Association of Chromosome and Enzyme Polymorphisms in Natural and Cage Populations of DROSOPHILA MELANOGASTER

The frequencies of a polymorphic inversion, In(2L)t, and of Adh and αGpdh alleles were analyzed in three natural populations of Drosophila melanogaster from Japan. Significant positive correlations between the frequencies of In(2L)t and Adh^S or αGpdh^F were detected due to tight linkage. An analysis of correlation with latitude showed that the negative cline of Adh^S frequency could be explained entirely by its linkage with In(2L)t; the frequency of Adh^S on the standard chromosome did not show a latitudinal cline. To the contrary, the cline of αGpdh^F frequency itself was positive, and its linkage with In(2L)t makes the positive cline unclear. These results suggest that the two allozymes themselves respond to latitudinal natural selection in different ways. When these populations were transferred to laboratory cages and maintained for a long time, they lost the chromosomal polymorphism but retained stable enzyme polymorphisms, although allele frequencies in the cage were not the same as in nature. The frequencies of Adh and αGpdh alleles were close to those in earlier cage populations of the same geographical origin.     

Linkage Disequilibrium in Natural Populations of DROSOPHILA MELANOGASTER

Two large, stable populations (Texas and Japan) of Drosophila melanogaster were surveyed at 21 allozyme loci on the second and third chromosomes and for chromosomal gene arrangements on those two chromosomes. Over 220 independent gametes were sampled from each population. The types and frequencies of the surveyed genetic variation are similar to those observed previously and suggest only slight differentiation among geographically distant populations. Linkage disequilibrium among linked allozymes loci is only slightly, if at all, detectable with these sample sizes. Linkage disequilibrium between linked inversions and allozymes loci is common especially when located in the same arm. These disequilibria appear to be in the same direction for most comparisons in the two population samples. This result is interpreted as evidence of similar selective environments (ecological and genetic) in the two populations. It is also noted that the direction of these linkage disequilibria appears to be oriented with respect to the gene frequencies at the component loci.     

Genes Affecting Productivity in Natural Populations of DROSOPHILA MELANOGASTER

Two hundred second chromosomes were extracted from a Japanese population in October of 1972, and the viabilities and productivities of homozygotes and heterozygotes from them were examined. Viability was measured by the Cy method and productivity by the number of progeny produced per female. The frequency of lethal-carrying chromosomes was 0.315. When the average heterozygote viability was standardized as 1.000, the average homozygote viability was 0.595 including the lethal lines, and 0.866 excluding them. The frequency of recessive sterile chromosomes among 131 non-lethal lines was 0.092 in females and 0.183 in males. There were two instances in which homozygosis for the second chromosome caused sterility in both sexes, which was close to the number expected (2.2) on a random basis of 0.092 x 0.183 x 131. When the average heterozygote productivity of 200 lines was standardized as 1.000, the average homozygote productivity was 0.532 including female steriles, and 0.584 excluding them. The ratio of detrimental load to lethal load was 0.383, while the ratio of partial sterility load to complete sterility load was 5.767. The average viability of lethal heterozygotes was slightly, but not significantly, lower than that of lethal-free heterozygotes, while the average productivity of lethal heterozygotes was significantly lower than that of lethal-free heterozygotes. There was a significant association of sterility in either sex with low viability of homozygotes. However, no statistically significant differences in viability and productivity were detected between sterile heterozygotes and non-sterile heterozygotes. The heterozygous effects of viability and productivity polygenes were examined by regressions of the heterozygotes on the sum of corresponding homozygotes. The regression coefficients were slightly positive for both viability and productivity if lethal and sterile chromosomes were excluded. The correlation between viability and productivity in homozygotes was significantly positive when sterile chromosomes were included, but the significance disappeared when the sterile chromosomes were excluded. In the heterozygotes there were no detectable correlations between them.     

