X-Y Exchange and the Coevolution of the X and Y rDNA Arrays in Drosophila melanogaster

Genetics ◽  
1987 ◽  
Vol 116 (2) ◽  
pp. 241-251
Author(s):  
M R Gillings ◽  
R Frankham ◽  
J Speirs ◽  
M Whalley
Keyword(s):  
Drosophila Melanogaster ◽  
X Chromosome ◽  
Molecular Genetic ◽  
Nucleolus Organizer ◽  
Rrna Genes ◽  
X Chromosomes ◽  
Normal Morphology ◽  
Y Chromosomes ◽  
Nucleolus Organizers

ABSTRACT The nucleolus organizers on the X and Y chromosomes of Drosophila melanogaster are the sites of 200-250 tandemly repeated genes for ribosomal RNA. As there is no meiotic crossing over in male Drosophila, the X and Y chromosomal rDNA arrays should be evolutionarily independent, and therefore divergent. The rRNAs produced by X and Y are, however, very similar, if not identical. Molecular, genetic and cytological analyses of a series of X chromosome rDNA deletions (bb alleles) showed that they arose by unequal exchange through the nucleolus organizers of the X and Y chromosomes. Three separate exchange events generated compound X·YL chromosomes carrying mainly Y-specific rDNA. This led to the hypothesis that X-Y exchange is responsible for the coevolution of X and Y chromosomal rDNA. We have tested and confirmed several of the predictions of this hypothesis: First, X·YL chromosomes must be found in wild populations. We have found such a chromosome. Second, the X·YL chromosome must lose the YL arm, and/or be at a selective disadvantage to normal X+ chromosomes, to retain the normal morphology of the X chromosome. Six of seventeen sublines founded from homozygous X·YLbb stocks have become fixed for chromosomes with spontaneous loss of part or all of the appended YL. Third, rDNA variants on the X chromosome are expected to be clustered within the X+ nucleolus organizer, recently donated ("Y") forms being proximal, and X-specific forms distal. We present evidence for clustering of rRNA genes containing Type 1 insertions. Consequently, X-Y exchange is probably responsible for the coevolution of X and Y rDNA arrays.

scholarly journals MALE-STERILIZING INTERACTIONS BETWEEN DUPLICATIONS AND DEFICIENCIES FOR PROXIMAL X-CHROMOSOME MATERIAL IN DROSOPHILA MELANOGASTER

Genetics ◽  
1981 ◽  
Vol 99 (1) ◽  
pp. 49-64
Author(s):  
Rezaur Rahman ◽  
Dan L Lindsley
Keyword(s):  
Drosophila Melanogaster ◽  
X Chromosome ◽  
Male Fertility ◽  
Primary Spermatocyte ◽  
Single Factor ◽  
X Chromosomes ◽  
Recessive Lethal ◽  
Lethal Mutations ◽  
Y Chromosomes ◽  
Chromosome Material

ABSTRACT The genetic limits of sixty-four deficiencies in the vicinity of the euchromatic-heterochromatic junction of the X chromosome were mapped with respect to a number of proximal recessive lethal mutations. They were also tested for male fertility in combination with three Y chromosomes carrying different amounts of proximal X-chromosome-derived material (BSYy+, y+Ymal126 and y  +  Ymal  +). All deficiencies that did not include the locus of bb and a few that did were male-fertile in all male-viable Df(1)/Dp(1;Y) combinations. Nineteen bb deficiencies fell into six different classes by virtue of their male-fertility phenotypes when combined with the duplicated Y chromosomes. The six categories of deficiencies are consistent with a formalism that invokes three factors or regions at the base of the X, one distal and two proximal to bb, which bind a substance critical for precocious inactivation of the X chromosome in the primary spermatocyte. Free duplications carrying these regions or factors compete for the substance in such a way that, in the presence of such duplications, proximally deficient X chromosomes are unable to command sufficient substance for proper control of X-chromosome gene activity preparatory to spermatogenesis. We conclude that there is no single factor at the base of the X that is required for the fertility of males whose genotype is otherwise normal.

scholarly journals Clustering of rDNA containing type 1 insertion sequence in the distal nucleolus organiser of Drosophila melanogaster: implications for the evolution of X and Y rDNA arrays

1988 ◽  
Vol 51 (3) ◽  
pp. 209-215 ◽  
Author(s):  
P. R. England ◽  
H. W. Stokes ◽  
R. Frankham
Keyword(s):  
Drosophila Melanogaster ◽  
Gene Clusters ◽  
Distal Portion ◽  
Nucleolus Organizer ◽  
Rrna Genes ◽  
Alternative Hypotheses ◽  
Nucleolus Organiser ◽  
Or Gene ◽  
Exchange Hypothesis

