Polygalacturonases (PGs) cleave runs of unesterified GalUA that form homogalacturonan regions along the backbone of pectin. Homogalacturonan-rich pectin is commonly found in the middle lamella region of the wall where two adjacent cells abut and its integrity is important for cell adhesion. Transgenic apple (Malus domestica Borkh. cv Royal Gala) trees were produced that contained additional copies of a fruit-specific apple PG gene under a constitutive promoter. In contrast to previous studies in transgenic tobacco (Nicotiana tabacum) where PG overexpression had no effect on the plant (K.W. Osteryoung, K. Toenjes, B. Hall, V. Winkler, A.B. Bennett [1990] Plant Cell 2: 1239-1248), PG overexpression in transgenic apple led to a range of novel phenotypes. These phenotypes included silvery colored leaves and premature leaf shedding due to reduced cell adhesion in leaf abscission zones. Mature leaves had malformed and malfunctioning stomata that perturbed water relations and contributed to a brittle leaf phenotype. Chemical and ultrastructural analyses were used to relate the phenotypic changes to pectin changes in the leaf cell walls. The modification of apple trees by a single PG gene has offered a new and unexpected perspective on the role of pectin and cell wall adhesion in leaf morphology and stomatal development.Polygalacturonases (PGs) are expressed in a wide range of tissues and developmental stages in plants and are encoded by relatively large gene families (e.g. approximately 52 genes in Arabidopsis; The Arabidopsis Genome Initiative, 2000). PGs are associated with fruit ripening, cell separation processes such as leaf and flower abscission, pod and anther dehiscence, pollen grain maturation, pathogen defense, plant-host interactions, and processes of cell expansion, growth, and xylogenesis (for review, see Hadfield and Bennett, 1998; Bergey et al., 1999;Sitrit et al., 1999; Torki et al., 1999; Wang et al., 2000). Endo-PGs cleave runs of unesterified GalUA that form homogalacturonan regions along the backbone of pectin. Homogalacturonan-rich pectin is commonly found in the middle lamella region of the wall where two adjacent cells abut (for example, see Knox et al., 1990;Steele et al., 1997). The best characterized (fungal) endo-PG enzyme requires four to five consecutive runs of unesterified GalUA residues for cleavage; however, little is known about the enzyme activities and specificities of most cloned PGs from plants.Transgenic plants have been used to study the role of endo-PGs in vivo. In tomato (Lycopersicon esculentum), down-regulation of the fruit-specific PG gene pTOM6 under the control of the constitutive cauliflower mosaic virus 35S promoter showed reduced depolymerization of pectin polymers in fruit . Overexpression of PG in the ripeninginhibited mutant rin background restored PG activity and pectin degradation in fruit (Giovannoni et al., 1989). In both cases, only the fruit was affected by the transgene expression; therefore, the gene product isolated from tomato fruit appeared to have frui...