Chromosome Inversion Polymorphisms in DROSOPHILA MELANOGASTER. I. Latitudinal Clines and Associations between Inversions in Australasian Populations

Nineteen Australasian populations of Drosophila melanogaster have been screened for chromosome inversion polymorphisms. All 15 of the inversion types found are paracentric and autosomal, but only four of these, one on each of the major autosome arms, are common and cosmopolitan. North-south clines occur, with the frequencies of all four of the common cosmopolitan inversions increasing toward the equator. These clines in the Southern Hemisphere mirror north-south clines in the Northern Hemisphere, where the frequencies of all four of the common cosmopolitan inversions again increase towards the equator.-While few of the Australasian populations show significant disequilibrium between linked common cosmopolitan inversions, those that do invariably have excesses of coupling gametes, which is consistent with other reports. We also find nonrandom associations between the two major autosomes, with the northern populations in Australasia (those with high inversion frequencies) tending to be deficient in gametes with common cosmopolitan inversions on both major autosomes, while the southern populations in Australasia (low inversion frequencies) tend to have an excess of this class of gametes.-The clines and the nonrandom associations between the two major autosomes are best interpreted in terms of selection operating to maintain the common cosmopolitan inversion polymorphisms in natural populations of D. melanogaster.     

Concurrent Multiple Paternity in Natural and Laboratory Populations of DROSOPHILA MELANOGASTER

Electrophoretic analysis of two or three linked enzyme loci in progenies of homozygous females from cage and natural populations demonstrates frequent concurrent multiple paternity in both cases.     

The Response of Enzyme Polymorphisms to Developmental Rate Selection in DROSOPHILA MELANOGASTER

Two strains of Drosophila melanogaster derived from different geographical localities were subjected to selection for fast and slow developmental rate. Four enzyme polymorphisms were monitored for their response to such selection. Significant changes in allele frequencies were effected for the αGpdh and 6Pgd polymorphisms under fast developmental rate selection. The strain from Rhode Island exhibited a correlated change in body weight during developmental rate selection, whereas the Georgia strain did not.     

Linkage Disequilibrium in Natural Populations of DROSOPHILA MELANOGASTER. Seasonal Variation

Linkage disequilibrium among ten polymorphic allozyme loci and polymorphic inversions on chromosomes 2 and 3 in a natural population of Drosophila melanogaster was examined early and late in the annual season. Similar to previous studies, little linkage disequilibrium was observed among allozymes. The two significant cases that were observed in the first sample behaved in a contradictory way. One declined much more rapidly than expected due simply to recombination; the other declined slowly as expected. There was little change in allozyme or inversion frequencies during the season.     

Genetic and Biochemical Basis of Enzyme Activity Variation in Natural Populations. I. Alcohol Dehydrogenase in DROSOPHILA MELANOGASTER

Recent studies by various authors suggest that variation in gene regulation may be common in nature, and might be of great evolutionary consequence; but the ascertainment of variation in gene regulation has proven to be a difficult problem. In this study, we explore this problem by measuring alcohol dehydrogenase (ADH) activity in Drosophila melanogaster strains homozygous for various combinations of given second and third chromosomes sampled from a natural population. The structural locus (Adh) coding for ADH is on the second chromosome. The results show that: (1) there are genes, other than Adh, that affect the levels of ADH activity; (2) at least some of these "regulatory" genes are located on the third chromosome, and thus are not adjacent to the Adh locus; (3) variation exists in natural populations for such regulatory genes; (4) the effect of these regulatory genes varies as they interact with different second chromosomes; (5) third chromosomes with high-activity genes are either partially or completely dominant over chromosomes with low-activity genes; (6) the effects of the regulatory genes are pervasive throughout development; and (7) the third chromosome genes regulate the levels of ADH activity by affecting the number of ADH molecules in the flies. The results are consistent with the view that the evolution of regulatory genes may play an important role in adaptation.     

Polymorphism of the Hereditary Sigma Virus in Natural Populations of DROSOPHILA MELANOGASTER

Previous studies have shown that, in natural French populations of Drosophila melanogaster, 10 to 20% of the flies are infected by the noncontagious, hereditary rhabdovirus sigma responsible for CO₂ sensitivity. These populations are also polymorphic for two alleles [ref(2)P^o and ref(2)P^p] of a gene for resistance to the sigma virus. Evidence is given here that two viral genetic types, differing in their response to the ref(2)P^p allele, are present in these populations of flies; the most common type is only slightly sensitive to the ref(2)P^p allele.     

Inversion Clines in Populations of DROSOPHILA MELANOGASTER

Twenty different natural populations of Drosophila melanogaster were sampled to determine the frequencies of inversions. Based on their frequencies and geographical distributions, the inversions could be classified as follows: (1) Common cosmopolitan inversions that are present in many populations in frequencies exceeding five percent and that may exhibit frequency clines over large geographical regions; (2) Rare cosmopolitan inversions that occur throughout the species range but usually at frequencies below five percent and that may be absent in many populations; (3) Recurrent endemic inversions that are found in several adjacent populations in frequencies usually not exceeding one or two percent; and (4) Unique endemic inversions that are found only among the progeny of a single individual and that may represent one aspect of the syndrome termed "hybrid dysgenesis". Four common cosmopolitan inversions that exhibit highly significant clines in populations in the eastern United States are In(2L)t, In(2R)NS, In(3L)P and In(3R)P.     

