Biological parameters and the segregation distortion (SD) phenomenon in Drosophila melanogaster

The relationship between some biological parameters (mortality, longevity, fertility, fecundity and sex ratio) and segregation of second chromosomes in heterozygous and homozygous SD males has been analyzed. The results obtained in SD/SD+ heterozygous males show: (1) their reduced fertility with respect to that of control males, (2) an alteration in the sex ratio in the SD+ progeny only, and (3) inversely related sex-ratio and segregation distortion values. In SDi/SDj combinations: (1) surprisingly, fertility is intermediate between that of SD/SD+ heterozygous males and that of control males, (2) the segregation ratios of the second chromosomes are normal (0.50), and (3) the sex ratio = 0.50 in both classes of SD progeny. The relationship between mortality (and therefore longevity) and fertility of the different genotypes and fecundity per male indicates that the total productivity of heterozygous males is less than that so far claimed. Indeed, their productivity depends not only on the mechanism of nonformation of the SD+ sperm, but also on their reduced longevity. The k = 0.50 and the high fecundity of SDi/SDj combinations indicated that in these males the SD phenomenon is partially suppressed, the SD chromosomes being insensitive to each other, thus implying that particular Rsp alleles are sensitive to given Sd alleles. The complementation pattern for male fertility of SD homozygous males again supports previous evidence that Sd factors from natural populations are, in effect, different Sd genes.     

Linkage and segregation distortion in Drosophila melanogaster

Segregation distortion is caused by a group of genetic elements in and near the centric heterochromatin of chromosome 2 of Drosophila melanogaster. These elements promote their preferential recovery in heterozygous males by rendering sperm bearing the homologous chromosome dysfunctional. Previous work has shown that numerous Y-autosome translocations are associated with the suppression of the segregation distorter phenotype. The present study examined the effects of translocations between the major autosomes upon the expression of segregation distortion. Autosomal translocations involving either the segregation distorter chromosome or its sensitive homologue had no significant effect upon the expression of segregation distortion. These results argue that linkage arrangement per se may not have a major effect on segregation distortion. The suppression of SD by specific Y-autosomal translocations may be due to the disruption of elements on the Y chromosome that are important for the expression of SD.     

Meiotic segregation of chromosomes in Drosophila melanogaster oocytes

A new technique was developed for a light microscopic analysis of meiosis in Drosophila oocytes. — When the nuclear envelope breaks down the bivalents, till then compressed into a karyosome, separate in early prometaphase. The homologues remain associated by chiasmata except for the fourth chromosomes which are no longer associated. Non-homologous chromosomes regularly segregating from each other in genetic experiments are also unconnected after karyosome disintegration but during metaphase I the fourth chromosomes and the heterologous pairs coorient on the same arc of the spindle and move precociously towards opposite poles. Nondisjunction and other irregularities are not infrequent in oocytes having an uneven number of achiasmatic elements. The fourth chromosomes and the Xs or the large autosomes, when lacking chiasmata, may be involved in non-homologous segregation. In c3G homozygotes all chromosomes appear as univalents in prometaphase. Segregation is variable but the observations suggest the polar distribution of equal numbers of chromosomes in variable combinations irrespective of the size. — Coorientation of univalents may be accounted for if the centromeres, whether homologous or non-homologous, are associated in pairs during early meiotic prophase, and that in the karyosome these pairing relationships are preserved until spindle organization at the onset of prometaphase.     

Meiotic segregation of compound-3 chromosomes in Drosophila

Analysis of crossos between genetically marked stocks of Drosophila melanogaster showed, that the compound-3 chromosomes C(3L)RM and C(3R)RM segregate preterentially in female meiosis, and the following two types of eggs are formed predominantly: C(3L)RM; 0 and 0; C(3R)RM. In male meiosis segregation is almost random and four types of sperm are formed: 1. C(3L)RM; C(3R)RM, 2. 0; 0, 3. C(3L)RM; 0, 4. 0; C(3R)RM. The frequencies of these sperm types vary with the genotypes tested. In the stock C(3L)RM, st; C(3R)RM, p^p, males produce 76.8% type 1 and 2, and 23.2% type 3 and 4; males of the stock C(3L)RM, ri; C(3R)RM, sr form 63.2% type 1 and 2, and 36.8% type 3 and 4.The segregational behaviour of compound-3 chromosomes found in female meiosis is expected according to the distributive pairing hypothesis. In the male however, where there is no distributive pairing, the stock-specific segregation of compound-3 chromosomes may be due to the presence of small homologous chromosome segments near the centromere which influence chromosome distribution.     

