Experimental Population Genetics of Meiotic Drive Systems I. Pseudo-Y Chromosomal Drive as a Means of Eliminating Cage Populations of DROSOPHILA MELANOGASTER

The experimental population genetics of Y-chromosome drive in Drosophila melanogaster is approximated by studying the behavior of T(Y;2),SD lines. These exhibit "pseudo-Y" drive through the effective coupling of the Y chromosome to the second chromosome meiotic drive locus, Segregation distorter (SD). T(Y;2),SD males consequently produce only male offspring. When such lines are allowed to compete against structurally normal SD⁺ flies in population cages, T(Y;2),SD males increase in frequency according to the dynamics of a simple haploid selection model until the cage population is eliminated as a result of a deficiency in the number of adult females. Cage population extinction generally occurs within about seven generations.—Several conclusions can be drawn from these competition cage studies:

(1) Fitness estimates for the T(Y;2),SD lines (relative to SD⁺ ) are generally in the range of 2–4, and these values are corroborated by independent estimates derived from studies of migration-selection equilibrium.

(2) Fitness estimates are unaffected by cage replication, sample time, or the starting frequency of T(Y;2),SD males, indicating that data from diverse cages can be legitimately pooled to give an overall fitness estimate.

(3) Partitioning of the T(Y;2),SD fitnesses into components of viability, fertility, and frequency of alternate segregation (Y + SD from X + SD⁺) suggests that most of the T(Y;2),SD advantage derives from the latter two components. Improvements in the system might involve increasing both the viability and the alternate segregation to increase the total fitness.

While pseudo-Y drive operates quite effectively against laboratory stocks, it is less successful in eliminating wild-type populations which are already segregating for suppressors of SD action. This observation suggests that further studies into the origin and rate of accumulation of suppressors of meiotic drive are needed before an overall assessment can be made of the potential of Y-chromosome drive as a tool for population control.

     

木兰科3个杂交组合及其亲本的核型研究

研究了木兰科（Magaoliaceae）3个杂交组合的亲本和杂交后代的核型。结果表明,云南含笑（Michelia yunnanensis）、灰岩含笑（Michelia calcicola）及其杂交组合A的核型分别为2n=2x=38=36m＋2sm、2n=2x=38=34m＋4sm和2n=2x=38=26m＋12sm;紫花含笑（Michelia crassipes）及其与云南含笑杂交组合C的核型分别为2n=2x=38=32m＋6sm和2n=2x=38=24m＋12sm＋2st;山玉兰（Mognolia delavayi）、广玉兰（Mognolia grondiflora）及其杂交组合U的核型分别为2n=2x=38=28m＋10sm、2n=6x=114=88m＋lOsm＋16st和2n=4x=76=57m＋15sm＋4st。杂交组合的核型与理论核型存在明显的差异,可能是在杂交组合的形成过程中,来自亲本的染色体发生了结构变异。     

The Genetics of DROSOPHILA SUBOBSCURA Populations. IX. Studies on Linkage Disequilibrium in Four Natural Populations

Gametic frequencies were obtained in four natural populations of D. sub-obscura by extracting wild chromosomes and subsequently analyzing them for inversions and allozymes. The genes Lap and Pept-1, both located within the same inversions of chromosome O, were found in striking nonrandom associations with them of the same kind and degree in all populations studied. On the contrary, the gene Acph, also located within the previously mentioned inversions, was found in linkage disequilibrium with them only in two populations and of opposite directions. This is also the case for the genes Est-9 and Hk, both located within chromosome E inversions. While the gene Est-9 was in strong linkage disequilibrium with the inversions, of the same kind and degree in all populations studied, Hk was found to be in linkage equilibrium. Allele frequencies for the 29 genes studied do not show geographical variation except for the genes Lap, Pept-1 and Est-9, the ones found in linkage disequilibria with the geographically varying gene arrangements. Although mechanical or historical explanations for these equilibria cannot be ruled out, these data cannot be explained satisfactorily by the "middle gene explanation," which states that loci displaying such linkage disequilibria are the ones located near the break points of inversions, while the ones displaying linkage equilibria with them are located in the middle of them. There is no evidence for consistent linkage disequilibria between pairs of loci, except for the closely linked genes of the complex locus, Est-9. This would imply, if it is not a peculiarity of the Est-9 complex, that the linkage disequilibria are found only between very closely linked loci or that, for less closely linked genes, the associations are too weak to be detected by the usual samples sizes.     

The Genetics of DROSOPHILA SUBOBSCURA Populations. Xiv. Further Data on Linkage Disequilibria

Data coming from one natural population of D. subobscura, that of Crete, are presented in detail and examined for nonrandom associations of genes and gene arrangements. This population and four others previously studied are reanalyzed for the detection of higher than first-order interactions. Only first-order interactions are important and statistically significant, especially those concerning genes and inversions in which these genes are included. The paucity of linkage disequilibria detected is remarkable, and we argue that it does not depend on the methods of study, rather it is genuine. We further argue that most of the disequilibria detected are probably due to mechanisms based on epistatic selection.     

