Factors Affecting Recognition and Disjunction of Chromosomes at Distributive Pairing in Female DROSOPHILA MELANOGASTER. I. Total Length VS. Arm Length

The behavior of a compound metacentric fourth chromosome (see PDF) has been examined to determine whether arm length or total length is the basis for recognition in distributive pairing. Recognition was judged by the frequency with which the (see PDF) nondisjoined from a series of X duplications (Dp), ranging in size from ≤ 0.3 to > 4 times the size of a single fourth chromosome. Dp, (see PDF) nondisjunction was measured in the absence and in the presence of a competitor, a compound metacentric X. In both situations, total length and not arm length, was found to confer the characteristic recognition property to the (see PDF). A comparison of Dp, (see PDF) nondisjunction curves for both the noncompetitive and competitive situations with analogous Dp, 4 curves previously obtained, show the Dp, (see PDF) curves to be similar in shape to those obtained earlier but displaced one unit to the right, corresponding precisely to the difference in size between the (see PDF) and the 4. Rules governing chromosome recognition for acrocentrics were found completely applicable to metacentrics; disjunctive behavior of metacentrics differed from that of acrocentrics in that two arms conferred on a chromosome the capacity to act as the intermediate of a trivalent when size no longer warranted this attribute. This capacity, itself, is size-dependent.     

Centromeric Effect on the Degree of Nonrandom Disjunction in the Female DROSOPHILA MELANOGASTER

From crosses of females possessing a heteromorphic X-chromosome bivalent, FR1/+, the shorter crossover products were recovered on the average more frequently than the longer reciprocals as predicted by Novitski's (1951) hypothesis of nonrandom disjunction (NRD). The present study stemmed from an unexpected result of these crosses. Evidence for a centromeric effect on NRD was obtained, suggested by a negative correlation between the degree of NRD, c, and the distance between the region of exchange and the centromere as inferred from SET's (single exchange tetrads). Studies on sex chromosome systems other than FR1 confirmed these results. An analogous centromeric effect on preferential segregation had been clearly demonstrated in maize (Kikudome 1958, 1959; Rhoades and Dempsey 1966). However, prior to the present investigation, no such effect of the centromere on NRD in Drosophila had been described, although reanalysis of part of the data of Novitski (1951) and Novitski and Sandler (1956) suggests some evidence of a seriation of increasing c values extending from the most distal region of the chromosome toward the centromere. A suggestion that the effect in Drosophila may be related in some way to the time required for chiasma terminalization, i.e., those terminalizing earlier (distally located crossovers) permitting more random disjunction of the chromatids from the asymmetric dyad and those terminalizing later, progressively less random, is considered and rejected since in general the expected pattern of c values for the various double exchange tetrads (DET's) is inconsistent with that prediction and provides evidence suggesting the possibility of reversals, in part, of c values obtained for SET's.     

Meiosis in Male DROSOPHILA MELANOGASTER I. Isolation and Characterization of Meiotic Mutants Affecting Second Chromosome Disjunction

Two second chromosome, EMS-induced, meiotic mutants which cause an increase in second chromosome nondisjunction are described. The first mutant is recessive and causes an increase in second chromosome nondisjunction in both males and females. It causes no increase in nondisjunction of the sex chromosomes in either sex, nor of the third chromosome in females. No haplo-4-progeny were recovered from either sex. Thus, it appears that this mutant, which is localized to the second chromosome, affects only second chromosome disjunction and acts in both sexes.-The other mutant affects chromosome disjunction in males and has no effect in females. Nondisjunction occurs at the first meiotic division. Sex chromosome disjunction in the presence of this mutant is similar to that of sc(4)sc(8), with an excess of X and nullo-XY sperm relative to Y and XY sperm. In some lines, there is an excess of nullo-2 sperm relative to diplo-2 sperm, which appears to be regulated, in part, by the Y chromosome. A normal Y chromosome causes an increase in nullo-2 sperm, where B(s)Y does not. There is also a high correlation between second and sex chromosome nondisjunction. Nearly half of the second chromosome exceptions are also nondisjunctional for the sex chromosomes. Among the double exceptions, there is an excess of XY nullo-2 and nullo-XY diplo-2 gametes. Meiotic drive, chromosome loss and nonhomologous pairing are considered as possible explanations for the double exceptions.     

