Self-incompatibility (SI) is a genetic mechanism that restricts inbreeding in flowering plants. In the nightshade family (Solanaceae) SI is controlled by a single multiallelic S locus. Pollen rejection in this system requires the interaction of two S locus products: a stylar (S)-RNase and its pollen counterpart (pollen S). pollen S has not yet been cloned. Our understanding of how this gene functions comes from studies of plants with mutations that affect the pollen but not the stylar SI response (pollen-part mutations). These mutations are frequently associated with duplicated S alleles, but the absence of an obvious additional allele in some plants suggests pollen S can also be deleted. We studied Nicotiana alata plants with an additional S allele and show that duplication causes a pollen-part mutation in several different genetic backgrounds. Inheritance of the duplication was consistent with a competitive interaction model in which any two nonmatching S alleles cause a breakdown of SI when present in the same pollen grain. We also examined plants with presumed deletions of pollen S and found that they instead have duplications that included pollen S but not the S-RNase gene. This finding is consistent with a bipartite structure for the S locus. The absence of pollen S deletions in this study and perhaps other studies suggests that pollen S might be required for pollen viability, possibly because its product acts as an S-RNase inhibitor.S elf-incompatibility (SI) in many plant families is controlled by a multiallelic S locus that enables a style to reject any pollen expressing the same S allelic specificity as itself (1). In the Solanaceae, the family that includes tobacco, tomato, and petunia, SI is described as gametophytic because the allelic specificity of each pollen grain is determined by its own haploid genotype. The S locus in this family encodes a secreted extracellular RNase [stylar (S)-RNase] that accumulates in the style (2). Recognizing which S allele each pollen grain expresses is thought to require an interaction between the S-RNase and an unknown product(s) of a second S locus gene called pollen S (3, 4).As part of a strategy to identify pollen S, we isolated Nicotiana alata plants with gamma ray induced mutations that specifically affect the SI phenotype of pollen but not the SI phenotype of the style (5). Such plants are called pollen-part mutants (PPMs). Because ionizing radiation can cause either the deletion of part of a chromosome or chromosomal aberrations such as translocations, inversions, and fragments (6), the mutations in PPMs are likely to be complex because they can arise through one of a few different types of lesion.Among the PPMs described so far, the most frequent types of lesion are either translocations or small ''centric'' fragments (short extra chromosomes) that carry a duplicated copy of an S allele (5, 7-10). Breakdown of the pollen SI response in these plants occurs because of a ''competitive interaction'' that enables pollen with two different S alleles (but not two i...