Conflicting demands on angiosperm xylem: Tradeoffs among storage, transport and biomechanics

The secondary xylem of woody plants transports water mechanically supports the plant body and stores resources. These three functions are interdependent giving rise to tradeoffs in function. Understanding the relationships among these functions and their structural basis forms the context in which to interpret xylem evolution. The tradeoff between xylem transport efficiency and safety from cavitation has been carefully examined with less focus on other functions, particularly storage. Here, we synthesize data on all three xylem functions in angiosperm branch xylem in the context of tradeoffs. Species that have low safety and efficiency, examined from a resource economics perspective, are predicted to be adapted for slow resource acquisition and turnover as characterizes some environments. Tradeoffs with water storage primarily arise because of differences in fibre traits, while tradeoffs in carbohydrate storage are driven by parenchyma content of tissue. We find support for a tradeoff between safety from cavitation and storage of both water and starch in branch xylem tissue and between water storage capacity and mechanical strength. Living fibres may facilitate carbohydrate storage without compromising mechanical strength. The division of labour between different xylem cell types allows for considerable functional and structural diversity at multiple scales. The vascular system of woody plants performs multiple functions related to transport, mechanical support and storage. No one vascular system can maximize performance of all three functions because of structural and evolutionary limitations, and this gives rise to tradeoffs. Understanding these tradeoffs and their structural basis yields insight into the evolution of this ecologically and commercially important tissue.