Genetic and molecular investigations were carried out with Eurasian Drosophila melanogaster populations on the P-M system of hybrid dysgenesis. In 27 strains sampled from France to Middle Asia, a clear gradient exists between Western Europe, in which most modern strains are of the Q type, and eastern areas, in which M-cytotype strains predominate. Molecular analysis on individual flies was performed with two complementary probes of the cloned 2.9-kilobase P element. The results provide evidence for a gradually decreasing frequency of P elements from west to east, but the presence of P-homologous sequences has been ascertained in all of the wild M-cytotype populations analyzed. Moreover, some active P elements with GD sterility potential were revealed in the majority of M-cytotype populations when tested with a highly sensitive reference line. The gradual change in distribution of the polymorphic P family in Eurasia is discussed in relation to the structure of the elements together with the theories of P-M evolution and is interpreted as the present invasion of Eurasian populations by these elements.About 10% of the genome ofDrosophila melanogaster exists as dispersed moderately repetitive sequences belonging to different families (1). The P family is composed of mobile dispersed genetic elements implicated in the P-M system of hybrid dysgenesis. This phenomenon, which is manifested in certain interstrain hybrids, results in a number of correlated aberrant genetic traits-e.g., high frequencies of gonadal sterility (GD sterility), mutation, and male recombination (2). Three types of individuals, P, Q, and M, have been described on the basis of their cross-effect properties. Hybrids between P males and M females show dysgenic traits that are reduced or absent in the reciprocal hybrids. Q individuals do not exhibit GD sterility in any cross-combinations but produce mutations and male recombinations in crosses with M females (3, 4).In the P-M system, hybrid dysgenesis results from interactions between chromosomally linked factors (P factors) and a particular type of cellular state referred to as the M cytotype (5). The P factors are active genetic elements of the P family, whose members are heterogeneous in size [0.5-2.9 kilobases (kb)], but which share substantial sequence homology (6-8). All P and Q strains thus far examined bear 30-50 copies of the P family (7). Q individuals are thought to carry a subset of the P-element family that apparently lacks sterility potential while retaining mutator activity and other Pelement functions (7,9,10). Conversely, all long-established laboratory M strains that have been examined completely lack homology with the P-element family (7). Some strains showing the M cytotype but with some homology to P sequences have also been found in laboratory collections (7).In this paper, such strains will be called M', the term M strain being reserved for strains of the M cytotype with no P homology at all.The M-cellular state component of the P-M interaction may be considered as a "s...