SummaryThe ribosomal RNAs produced by the multigene families on the X and Y chromosomes of Drosophila melanogaster are very similar despite the apparent evolutionary isolation of the X and Y chromosomal rDNA. X–Y exchange through the rDNA is one mechanism that may promote co-evolution of the two gene clusters by transferring Y rDNA copies to the X chromosome. This hypothesis predicts that the proximal rDNA of X chromosomes will be Y-like. Consequently, rDNA variants found only on the X chromosome (such as those interrupted by type 1 insertions) should be significantly clustered in the distal X nucleolus organizer. Proximal and distal portions of the X chromosome nucleolus organizer were separated by recombination between the inverted chromosomes In(1)scv2 (breakpoint in the centre of the rDNA) and In(1)sc4Lsc8R (no rDNA). Molecular analyses of the resulting stocks demonstrated that rRNA genes containing type 1 insertions were predominantly located on the chromosome carrying the distal portion of the X rDNA, thus confirming a prediction of the X–Y exchange hypothesis for the co-evolution of X and Y chromosomal rDNA. Distal clustering is not predicted by the alternative hypotheses of selection or gene conversion.

scholarly journals VARIATIONS IN THE NUMBER OF THE Y CHROMOSOMAL rRNA GENES IN DROSOPHILA HYDEI

Genetics ◽  
1976 ◽  
Vol 82 (1) ◽  
pp. 25-34
Author(s):  
W Kunz
Keyword(s):  
Drosophila Melanogaster ◽  
Y Chromosome ◽  
X Chromosome ◽  
Nucleolus Organizer ◽  
Rrna Genes ◽  
Wild Type ◽  
Drosophila Hydei

ABSTRACT The number of rRNA cistrons is measured by filter saturation hybridization in different stocks of D. hydei, where the wild-type X chromosome has one nucleolus organizer (NO) and the wild-type Y has two separated NO's. (see PDF) females having no X chromosomal NO show an rDNA content exceeding that of a Y chromosome. An even greater increase in the rRNA cistron number is measured in two translocation stocks where the (see PDF) is combined with one half of a Y and, therefore, each stock contains only one of the two Y chromosomal NO's. But when the same Y fragments are brought together with a wild-type X chromosome they lose about one-half of their rRNA cistrons within one generation. Males with two complementary Y fragments but having no X chromosomal NO show a considerably higher rDNA content than the (see PDF) females, although both are equal in respect of their NO number. Consideration is given to related phenomena in Drosophila melanogaster.

scholarly journals SITE-SPECIFIC INSTABILITY IN DROSOPHILA MELANOGASTER: EVIDENCE FOR TRANSPOSITION OF DESTABILIZING ELEMENT

Genetics ◽  
1982 ◽  
Vol 101 (3-4) ◽  
pp. 461-476
Author(s):  
Todd R Laverty ◽  
J K Lim
Keyword(s):  
Drosophila Melanogaster ◽  
X Chromosome ◽  
Lethal Mutation ◽  
Chromosome Rearrangements ◽  
Chromosome Break ◽  
Secondary Mutation ◽  
X Chromosomes ◽  
Site Specific ◽  
Normal Sequence ◽  
Lethal Mutations

ABSTRACT In this study, we show that at least one lethal mutation at the 3F-4A region of the X chromosome can generate an array of chromosome rearrangements, all with one chromosome break in the 3F-4A region. The mutation at 3F-4A (secondary mutation) was detected in an X chromosome carrying a reverse mutation of an unstable lethal mutation, which was mapped in the 6F1-2 doublet (primary mutation). The primary lethal mutation at 6F1-2 had occurred in an unstable chromosome (Uc) described previously (Lim 1979). Prior to reversion, the 6F1-2 mutation had generated an array of chromosome rearrangements, all having one break in the 6F1-2 doublet (Lim 1979, 1980). In the X chromosomes carrying the 3F-4A secondary lethal mutation the 6F1-2 doublet was normal and stable, as was the 3F-4A region in the X chromosome carrying the primary lethal mutation. The disappearance of the instability having a set of genetic properties at one region (6F1-2) accompanied by its appearance elsewhere in the chromosome (3F-4A) implies that a transposition of the destabilizing element took place. The mutant at 3F-4A and other secondary mutants exhibited all but one (reinversion of an inversion to the normal sequence) of the eight properties of the primary lethal mutations. These observations support the view that a transposable destabilizing element is responsible for the hypermutability observed in the unstable chromosome and its derivaties.

scholarly journals Hybrid dysgenesis-induced quantitative variation on the X chromosome of Drosophila melanogaster.

