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Transcript
Editorial Blueprints for Building Plant Cell Walls Elucidating the structure and functional properties of plant cell walls remains a daunting scientific challenge. While some experimental hurdles are common to many areas of plant biology (e.g. the design of novel assays for gene products of unannotated function), others are compounded in cell wall research because it involves the study of highly complex architectures built from secondary gene products and assembled at the cell’s frontier. Full genome sequences are providing inventories of an estimated 10% of genes devoted to the synthesis, secretion, targeting, assembly, and remodeling of diverse wall structures made in different cell types and deposited in distinct regions even around a single cell. However, we still lack fundamental and complete knowledge of sequences of wall polysaccharides beyond their unit structures of 10 sugars or so, as well as the nature, number, and spacing of cross-links that connect them. Similarly, many aspects of the composition and organization of the constituents of cuticularized and lignified walls remain mysterious. For example, the targeted deposition of monolignols, sites of polymerization, and assembly into a crosslinked macromolecule are not at all understood, and most of our knowledge is inferred from examining products once the macromolecular structure is dissolved. Furthermore, we know little about the pathways by which polysaccharides and other wall components transit from their sites of synthesis to the cell surface. The availability of loss-of-function and gain-of-function genetic variants provides useful tools to address function, but our ability to perturb structures and understand the consequences are severely limited by our knowledge of functional molecular architectures. The diversity of wall composition and architecture found in distinct regions or domains around a cell, in different tissues, and across taxonomic groups underscores the heterogeneous and dynamic nature of this cellular compartment and its key importance in cell growth and differentiation. Not surprisingly, then, in addition to the structural components of the wall, the apoplastic environment as a whole and the wallplasma membrane interface are host to large numbers of proteins with a multitude of functions. These include many families of receptors, whose responses may be conditioned by an environment with numerous complex elements, and new components of signal transduction pathways related to sensing the presence of pathogens, abiotic conditions, metabolic status, and structural integrity of the wall are just beginning to emerge. Plant cell walls, the source of lignocellulosic biomass, are currently in the spotlight for their value as feedstocks in the production of ethanol, butanol, and other advanced biofuels. As global challenges of energy and food security emerge, use-inspired research into deconstructing cell wall polysaccharides for fermentation by microbes, digestibility of animal feed, and nutritional content and quality of grains for human consumption is a critical component of the response. We wear cell wall products as natural fibers, use them as wood and paper products, and value the textural properties of fruits and vegetables that they impart. However, this Focus Issue touches only peripherally on the “four F’s” of food, feed, fiber, and fuel and rather aims to provide an update on our fundamental understanding of aspects of cell wall biology from polymer biosynthesis to their evolutionary history. In the Update articles presented in this Focus Issue, the contributors have synthesized current knowledge of cell wall biogenesis; the secretion and targeting mechanisms of individual components such as cellulose synthase; the catalytic mechanisms of some of the myriad enzyme activities involved in wall remodeling; the intersection between cell wall molecules, receptors, and signaling pathways; and the inference of structure-function relationships from an evolutionary-developmental perspective. These overviews are complemented by several highlighted research articles that showcase new tools to probe the complexity of cell wall architecture as well as bioinformatic, genetic, and biochemical analyses of members of large cell wallrelated gene families. The response to the call for this Focus Issue was tremendous and will spill over into the next few volumes of Plant Physiology. These other Updates and Research Articles will be associated electronically with this Focus Issue. We believe that this response is symptomatic of a reenergized research community for cell wall biology, stimulated but not displaced by useinspired research. Thanks to all of the contributors for their insights and for showcasing the progress and challenges of this area. These are exciting times! Maureen McCann and Jocelyn Rose Guest Editors Plant Physiology www.plantphysiol.org/cgi/doi/10.1104/pp.110.900324 Plant PhysiologyÒ, June 2010, Vol. 153, p. 365, www.plantphysiol.org Ó 2010 American Society of Plant Biologists Downloaded from on June 16, 2017 - Published by www.plantphysiol.org Copyright © 2010 American Society of Plant Biologists. All rights reserved. 365