Scholarly Literature
This is an aggregation of scholarly literature from peer-reviewed journals, bioRxiv , medRxiv and arXiv preprint servers focused on natural and engineered selfish genetic elements and genetic biocontrol. New citations are added as they appear. This content is intended for anyone interested in the scholarly literature on gene drive and genetic biocontrol.
Defects in nuclear transport enhance segregation distortion
4242McElroy, JMM, R. A.; McLean, J. R., Fly, 2:280-290. 2008-01-20 00:00:00.
The equal segregation of chromosomes into gametes is a central tenet of Mendelian genetics. It is this process that is responsible for generating predictable outcomes of crosses, as well as subjecting all chromosomes to the natural selective pressures that exert themselves on a population. Meiotic drive systems violate the principle, and result in a distortion of normal Mendelian ratios. Examples of such meiotic drive systems exist in many organisms, including insects, mammals and plants (reviewed in Lyttle).1 The most clearly understood of these is the Segregation Distorter (SD) system in Drosophila melanogaster. SD was first observed because males carrying one SD chromosome and one normal (SD+) chromosome failed to transmit the normal chromosome to their offspring, causing transmission ratios of near 100% of the SD chromosome.2 T
Transmission ratio distortion in Arabidopsis lyrata: effects of population divergence and the S-locus
4241Leppala, JB, J. S.; Schierup, M. H.; Savolainen, O., Heredity, 100:71-78. 2008-01-19 00:00:00.
We investigated transmission ratio distortion within an Icelandic population of Arabidopsis lyrata using 16 molecular markers unlinked to the S-locus. Transmission ratio distortion was found more often than expected by chance at the gametic level, but not at the genotypic or zygotic level. The gametic effect may be due to meiotic drive or selection acting postmeiotically. At the gametic level, 10.9% of the tests were significant, which is substantially lower than earlier observed in an interpopulation cross (allowing for differences in power)-suggesting that the high level of transmission ratio distortion in the interpopulation cross is due to population divergence. It is also substantially lower than previously observed in intrapopulation crosses at the self-incompatibility locus, suggesting inherent fitness differences of the self-incompatibility alleles. We discuss the possible role of deleterious alleles accumulating at loci under balancing selection. Zygotic effects play a larger role in the interpopulation cross than in the intrapopulation crosses suggesting that Dobzhansky-Muller incompatibilities may be accumulating between the widely diverged populations.
The evolution of sex-independent transmission ratio distortion involving multiple allelic interactions at a single locus in rice
4240Koide, YI, M.; Sawamura, N.; Nishimoto, D.; Matsubara, K.; Onishi, K.; Kanazawa, A.; Sano, Y., Genetics, 180:409-420. 2008-01-18 00:00:00.
Transmission ratio distortion (TRD) is frequently observed in inter-and intraspecific hybrids of plants, leading to a violation of Mendelian inheritance. Sex-independent TRD (siTRD) was detected in a hybrid between Asian cultivated rice and its wild ancestor. Here we examined how siTRD is controlled by the S-6 locus via a mechanism in which the S-6 allele acts as a gamete eliminator, and both the male and female gametes possessing the opposite allele (S-6(a)) are aborted only in heterozygotes (S-6/S-6(a)). Fine mapping revealed that the S-6 locus is located neaar the centromere of chromosome 6. Tescross experiments using near-isogenic lines (NILs) carrying either the S-6 or S-6(a) alleles revealed that Asian rice strains frequently harbor an additional allele (S-6(n)) the presence of which, in heterozygotic states (S-6/S-6(n) and S-6(a)/S-6(n)), does not result in siTRD. A prominent reduction in the nucleotide diversity of S-6 or S-6(a) carries relative to that of S-6(n) carries was detected in the chromosomal region. These results suggest that the two incompatible alleles (S-6/S-6(a)) arose independently from S-6(n) and established genetically discontinuous relationships between limited constituents of the Asian rice population.
X chromosome drive
4239Jaenike, J, Current Biology, 18:R508-R511. 2008-01-17 00:00:00.
In the past 10 years, the world record for the men's 100 meter dash has declined from 9.79 to 9.74 seconds, the detection of such small differences being made possible by sophisticated electronic timing devices. If someone were to run the 100 meters in 9.73999 seconds in the 2008 Olympics, would the timers be sensitive enough to show him to be the world's fastest human? Natural selection could, as differences in fitness of that magnitude (10?6) can be detected in species with large effective population sizes. In this way, natural selection can bring about the evolution of exquisitely well-adapted creatures.; Getting back to the race, imagine that one of the runners somehow manages to get a 50-meter head start. With such an advantage, even an overweight, out of shape, or injured athlete could win the gold. An analogous situation applies in evolution. Mendelian segregation, in which the two alleles carried by a heterozygous individual are passed to equal numbers of gametes, ensures that alleles compete fairly and that they succeed (or fail) on the basis of their effects on survival and fertility. Meiotic drive — the process by which alleles are not represented equally in an individual's gametes — subverts the entire process. In the best documented examples of drive, one allele may be passed on to ?100% of an organism's gametes, equivalent to a runner getting a 50-meter head start in the 100 meter dash. The overrepresentation of such alleles in gametes can more than make up for any associated deficiencies in survival and fertility. As a result, these alleles — and those closely linked to them — can rapidly spread through a population, and thus actually cause a decline in the adaptation of a species to its environment. If such an allele drives all the way to fixation, the species may end up a little less well adapted, but there would be no evidence that meiotic drive was the cause, as drive would no longer be expressed.
A Killer–Rescue system for self-limiting gene drive of anti-pathogen constructs
4238Gould, FH, Yunxin; Legros, Mathieu; Lloyd, Alun L., Proceedings of the Royal Society B: Biological Sciences, 275:2823-2829. 2008-01-16 00:00:00.
A number of genetic mechanisms have been suggested for driving anti-pathogen genes into natural populations. Each of these mechanisms requires complex genetic engineering, and most are theoretically expected to permanently spread throughout the target species' geographical range. In the near term, risk issues and technical limits of molecular methods could delay the development and use of these mechanisms. We propose a gene-drive mechanism that can be self-limiting over time and space, and is simpler to build. This mechanism involves one gene that codes for toxicity (killer) and a second that confers immunity to the toxic effects (rescue). We use population-genetic models to explore cases with one or two independent insertions of the killer gene and one insertion of the rescue gene. We vary the dominance and penetrance of gene action, as well as the magnitude of fitness costs. Even with the fitness costs of 10 per cent for each gene, the proportion of mosquitoes expected to transmit the pathogen decreases below 5 per cent for over 40 generations after one 2?:?1 release (engineered?:?wild) or after four 1?:?2 releases. Both the killer and rescue genes will be lost from the population over time, if the rescue construct has any associated fitness cost. Molecular approaches for constructing strains are discussed.
Broadening the application of evolutionarily based genetic pest management
4237Gould, F, Evolution, 62:500-510. 2008-01-15 00:00:00.
Insect- and tick-vectored diseases such as malaria, dengue fever, and Lyme disease cause human suffering, and current approaches for prevention are not adequate. Invasive plants and animals such as Scotch broom, zebra mussels, and gypsy moths continue to cause environmental damage and economic losses in agriculture and forestry Rodents transmit diseases and cause major pre- and postharvest losses, especially in less affluent countries. Each of these problems might benefit from the developing field of Genetic Pest Management that is conceptually based on principles of evolutionary biology This article briefly describes the history of this field, new molecular tools in this field, and potential applications of those tools. There will be a need for evolutionary biologists to interact with researchers and practitioners in a variety of other fields to determine the most appropriate targets for genetic pest management, the most appropriate methods for specific targets, and the potential of natural selection to diminish the effectiveness of genetic pest management. In addition to producing environmentally sustainable pest management solutions, research efforts in this area could lead to new insights about the evolution of selfish genetic elements in natural systems and will provide students with the opportunity to develop a more sophisticated understanding of the role of evolutionary biology in solving societal problems.