Factors Influencing the Effect of Segregation Distortion in Natural Populations of DROSOPHILA MELANOGASTER

The major components of the SD system have been examined in two natural populations of D. melanogaster to investigate how SD behaves and is maintained in nature and to estimate its impact and efficiency. A twofold approach was used: (1) direct measurements of segregation distortion in wild males and (2) measurement of sensitivity of wild SD (+) chromosomes to SD action. Characterization of newly isolated SD chromosomes and of a large number of SD( +) chromosomes from nature demonstrated that (1) SD can operate efficiently in the wild genome: 45% of SD/SD(+) males collected from nature had k values larger than 0.70. (2) Forty-three of 44 newly recovered SD chromosomes are of the SD-72 type, having a small pericentric inversion that maintains tight linkage among the Sd, E(SD) and Rsp loci in the SD complex. In 1956, most SD chromosomes in Madison lacked this inversion. (3) Only 12 of the 44 SD chromosomes carried a recessive lethal (compared with five of six in 1956), and many of the viable SD chromosomes were fertile as homozygotes, indicating that SD homozygotes need not have obvious reductions in fitness. (4) Among more than 500 wild chromosomes assayed for response to distortion by a strong SD, at least 40-50% were sensitive, about 33% were partially sensitive and 17% were insensitive. This frequency of sensitives is higher than in reports from some other populations. An estimated 12% of the wild chromosomes were classified as true Rsp(i) by their constellation of effects, including a special test of ability to cause self-distortion of a "suicide" chromosome, R(cn)-10. In a direct assay with R(cn)-10, an independent sample of 99 chromosomes from nature gave 30% putative Rsp(i). Thus, these populations contain in the range of 12-30% Rsp(i). (5) Chromosomes supersensitive to SD, previously described for certain laboratory stocks, were also found to coexist in nature with SD. (6) Profiles of wild chromosomes with a panel of three or four different SD testers suggest a series of allelic alternatives at the Rsp locus including supersensitive, sensitive, semisensitive and insensitive, and that loci other than Rsp may also be important in determining the effect of SD in nature.     

Cytogenetic Analysis of an SD Chromosome from a Natural Population of DROSOPHILA MELANOGASTER

The discovery and the cytogenetic characterization of a new SD (Segregation Distorter) chromosome 2 from a natural population in Ranna (Sicily, Italy), SD(Ra), are reported. The main features of this chromosome are as follows: (a) it contains an Sd(Ra) gene with a moderate degree of segregation distortion (k = 0.72), (b) a recessive female sterile gene, fs(2)(TLM), responsible for modifications of the morphology and structure of the tests and ovaries is located at 89.7, (c) SD(Ra)/SD(Ra) males and females are viable but sterile, the females due to homozygosis of fs(2)(TLM) and the males because of homozygosis of a region containing the Sd locus, and (d) SDi/SDj combinations are fertile, thus suggesting that the different Sd factors found in natural populations constitute a multiple allelic series.-These data may indicate that each population containing SD chromosomes has evolved its own genetic architecture for the complex SD system, with specific modifiers and perhaps different Sd genes. The possibility of reconstructing the evolutionary pattern of the SD(Ra) chromosome in the natural Ranna population after the model of Charlesworth and Hartl (1978) and Crow (1979) is considered.     

Reproduction in Cage Populations of a Polymorphism Regularly Observed in the Natural Populations of DROSOPHILA MELANOGASTER in France

Polymorphism for both alleles of a gene ref(2)P, which is a usual trait of French natural populations of Drosophila melanogaster , can be reproduced in experimental conditions. ref(2)P is a gene for resistance to the hereditary, noncontagious Rhabdovirus σ, responsible for CO₂ sensitivity in Drosophila melanogaster . The equilibrium frequencies observed in cages are the same as in the wild, whether σ virus is present or not. The rapid rate of return to these equilibrium frequencies indicates that strong forces, which remain to be determined, are responsible for the maintenance of this polymorphism.