In land plants, xyloglucans (XyGs) tether cellulose microfibrils into a strong but extensible cell wall. The MUR2 and MUR3 genes of Arabidopsis encode XyG-specific fucosyl and galactosyl transferases, respectively. Mutations of these genes give precisely altered XyG structures missing one or both of these subtending sugar residues. Tensile strength measurements of etiolated hypocotyls revealed that galactosylation rather than fucosylation of the side chains is essential for maintenance of wall strength. Symptomatic of this loss of tensile strength is an abnormal swelling of the cells at the base of fully grown hypocotyls as well as bulging and marked increase in the diameter of the epidermal and underlying cortical cells. The presence of subtending galactosyl residues markedly enhance the activities of XyG endotransglucosylases and the accessibility of XyG to their action, indicating a role for this enzyme activity in XyG cleavage and religation in the wall during growth for maintenance of tensile strength. Although a shortening of XyGs that normally accompanies cell elongation appears to be slightly reduced, galactosylation of the XyGs is not strictly required for cell elongation, for lengthening the polymers that occurs in the wall upon secretion, or for binding of the XyGs to cellulose.The plant cell wall is continually modified during cell growth and differentiation. The tensile strength of the wall is provided by a dense spool of cellulose microfibrils interlaced with cross-linking glycans (Carpita and Gibeaut, 1993). When plant cells grow, the wall is biochemically "loosened" to permit turgor-driven cell expansion (Cosgrove, 2000). One of the mysteries of cell growth in plants that researchers have pondered for decades is how these interlaced glycans loosen without compromising the tensile strength of the pliant wall.In all dicots and certain monocots, xyloglucans (XyGs) are the principal cellulose tethering molecules, and the loosening of these tethers from the microfibrils provides a physical control point of cell expansion (Cosgrove, 2000). The unique ability of XyG endotransglucosylases (XETs) to cleave XyGs and rejoin the cut ends with new partners suggested a role for these enzyme activities in wall loosening during growth and in the restructuring of cell walls after extension (Nishitani and Tominaga, 1992). (A revised nomenclature has been adopted by consensus of researchers of the XyG endotransglucosylase/ hydrolase gene/protein family [Rose et al., 2002]. The abbreviation XTH refers to any gene/protein of the family regardless of activity of the protein. However, when referring to endotransglucosylase activities, the abbreviation remains XET, and hydrolase activities are abbreviated XEH.) However, the only cell wall proteins proven to be capable of causing extension of isolated walls in vitro under mechanical stress are expansins, which increase wall extensibility under constant stress (McQueen-Mason et al., 1992), and yieldins, which lower the "yield threshold," the minimum stress required that p...