Centromere-associated female meiotic drive entails male fitness costs in Monkeyflowers
4236Fishman, LS, A., Science, 322:1559-1562. 2008-01-14 00:00:00.
Female meiotic drive, in which paired chromosomes compete for access to the egg, is a potentially powerful but rarely documented evolutionary force. In interspecific monkeyflower ( Mimulus) hybrids, a driving M. guttatus allele ( D) exhibits a 98: 2 transmission advantage via female meiosis. We show that extreme interspecific drive is most likely caused by divergence in centromere- associated repeat domains and document cytogenetic and functional polymorphism for drive within a population of M. guttatus. In conspecific crosses, D had a 58: 42 transmission advantage over nondriving alternative alleles. However, individuals homozygous for the driving allele suffered reduced pollen viability. These fitness effects and molecular population genetic data suggest that balancing selection prevents the fixation or loss of D and that selfish chromosomal transmission may affect both individual fitness and population genetic load.
Selective sweeps in a 2-locus model for sex-ratio meiotic drive in Drosophila simulans
4235Derome, NB, E.; Ogereau, D.; Veuille, M.; Montchamp-Moreau, C., Molecular Biology and Evolution, 25:409-416. 2008-01-13 00:00:00.
A way to identify loci subject to positive selection is to detect the signature of selective sweeps in given chromosomal regions. It is revealed by the departure of DNA polymorphism patterns from the neutral equilibrium predicted by coalescent theory. We surveyed DNA sequence variation in a region formerly identified as causing "sex-ratio" meiotic drive in Drosophila simulans. We found evidence that this system evolved by positive selection at 2 neighboring loci, which thus appear to be required simultaneously for meiotic drive to occur. The 2 regions are approximately 150-kb distant, corresponding to a genetic distance of 0.1 cM. The presumably large transmission advantage of chromosomes carrying meiotic drive alleles at both loci has not erased the individual signature of selection at each locus. This chromosome fragment combines a high level of linkage disequilibrium between the 2 critical regions with a high recombination rate. As a result, 2 characteristic traits of selective sweeps-the reduction of variation and the departure from selective neutrality in haplotype tests-show a bimodal pattern. Linkage disequilibrium level indicates that, in the natural population from Madagascar used in this study, the selective sweep may be as recent as 100 years.
The population genetics of using homing endonuclease genes in vector and pest management
4234Deredec, AB, A.; Godfray, H. C. J., Genetics, 179:2013-2026. 2008-01-12 00:00:00.
Homing endonuclease genes (HEGs) encode proteins that in the heterozygous state cause double- strand breaks in the homologous chromosome at the precise position opposite the HFG. If the double-strand break is repaired using the homologous chromosome, the HEG becomes homozygous, and this represents it Powerful genetic drive mechanism that might he used as it tool managing vector or pest populations. HEGs may be used to decrease population fitness to drive down population densities (possibly causing local extinction) or, in disease vectors, to knock out a gene required for pathogen transmission. The relative advantages of HEGs thia target viability or fecundity, that are active in one sex or both, and whose target. is expressed before or after homing are explored. The conditions under which escape mutants arise are also analyzed. A different strategy is to place HEGs on the Y chromosome that cause one, or more breaks on the X chromosome and so disrupt. sex ratio. This strategy can cause severe sex-ratio biases with efficiencies that depend on the details of sperm Competition and zygote mortality. This strategy is probably less susceptible to escape mutants, especially when multiple X shredders are used.
Rapid evolution of yeast centromeres in the absence of drive
4233Bensasson, DZ, M.; Burt, A.; Koufopanou, V., Genetics, 178:2161-2167. 2008-01-11 00:00:00.
To find the most rapidly evolving regions in the yeast genome we compared most of chromosome III from three closely related lineages of the wild yeast Saccharomyces paradoxits. Unexpectedly, the centromere appears to be the fastest-evolving part of the chromosome, evolving even faster than DNA sequences unlikely to be under selective constraint (i.e., synonymous sites after correcting for codon usage bias and remnant transposable elements). Centromeres on other chromosomes also show an elevated rate of nucleotide substitution. Rapid centromere evolution has also been reported for some plants and animals and has been attributed to selection for inclusion in the egg or the ovule at female meiosis. But Saccharomyces yeasts have symmetrical meioses with all four products surviving, thus providing no opportunity for meiotic drive. In addition, yeast centromeres show the high levels of polymorphism expected tinder a neutral model of molecular evolution. We suggest that yeast centromeres suffer an elevated rate of mutation relative to other chromosomal regions and they change through a process of "centromere drift," not drive.
Guidance for contained field trials of vector mosquitoes engineered to contain a gene drive system: Recommendations of a scientific working group
4232Benedict, MDA, P.; Dobson, S.; Gottlieb, M.; Harrington, L.; Higgs, S.; James, A.; James, S.; Knols, B.; Lavery, J.; O'Neill, S.; Scott, T.; Takken, W.; Toure, Y.; Core Working Grp Guidance, Containe, Vector-Borne and Zoonotic Diseases, 8:127-166. 2008-01-10 00:00:00.
The following recommendations represent the response of a group of involved scientists to the need for guidance to aid researchers, government authorities, and community leaders as they consider the design and implementation of field trials to assess the safety and efficacy of genetic strategies for reducing the transmission of diseases by mosquito vectors. Guidance is provided for contained (caged) field trials of genetically-engineered (GE) vector mosquitoes that are fertile and contain novel genetic constructs designed to spread through natural mosquito populations (“gene drive systems”). An effort is made to raise the practical issues that must be considered in advance of such testing, provide generalized recommendations based on currently available information, and identify “points to consider” regarding additional information that may be required in order to make informed decisions on a case-by-case basis. This guidance is intended to clarify the pathway for further assessment of the potential utility of such GE mosquitoes as a tool for the improvement of public health in disease-endemic regions.
Targeting the X chromosome during spermatogenesis induces Y chromosome transmission ratio distortion and early dominant embryo lethality in Anopheles gambiae
4247Windbichler, NP, P. A.; Crisanti, A., PLOS Genetics, 4:1-9. 2008-01-05 00:00:00.
We have exploited the high selectivity of the homing endonuclease I-PpoI for the X-linked Anopheles gambiae 28S ribosomal genes to selectively target X chromosome carrying spermatozoa. Our data demonstrated that in heterozygous males, the expression of I-PpoI in the testes induced a strong bias toward Y chromosome-carrying spermatozoa. Notably, these male mosquitoes also induced complete early dominant embryo lethality in crosses with wild-type females. Morphological and molecular data indicated that all spermatozoa, irrespectively of the inheritance of the transgene, carried a substantial amount of I-PpoI protein that could attack the maternally inherited chromosome X of the embryo. Besides the obvious implications for implementing vector control measures, our data demonstrated the feasibility of generating synthetic sex distorters and revealed the intriguing possibility of manipulating maternally inherited genes using wild-type sperm cells carrying engineered endonucleases.
Meiotic drive and sex determination: molecular and cytological mechanisms of sex ratio adjustment in birds
4246Rutkowska, JB, A. V., Philosophical Transactions of the Royal Society B-Biological Sciences, 363:1675-1686. 2008-01-04 00:00:00.