Genetics of Factors Affecting the Life History of DROSOPHILA MELANOGASTER. I. Female Productivity

Starting from four basic strains of Drosophila melanogaster, two laboratory strains (cn bw, Tokyo) and two isofemale lines (B-102, B-103) originated from a wild population in Texas, we constructed by repeated backcrosses through females for 20 or more generations a total of 16 strains of all possible combinations between the chromosome sets and cytoplasmic classes. Females from these 16 synthesized strains were then examined for their reproductive performance during their entire life span.---The chromosome set from the cn bw strain was found to associate with the highest female productivity when the age of females was very young, but these females ceased their reproduction and died relatively earlier, resulting in a smaller number of total progeny. The B-102 and B-103 chromosome sets, on the other hand, were associated with the lowest productivity when the females were young, but they lived and continued reproduction longer, resulting in a larger number of total progeny. The Tokyo chromosome set was associated with female productivity intermediate between the other two groups.---Cytoplasmic factors were found to affect the productivity of young females, with the cytoplasm from the cn bw strain associated with the highest productivity. Longevity was not cytoplasmically affected.---There was a clear interaction in female productivity between the Tokyo chromosome set and the cytoplasm from the Texas isofemale lines; the lifetime female productivity, as well as longevity, associated with the Tokyo chromosome set was found to increase considerably when it was substituted into the cytoplasm of the Texas isofemale lines. This chromosome-cytoplasm interaction appeared to be independent of the two systems of hybrid dysgenesis.     

Meiosis in Male DROSOPHILA MELANOGASTER. II. Nonrandom Segregation of Compound-Second Chromosomes

The segregation of compound-second chromosomes in males from two different stocks has been examined. Segregation is random in males from the C(2L)RM4, dp; C(2R)RM4, px stock. Gametes containing only one of the two compound chromosomes comprise 50% of the gametes, and gametes containing either both elements or neither element make up the other 50% of the gametes.——In males from the C(2L)RM, b; C(2R)RM, cn stock, gametes containing either C(2L)RM, b or C(2R)RM, cn make up the majority of the gametes. Gametes containing both chromosomes or neither chromosome account for only 2-3% of the gametes. The nonrandom segregation is due to the C(2R)RM, cn chromosome.——Viability is reduced in flies carrying the C(2R)RM, cn chromosome. This includes larval lethality, delayed development and premature adult lethality. Cytologically, this chromosome contains a large duplication of 2L material, which includes material proximal to region 38 or 39. It is suggested that the viability and segregational properties associated with this chromosome are due to the duplicated 2L material.     

Non-Mendelian Female Sterility in DROSOPHILA MELANOGASTER: Characterization of the Noninducer Chromosomes of Inducer Strains

In relation to non-Mendelian female sterility, Drosophila melanogaster strains can be divided into two main classes, inducer and reactive. The genetic element responsible for the inducer condition (I factor) is chromosomal and may be linked to any inducer-strain chromosome. Each chromosome carrying the I factor (i(+) chromosome) can, when introduced by the paternal gamete into a reactive oocyte, give rise to females (denoted SF) showing more-or-less reduced fertility. As long as i(+) chromosomes are transmitted through heterozygous males with reactive originating chromosomes (r chromosomes), I factor follows Mendelian segregation patterns. In contrast, in heterozygous i(+)/r females, a varying proportion of r chromosomes may irreversibly acquire I factor, independently of classical genetic recombination, by a process called chromosomal contamination. The contaminated reactive chromosomes behave as i(+) chromosomes.-In the present paper, evidence is given that the Luminy inducer strain displays a polymorphism for two kinds of second chromosomes. Some of them are i(+), while others, denoted i(o), are unable to induce any SF sterility when introduced by paternal gametes into reactive oocytes. They are also unable to induce contamination of r chromosomes, but, like r chromosomes, they may be contaminated by i(+) chromosomes in SF or RSF females. The study of the segregation of i(+) and i(o) second chromosomes in the progeny of heterozygous Luminy males and females leads to the conclusion that on chromosome 2 of the Luminy stock the I factor is at a single locus. -X, second and third i(o) chromosomes have been found in several inducer strains. Since these chromosomes can be maintained with i(+) chromosomes in inducer strains in spite of their ability to be contaminated in RSF females, it can be concluded that chromosomal contamination does not take place in females of inducer strains. This implies that contamination occurs only in cells having cytoplasm in a reactive state.     

