Cope's Rule and the evolution of long-distance transport in vascular plants: allometric scaling, biomass partitioning and optimization

Recent advances in allometric theory have proposed a novel quantitative framework by which to view the evolution of plant form and function. This general theory has placed strong emphasis on the importance of long-distance transport in shaping the evolution of many attributes of plant form and function. Specifically, it is hypothesized that with the evolutionary increase in plant size natural selection has also resulted in vascular networks that minimize scaling of total hydrodynamic resistance associated with increasing transport distances. Herein the central features of this theory are reviewed and a broad sampling of supporting but yet preliminary empirical data are analysed. In particular, subtle attributes of the scaling of tracheid and vessel anatomy are hypothesized to be crucial for the evolution of increased plant size. Furthermore, the importance of minimizing hydrodynamic resistance associated with increased transport distances is also hypothesized to be reflected in an isometric scaling relationship between stem mass, M(S) and root mass, M(R) (i.e. M(S) proportional to M(R)). Preliminary data from multiple extant and fossil plant taxa provide tantalizing evidence supporting the predicted relationships. Together, these results suggest that selection for the minimization of the scaling of hydrodynamic resistance within plant vascular networks has in turn allowed for the enormous diversification in vascular plant size.