Differences in relative fitness of male and female offspring across ecological and social environments should favour the evolution of sex-determining mechanisms that enable adjustment of brood sex ratio to the context of breeding. Despite the expectation that genetic sex determination should not produce consistent bias in primary sex ratios, extensive and adaptive modifications of offspring sex ratio in relation to social and physiological conditions during reproduction are often documented. Such discordance emphasizes the need for empirical investigation of the proximate mechanisms for modifying primary sex ratios, and suggests epigenetic effects on sex-determining mechanisms as the most likely candidates. Birds, in particular, are thought to have an unusually direct opportunity to modify offspring sex ratio because avian females are heterogametic and because the sex-determining division in avian meiosis occurs prior to ovulation and fertilization. However, despite evidence of strong epigenetic effects on sex determination in pre-ovulatory avian oocytes, the mechanisms behind such effects remain elusive. Our review of molecular and cytological mechanisms of avian meiosis uncovers a multitude of potential targets for selection on biased segregation of sex chromosomes, which may reflect the diversity of mechanisms and levels on which such selection operates in birds. Our findings indicate that pronounced differences between sex chromosomes in size, shape, size of protein bodies, alignment at the meiotic plate, microtubule attachment and epigenetic markings should commonly produce biased segregation of sex chromosomes as the default state, with secondary evolution of compensatory mechanisms necessary to maintain unbiased meiosis. We suggest that it is the epigenetic effects that modify such compensatory mechanisms that enable context-dependent and precise adjustment of primary sex ratio in birds. Furthermore, we highlight the features of avian meiosis that can be influenced by maternal hormones in response to environmental stimuli and may account for the precise and adaptive patterns of offspring sex ratio adjustment observed in some species.
Sexually antagonistic “Zygotic Drive” of the sex Chromosomes
4245Rice, WRG, S.; Friberg, U., PLOS Genetics, 4:e1000313. 2008-01-03 00:00:00.
Genomic conflict is perplexing because it causes the fitness of a species to decline rather than improve. Many diverse forms of genomic conflict have been identified, but this extant tally may be incomplete. Here, we show that the unusual characteristics of the sex chromosomes can, in principle, lead to a previously unappreciated form of sexual genomic conflict. The phenomenon occurs because there is selection in the heterogametic sex for sex-linked mutations that harm the sex of offspring that does not carry them, whenever there is competition among siblings. This harmful phenotype can be expressed as an antagonistic green-beard effect that is mediated by epigenetic parental effects, parental investment, and/or interactions among siblings. We call this form of genomic conflict sexually antagonistic "zygotic drive'', because it is functionally equivalent to meiotic drive, except that it operates during the zygotic and postzygotic stages of the life cycle rather than the meiotic and gametic stages. A combination of mathematical modeling and a survey of empirical studies is used to show that sexually antagonistic zygotic drive is feasible, likely to be widespread in nature, and that it can promote a genetic "arms race'' between the homo- and heteromorphic sex chromosomes. This new category of genomic conflict has the potential to strongly influence other fundamental evolutionary processes, such as speciation and the degeneration of the Y and W sex chromosomes. It also fosters a new genetic hypothesis for the evolution of enigmatic fitness-reducing traits like the high frequency of spontaneous abortion, sterility, and homosexuality observed in humans.
Selfish genetic elements promote polyandry in a fly
4244Price, TARH, D. J.; Lewis, Z.; Hurst, G. D. D.; Wedell, N., Science, 322:1241-1243. 2008-01-02 00:00:00.
It is unknown why females mate with multiple males when mating is frequently costly and a single copulation often provides enough sperm to fertilize all a female's eggs. One possibility is that remating increases the fitness of offspring, because fertilization success is biased toward the sperm of high- fitness males. We show that female Drosophila pseudoobscura evolved increased remating rates when exposed to the risk of mating with males carrying a deleterious sex ratio- distorting gene that also reduces sperm competitive ability. Because selfish genetic elements that reduce sperm competitive ability are generally associated with low genetic fitness, they may represent a common driver of the evolution of polyandry.
Control of introduced species using Trojan sex chromosomes
11543S. Cotton and C. Wedekind, Trends in Ecology & Evolution, 22:441-443. 2007-07-20 17:25:46.
To control introduced exotic species that have predominantly genetic, but environmentally reversible, sex determination (e.g. many species of fish), Gutierrez and Teem recently modeled the use of carriers of Trojan Y chromosomes - individuals who are phenotypically sex reversed from their genotype. Repeated introduction of YY females into wild populations should produce extreme male-biased sex ratios and eventual elimination of XX females, thus leading to population extinction. Analogous dynamics are expected in systems in which sex determination is influenced by one or a few major genes on autosomes.
Homing endonuclease mediated gene targeting in Anopheles gambiae cells and embryos
4259Windbichler, NP, P. A.; Catteruccia, F.; Ranson, H.; Burt, A.; Crisanti, A., Nucleic Acids Research, 35:5922-5933. 2007-01-17 00:00:00.
Homing endonuclease genes (HEGs) are selfish genetic elements that combine the capability to selectively disrupt specific gene sequences with the ability to rapidly spread from a few individuals to an entire population through homologous recombination repair events. Because of these properties, HEGs are regarded as promising candidates to transfer genetic modifications from engineered laboratory mosquitoes to wild-type populations including Anopheles gambiae the vector of human malaria. Here we show that I-SceI and I-PpoI homing endonucleases cleave their recognition sites with high efficiency inA. gambiae cells and embryos and we demonstrate HEG-induced homologous and non-homologous repair events in a variety of functional assays. We also propose a gene drive system for mosquitoes that is based on our finding that I-PpoI cuts genomic rDNA located on the X chromosome in A. gambiae, which could be used to selectively incapacitate X-carrying spermatozoa thereby imposing a severe male-biased sex ratio.
Viability effects and not meoitic drive cause dramatic departures from Mendelian inheritance for malic enzyme in hybrids of Tigriopus californicus populations
4258Willett, CSB, J. N., Journal of Evolutionary Biology, 20:1196-1205. 2007-01-16 00:00:00.
The genetic basis of post-zygotic reproductive isolation is beginning to be untangled in closely related species, but less is known about the genetics of reproductive isolation between divergent populations. Here, two genes encoding malic enzyme (ME) are isolated from the copepod Tigriopus californicus and their influence upon lowered viability in F-2 hybrids of genetically divergent populations is determined. Each ME gene has diverged extensively between T. californicus populations and one gene shows evidence for a recent selective sweep. Segregation patterns of genotypes for both ME genes in adult F-2 hybrids reveal dramatic departures from Mendelian inheritance, deviations that are not seen in F-2 nauplii implying that selection is acting during development based upon the genotype at these ME genes. These results imply that selection against deleterious gene combinations and not aberrant segregation (i.e. meiotic drive) is likely to lead to dramatic departures from Mendelian inheritance observed in these crosses.
Small steps or giant leaps for male-killers? Phylogenetic constraints to male-killer host shifts
4257Tinsley, MCM, M. E. N., BMC Evolutionary Biology, 7:e1000313. 2007-01-15 00:00:00.