Variation of Spontaneous Occurrence Rates of Chromosomal Aberrations in the Second Chromosomes of DROSOPHILA MELANOGASTER

After accumulating mutations by the aid of marked inversions, spontaneous occurrence rates of chromosome aberrations were estimated for 1148 chromosome lines that originated from five stem line second chromosomes of Drosophila melanogaster. In chromosome lines originating from three stem chromosomes (CH, PQ, and RT), mutations were accumulated for 7550, 7252, and 7256 chromosome generations, respectively, but no structural change was detected. For the chromosome lines that originated from the other two stem chromosomes, the situation was different: Twenty aberrations (19 paracentric inversions and 1 translocation between the second and the third chromosomes) during 45990 chromosome generations took place in the 500 chromosome lines derived from stem line chromosome (AW), and 92 aberrations (83 paracentric inversions, 6 pericentric inversions, 2 translocations between the second and the third chromosomes and 1 transposition) arose during 45006 chromosome generations in the 500 chromosome lines derived from stem line chromosome (JH). For the AW group the occurrence rate becomes 0.00043 per chromosome per generation for all aberrations and 0.00041 for inversions. For the JH group the corresponding rates are 0.00204 and 0.00198, respectively.-A non-random distribution of the breakpoint on the salivary gland chromosome was observed and the breakpoints were concentrated in the regions 26, 29, 33, and 34.-The cytoplasms and the chromosomes (other than the second chromosomes) were made approximately uniform throughout the experiments. Thus, this remarkable variability in the occurrence rate is most probably due to the differences in one or more chromosomal elements on the original five stem chromosomes. The mutable chromosomes (AW and JH) appear to carry a kind of mutator factor such as hi (Ives 1950).     

Gametic Frequency of Second Chromosomes of the T-007 Type in a Natural Population of DROSOPHILA MELANOGASTER in Texas

The T-007 second chromosome, which was isolated from a natural population of Drosophila melanogaster in south Texas in 1970, is known to show, when made heterozygous in males with a standard cn bw second chromosome, a transmission frequency (k) of 0.35—much lower than the theoretically expected 0.5. Natural populations of this species in Texas contain second chromosomes that, against the standard cn bw genetic background, are associated with distorted transmission frequencies comparable to that of the T-007 chromosome. In order to explain how such chromosomes can persist in natural populations in nontrivial frequencies, it has been postulated that, although such chromosomes show reduced k values when tested under the genetic background of a laboratory stock such as cn bw, they may show, on the average, k values larger than 0.5 under natural genetic backgrounds. If this were true, the frequency of chromosomes of the T-007 type (T chromosomes) should be higher in male than in female gametes under natural genetic backgrounds. The present study was conducted to examine this possibility. The results clearly showed that the frequency of such chromosomes was much higher among male than among female gametes, and that the transmission frequency of this type of chromosome was higher than 0.5 under natural genetic backgrounds. These results suggest that T chromosomes behave like Segregation Distorter (SD) chromosomes in natural populations of this species in Texas. A possible relationship between T-007 and SD chromosomes is suggested.     

