ACT7 encodes one of the six distinct and ancient subclasses of actin protein in the complex Arabidopsis actin gene family. We determined the sequence and structure of the Arabidopsis tbaliana ACT7 actin gene and investigated its tissue-specific expression and regulation. The ACT7 mRNA levels varied by 128-fold among severa1 different tissues and organs. The highest levels of ACT7 mRNA were found in rapidly expanding vegetative organs, the lowest in pollen. A translational fusion with the 5 ' end of ACT7 (1.9 kb) joined to the P-glucuronidase reporter gene was strongly and preferentially expressed in all young, developing vegetative tissues of transgenic Arabidopsis plants. ACT7 was the only Arabidopsis actin gene strongly expressed in the hypocotyl and seed coat. Although no p-glucuronidase expression was seen in developing ovules or immature seeds, strong expression was seen in dry seeds and immediately after imbibition in the entire seedling. ACT7was the only Arabidopsis actin gene to respond strongly to auxin, other hormone treatments, light regime, and wounding, and may be the primary actin gene responding to externa1 stimuli. The ACT7 promoter sequence contains a remarkable number of motifs with sequence similarity to putative phytohormone response elements.Cellular morphogenesis in plants is constrained by rigid cell walls that limit expansion, restrict shape, and prevent migration. Thus, morphogenesis is in part a function of asymmetric cell division and expansion. A number of studies have suggested that these processes are controlled by a dynamic cytoskeleton that consists of dozens of interacting proteins and responds to both developmental and environmental cues (Lloyd, 1991). Actin is a major component of the plant cytoskeleton and is thought to be required for correct cell-division plate alignment and synthesis, cellshape determination, cell-polarity establishment, cytoplasmic streaming, organelle movement, and tip growth (Staiger and Schliwa, 1987;Staiger and Lloyd, 1991;Meagher and Williamson, 1994