Background: Arthropods are infected by a wide diversity of maternally transmitted microbes. Some of these manipulate host reproduction to facilitate population invasion and persistence. Such parasites transmit vertically on an ecological timescale, but rare horizontal transmission events have permitted colonisation of new species. Here we report the first systematic investigation into the influence of the phylogenetic distance between arthropod species on the potential for reproductive parasite interspecific transfer. Results: We employed a well characterised reproductive parasite, a coccinellid beetle male-killer, and artificially injected the bacterium into a series of novel species. Genetic distances between native and novel hosts were ascertained by sequencing sections of the 16S and 12S mitochondrial rDNA genes. The bacterium colonised host tissues and transmitted vertically in all cases tested. However, whilst transmission efficiency was perfect within the native genus, this was reduced following some transfers of greater phylogenetic distance. The bacterium's ability to distort offspring sex ratios in novel hosts was negatively correlated with the genetic distance of transfers. Male-killing occurred with full penetrance following within-genus transfers; but whilst sex ratio distortion generally occurred, it was incomplete in more distantly related species. Conclusion: This study indicates that the natural interspecific transmission of reproductive parasites might be constrained by their ability to tolerate the physiology or genetics of novel hosts. Our data suggest that horizontal transfers are more likely between closely related species. Successful bacterial transfer across large phylogenetic distances may require rapid adaptive evolution in the new species. This finding has applied relevance regarding selection of suitable bacteria to manipulate insect pest and vector populations by symbiont gene-drive systems.
A sex-ratio meiotic drive system in Drosophila simulans. I: An autosomal suppressor
4256Tao, YM, J. P.; Araripe, L.; Ke, Y.; Hartl, D. L., PLOS Biology, 5:2560-2575. 2007-01-14 00:00:00.
Sex ratio distortion (sex-ratio for short) has been reported in numerous species such as Drosophila, where distortion can readily be detected in experimental crosses, but the molecular mechanisms remain elusive. Here we characterize an autosomal sex-ratio suppressor from D. simulans that we designate as not much yang (nmy, polytene chromosome position 87F3). Nmy suppresses an X-linked sex-ratio distorter, contains a pair of near-perfect inverted repeats of 345 bp, and evidently originated through retrotransposition from the distorter itself. The suppression is likely mediated by sequence homology between the suppressor and distorter. The strength of sex-ratio is greatly enhanced by lower temperature. This temperature sensitivity was used to assign the sex-ratio etiology to the maturation process of the Y-bearing sperm, a hypothesis corroborated by both light microscope observations and ultrastructural studies. It has long been suggested that an X-linked sex-ratio distorter can evolve by exploiting loopholes in the meiotic machinery for its own transmission advantage, which may be offset by other changes in the genome that control the selfish distorter. Data obtained in this study help to understand this evolutionary mechanism in molecular detail and provide insight regarding its evolutionary impact on genomic architecture and speciation.
A sex-ratio meiotic drive system in Drosophila simulans. II: An X-linked distorter
4255Tao, YA, L.; Kingan, S. B.; Ke, Y.; Xiao, H.; Hartl, D. L., PLOS Biology, 5:2576-2588. 2007-01-13 00:00:00.
The evolution of heteromorphic sex chromosomes creates a genetic condition favoring the invasion of sex-ratio meiotic drive elements, resulting in the biased transmission of one sex chromosome over the other, in violation of Mendel's first law. The molecular mechanisms of sex-ratio meiotic drive may therefore help us to understand the evolutionary forces shaping the meiotic behavior of the sex chromosomes. Here we characterize a sex-ratio distorter on the X chromosome (Dox) in Drosophila simulans by genetic and molecular means. Intriguingly, Dox has very limited coding capacity. It evolved from another X-linked gene, which also evolved de nova. Through retrotransposition, Dox also gave rise to an autosomal suppressor, not much yang (Nmy). An RNA interference mechanism seems to be involved in the suppression of the Dox distorter by the Nmy suppressor. Double mutant males of the genotype dox; nmy are normal for both sex-ratio and spermatogenesis. We postulate that recurrent bouts of sex-ratio meiotic drive and its subsequent suppression might underlie several common features observed in the heterogametic sex, including meiotic sex chromosome inactivation and achiasmy.
Neurospora spore killers Sk-2 and Sk-3 suppress meiotic silencing by unpaired DNA
4254Raju, NBM, R. L.; Shiu, P. T., Genetics, 176:43-52. 2007-01-12 00:00:00.
In Neurosphora, crassa., pairing of homologous DNA segments is monitored during meiotic prophase I. Any genes not paired with a homolog, as well as any paired homologs of that gene, are silenced during the sexual phase by a mechanism known as meiotic silencing by unpaired DNA (MSUD). Two genes required for MSUD have been described previously: Sad-1 (Suppressor of ascus dominance), encoding an RNA-directed RNA polymerase, and sad-2, encoding a protein that controls the perinuclear localization of SAD-1. Inactivation of either sad-1 or sad-2 suppresses MSUD. We have now shown that MSUD is also suppressed by either of two Spore killer strains, Sk-2 and Sk-3. These were both known to contain a haplotype segment that behaves as a meiotic drive element in heterozygous crosses of killer X sensitive. Progeny ascospores not carrying the killer element fail to mature and are inviable. Crosses homozygous for either of the killer haplotypes suppress MSUD even though ascospores are not killed. The killer activity maps to the same 30-unit-long region within which recombination is suppressed in killer X sensitive crosses. We suggest that the region contains a suppressor of MSUD.
Meiotic drive by the Y-linked D gene in Aedes aegypti (L.) (Diptera : Culicidae) is associated with disruption of spermiogenesis, leading to premature senescence of spermatozoa
4253Owusu-Daaku, KOB, R. D.; Wood, R. J., Arthropod Structure & Development, 36:233-243. 2007-01-11 00:00:00.
Y chromosome meiotic drive in the mosquito Aedes aegypti, due to the gene D (Distorter) in coupling with M (male determination) [the MD haplotype], is associated with spermiogenic disruption, leading to senescence, at a rate Proportionate to male excess. Spermiogenesis was compared between 'Enhanced Mutant' males with a strongly female-depleted sex ratio (8.9% females), 'Mutant' males showing a lesser degree of distortion (38.3% females), and two controls with normal sex ratios (51.2% and 49.2% females). Sections of testes dissected front mature pupae and adults aged 0, 4, 8, 12 and 16 days were examined by transmission electron microscopy. A difference between Mutant and control spermiogenesis was apparent as early as the pupal stage when some Mutant spermatids showed extra tail elements (axonemes and/or mitochondrial derivatives). The same was true of Enhanced Mutant males but to a more extreme degree. Sperm senescence was evident in Enhanced Mutant testes from day 0 of adult life but in Mutant testes not until day 4. Progressive disorganisation was associated with many loose organelles, and disturbance of the anterior-posterior axis of gamete differentiation within the testis. Degenerative changes of a similar kind in the controls did not become apparent until day 8. These findings are discussed with respect to other characteristics of this meiotic drive system, in terms of a theory of inhibition of reduction division in spermatogenesis associated with fragmentation of the X chromosome, leading to the formation of a restitution nucleus as early as rnetaphase 1. (c) 2007 Elsevier Ltd. All rights reserved.
Identification and characterization of segregation distortion loci along chromosome 5B in tetraploid wheat
4252Kumar, SG, B. S.; Faris, J. D., Molecular Genetics and Genomics, 278:187-196. 2007-01-10 00:00:00.
Segregation distortion genes are widespread in plants and animals and function by their effect on competition among gametes for preferential fertilization. In this study, we evaluated the segregation distortion of molecular markers in multiple reciprocal backcross populations derived from unique cytogenetic stocks involving the durum cultivar Langdon (LDN) and wild emmer accessions that allowed us to study the effects of chromosome 5B in isolation. No segregation distortion of female gametes was observed, but three populations developed to analyze segregation of male gametes had genomic regions containing markers with skewed segregation ratios. One region of distortion was due to preferential transmission of LDN alleles over wild emmer alleles through male gametes. Another region required the presence of LDN 5B chromosomes in the female for preferential fertilization by male gametes harboring LDN alleles indicating that the corresponding genes in the female gametes can govern genes affecting segregation distortion of male gametes. A third region of distortion was the result of preferential transmission of wild emmer alleles over LDN alleles through male gametes. These results indicate the existence of different distorter/meiotic drive elements among different genotypes and show that distortion factors along wheat chromosome 5B differ in chromosomal location as well as underlying mechanisms.
