Nanos (Nos) is one of the evolutionarily conserved proteins known to direct germ-line development. In Drosophila, maternal Nos protein maintains transcriptional quiescence in the germ-line progenitors or pole cells to repress ectopic expression of somatic genes. Here we show that maternal Nos is required to establish and maintain germ-line identity by preventing apoptosis and somatic cell fate. The pole cells lacking maternal Nos were degraded by apoptosis during mid to late embryogenesis. When apoptosis was suppressed by Df(3L)H99, some pole cells lacking Nos adopted somatic cell fates. These pole cells expressed somatic markers ectopically and lost the germ-line marker Vasa. We further found that some Nos-negative pole cells were able to migrate into the gonads, but they failed to develop as functional germ cells during postembryonic stages. We propose a model in which Nos establishes germ-line͞soma dichotomy and is also required to maintain germ-line fate.T he mechanism underlying the segregation of germ line from somatic line is a century-old issue in developmental biology. In many animal groups, maternal factors required for germ-line formation are localized in a histologically remarkable region in egg cytoplasm, or germ plasm, and are inherited in the germ-line progenitors (1, 2). In Drosophila, the germ-line progenitors known as pole cells are first formed at the posterior pole of the blastoderm embryos. During later embryogenesis, the pole cells pass through midgut epithelium into hemocoel and migrate within the embryos to reach the embryonic gonads, where they differentiate as functional germ line (1-4). Previous works have demonstrated that none of the pole cells becomes incorporated into any somatic tissues and contributes to somatic development during embryogenesis (5, 6). However, it remains elusive how the developmental fate of pole cells is regulated by germ plasm components.One of the germ plasm components, Nanos (Nos), is evolutionarily conserved and has a widespread role in germ-line development (7-13). Maternal nos mRNA is enriched in germ plasm, and its protein product is partitioned into pole cells when they are formed and remains detectable in these cells throughout embryogenesis (14). Nos is required in pole cells for their proper migration into the embryonic gonads (15, 16). Within pole cells, Nos is involved in maintaining transcriptional quiescence (17) and is also required for maintenance of a germ-line-specific chromatin status that correlates with transcriptional inactivity (18). In the absence of maternal Nos activity, ectopic expression of somatic genes is detectable in pole cells (17). These results led us to speculate that pole cells lacking Nos may adopt somatic cell fate.Here, we show that pole cells are able to adopt both germ-line and somatic cell fates and undergo apoptosis. Nos is required to repress the pathway leading to somatic differentiation and apoptosis. Thus, we conclude that Nos is essential for germline͞soma dichotomy and for germ-line maintenance. H99-PLH⌬23͞TM3...