Hypothesis Of Genetic Drift Research Articles

Origin of Symphyotrichum anticostense (Asteraceae, Astereae), an endemic, high polyploid species of the Gulf of St. Lawrence region, based on morphological and nrDNA evidence

Symphyotrichum anticostense (Fernald) G.L.Nesom (Asteraceae, Astereae), a rare endemic of the Gulf of St. Lawrence region, is a high allopolyploid (2n = 10x = 80). It has been hypothesized to be derived from the hybrid between tetraploid (2n = 4x = 32) individuals of Symphyotrichum boreale (Torr. & A.Gray) Á.Löve & D.Löve and the hexaploid (2n = 6x = 48) Symphyotrichum novi-belgii (L.) G.L.Nesom. We investigated this hypothesis using morphological and molecular internal transcribed spacer sequence data, and we tried to determine the potentially geographic origins of the taxon. Univariate morphological analyses show that 67.5% of the S. anticostense characters are parent-like, 43.5% from S. novi-belgii and 13% from S. boreale, the remainder not differing statistically from either parent; 23.5% are intermediate and 9% transgressive. Multivariate analyses show that S. anticostense is intermediate between its putative parents. The molecular results support the morphological data, but due to the insufficient resolution among ribotypes on the tree, a more rapidly evolving marker will be needed to more reliably ascertain the origin of S. anticostense. Besides the hypothesis of genetic drift and allele fixation following long-distance dispersals, at least three independent geographic origins may be suggested for S. anticostense; Anticosti Island, Lake St. John, and Gaspé Peninsula-New Brunswick-Maine.

Common evolutionary patterns in the human nasal region across a worldwide sample.

Variation in the external nasal region among human populations has long been proposed in the literature to reflect adaptations to facilitate thermoregulation, air conditioning, and moisture retention in local climates and environments. More specifically, adaptations in populations living in colder climates have often been assumed due to correlational relationships found between variation in the nasal region and climatic variables. Here, we test this hypothesis by applying a quantitative genetics approach based on the Lande model to assess whether variation in the nasal region can be explained by random neutral processes (e.g., genetic drift) or if non-random forces (i.e., adaptation) have contributed significantly to its diversity. A mixed-sex sample representing 28 population groups from Howells' craniometric dataset were analyzed (n=2504). Twenty standard measurements were chosen to reflect the external skeletal morphology of the nasal region. We apply statistical tests developed from evolutionary quantitative genetics theory to analyze patterns of within- and between-population divergence under a null hypothesis of genetic drift. This study finds a rejection of genetic drift in all analyses, across tests that involve all 28 populations, exclusively cold-climate populations, and with cold-climate populations excluded, indicating that non-random evolutionary forces have contributed significantly to variation in the nasal region overall. These results show that nasal region adaptation is not exclusive to cold-climate populations, which have often been implicated in the literature to drive nasal variation, instead suggesting that the propensity for adaptation in the nasal region is shared among all human populations.

First analysis of ancient mtDNA genetic diversity in Northern coast of Argentinean Patagonia

In the past decades, archaeological research advances in different Patagonian sectors have contributed to the knowledge of lifestyle and evolution of pre-Columbian populations in these regions. The San Matías Gulf archaeological record (Río Negro province, Northern Patagonia, Argentina) shows that humans occupied the area from Middle Holocene to contact period, where palaeodietary and technological changes are evident from around 1700 14C BP. Nevertheless, few studies have been done on biological origin and evolution of these populations. For this reason and in order to investigate mitochondrial genetic variability in comparison with other ancient and contemporary Patagonian populations, we analyzed ancient DNA on human skeletal remains belonging to different chronologies of this area. We found a prevalence of founding lineages C1, D1 (D1g and D1j subclades) and D4h3a, low frequency of A2 and absence of B2. This supports the hypothesis of genetic drift acting on populations of the Southern Cone of South America. Individuals analyzed present similar haplotypes, indicating a possible temporal and regional continuity or a common population origin. Those lineages are also present in Tehuelche and Mapuche contemporary populations. However, B2 is highly represented in these extant groups possibly as a consequence of recent migrations related with Araucanization process.

Extending Genome-Wide Association Study Results to Test Classic Anthropological Hypotheses: Human Third Molar Agenesis and the "Probable Mutation Effect".

