[Induction of MGE 412 transposition in an isogenic strain of Drosophila melanogaster by different doses of ethanol fumes]. / Induktsiia transpozitsii MGE 412 razlichnymi dozami parov étanola v izogennoi linii Drosophila melanogaster.

The effect of treatment of males from an isogenic Drosophila melanogaster strain by limiting doses of ethanol fumes on transpositions of MGE 412 was examined. Validity of the phenomenon of transposition induction was demonstrated. We estimated rates of induced transposition (approximately 10(-2) events per site, per sperm, per generation versus < 10(-3) in control) and showed dose dependence of the rate on the exposure time of the males to ethanol fumes. Experiments with alcohol treatment at limiting doses must end either in death of the individuals or bursts of genetic variability in their progeny. In terms of genetics of an individual, this may mean loss of vital hereditary basis followed by mass degradation of the progeny of the "hard drinkers." In terms of populations genetics, this mode of MGE transposition induction can rapidly create a burst of novel genetic variation, which, apart of great losses, may generate a number of advantageous individuals, i.e., be significant for population survival in new, stressful environments.

[Induction of MGE 412 transposition individually by heat and cold shock in spermatogenesis in Drosophila males]. / Induktsiia transpozitsii MGE 412 razdel'no teplovym i kholodovym shokom v spermatogeneze u samtsov drozofily.

Effects of temperature treatment (heavy heat shock, HHS; heat shock, HS; and cold shock, CS) on the daily productivity of treated males in different spermatogenesis stages have been studied in isogenic line 51 of Drosophila melanogaster. The average productivity was shown to substantially decrease in all cases. The sum of the HS and CS contributions to this decrease was nearly equal to the HHS (the combined HS and CS) contribution, i.e., these contributions were almost additive. The temperature treatments did not kill mature sperm. In the control, mating productivity of day 1 exceeded that of the next day at least by 10-20%. Each day, most sperm in matings was new, i.e., matured during that day. Transposition induction of MGE 412 was studied at four spermatogenesis stages after HS and CS. Both temperature treatments were effective but CS had a more pronounced inducing effect. Most temperature-induced transpositions occurred at stage 3 (meiosis) and 4 (spermiogenesis). The day rates of transpositions at different stages were estimated. After HS at the meiosis stage, lambda = 0.11 events per initial MGE copy per sperm per day of mating, which is approximately equal to the previous estimates after HHS. After CS at the meiosis stage, lambda = 0.51. The transposition hot sites (including the previously known 43B and 97DE as well as a number of new sites) were detected. The lists of transpositions after CS completely included the corresponding lists after HS, which suggests similarity of induction mechanisms underlying CS and HS.

[Prolongation of MGE 412 transposition induction after gamma-irradiation in an isogenic line of Drosophila melanogaster]]. / Prolongatsiia induktsii transpozitsii MGE 412 posle gamma-oblucheniia v izogennoi linii Drosophila melanogaster.

The dose dependence of the rate of gamma-induced transpositions and consequent dynamics of the MGE 412 pattern after gamma-irradiation were investigated in isogenic line 49 in generations F1, F12, F140, and F170. It was shown that the results on dose dependence of transpositions was very similar with the corresponding results of the classic works by Timofeeff-Ressovsky et al. (1935). It is suggested that the transcribed copies of retrotransposon 412 "cure" gamma-radiation-induced double-strand DNA breaks. The phenomenon of prolongation of MGE transposition induction during early generations after treatment was shown. In this period (F1-F12), the maximum transposition rate (lambda approximately equal to 2 x 10(-2) events per MGE copy, per haploid genome, per generation) and the maximum number of heterozygous MGE copies were achieved. In the late generations (F140 and F170), the reduced induction level (lambda approximately 10(-3) was established. In the population of effective size Ne = 2000 individuals, this corresponds to the state when lambda >> 1/4Ne, i.e., when the transposition flow prevails over the MGE copy loss by genetic drift. These data together with some indirect evidence argue for the hypothesis that the spontaneous transposition rate is proportional to the average number of heterozygous MGE copies per diploid genome.