Latitudinal variation in wild populations of Drosophila melanogaster: heritabilities and reaction norms

Large amounts of genetic variation for wing length and wing area were demonstrated both within and between Drosophila melanogaster populations along a latitudinal gradient in South America. Wing length and wing area showed a strong positive correlation with latitude in both wild flies and laboratory-raised descendants. Large population differences were observed for heritability and coefficient of variation of these two traits, whereas relatively small population differences were found for development time, viability, pupal mortality, sex ratio and their norms of reaction to four developmental temperatures. No clear-cut latitudinal clines were established for these life-history characters. These results are discussed in the light of Bergmann's Rule and the relation between larval development and adult body size.     

Vital genes in the heterochromatin of chromosomes 2 and 3 of Drosophila melanogaster

Heterochromatin has been traditionally regarded as a genomic wasteland, but in the last three decades extensive genetic and molecular studies have shown that this ubiquitous component of eukaryotic chromosomes may perform important biological functions. In D. melanogaster, about 30 genes that are essential for viability and/or fertility have been mapped to the heterochromatin of the major autosomes. Thus far, the known essential genes exhibit a peculiar molecular organization. They consist of single-copy exons, while their introns are comprised mainly of degenerate transposons. Moreover, about one hundred predicted genes that escaped previous genetic analyses have been associated with the proximal regions of chromosome arms but it remains to be determined how many of these genes are actually located within the heterochromatin. In this overview, we present available data on the mapping, molecular organization and function of known vital genes embedded in the heterochromatin of chromosomes 2 and 3. Repetitive loci, such as Responder and the ABO elements, which are also located in the heterochromatin of chromosome 2, are not discussed here because they have been reviewed in detail elsewhere.     

The role of the chromocenter in nonrandom meiotic segregation of nonhomologous chromosomes in Drosophila melanogaster females

The evidence supporting universal significance of physical links between pericentromeric regions of homologous chromosomes for their bipolar orientation during the first meiotic division is discussed. The pericentromeric chiasmata between homologs or (in the absence of the latter) chromocentric links between nonhomologs, which are preserved until prometaphase, compensate for the disturbed binding between homologous pericentromeric regions in both structural or locus mutants. When the links between nonhomologs are involved, interchromosomal effects on chromosome disjunction and nonhomologous pairing were revealed by the genetic methods. An explanation suggested for genetic events observed during Drosophila meiosis conforms with the original, cytogenetically proved model of the orderly two-ring chromocenter formation and reorganization.     

Phenotypic stability of the P-M system in wild populations of Drosophila melanogaster

The P element appears to be one of the most recently invaded transposons of D. melanogaster. To study the dynamics and long-term fate of P elements in natural populations of D. melanogaster, 472 isofemale lines newly collected from 27 localities of Japan were examined for the P element-associated characteristics (abilities to induce and repress of P element transposition) and genomic P element composition (size classes and their numbers). There was variation in the P element-related phenotypes among local populations, but genomic P composition did not correlate strongly with the phenotype of each line: full-size P and KP elements predominated in their genomes (FP+ KP predominance). Comparison with previous results suggests a stability in the P-M system in local populations over about 15 years. In some populations, phenotypic stability for particularly long times was found: for 30 years or more Q strains predominated in Hikone and Tanushimaru, P or Q strains around Inakadate, and M' or Q strains around Tozukawa. There was no clear evidence of structural destruction underlying functional variation of P elements during this period. These results suggest that the current evolutionary status of P elements in the gene pool of D. melanogaster is not intermediary stage predicted by the original recent invasion hypothesis, and that several other factors such as the position effect play important roles.     

