Cytogenetic mapping of marker genes on the chromosome elements C and E of Drosophila pseudoobscura and D. subobscura

Enzyme loci, visible marker genes and -cloned DNA-sequences from a D. miranda library were mapped cytologically on the chromosome elements C and E of D. pseudoobscura and D. subobscura. New data are incorporated into the linkage maps of the two species. Homologous segments can now be localized in the polytene chromosomes with these markers. A comparison of the chromosome elements E of D. melanogaster and D. subobscura shows 12 conserved subsections which have been rearranged by paracentric inversions in the evolution of the two lineages.     

Cytogenetic Mapping of Enzyme Loci on Chromosomes J and U of DROSOPHILA SUBOBSCURA

Enzyme loci located on chromosome J and U were mapped cytologically by means of a Y translocation technique. A linkage map of the two chromosomes was established in a parallel experiment and the recombination frequency in different regions of the chromosomes determined. A comparison of the cytogenetic localization of the enzyme genes in D. subobscura and D. melanogaster indicates that many paracentric inversions must have taken place in the course of divergent evolution. However, no displacements of genes from one element to another due to pericentric inversions, reciprocal translocations or transposing elements can be observed. In spite of the large number of structural rearrangements that have occurred in the phylogeny of the genus Drosophila, gross similarities of banding pattern in homologous regions of the chromosomes of the two species become apparent.     

Inversionen und chromosomale Strukturtypen bei Drosophila subobscura Coll

Zusammenfassung 1. Auf Grund der Untersuchung von zahlreichen Stämmen von Drosophila subobscura, die aus Köderfängen in der Natur an verschiedenen Orten in Österreich, Oberitalien, der Schweiz, England und einigen außereuropäischen Ländern abgeleitet worden waren, wurde ein Verzeichnis aller bisher bei dieser Art festgestellten natürlichen Inversionen gegeben, wobei die Anordnung des strukturell homozygoten Stammes Küsnacht als Standardanordnung diente.2. Durch den Vergleich von Stämmen, durch deren Kreuzung mit dem Standardstamm, durch die Ableitung von Inzuchtlinien aus verschiedenem Ausgangsmaterial und deren Kreuzung mit bestimmten Stämmen wurde die Zahl und der Bau der chromosomalen Strukturtypen ermittelt, auf deren Koexistenz in der Population die Mannigfaltigkeit der beobachteten Inversionsbilder beruht.3. Die Verteilung der Inversionen auf die verschiedenen Chromosomen des Satzes ist nicht zufällig. Auch die relative Länge der Inversionen scheint nicht rein zufallsmäßig zu sein.4. Bei den Bruchstellen der Inversionen finden sich auffallend oft Koinzidenzen, bis zu 5 Bruchstellen an einem Ort eines Chromosoms. Wie eine statistische Überlegung zeigt, können diese nicht zufallsmäßig bedingt sein. Es werden einige hypothetische Deutungsmöglichkeiten dieses Phänomens vorgelegt.5. Es wird die phylogenetische Ableitung der Strukturtypen der verschiedenen Chromosomen diskutiert.Mit 7 Textabbildungen.     

Wild O chromosomes of Drosophila subobscura from different geographic regions have different effects on viability

By means of the marker strain Va/Ba wild chromosomes O of Drosophila subobscura were extracted from eight natural populations situated on a north-south gradient from Sweden or Scotland to Tunesia. Lethal frequencies and viability effects of the wild chromosomes O were studied in homozygous and random heterozygous combinations. In accordance with results from other Drosophila species random heterozygotes were always more viable than homozygotes. The viability-determining polygene system proved, however, dominant to some degree. Geographic differences became apparent especially with respect to three different characteristics: (1) The lethal frequencies for chromosomes O from central populations are higher than for those from northern and southern marginal populations; (2) Mean viabilities of non-lethal homozygotes and random heterozygotes are lower for central than for marginal populations; (3) The increase of viability through heterozygosity is more pronounced in the northern populations than in the others. The differences are thought to be mainly due to differences in the adaptation strategy of marginal and central populations. The viability fitness components seem of more importance for the marginal populations while fertility components may be of greater weight under central conditions. The geographic variability of the viability polygene system is finally compared with that of other genetic traits in D. subobscura.     

Geographic pattern of allozyme and inversion polymorphism on chromosome O of Drosophila subobscura and its evolutionary origin

The chromosome O of Drosophila subobscura was studied with respect to genetic variability at three enzyme loci (Odh, Me, and Lap-4) and with respect to inversion polymorphism. Population samples were taken from seven localities along a north-south gradient from Sweden and Scotland to Tunisia.The chromosomal analysis revealed clinal frequency changes for gene arrangements from north to south. With the enzyme loci Odh and Me allele frequencies are similar throughout the distribution range. Both loci are located outside the common inversion complex O₃₊₄. On the other hand, frequency changes parallel to those of the gene arrangements were observed for the alleles of the Lap-locus. Nonrandom associations between Lap-alleles and the superimposed gene arrangements O_ST, O₃₊₄, O₃₊₄₊₈, and O₃₊₄₊₂₃ were found. These gene arrangements differ from each other with respect to allele frequencies at the Lap-locus but for a given gene arrangement the relative frequencies of Lap-alleles remain relatively constant along the north-south gradient. Thus allele frequencies at the Lap-locus can be predicted from inversion frequencies.These observations can be interpreted in such a way that the pattern of allozyme variation within gene arrangements is due to founder effects caused by the unique origin of inversions. The gene blocks in the different inversions seem to represent more or less separated gene pools. In polymorphic populations the coexistence of genetically differentiated inversions presumably gives rise to heterotic interaction.