Genetics ◽  
1990 ◽  
Vol 124 (3) ◽  
pp. 627-636
Author(s):  
C Q Lai ◽  
T F Mackay
Keyword(s):  
Drosophila Melanogaster ◽  
X Chromosome ◽  
Quantitative Traits ◽  
Natural Populations ◽  
Quantitative Variation ◽  
Hybrid Dysgenesis ◽  
X Chromosomes ◽  
Bristle Number ◽  
Seven Generations ◽  
Polygenic Variation

Abstract To determine the ability of the P-M hybrid dysgenesis system of Drosophila melanogaster to generate mutations affecting quantitative traits, X chromosome lines were constructed in which replicates of isogenic M and P strain X chromosomes were exposed to a dysgenic cross, a nondysgenic cross, or a control cross, and recovered in common autosomal backgrounds. Mutational heritabilities of abdominal and sternopleural bristle score were in general exceptionally high-of the same magnitude as heritabilities of these traits in natural populations. P strain chromosomes were eight times more mutable than M strain chromosomes, and dysgenic crosses three times more effective than nondysgenic crosses in inducing polygenic variation. However, mutational heritabilities of the bristle traits were appreciable for P strain chromosomes passed through one nondysgenic cross, and for M strain chromosomes backcrossed for seven generations to inbred P strain females, a result consistent with previous observations on mutations affecting quantitative traits arising from nondysgenic crosses. The new variation resulting from one generation of mutagenesis was caused by a few lines with large effects on bristle score, and all mutations reduced bristle number.

Studies on Nucleolar RNA Synthesis in Drosophila Melanogaster

1972 ◽  
Vol 11 (3) ◽  
pp. 689-697
Author(s):  
H. M. KRIDER ◽  
W. PLAUT
Keyword(s):  
Drosophila Melanogaster ◽  
Salivary Glands ◽  
Rna Synthesis ◽  
First Generation ◽  
Nucleolus Organizer ◽  
Fifth Generation ◽  
Autoradiographic Analysis ◽  
Temporally Correlated ◽  
Nucleolar Rna ◽  
Nucleolus Organizers

The influence of conditions resulting in bobbed phenotypes on nucleolar RNA synthesis and the formation of constrictions at nucleolus organizers was examined in larval tissues of Drosophila melanogaster. By means of [3H]uridine incorporation and autoradiographic analysis, a mutation at the bobbed locus was shown to limit the rate of nucleolar RNA synthesis in salivary glands of XO larvae. The formation of constrictions at the organizer sites of a 4-nucleolus-organizer stock was monitored in dividing neuroblast cells stained with acridine orange. Loss of the ribosomal cistrons had been reported by other workers when such stocks were maintained for several generations. In the first generation in our work, constrictions were visible at only 2 of the 4 nucleolus organizers. This situation persisted until the fifth generation, when constrictions appeared at all 4 of the organizer sites. An increase in the rate of nucleolar RNA synthesis in the salivary glands was temporally correlated with the appearance of the extra constrictions. We interpret these observations to mean that 2 of the organizers of the 4-nucleolus-organizer stock were caused to function through the loss of ribosomal RNA cistrons; thus the functional status of an organizer would appear to be subject to control.

Spontaneous formation of compound X chromosomes in Drosophila melanogaster.

Genetics ◽  
1988 ◽  
Vol 119 (1) ◽  
pp. 95-103
Author(s):  
R J Morrison ◽  
J D Raymond ◽  
J R Zunt ◽  
J K Lim ◽  
M J Simmons
Keyword(s):  
Drosophila Melanogaster ◽  
X Chromosome ◽  
The Other ◽  
X Chromosomes ◽  
Multiple Factors ◽  
Spontaneous Formation ◽  
Chromosome Attachment ◽  
Recombination Experiments ◽  
Attachment Process

Abstract Males carrying different X chromosomes were tested for the ability to produce daughters with attached-X chromosomes. This ability is characteristic of males carrying an X chromosome derived from 59b-z, a multiply marked X chromosome, and is especially pronounced in males carrying the unstable 59b-z chromosomes Uc and Uc-lr. Recombination experiments with one of the Uc-lr chromosomes showed that the formation of compound chromosomes depends on two widely separated segments. One of these is proximal to the forked locus and is probably proximal to the carnation locus. This segment may contain the actual site of chromosome attachment. The other essential segment lies between the crossveinless and vermilion loci and may contain multiple factors that influence the attachment process.

scholarly journals MEIOTIC BEHAVIOR OF COMPOUND AUTOSOMES IN FEMALES OF DROSOPHILA MELANOGASTER: INTERCHROMOSOMAL EFFECTS AND THE SOURCE OF SPONTANEOUS NONSEGREGATION