Transient equilibrium at the est-6 locus in wild populations of Drosophila melanogaster

Est-6 polymorphism was studied by extensive sampling of Drosophila melanogaster populations in different years and in different places in the Venetia region of North-East Italy. In many cases an unexpected excess of homozygous genotypes was found. The excess is relatively high in late August and early September and it is followed by a progressive decrease with time toward late autumn, when the population size undergoes a progressive increase.It is postulated that in late August an apparent natural population of D. melanogaster virtually consists of several micropopulations; in each of which geographic isolation induces a certain degree of inbreeding. The excess of homozygous genotypes decreases with time since the progressive increase of the effective population size gives progressively more opportunities to the original micropopulations of coming into contact and of intermizing.     

The transmission of fragmented chromosomes in Drosophila melanogaster.

We investigated the fate of dicentric chromosomes in the mitotic divisions of Drosophila melanogaster. We constructed chromosomes that were not required for viability and that carried P elements with inverted repeats of the target sites (FRTs) for the FLP site-specific recombinase. FLP-mediated unequal sister-chromatid exchange between inverted FRTs produced dicentric chromosomes at a high rate. The fate of the dicentric chromosome was evaluated in the mitotic cells of the male germline. We found that dicentric chromosomes break in mitosis, and the broken fragments can be transmitted. Some of these chromosome fragments exhibit dominant semilethality. Nonlethal fragments were broken at many sites along the chromosome, but the semilethal fragments were all broken near the original site of sister-chromatid fusion, and retained P element sequences near their termini. We discuss the implications of the recovery and behavior of broken chromosomes for checkpoints that detect double-strand break damage and the functions of telomeres in Drosophila.     

Heterosis in crosses between geographically separated populations of Drosophila melanogaster

Summary An experiment was performed to test the hypothesis that the genetic distance between populations estimated from enzyme loci could be used to predict the amount of heterosis that would occur in crosses between these populations. A partial diallel cross using 11 populations of Drosophila melanogaster from the AustralianPacific region and from England was carried out. Heterosis for larval viability, fecundity, cold shock mortality, and an index of these three traits was recorded. When two populations originating from the same location were crossed, no heterosis occurred, but otherwise heterosis was significant for all traits. For larval viability, a similar low level of heterosis occurred in all crosses. For cold shock mortality, the level of heterosis varied widely and fecundity showed a pattern intermediate between these two. The geographic distance between the sites from which populations originated was not correlated with the amount of heterosis in their crosses. There was a tendency for populations from ecologically different environments to show heterosis in crosses. Genetic distance based on ten enzyme loci was correlated with heterosis for cold shock mortality and the combined trait index. These results can be explained by the hypothesis that genes affecting larval viability are subject to strong, uniform selection in all populations, which limits the extent to which gene frequencies can drift apart. However, genes affecting cold shock mortality and the enzyme loci are subject to different selection pressures in different environments. This divergent selection combined with genetic drift causes divergence in gene frequency and heterosis.     

The association between inversion In(3R)Payne and clinally varying traits in Drosophila melanogaster

In Drosophila melanogaster, inversion In(3R)Payne increases in frequency towards low latitudes and has been putatively associated with variation in size and thermal resistance, traits that also vary clinally. To assess the association between size and inversion, we obtained isofemale lines of inverted and standard karyotype of In(3R)Payne from the ends of the Australian D. melanogaster east coast cline. In the northern population, there was a significant association between In(3R)Payne and body size, with standard lines from this population being relatively larger than inverted lines. In contrast, the inversion had no influence on development time or cold resistance. We strengthened our findings further in a separate study with flies from populations from the middle of the cline as well as from the cline ends. These flies were scored for wing size and the presence of In(3R)Payne using a molecular marker. In females, the inversion accounted for around 30% of the size difference between cline ends, while in males the equivalent figure was 60%. Adaptive shifts in size but not in the other traits are therefore likely to have involved genes closely associated with In(3R)Payne. Because the size difference between karyotypes was similar in different populations, there was no evidence for coadaptation within populations.