␥-Tubulin localizes to microtubule-organizing centers in animal and fungal cells where it is important for microtubule nucleation. Plant cells do not have morphologically defined microtubule organizing centers, however, and ␥-tubulin is distributed in small, discrete structures along microtubules. The great difference in distribution has prompted speculation that plant ␥-tubulins function differently from animal and fungal ␥-tubulins. We tested this possibility by expressing Arabidopsis ␥-tubulin in the fission yeast Schizosaccharomyces pombe. At high temperatures, the plant ␥-tubulin was able to bind to microtubule-organizing centers, nucleate microtubule assembly, and support the growth and replication of S. pombe cells lacking endogenous ␥-tubulin. However, the distribution of microtubules was abnormal as was cell morphology, and at low temperatures, cells were arrested in mitosis. These results reveal that Arabidopsis ␥-tubulin can carry out essential functions in S. pombe and is, thus, functionally conserved. The morphological abnormalities reveal that it cannot carry out some nonessential functions, however, and they underscore the importance of ␥-tubulin in morphogenesis of fission yeast cells and in maintaining normal interphase microtubule arrays.Nucleation of microtubule assembly is a key step in the cellular organization of microtubules (for review, see Brinkley, 1985;Oakley, 1992;Joshi, 1994; Gislene and Schiebel, 1997;Oakley and Akkari, 1999), and ␥-tubulin is an indispensable component of microtubule nucleation complexes in a variety of organisms including fungi and animal cells (Oakley et al., 1990; Horio et al., 1991;Stearns et al., 1991;Zheng et al., 1991; Joshi et al., 1992; Felix et al., 1994). Previously, we have reported that human ␥-tubulin can replace the endogenous ␥-tubulin of Schizosaccharomyces pombe (Horio and Oakley, 1994). This finding demonstrated that ␥-tubulin is functionally conserved in phylogenetically distant organisms with microtubule-organizing centers (MTOCs) that differ vastly in size and appearance. However, the spindles of these species do share the common feature that microtubules extend from discrete MTOCs, spindle pole bodies (SPBs) in fungi, and centrosomes in higher animal cells.In contrast, in plant cells, spindle microtubules are less focused than in typical animal cells and no discrete MTOCs equivalent to the centrosomes in animal cells are present (for review, see Vaughn and Harper, 1998; Canaday et al., 2000;Wasteneys, 2002). The localization of ␥-tubulin in plant cells has been examined extensively (Liu et al., 1993(Liu et al., , 1994(Liu et al., , 1995 Binarova et al., 2000;Kumagai et al., 2003). In most microtubule arrays of plant cells, ␥-tubulin is distributed along the microtubules, often in very small, punctate structures. Although ␥-tubulin was sometimes enriched in regions containing the minus ends of the microtubules, it did not localize to, or help to define, MTOCs (Liu et al., 1993(Liu et al., , 1994 for review, see Joshi and Palevitz, 1996;Vaug...