydraulic conductance (K(x) and K(leaf), respectively), important traits in determining photosynthesis and growth. We tested the sensitivity of leaves to altered vein order conductivities (1) in the absence or (2) presence of hierarchical vein architecture, (3) to major vein tapering, and (4) to modification of vein densities (length/leaf area). The K(x) and K(leaf) increased with individual vein order conductivities and densities; for hierarchical venation systems, the greatest impact was from increases in vein conductivity for lower vein orders and increases in density for higher vein orders. Individual vein order conductivities were colimiting of K(x) and K(leaf), as were their densities, but the effects of vein conductivities and densities were orthogonal. Both vein hierarchy and vein tapering increased K(x) relative to xylem construction cost. These results highlight the important consequences of venation traits for the economics, ecology, and evolution of plant transport capacity.