The Arabidopsis wood model-the case for the inflorescence stem.

Arabidopsis thaliana has successfully served as a model to discover genes and proteins that have roles in a wide range of plant traits, including wood-related traits, such as lignin, cellulose and hemicellulose biosynthesis, secondary growth regulation, and secondary cell wall synthesis. Both the radially thickened hypocotyl and the inflorescence stem (flower stalk) have been studied. In this review, we address lingering doubts regarding the utility of Arabidopsis as a model for wood development by highlighting studies that provide new biochemical and biophysical evidence that extend support for the Arabidopsis inflorescence stem as a model for wood development beyond what is currently thought. We describe different aspects of Arabidopsis that make it a highly versatile tool for the study of wood development. One would likely utilise the radially thickened hypocotyl because of its more fully developed vascular cambium for traits related specifically to secondary (i.e. cambial) growth. It is more productive to utilise the inflorescence stem for wood-like biophysical traits. Accession variation has been underexploited as a powerful method to discover genes governing wood-like traits. We discuss recent findings that survey the accession variation in Arabidopsis for biochemical and biophysical properties of various wood traits, such as microfibril angle, tensile strength and cellulose/hemicellulose content. Furthermore we discuss how larger-scale studies of this nature using plants grown in long days (as opposed to the current short-day paradigm) could accelerate gene discovery and our understanding of cell wall and wood development. We highlight some relatively unexplored areas of research relating to the secondary cell wall composition, architecture and biophysical properties of the inflorescence stem, and how these traits are relevant to wood formation. The Arabidopsis inflorescence stem has other characteristics, expressed genes and traits held in common with woody species that have not been widely characterised or discussed to date. We discuss how this conservation may indicate the more general potential for "true" woodiness in herbaceous species, in the context of so-called secondary woodiness.