[Response of the MGE 412 pattern to truncation selection of a quantitative trait in an isogenic line of drosophila after severe heat shock (SHS)]. / Otklik patterna MGE 412 na otsekaiushchii otbor kolichestvennogo priznaka v izogennoi linii drozofil posle tiazhelogo teplovogo shoka (TTSh).

Positive and negative selection on the total length of two fragments of an interrupted longitudinal wing vein in an isogenic line of Drosophila melanogaster was accompanied by changes in the genomic localization pattern of MGE 412. Strong truncation selection was conducted in the population of effective size Ne = 160 for 50 generations. Twenty-six out of 35 polymorphic HHS-induced segments of MGE localization behaved as independent copies and markers, whereas 9 segments proved to be selective. The second group included "hot" segments of HHS transposition induction (43B, 97E, etc.). Thus, final consensus patterns of induced MGE transpositions have a random and an adaptive component in generation 50 of positive and negative selection. Selective patterns probably include modifier MGEs, which generate induced genetic regulatory variation of polygenes controlling the selected quantitative trait in the isogenic line after HHS.

Heavy heat shock induced retrotransposon transposition in Drosophila.

The phenomenon of transposition induction by heavy heat shock (HHS) was studied. Males of a Drosophila isogenic line with a mutation in the major gene radius incompletus (ri) were treated by HHS (37 degrees C for 1 h followed by 4 degrees C for 1 h, with the cycle repeated three times) and crossed to untreated females of the same line. The males were crossed 5 d after heat shock, and also 9 d after HHS. Many transpositions were seen in the F1 larvae by in situ hybridization. The rate of induced transposition was at least 2 orders of magnitude greater than that of the control sample, and was estimated to be 0.11 events per transposable element copy per sperm. Two 'hot' subdivisions for transpositions, induced probably during the post-meiotic stage of spermiogenesis, were found: 43B and 97DE. Three-quarters of all transpositions were localized in these positions. In other sites the rates of induced transpositions were (1.3-3.2) x 10(-2) events per occupied segment per sperm, 1 order of magnitude greater than those of the control.

[Population dynamics of the response of the genomic pattern of the mobile genetic element Dm412 in Drosophila on selection for a quantitative trait]. / Populiatsionnaia dinamika otklika genomnogo patterna MGE Dm412 drozofily na otbor po kolichestvennomu priznaku.

In an isogenic line of Drosophila melanogaster carrying the Mendelian mutation radius incompletus, selection for the total length of two segments of the disrupted longitudinal wing vein was conducted. After gamma-irradiation at a dose of 13 Gy, positive and negative truncation selection became highly effective and was completed in 50 generations. The pattern of mobile genetic element Dm412 was almost completely fixed in the course of selection. In the positive direction of selection, fixations of mobile genetic element (MGE) sites exceeded losses; in the negative direction, this relationship was reversed. The number of MGE sites in the pattern increased from 23 to 33 and to 26 in the positive and negative directions, respectively. The mean heterozygosity of MGE sites decreased respectively ten and six times. The dynamics of some sites (6F, 43B, 66A, 69E, and others) corresponded to that expected with an adaptive response to selection. Two out of these sites (43B and 66A) were previously assigned to hot sites of Dm412 transposition induced by heat shock. Fixation and loss of sites continued on average for tens of generations. Four hypotheses describe the relationship between patterns of polygenes and MGE in the context of explanation of the above facts: (1) genetic drift; (2) the linkage of MGE and polygenes without modification of the latter (hitchhiking); (3) the linkage and modifying effect of MGE on polygenes linked with them; (4) the selection of the "champion" pattern of polygenes and a random or adaptive MGE pattern linked with it. Hypotheses 1 and 2 are unlikely, hypothesis 3 is possible in the case of other selection modes, whereas hypothesis 4 seems to be most plausible.

[New evidence for the induction of mobile genetic element transpositions by severe heat shock]. / Novoe podtverzhdenie iavleniia induktsii transpozitsii MGE tiazhelym teplovym shokom.

