Growth and maturation of the edible cortical cells of apples (Malus domestica Borkh) are accompanied by a selective loss of pectin-associated (1/4)-b-D-galactan from the cell walls, whereas a selective loss of highly branched (1/5)-a-L-arabinans occurs after ripening and in advance of the loss of firm texture. The selective loss of highly branched arabinans occurs during the overripening of apples of four cultivars (Gala, Red Delicious, Firm Gold, and Gold Rush) that varied markedly in storage life, but, in all instances, the loss prestages the loss of firm texture, measured by both breaking strength and compression resistance. The unbranched (1/5)-linked arabinans remain associated with the major pectic polymer, rhamnogalacturonan I, and their content remains essentially unchanged during overripening. However, the degree of rhamnogalacturonan I branching at the rhamnosyl residues also decreases, but only after extensive loss of the highly branched arabinans. In contrast to the decrease in arabinan content, the loss of the rhamnogalacturonan I branching is tightly correlated with loss of firm texture in all cultivars, regardless of storage time. In vitro cell separation assays show that structural proteins, perhaps via their phenolic residues, and homogalacturonans also contribute to cell adhesion. Implications of these cell wall modifications in the mechanisms of apple cortex textural changes and cell separation are discussed.Loss of firm texture and cell adhesion are ripeningassociated processes that affect fruit quality and postharvest storage. Because the pectin substances are enriched in the walls of fruit cells compared to primary walls of other growing cells and because they constitute major components of the middle lamellae, these acidic polysaccharides have long been suspected to play key roles in fruit ripening (Jarvis, 1984;Brummel and Harpster, 2001). Over the years changes in pectin composition during fruit ripening have been studied extensively in many species, but the association of biochemical changes with changes in fruit texture is still unclear. Pectin depolymerization has little impact on wall swelling and softening during tomato (Lycopersicon esculentum) fruit ripening (Smith et al., 1988), and in other instances the swelling state of several fruit walls may change markedly without significant pectin depolymerization (Redgwell et al., 1997). Rose et al. (1998) show that wall swelling and softening may be accompanied by pectin depolymerization events, but the events are either not the only requirements or they may be unrelated to ripening. One of the surprises that emerged from the completion of the Arabidopsis genome sequence is the large number of genes that putatively encode polysaccharide degrading enzymes (Arabidopsis Genome Initiative, 2000). For example, Arabidopsis contains more than 250 genes that encode enzymes that are related to those that depolymerize pectins (Henrissat et al., 2001). This significant allotment of the genome toward pectin hydrolysis forces plant biologists to rec...