Introducing desirable transgenes into insect populations using Y-linked meiotic drive – A theoretical assessment
4251Huang, YXM, K.; Lloyd, A. L.; Gould, F., Evolution, 61:717-726. 2007-01-09 00:00:00.
The use of genetic drive mechanisms to replace native mosquito genotypes with individuals bearing antipathogen transgenes is a potential strategy for repressing insect transmission of human diseases such as malaria and dengue. Antipathogen transgenes have been developed and tested, but efficient gene drive mechanisms are lacking. Here we theoretically assess the feasibility of introducing antipathogen genes into wild Aedes aegypti populations by using a naturally occurring meiotic drive system. We consider the release of males having both a Winked meiotic drive gene and an X-linked drive-insensitive response allele to which an antipathogen gene is linked. We use mathematical models and computer simulations to determine how the post-introduction dynamics of the antipathogen gene are affected by specific genetic characteristics of the system. The results show that when the natural population is uniformly sensitive to the meiotic drive gene, the antipathogen gene may be driven close to fixation if the fitness costs of the drive gene, the insensitive response allele, and the antipathogen gene are low. However, when the natural population has a small proportion of an X-linked insensitive response allele or an autosomal gene that strongly reduces the effect of the drive gene, the antipathogen gene does not spread if it has an associated fitness cost. Our modeling results provide a theoretical foundation for further experimental tests.
Introducing transgenes into insect populations using combined gene-drive strategies: Modeling and analysis
4250Huang, YXM, K.; Lloyd, A. L.; Gould, F., Insect Biochemistry and Molecular Biology, 37:1054-1063. 2007-01-08 00:00:00.
Engineered underdominance (EU), meiotic drive (MD) and Wolbachia have been proposed as mechanisms for driving anti-pathogen transgenes into natural populations of insect vectors of human diseases. EU can drive transgenes to high and stable frequencies but requires the release of sizeable numbers of engineered insects. MD and Wolbachia either cannot maintain high frequencies of transgenes or lack appropriate expression in critical tissues, but both can drive the transgenes to spread from very low initial frequencies. Here we use mathematical models to assess the utility of combining EU with MD or with Wolbachia. Under some conditions, the combination of EU and MD results in a more efficient transgene-drive strategy than either mechanism alone. This combined strategy could drive the transgenes to stable fixation and would require fewer released insects than EU alone, especially when only males are released. However, a combination of EU and Wolbachia does not work better than EU alone because it requires the release of even more engineered insects. (C) 2007 Elsevier Ltd. All rights reserved.
Chromosome-wide linkage disequilibrium as a consequence of meiotic drive
4249Dyer, KAC, B.; Jaenike, J., Proceedings of the National Academy of Sciences of the United States of America, 104:1587-1592. 2007-01-07 00:00:00.
Adaptation by natural selection proceeds most efficiently when alleles compete solely on the basis of their effects on the survival and reproduction of their carriers. A major condition for this is equal Mendelian segregation, but meiotic drive can short-circuit this process. The evolution of drive often involves multiple, interacting genetic components, together with enhancers and suppressors of drive. Chromosomal inversions that suppress crossing over are also frequently associated with drive systems. This study investigates the effects of these processes on patterns of molecular evolution in the fly Drosophila recens, which is polymorphic for a driving X chromosome (X-D). Whereas standard wild-type chromosomes exhibit high levels of polymorphism at multiple loci, all of the X-D chromosomes effectively carry a single multilocus haplotype that spans at least 130 cM. The X-D is associated with a complex set of inversions that completely suppresses recombination between the standard wild-type chromosome and X-D in heterozygous females, which maintain nonrandom associations among loci that presumably interact epistatically for the expression of drive. The long-term costs of foregoing recombination may be substantial; in combination with its low equilibrium frequency, this makes the XD chromosome susceptible to the accumulation of deleterious mutations. Consistent with this, X-D chromosomes are apparently fixed for a recessive mutation that causes female sterility. Thus, the X-D in D. recens appears to be in chromosome-wide linkage disequilibrium and in the early stages of mutational degradation.
A synthetic maternal-effect selfish genetic element drives population replacement in Drosophila
4248Chen, CHH, H. X.; Ward, C. M.; Su, J. T.; Schaeffer, L. V.; Guo, M.; Hay, B. A., Science, 316:597-600. 2007-01-06 00:00:00.
One proposed strategy for controlling the transmission of insect-borne pathogens uses a drive mechanism to ensure the rapid spread of transgenes conferring disease refractoriness throughout wild populations. Here, we report the creation of maternal-effect selfish genetic elements in Drosophila that drive population replacement and are resistant to recombination-mediated dissociation of drive and disease refractoriness functions. These selfish elements use microRNA-mediated silencing of a maternally expressed gene essential for embryogenesis, which is coupled with early zygotic expression of a rescuing transgene.
First Anopheles arabiensis germline transformation: Toward the development of a transgenic genetic sexing strain
16649H. C. Bossin, J. Thailayil, F. Catteruccia, J. P. Benton, A. Crisanti, M. Q. Benedict, B. G. Knols and A. S. Robinson, American Journal of Tropical Medicine and Hygiene, 75:66-66. 2006-11-01 20:17:39.
The ability to genetically engineer mosquitoes is likely to have major implications for the development and implementation of genetic control systems against mosquito disease vectors such as the Sterile Insect Technique (SIT). In particular, genetically transformed mosquito strains can be created for genetic marking and sexing, two key factors known to influence the effectiveness of SIT programmes. In addition, the removal of biting females before releasing sterile males in the field will be of critical importance as they contribute to disease transmission and reduce the efficiency of the release campaign. Parallel to the creation of a conventional genetic sexing strain (Y-translocation of a resistance marker), our group is undertaking a transgenic approach to the development of an A. arabiensis genetic sexing strain (GSS). The sex separation strategy under investigation relies on the sex-specific properties of the A. gambiae B2tubulin gene regulatory regions. It is hoped this approach will achieve the high sex separation efficiency (above 99%) and strain stability required for safe and efficient male-only SIT releases. We report here the successful development of transgenic A. arabiensis lines using the pPB[DsRed]B2EGFP construct. Wild-type A. arabiensis embryos were injected with a mixture of pPB[DsRed]B2EGFP and helper plasmid phsppBac (700 and 300 ng/µI respectively) following an appropriate protocol. Injections generated several transgenic sexing lines. The effectiveness of the transgenic-based sex-separation procedure, the stability of transgenic mosquito GSS under various (mass-)rearing regimes, as well as the viability and reproductive competitiveness of transgenic sterile males are being assessed.
The Sterile Insect Technique: can established technology beat malaria?
25843M. E. H. Helinski, B. El-Sayed and B. G. J. Knols, Entomologische Berichten, 66:13-20. 2006-06-06 14:48:37.
The Sterile Insect Technique (SIT) is the mass production, sterilisation and subsequent release of sterile insects into a target population in an area-wide integrated approach. The released sterile males mate with wild females; they thus no longer produce offspring and therefore the size of the target population is reduced. Over the years, SIT has proven to be a safe, effective and environmentally sound method to suppress, eliminate or contain pest populations. The International Atomic Energy Agency (IAEA) has a long history of supporting SIT programmes against key insect pests, including fruit flies, tsetse flies and moths. Recently, an integrated five year study to assess the feasibility of SIT to control African malaria mosquitoes has been initiated. In this article, we discuss the components and research requirements for such a feasibility study including sexing, mass production, sterilisation and release methodologies.