The Population Genetics of Parthenogenetic Strains of DROSOPHILA MERCATORUM. II. the Capacity for Parthenogenesis in a Natural, Bisexual Population

Drosophila mercatorum is a bisexual species, but certain strains are capable of parthenogenetic reproduction in the laboratory. We investigated the parthenogenetic capacity of the virgin daughters of females captured from a natural, bisexual population in Hawaii. An isozyme survey indicated the natural population is polymorphic at about 50% of its loci, and its individuals heterozygous at 18% of their loci. The predominant mode of parthogenesis in D. mercatorum causes homozygosity for all loci in a single generation. Despite this radical change in genetic state, 23% of the virgin female lines produced adult parthenogenetic progeny, and 16% produced parthenogenetic progeny themselves capable of parthenogenetic reproduction. The parthenogenetic rats as measured by the number of parthenogenetic progeny themselves capable of parthenogenesis divided by the number of eggs laid is arougn 10(-5) for the virgin female lines. We argue that one of the major reasons for this low rate is that very few of the impaternate zygotes have a genotype that can survive and reproduce under the genetic conditions imposed by parthenogenetic reproduction. This intense selective bottleneck can be passed in a single generation if enough unfertilized eggs are laid, and once passed is accompanied by a large (perhaps a thousandfold) increase in the rate of parthenogenesis and by modifications in many phenotypic traits such as morphology and behavior.     

The Molecular through Ecological Genetics of Abnormal Abdomen in DROSOPHILA MERCATORUM. I. Basic Genetics

The abnormal abdomen syndrome (aa) in Drosophila mercatorum is characterized by the persistence of juvenilized cuticle on the adult abdomen. The aa phenotype is shown to depend on at least two X-linked genetic elements that are about one map unit apart near the centromeric end of the X chromosome. These two genetic elements are necessary for aa expression; one behaves as a dominant element and the other as a recessive. Overlaying these genetic studies upon molecular work reported elsewhere, it is argued that the dominant element is the presence of a 5 kb insertion in a majority of the X-linked repeats coding for the 28S ribosomal RNA. The recessive element appears to be a locus controlling differential replication of noninserted over inserted 28S genes during polytenization. The aa syndrome requires both the presence of the inserted repeats and the failure to preferentially amplify noninserted repeats. Given the necessary X-linked elements for aa, a variety of modifiers are revealed. First, aa expression in males is Y-linked, apparently corresponding to a deletion of the 18S/28S rDNA gene cluster normally found on the Y. Moreover, all major autosomes can modify the penetrance of aa.     

Allozyme Genotype-Environment Relationships. I. Variation in Natural Populations of DROSOPHILA PACHEA

Remarkable geographic and seasonal uniformities in gene and genotype frequencies were found for four enzyme loci. Principal component analysis revealed two patterns of allele frequency changes in three of the allozyme loci (Est-b, Est-c and Acph) and one pattern in the Mdh-d locus. These patterns accounted for 90% to 100% of the variability at these loci. Significant positive correlations were obtained between several of the components and the variability in the non-saponifiable and alkaloid fractions of the senita cactus host plant. The genetic variance within a single locality (Guaymas, Mexico) was as great as the variance between all localities sampled. This is interpreted as a local founder effect. The temporary nature of the rotting cactus breeding site, the delayed maturation and the differential maturation rate of females and males are three factors proposed to cause forced outbreeding and counter the local founder effect. A combination of selection resulting from a direct interaction of the genotype and the environment, in addition to migration, is proposed to maintain the uniform gene frequencies.     

Studies in the Dynamics of Genetically Variable Populations. I. Frequency- and Density-Dependent Selection in Experimental Populations of DROSOPHILA MELANOGASTER

Changes in genetic composition and in fitness of karyotypes in synthetic populations of Drosophila melanogaster carrying fourth chromosomes marked with recessive mutants in repulsion were measured as functions of initial population composition and density. There was no detectable influence of initial population density on the magnitude of chromosome frequency change or on adaptive values of karyotypes, but both measures proved sensitive to population composition. That fitness be measured zygote-to-zygote is argued to be of dubious necessity in all contexts except demonstration of frequency-dependent selection. Some prior reports cited as evidence of the operation of density-dependent selection, like the present study, contain no evidence that the rate of natural selection is influenced by population crowding. Reports that do demonstrate density-dependent selection suggest that the relationship between fitness and crowding differs from that assumed in most theoretical treatments. Further work is needed to elucidate the interaction between fitness, population crowding, and population composition.     

The Genetics of DROSOPHILA SUBOBSCURA Populations. V. a Study of Linkage Disequilibrium in Natural Populations between Genes and Inversions of the E Chromosome

The genetics of Hk and Est-9 complex gene have been studied in Drosophila subobscura. While Hk alleles mendelize normally, Est-9 is a complex locus consisting of several very closely linked genes with active and silent alleles. Both genes are located on chromosome E; a detailed genetic map was constructed with the help of visible markers and inversions. Both Hk and Est-9 are included in or are very near to inversions of the E chromosome. While Hk does not show linkage disequilibrium either with Est-9 or inversions, Est-9 does show disequilibrium in two ways: both with inversions and between different Est-9 genes. All natural populations studied show the same kind of association between Est-9 gene combinations and inversions. It is argued that these results are better explained by selection than by neutrality.