Height-related scaling of phloem anatomy and the evolution of sieve element end wall types in woody plants.

ABSTRACT

In the sieve elements (SEs) of the phloem, carbohydrates are transported throughout the whole plant from their site of production to sites of consumption or storage. SE structure, especially of the pore-rich end walls, has a direct effect on translocation efficiency. Differences in pore size and other features were interpreted as an evolutionary trend towards reduced hydraulic resistance. However, this has never been confirmed. Anatomical data of 447 species of woody angiosperms and gymnosperms were used for a phylogenetic analysis of end wall types, calculation of hydraulic resistance and correlation analysis with morphological and physiological variables. end wall types were defined according to pore arrangement either grouped into a single area (simple) or into multiple areas along the end wall (compound). Convergent evolution of end wall types was demonstrated in woody angiosperms. In addition, an optimization of end wall resistance with plant height was discovered, but found to be independent of end wall type. While physiological factors also showed no correlation with end wall types, the number of sieve areas per end wall was found to scale with SE length. The results exclude the minimization of hydraulic resistance as evolutionary driver of different end wall types, contradicting this long-standing assumption. Instead, end wall type might depend on SE length.