Radiation-induced sterility for pupal and adult stages of the malaria mosquito Anopheles arabiensis
25750M. E. H. Helinski, A. G. Parker and B. G. J. Knols, Malaria Journal, 5:10. 2006-05-16 15:23:31.
The optimal dose for male insects to be released in an SIT programme depends on their level of sterility and competitiveness. The use of semi-sterilizing doses to produce more competitive insects is discussed. The most convenient developmental stage for mosquito irradiation on a mass-scale are pupae, but pupal irradiation resulted in a lower insemination rate at the highest dose compared to adult irradiation. On the basis of this study, a suitable dose range that includes semi-sterilizing doses is identified to initiate competitiveness experiments for males irradiated at both developmental stages.
Genetic strategies for controlling mosquito-borne diseases
6191F. Gould, K. Magori and Y. Huang, American Scientist, 94:238. 2006-05-03 19:19:14.
Malaria kills more than a million people each year, primarily children under the age of six. Dengue fever is less deadly, but an outbreak can debilitate millions of people and easily overwhelm doctors and hospitals in tropical cities
The hitchhiking effect of an autosomal meiotic drive gene
4263Chevin, LMH, F., Genetics, 173:1829-1832. 2006-01-21 00:00:00.
Transmission-ratio distortion is a departure from a 1:1 segregation of alleles in the gametes of a heterozygous individual. The so-called driving allele is strongly selected regardless of its effect on the fitness of the carrying individual. It may then have an important impact on neutral polymorphism due to the genetic hitchhiking effect. We study this hitchhiking effect in the case of true meiotic drive in autosomes and show that it is more dependent on the recombination rate than in the classical case of a gene positively selected at the organism level.
Cage trials using an endogenous meiotic drive gene in the mosquito Aedes aegypti to promote population replacement
4262Cha, SJM, A.; Chadee, D. D.; Severson, D. W., American Journal of Tropical Medicine and Hygiene, 74:62-68. 2006-01-20 00:00:00.
Control of arthropod-borne diseases based on population replacement with genetically modified noncompetent vectors has been proposed as a promising alternative to conventional control strategies. Due to likely fitness costs associated with vectors manipulated to carry anti-pathogen effector genes, the effector genes will need to be coupled with a strong drive system to rapidly sweep them into natural populations. Endogenous meiotic drive systems have strong and stable population replacement potential, and have previously been reported in two mosquito species: Aedes aegypti and Culex pipiens. To investigate the influence of an endogenous meiotic drive gene on Ae. aegypti population dynamics, we established three experimental population types that were initiated with 100%, 10%, and 1% male mosquitoes carrying a strong meiotic driver (T37 strain) and 100% sensitive females (RED strain), respectively. Among the 100% and 10% populations, early generations were highly male biased, which reflected the effects of the meiotic driver, and remained more than 60% male by the F-15. A genetic marker tightly linked with the meiotic driver on chromosome 1 showed strong selection for the T37 strain-specific allele. Similar but reduced effects of the meiotic driver were also observed in the 1% populations. These results suggest that release of A e. aegypti males carrying a strong meiotic driver into drive sensitive populations can be an effective tool for population replacement, and provide a foundation for additional studies including both experimental populations and simulations by mathematical modeling.
Isolation and characterization of the RanGAP gene in the mosquito Aedes aegypti
4261Cha, SJL, N.; Debruyn, B.; Severson, D. W., DNA Sequence, 17:223-230. 2006-01-19 00:00:00.
A duplicated 3'-truncated version of RanGAP was previously identified as Segregation distorter (Sd), the meiotic drive gene in Drosophila melanogaster. Here we report the cloning and characterization of the complete gene sequence for the RanGAP homolog from the mosquito Aedes aegypti. The 1995 bp cDNA sequence consists of a 113 bp 5'UTR and 130 bp 3'UTR, and encodes a 583 amino acid protein with high sequence identity with RanGAP homologues of several species. A 20,125 bp genomic DNA sequence contains the complete RanGAP gene, consisting of three exons and two introns. Intron 2 comprises 18,082 bp and contains multiple repetitive elements as well as putative coding regions. The RanGAP locus was mapped to the q-arm of chromosome 2. Because the meiotic drive gene (M-D) in A. aegpti was previously shown to be tightly linked with the sex determining locus on chromosome 1, RanGAP is likely not the M-D gene.
Population dynamics of an endogenous meiotic drive system in Aedes aegypti in Trinidad
4260Cha, SJC, D. D.; Severson, D. W., American Journal of Tropical Medicine and Hygiene, 75:70-77. 2006-01-18 00:00:00.
An endogenous meiotic drive system was previously reported to be segregating in the yellow fever mosquito Aedes aegypti L. (Diptera: Culicidae) population in Trinidad. The meiotic driver (M-D) is tightly linked to the male determining locus and selectively targets sensitive responders linked to the female determining allele, causing fragmentation of female gametes. This results in highly male-biased progeny. The M-D system was initially studied as a genetic tool for population control with limited success, but recently interest has focused on its potential for population replacement. This study examines the distribution and dynamics of the M-D system in Trinidad natural populations. We obtained ovitrap samples from seven geographically distinct regions and determined the allele frequencies of the driver (M-D) and sensitive (m(s)) versus insensitive (m(i)) responders, respectively. Frequencies of the M-D allele ranged from 0.1 to 0.5 and were low at the two major port cities, Port of Spain and San Fernando, suggesting the effects of frequent immigration by non-driving genotypes. Frequencies of the m(i) allele ranged from 0.4 to 0.7, suggesting the effects of strong selection by the driver. In addition, our results show that the driver and sensitivity of responders in the Trinidad populations are highly polymorphic. Continued studies of the dynamics of the M-D system in natural populations are critical to considerations of its use in population replacement.
Fitness effects of X chromosome drive in the stalk-eyed fly, Cyrtodiopsis dalmanni
4271Wilkinson, GSJ, P. M.; Kelleher, E. S.; Muscedere, M. L.; Lorsong, A., Journal of Evolutionary Biology, 19:1851-1860. 2006-01-09 00:00:00.
Sex-ratio (SR) males produce predominantly female progeny because most Y chromosome sperm are rendered nonfunctional. The resulting transmission advantage of X-SR chromosomes should eventually cause population extinction unless segregation distortion is masked by suppressors or balanced by selection. By screening male stalk-eyed flies, Cyrtodiopsis dalmanni, for brood sex ratio we found unique SR alleles at three X-linked microsatellite loci and used them to determine if SR persists as a balanced polymorphism. We found that X-SR/X-ST females produced more offspring than other genotypes and that SR males had lower sperm precedence and exhibited lower fertility when mating eight females in 24 h. Adult survival was independent of SR genotype but positively correlated with eye span. We infer that the SR polymorphism is likely maintained by a combination of weak overdominance for female fecundity and frequency dependent selection acting on male fertility. Our discovery of two SR haplotypes in the same population in a 10-year period further suggests that this SR polymorphism may be evolving rapidly.
Segregation distortion in Arabidopsis C24/Col-0 and Col-0/C24 recombinant inbred line populations is due to reduced fertility caused by epistatic interaction of two loci
4270Torjek, OW-W, H.; Meyer, R. C.; von Korff, M.; Kusterer, B.; Rautengarten, C.; Altmann, T., Theoretical and Applied Genetics, 113:1551-1561. 2006-01-08 00:00:00.