The induction of retrotransposon 412 transpositions by stress was studied in detail. Males of an isogenic line carrying the radius incompletus (ri) mutation of the Mendelian gene were exposed to heavy heat shock (HHS). The procedure consisted of treatment at 37 degrees C for 1 h and at 4 degrees C for 1 h, with reciprocal changes of developmental temperature 3 times, sequentially; the males were then crossed with untreated females. The same males were crossed both on the fifth and ninth day after the HHS treatment. On the basis of in situ hybridization in 85 F1 larvae, 193 transpositions were identified. After treatment, the transposition rate increased by two orders of magnitude (compared to control) and amounted to 0.11 events per site of the original isogenic line per spermium per generation. Two hot sites (segments) of preferential transposition localization, 43B and 97CD, were detected after the first cross; these sites comprised more than 3/4 of all transpositions. Sperm from the first cross were exposed to HHS during the time period of 120 to 244 h after the appearance of the corresponding germline cells, probably at the stage of spermatid maturation. The overinduction of transpositions was shown to occur in these sites and at this stage. In the remaining sites, after the first cross, and in all sites, after the second cross, the rate of induced transpositions was (1.3-3.1) x 10(-2) events per site per spermium per generation, which is higher than in the control by an order of magnitude. This basic induction level was observed at all stages of spermatogenesis. The induction of transposition by heavy heat shock may be considered established.

[Comparative contribution of different genetic factors in induction of MGE transpositions under isogenization]. / Sravnitel'nye vklady razlichnykh eneticheskikh faktorov v induktsiiu transpozitsii MGE pri izogenizatsii.

Localization patterns of mobile genetic element 412 in polytene chromosomes of larvae from the control (riC) line, the balancer line, the F1 and F2 generations of the isogenization scheme, and 10 final isogenic lines were obtained and compared. The contributions of the recombination transfer of mobile genetic element copies from the balancer line, the outbreeding of control and balancer lines, and the inbreeding of isogenized lines to the rate of transposition were determined and estimated. These constituted < 0.187, < 0.30, and > 0.207 events per initial mobile genetic element copy per isogenized haploid genome per isogenization, respectively. During consecutive steps of isogenization (F1-F2-isogenic lines), the total transposition rate decreased: 2.09, 1.78, and 0.69. This was explained in terms of the existence of large selective and random losses in the variability of mobile genetic elements within the sites of their patterns during isogenization. The existence of a recombination transfer does not change the main conclusions and estimates regarding isogenization-induced transpositions.

[Stress induction of retrotransposon transposition in Drosophila: reality of the phenomenon, characteristic features, possible role in rapid evolution]. / Stressovaia induktsiia transpozitsii retrotranspozonov drozofily: real'nost' iavleniia, kharakternye osobennosti i vozmozhnaia rol' v bystroi évoliutsii.

A polygenic system of expression of the quantitative character radius incompletus was shown to be sensitive to external and physiological stresses: heat shock, gamma-irradiation, isogenization, etc. This stress response involved mobilization of retrotransposons. Heavy heat shock induced transpositions of Dm412 and B104 in three and one isogenic lines, respectively. The induced transposition rate was (2.5-11.0) x 10(-2) per site per sperm per generation, i.e., 1-2 orders of magnitude higher than for spontaneous transpositions. Induction of transpositions by gamma-radiation yielded similar estimates. Recently, induction of transpositions and excisions by isogenization was demonstrated; transposition and excision rates were, respectively, 0.35 and 0.13 per site per sperm per generation, which was 2-3 orders of magnitude higher than in control lines. In all these cases, stress induction of retrotransposon transpositions was mediated by molecular mechanisms of the heat shock system-the general system of cell resistance to external and physiological stress factors. From the viewpoint of evolution, stress induction of transpositions is a powerful factor generating new genetic variation in populations under stressful environmental conditions. Passing through a "bottleneck," a population can rapidly and significantly alter its population norm and become the founder of new, normal forms.