An Effect of Centromere Function on the Behavior of Ring-X Chromosomes in DROSOPHILA MELANOGASTER

It is shown that under the influence of an autosomal meiotic mutant that causes abnormalities in meiotic centromere function (mei-S332), ring-X chromosomes are frequently nonrecoverable. Evidence is presented that this nonrecoverability is caused by a failure of sister ring-chromatids to successfully effect an equational separation with resultant dominant lethality. Because mei-S332 results in meiotic abnormalities only after replication has been completed, and because ring chromosomes are normally transmitted with approximately the same efficiency as rod chromosomes, it is suggested that during replication in normal meioses, sister ring-chromatids form mutually interlocked ring complexes that are resolved without genetic consequences at anaphase II, with the resolution owing at least in part to normal centromere function.     

Hybrid Dysgenesis in DROSOPHILA MELANOGASTER: Factors Affecting Chromosomal Contamination in the P-M System

The two interacting components of the P-M system of hybrid dysgenesis are chromosomally associated elements called P factors and a susceptible cytoplasmic state referred to as M cytotype. Previous experiments have indicated that P factors are a family of multiple-copy transposable genetic elements dispersed throughout the genome of P strains but absent in long-established M strains.—Evidence is presented that the sterility and male recombination-inducing potential of P elements may be acquired by X chromosomes, derived from M strains, through nonhomologous association with P strain autosomes, a process referred to as "chromosomal contamination." The frequencies of chromosomal contamination of X chromosomes by P strain autosomes were highly variable and depended on a number of factors. M cytotype (as opposed to P cytotype) was essential for high frequencies of P factor contamination. There were large differences in contamination potential among individual female families, and a weak negative correlation existed between family size and contamination frequency. Chromosomal contamination in the P-M system was shown to be independent of that in the I-R system.—Frequency distributions suggested that the relationship between sterility production and P factor insertion is complex. The majority of P element transpositions, identified by in situ hybridization in one X chromosome, were not associated with gonadal sterility. However, high sterility potential was found to be associated with the presence of at least one P element inserted into the X chromosome. This potential was lost at a rate of about one-sixth per generation in M cytotype but was stabilized in P cytotype. Various hypotheses concerning the relationship between transposition and chromosomal contamination are discussed.     

Magnification of the Ribosomal Genes in Female DROSOPHILA MELANOGASTER

The genetically induced increase in the number of 18S + 28S ribosomal genes known as magnification has been reported to occur in male Drosophila but has not previously been observed in females. We now report that bobbed magnified (bbm) is recovered in progeny of female Drosophila carrying three different X bobbed (Xbb) chromosomes and the helper XYbb chromosome, which is a derivative of the Ybb- chromosome. Using different combinations of bb or bb+ X and Y chromosomes, we show that magnification in females requires both a deficiency in ribosomal genes and the presence of a Y chromosome: X/X females that are rDNA-deficient but do not carry a Y chromosome do not produce bbm; similarly, X/X/Y females that carry a Y chromosome but are not rDNA-deficient do not produce bbm. Bobbed magnified is only recovered from rDNA-deficient X/XY, X/X/Y or XX/Y females. We have also found that females carrying a ring Xbb chromosome together with the XYbb- chromosome do not produce bbm, indicating that ring X chromosomes are inhibited to magnify in females as in males. We postulate that the requirement for a Y chromosome is due to sequences on the Y chromosome that regulate or encode factor(s) required for magnification, or alternatively, affect pairing of the ribosomal genes.--These studies demonstrate that magnification is not limited to males but also occurs in females. Magnification in females is induced by rDNA-deficient conditions and the presence of a Y chromosome, and probably occurs by a mechanism similar to that in males.     

Mitotic Recombination in the Heterochromatin of the Sex Chromosomes of DROSOPHILA MELANOGASTER

The frequency of spontaneous and X-ray-induced mitotic recombination involving the Y chromosome has been studied in individuals with a marked Y chromosome arm and different XY compound chromosomes. The genotypes used include X chromosomes with different amounts of X heterochromatin and either or both arms of the Y chromosome attached to either side of the centromere. Individuals with two Y chromosomes have also been studied. The results show that the bulk of mitotic recombination takes place between homologous regions.