Imperforate tracheary element classification for studies of xylem structure-function relations

Abstract

Discovering how xylem function emerges from xylem structure requires approaches that are distinct from those invented for wood identification. I exemplify this need for different approaches by discussing the classification of imperforate tracheary elements, the cells that go by names such as "fibers,"tracheids, fiber-tracheids, and libriform fibers. Currently, there are two classification systems, one for wood identification and one championed by Sherwin Carlquist for research on xylem function. I emphasize that neither is globally "correct"for all applications, but instead each is appropriate for the aims of its field. Wood identification uses an easily-applied classification of imperforate tracheary elements, designed to remain stable over the long term. This stability allows anatomists to build large databases of wood data that are maximally useful for wood identification. In contrast, functional xylem biologists need flexible definitions that serve as hypotheses of imperforate tracheary element function to be tested, modified, and tested again, in an open-ended process of refining knowledge of xylem structure-function relations. I highlight some of the many open questions that the functional classification points to, such as the morphological correlates of the conductive/nonconductive imperforate tracheary element distinction, how imperforate tracheary element features are associated with vessel grouping, whether the libriform fiber-fiber tracheid distinction is arbitrary or not, and other topics. Recognizing that the purpose of a functional classification is to drive research shifts focus away from debates about whether a given cell type classification is "correct"and onto the empirical priorities that need study. While there is a solid tradition within comparative wood anatomy of training in wood identification, there is a need to train functional comparative wood anatomists, who can guide and interact with xylem physiologists to build more robust explanations of xylem structure and function.