A new large set of reciprocal recombinant inbred lines (RILs) was created between the Arabidopsis accessions Col-0 and C24 for quantitative trait mapping approaches, consisting of 209 Col-0 x C24 and 214 C24 x Col-0 F-7 RI lines. Genotyping was performed using 110 evenly distributed framework single nucleotide polymorphism markers, yielding a genetic map of 425.70 cM, with an average interval of 3.87 cM. Segregation distortion (SD) was observed in several genomic regions during the construction of the genetic map. Linkage disequilibrium analysis revealed an association between a distorted region at the bottom of chromosome V and a non-distorted region on chromosome IV. A detailed analysis of the RILs for these two regions showed that an SD occurred when homozygous Col-0 alleles on chromosome IV coincided with homozygous C24 alleles at the bottom of chromosome V. Using nearly isogenic lines segregating for the distorted region we confirmed that this genotypic composition leads to reduced fertility and fitness.
Gene drive systems for insect disease vectors
4269Sinkins, SPG, F., Nature Reviews Genetics, 7:427-435. 2006-01-07 00:00:00.
The elegant mechanisms by which naturally occurring selfish genetic elements, such as transposable elements, meiotic drive genes, homing endonuclease genes and Wolbachia, spread at the expense of their hosts provide some of the most fascinating and remarkable subjects in evolutionary genetics. These elements also have enormous untapped potential to be used in the control of some of the world's most devastating diseases. Effective gene drive systems for spreading genes that can block the transmission of insect-borne pathogens are much needed. Here we explore the potential of natural gene drive systems and discuss the artificial constructs that could be envisaged for this purpose.
A model describing the effect of sex-reversed YY fish in an established wild population: The use of a Trojan Y chromosome to cause extinction of an introduced exotic species
11545J. B. Gutierrez and J. L. Teem, Journal of Theoretical Biology, 241:333-341. 2006-01-06 17:28:07.
A novel means of inducing extinction of an exotic fish population is proposed using a genetic approach to shift the ratio of male to females within a population. In the proposed strategy, sex-reversed fish containing two Y chromosomes are introduced into a normal fish population. These YY fish result in the production of a disproportionate number of male fish in subsequent generations. Mathematical modeling of the system following introduction of YY fish at a constant rate reveals that female fish decline in numbers over time, leading to eventual extinction of the population. (c) 2005 Elsevier Ltd. All rights reserved.
Male biased sex ratio in the Mediterranean fruit fly Ceratitis capitata, an example of Y-chromosome meiotic drive
4268Shahjahan, RMR, P. A.; Cook, L. M.; Wood, R. J., Heredity, 96:464-470. 2006-01-06 00:00:00.
A case of Y-chromosome meiotic drive is reported in the Mediterranean fruit fly Ceratitis capitata. It arose in an irradiated male and results in excess of males. Male excess is inherited strictly from father to son. A Y-linked factor MP (male producer) is proposed. Higher drive can be selected, but distortion declines rapidly in the absence of selection. Hybrid males from crosses between driving males and nondriving females also show drive but to a reduced extent, suggesting the action of suppressors. Sex ratio distortion is independent of postzygotic mortality, and is not associated with an obvious chromosome arrangement. Spermiogenesis in driving males is characterised by abnormalities in sperm tails and reduced numbers in some sperm cysts, whereas neighbouring cysts of the same MP testis are essentially wild type. The average number of missing sperms plus deformed sperms approximates to the average depression in female recovery among the progenies of siblings, suggesting that most of the missing or abnormal sperms would have given rise to females, that is, they would have been X-bearing. To explain the heterogeneity between neighbouring cysts, a theory is proposed that links it to variation in X-chromosome sensitivity to MP, arising by random suppression of the genetic basis of sensitivity during the six mitotic divisions in the origin of the cyst from its stem cell before meiosis.
The maize Ab 10 meiotic drive system maps to supernumerary sequences in a large complex haplotype
4267Mroczek, RJM, J. R.; Luce, A. C.; Hiatt, E. N.; Dawe, R. K., Genetics, 174:145-154. 2006-01-05 00:00:00.
The meiotic drive system on maize abnormal chromosome 10 (Ab10) is contained within a terminal domain of chromatin that extends the long arm of Ab10 to similar to 1.3 times the size of normal chromosome 10L. Ab10 type I (Ab10-I) does not recombine with normal chromosome 10 (N10) over an similar to 32-cM terminal region of the long arm. Comparative RFLP mapping demonstrates that multiple independent rearrangements are responsible for the current organization of Ab10-I, including a set of nested inversions and at least one long supernumerary segment at the end of the chromosome. Four major meiotic drive functions, i.e., the recombination effect, smd3, 180-bp neocentromere activity, and the distal tip function, all map to the distal supernumerary segment. TR-1-mediated neocentromere activity (the fifth known drive function) is nonessential in the type II variant of Ab10 and maps to a central region that may include a second supernumerary insertion. Both neocentromere activity and the recombination effect behave as dominant gain-of-function mutations, consistent with the view that meiotic drive involves new or alien gene products. These and other data suggest that the Ab10 meiotic drive system was initially acquired from a related species and that a complex haplotype evolved around it.
Organization of the sex-ratio meiotic drive region in Drosophila simulans
4266Montchamp-Moreau, CO, D.; Chaminade, N.; Colard, A.; Aulard, S., Genetics, 174:1365-1371. 2006-01-04 00:00:00.
Sex-ratio meiotic drive is the preferential transmission of the X chromosome by XY males, which occurs in several Drosophila species and results in female-biased progeny. Although the trait has long been known to exist, its molecular basis remains completely unknown. Here we report a fine-mapping experiment designed to characterize the major drive locus on a sex-ratio X chromosome of Drosophila simulans originating from the Seychelles (X-SRG). This primary locus was found to contain two interacting elements at least, both of which are required for drive expression. One of them was genetically tracked to a tandem duplication containing six annotated genes (Trf2, CG32712, CG12125, CG1440, CG12123, org-1), and the other to a candidate region located similar to 110 kb away and spanning seven annotated genes. RT-PCR showed that all but two of these genes were expressed in the testis of both sex-ratio and standard males. In situ hybridization to polytene chromosomes revealed a complete association of the duplication with the sex-ratio trait in random samples of X chromosomes from Madagascar and Reunion.
Sex-ratio meiotic drive in Drosophila simulans: cellular mechanism, candidate genes and evolution
4265Montchamp-Moreau, C, Biochemical Society Transactions, 34:562-565. 2006-01-03 00:00:00.
The sex-ratio trait, reported in a dozen Drosophila species, is a type of naturally occurring meiotic drive in which the driving elements are located on the X chromosome. Typically, as the result of a shortage of Y bearing spermatozoa, males carrying a sex-ratio X chromosome produce a large excess of female offspring. The presence of sex-ratio chromosomes in a species can have considerable evolutionary consequences, because they can affect individual fitness and trigger extended intragenomic conflict. Here, I present the main results of the study performed in Drosophilosimulans. In this species, the loss of Y-bearing spermatozoa is related to the inability of the Y chromosome sister-chromatids to separate properly during meiosis II. Fine genetic mapping has shown that the primary sex-ratio locus on the X chromosome contains two distorter elements acting synergistically, both of which are required for drive expression. one element has been genetically mapped to a tandem duplication. To infer the natural history of the trait, the pattern of DNA sequence polymorphism in the surrounding chromosomal region is being analysed in natural populations of D. simulans harbouring sex-ratio X chromosomes. Initial results have revealed the recent spread of a distorter allele.
Transmission ratio distortion in the human body louse, Pediculus humanus (Insecta : Phthiraptera)
4264McMeniman, CJB, S. C., Heredity, 96:63-68. 2006-01-02 00:00:00.
