Tropomyosins were discovered as regulators of actomyosin contractility in muscle cells, making yeasts and other fungi seem unlikely to harbor such proteins. Fungal cells are encased in a rigid cell wall and do not engage in the same sorts of contractile shape changes of animal cells. However, discovery of actin and myosin in yeast raised the possibility for a role for tropomyosin in regulating their interaction. Through a biochemical search, fungal tropomyosins were identified with strong similarities to their animal counterparts in terms ofprotein structure and physical properties. Two particular fungi, the buddingyeast Saccharomyces cerevisiae and the fission yeast Schizosaccharomyces pombe, have provided powerful genetic systems for studying tropomyosins in nonmetazoans. In these yeasts, tropomyosins associate with subsets ofactin filamentous structures. Mutational studies oftropomyosin genes and biochemical assays of purified proteins point to roles for these proteins as factors that stabilize actin filaments, promote actin-based structures of particular architecture and help maintain distinct biochemical identities among different filament populations. Tropomyosin-enriched filaments are the cytoskeletal structures that promote the major cell shape changes of these organisms: polarized growth and cell division.