Genetics ◽  
1980 ◽  
Vol 96 (2) ◽  
pp. 455-470
Author(s):  
Hideh Harger ◽  
David G Holm
Keyword(s):  
Drosophila Melanogaster ◽  
X Chromosome ◽  
Inverse Correlation ◽  
Proximal Region ◽  
Crossing Over ◽  
X Chromosomes ◽  
Repulsion Phase ◽  
Base Level ◽  
Exchange Processes ◽  
Spontaneous Frequency

ABSTRACT In females of Drosophila melanogaster, compound autosomes enter the repulsion phase of meiosis uncommitted to a particular segregation pattern because their centromeres are not restricted to a bivalent pairing complex as a consequence of crossing over. Their distribution at anaphase, therefore, is determined by some meiotic property other than exchange pairing, a property that for many years has been associated with the concept of nonhomologous pairing. In the absence of heterologous rearrangements or a free Y chromosome, C(3L) and C(3R) are usually recovered in separate gametes, that is as products of meiotic segregation. Nevertheless, there is a regular, albeit infrequent, recovery of reciprocal meiotic products (the nonsegregational products) that are disomic and nullosomic for compound thirds. The frequency of these exceptions, which is normally between 0.5 and 5.0%, differs for the various strains examined, but remains constant for any given strain. Since previous studies have not uncovered a cause for this base level of nonsegregation, it has been referred to as the spontaneous frequency. In this study, crosses between males and females whose X chromosomes, as well as compound autosomes, are differentially marked reveal a highly significant positive correlation between the frequency of compound-autosome nonsegregation and the frequency of X-chromosome nondisjunction. However, an inverse correlation is found when the frequency of nondisjunction is related to the frequency of crossing over in the proximal region of the X chromosome. These findings have been examined with reference to the distributive pairing and the chromocentral models and interpreted as demonstrating (1) that nonsegregational meiotic events arise primarily as a result of nonhomologous interactions, (2) that forces responsible for the segregation of nonhomologous chromosomes are properties of the chromocentral region, and (3) that these forces come into expression after the exchange processes are complete.

scholarly journals POLYTENIZATION OF THE RIBOSOMAL GENES ON THE X AND Y CHROMOSOMES OF DROSOPHILA MELANOGASTER

Genetics ◽  
1982 ◽  
Vol 100 (3) ◽  
pp. 375-385 ◽  
Author(s):  
Sharyn A Endow
Keyword(s):  
Drosophila Melanogaster ◽  
Southern Blotting ◽  
Dominance Hierarchy ◽  
Ribosomal Genes ◽  
Wild Type ◽  
Y Chromosomes ◽  
Blotting Technique ◽  
X Cells ◽  
Y Cells ◽  
Nucleolus Organizers

ABSTRACT It has previously been shown (Endow and Glover 1979), that polytenization of the ribosomal genes in D. melanogaster Ore-R X/Y cells and in hybrid X/X cells (Endow 1980) involves replication of genes predominantly from one of the cell's two nucleolus organizers. This analysis takes advantage of strain-specific differences in X and Y chromosome rDNA hybridization patterns detected using the Southern blotting technique. In this report, I extend the previous observations by examining polytene rDNA patterns in wild-type and hybrid X/Y cells. A dominance hierarchy for the X and Y chromosomes from three strains of D. melanogaster is presented and possible mechanisms of replicative dominance are discussed.

scholarly journals FITNESS EFFECTS OF EMS-INDUCED MUTATIONS ON THE X CHROMOSOME OF DROSOPHILA MELANOGASTER. II. HEMIZYGOUS FITNESS EFFECTS

Genetics ◽  
1977 ◽  
Vol 87 (4) ◽  
pp. 775-783
Author(s):  
Joyce A Mitchell ◽  
Michael J Simmons
Keyword(s):  
Drosophila Melanogaster ◽  
X Chromosome ◽  
Induced Mutations ◽  
Fitness Effect ◽  
X Chromosomes ◽  
Fitness Effects ◽  
Induced Mutants ◽  
Time Changes

ABSTRACT X chromosomes mutagenized with EMS were tested for their effects on the fitness of hemizygous carriers. The tests were carried out in populations in which treated and untreated X chromosomes segregated from matings between males and attached-X females; the populations were maintained for several generations, during which time changes in the frequencies of the treated and untreated chromosomes were observed. From the rates at which the frequencies changed, the fitness effects of the treated chromosomes were determined. It was found that flies hemizygous for a mutagenized chromosome were 1.7% less fit per mm EMS treatment than those hemizygous for an untreated chromosome. Since the same flies were only 0.5% per m m less viable than their untreated counterparts, the total fitness effect of an X chromosome carrying EMS-induced mutants is three to four times greater than its viability effect. By comparing the heterozygous effect of a mutagenized X chromosome on fitness with the corresponding hemizygous effect, the dominance value for the chromosome is estimated to be about 0.25.

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