We studied inheritance at three microsatellite loci in eight F-1 and two F-2 families of the body (clothes) louse of humans, Pediculus humanus. The alleles of heterozygous female-parents were always inherited in a Mendelian fashion in these families. Alleles from heterozygous male-parents, however, were inherited in two different ways: (i) in a Mendelian fashion and (ii) in a non-Mendelian fashion, where males passed to their offspring only one of their two alleles, that is, 100% nonrandom transmission. In male body lice, where there was non-Mendelian inheritance, the paternally inherited set of alleles was eliminated. We interpret this pattern of inheritance as evidence for extreme transmission ratio distortion of paternal alleles in this species.
An Anopheles transgenic sexing strain for vector control
16647F. Catteruccia, J. P. Benton and A. Crisanti, Nature Biotechnology, 23:1414-1417. 2005-10-09 20:04:20.
Here we report on the development of transgenic sexing lines for the mosquito Anopheles stephensi, the principal vector of human malaria in Asia. Male mosquitoes, expressing enhanced green fluorescent protein (EGFP) under the control of the beta 2-tubulin promoter, are identified by their fluorescent gonads in as early as their 3(rd) instar larval stage, and can be efficiently separated from females using both manual methods and automated sorting machines. Importantly, beta 2-EGFP males are not impaired in their mating ability and viable fluorescent spermatozoa are also detected in spermathecae of wild-type females mated with transgenic males. The transgenic mosquito lines described here combine most of the features desired and required for a safe application of transgenic methodologies to malariacontrol programs.
History of the Sterile Insect Technique
6360Klassen, W. and C. F. Curtis, Sterile Insect Technique: Principles and Practice in Area-Wide Integrated Pest Managemen, 2005:3-36.. 2005-03-09 20:56:59.
During the 1930s and 1940s the idea of releasing insects of pest species to introduce sterility (sterile insect technique or SIT) into wild populations, and thus control them, was independently conceived in three extremely diverse intellectual environments. The key researchers were A. S. Serebrovskii at Moscow State University, F. L. Vanderplank at a tsetse field research station in rural Tanganyika (now Tanzania), and E. F. Knipling of the United States Department of Agriculture. Serebrovskii’s work on chromosomal translocations for pest population suppression could not succeed in the catastrophic conditions in the USSR during World War II, after which he died. Vanderplank used hybrid sterility to suppress a tsetse population in a large field experiment, but lacked the resources to develop this method further. Knipling and his team exploited H. J. Muller’s discovery that ionizing radiation can induce dominant lethal mutations, and after World War II this approach was applied on an area-wide basis to eradicate the New World screwworm Cochliomyia hominivorax (Coquerel) in the USA, Mexico, and Central America. Since then very effective programmes integrating the SIT have been mounted against tropical fruit flies, some species of tsetse flies Glossina spp., the pink bollworm Pectinophora gossypiella (Saunders), and the codling moth Cydia pomonella (L.). In non-isolated onion fields in the Netherlands, the onion maggot Delia antiqua (Meigen) has since 1981 been suppressed by the SIT. In the 1970s there was much research conducted on mosquito SIT, which then went into “eclipse”, but now appears to be reviving. Development of the SIT for use against the boll weevil Anthonomus grandis grandis Boheman and the gypsy moth Lymantria dispar (L.) has ended, but it is in progress for two sweetpotato weevil species, Cylas formicarius (F.) and Euscepes postfasciatus (Fairmaire), the false codling moth Cryptophlebia leucotreta (Meyrick), the carob moth Ectomyelois ceratoniae (Zeller), the cactus moth Cactoblastis cactorum (Berg), the Old World screwworm Chrysomya bezziana (Villeneuve), additional Glossina spp., other Anastrepha spp. and Bactrocera spp. fruit flies, and other pest insects.
Maternal transmission ratio distortion at the mouse Om locus results from meiotic drive at the second meiotic division
4283Wu, GMH, L. P.; Han, Z. M.; Gao, S. R.; Latham, K. E.; de Villena, F. P. M.; Sapienza, C., Genetics, 170:327-334. 2005-01-21 00:00:00.
We have observed maternal transmission ratio distortion (TRD) in favor of DDK alleles at the Ovum mutant (Om) locus on mouse chromosome I I among the offspring of (C57BL/6 X DDK) F, females and C57BL/6 males. Although significant lethality occurs in this backcross (similar to 50%), differences in the level of TRD found in recombinant vs. nonrecombinant chromosomes among offspring argue that TRD is due to nonrandom segregation of chromatids at the second meiotic division, i.e., true meiotic drive. We tested this hypothesis directly, by determining the centromere and Om genotypes of individual chromatids in zygote stage embryos. We found similar levels of TRD in favor of DDK alleles at Om in the female pronucleus and TRD in favor of C57BL/6 alleles at Om in the second polar body. In those embryos for which complete dyads have been reconstructed, TRD was present only in those inheriting heteromorphic dyads. These results demonstrate that meiotic drive occurs at MII and that preferential death of one genotypic class of embryo does not play a large role in the TRD.
Population dynamics of transposable elements: Copy number regulation and species invasion requirements
4282Struchiner, CJK, M. G.; Ribeiro, J. M. C., Journal of Biological Systems, 13:455-475. 2005-01-20 00:00:00.
A deterministic population dynamics model of the spread of transposable elements (TE) in sexually reproducing populations is presented. The population is modeled by a three-parameter equation describing host reproductive capacity, population size and the strength of the density dependence, while TE dynamics were modeled based also on three parameters, the maximum ability of the element to copy itself in the absence of regulation (T(0)), the regulatory effect of copy number decreasing transposition (C(0.5)), and the deleterious effect of each new transposition on host fitness (d). The mechanism of transposition control is therefore a function of the number of new TE copies. Our results indicate that non-regulated elements cannot fix in host populations, and that prediction of stable copy number following successful invasion is mainly a function of the combination of T(0) and C(0.5) values. Fitness reduction does not affect the final copy number after successful invasion of the element. Fitness reduction, however, will affect the surface of the {T(0) x C(0.5)} parameter space leading to successful invasion of the TE. Invasion of host populations by eight or more individuals containing elements with appropriate parameters will lead to successful element fixation at any size of the host population. Host population extinction due to the invasion of TE's is observed in a small area of the {T(0) x C(0.5)} parameter space. These results are qualitatively preserved under alternative choices for the shape of the functions defining regulation of transposition and distinct sets of parameters determining host population dynamics.
Evidence of susceptibility and resistance to cryptic X-linked meiotic drive in natural populations of Drosophila melanogaster
4281Reed, FAR, R. G.; Aquadro, C. F., Evolution, 59:1280-1291. 2005-01-19 00:00:00.
There is mounting evidence consistent with a general role of positive selection acting on the Drosophila melanogaster X-chromosome. However, this positive selection need not necessarily arise from forces that are adaptive to the organism. Nonadaptive meiotic drive may exist on the X-chromosome and contribute to forces of selection. Females from a reference D. melanogaster line, containing the X-linked marker white, were crossed to males from 49 isofemale lines established from seven African and five non-African natural populations to detect naturally occurring meiotic drive. Several lines exhibited a departure from expected Mendelian transmission of X-chromosomes to the third generation (F-2) offspring, particularly those from hybrid African male parents. F2 viability was not correlated with skewed chromosomal inheritance. However, a significant difference in viability between cosmopolitan and tropical African crosses was observed. Recombination analysis supports the presence of a male-acting meiotic drive element near the centromeric region of the X-chromosome and putative recessive autosomal drive suppression. There is also evidence of another female-acting drive element linked to white. The possible role meiotic drive may contribute in shaping levels of genetic variation in D. melanogaster, and additional ways to test this hypothesis are discussed.