A genome-wide association study (GWAS) identifies regions of the genome that likely affect the variable state of a phenotype of interest. These regions can then be studied with population genetic methods to make inferences about the evolutionary history of the trait. There are increasing opportunities to use GWAS results-even from clinically motivated studies-for tests of classic anthropological hypotheses. One such example, presented here as a case study for this approach, involves tooth development variation related to dental crowding. Specifically, more than 10% of humans fail to develop one or more permanent third molars (M3 agenesis). M3 presence/absence variation within human populations has a significant genetic component (heritability estimate h 2 = 0.47). The evolutionary significance of M3 agenesis has a long history of anthropological speculation. First, the modern frequency of M3 agenesis could reflect a relaxation of selection pressure to retain larger and more teeth following the origins of cooking and other food-softening behaviors (i.e., the genetic drift hypothesis or, classically, the "probable mutation effect"). Alternatively, commensurate with increasing hominin brain size and facial shortening, M3 agenesis may have conferred an adaptive fitness advantage if it reduced the risk of M3 impaction and potential health complications (i.e., the positive selection hypothesis). A recent GWAS identified 70 genetic loci that may play a role in human M3 presence/absence variation. To begin evaluating the contrasting evolutionary scenarios for M3 agenesis, we used the integrated haplotype score (iHS) statistic to test whether those 70 genetic regions are enriched for genomic signatures of recent positive selection. None of our findings are inconsistent with the null hypothesis of genetic drift to explain the high prevalence of human M3 agenesis. This result might suggest that M3 impaction rates for modern humans do not accurately retrodict those of the preagricultural past. Alternatively, the absence of support for the positive selection hypothesis could reflect a lack of power; this analysis should be repeated following the completion of more comprehensive GWAS analyses for human M3 agenesis.

Genetic and alarm call diversity across scattered populations of speckled ground squirrels (Spermophilus suslicus)

Alarm calls of ground squirrels are innate signals, showing substantial geographical variation across populations without the masking effects of sex and age- related variation. This makes them a convenient model for studying population genetic effects on the evolution of alarm communication. We compared data on the alarm call structure and the mitochondrial DNA (mtDNA) complete control region (C-region) (998–1002bp) polymorphism in the same 90 individual speckled ground squirrels (Spermophilus suslicus) across 6 populations (15 individuals per population), separated by distances from 12 to 1274km. We calculated acoustic distances between each pair of populations based on acoustic variables of alarm calls, averaged for each individual using Euclidean distances of population centroids, in the space of canonical axes of discriminant function analysis. Genetic distances ranged of 0–1.1% within populations and of 0.5–4.9% between populations. Prominent differences were found between eastern and western populations separated by the Dnieper River. Both genetic and acoustic distances showed a significant positive correlation with geographical among populations. Positive correlation between acoustic and genetic distances did not reach significance. These results support effects of ecological selection on the alarm call variables rather than the genetic drift hypothesis. In addition, these results support the current taxonomic separation between subspecies of speckled ground squirrels differing in diploid chromosome sets 2n=34 (Spermophilus suslicus guttatus) and 2n=36 (Spermophilus suslicus odessanus).

Identification of genomic loci associated with crown rust resistance in perennial ryegrass (Lolium perenne L.) divergently selected populations

The inheritance of crown rust resistance in perennial ryegrass is complex with both major and minor quantitative trait loci (QTL) being detected on all seven linkage groups. However, QTL mapping populations have only few segregating alleles, limiting the transferability of results to other materials. In this study, a synthetic population was developed from four crown rust resistant and susceptible parents as starting material for a divergent selection experiment of crown rust resistance to be closer to practice in plant breeding programs, and to identify genome regions relevant across a broader range of genotypes. Following three cycles of directional selection, perennial ryegrass populations were produced with a two-fold difference in average rust resistance. Divergently selected populations were genotyped at 7 resistance gene analog-derived expressed sequence tag (RGA-derived EST) as well as 15 simple sequence repeat (SSR) loci. A test for selective neutrality (Waples test), which tests the hypothesis of genetic drift versus selection, identified significant differences in allele frequencies for 7 loci (32%). The selection effect was bidirectional with the same loci showing significant response in both positively and negatively selected populations. A region under selection represented by markers LpSSR006 and EST13 on linkage group (LG) 4 was further confirmed by colocation with two separate QTL for crown rust resistance in a VrnA, a two-way pseudo-testcross mapping population. This suggests suitability of alleles identified for introgression into perennial ryegrass germplasm, where quantitative resistance to crown rust is desired.

Genomic analysis of bovine herpesvirus type 4 (BoHV-4) from Argentina: High genetic variability and novel phylogenetic groups

Bovine herpesvirus 4 (BoHV-4) is a γ-herpesvirus that has been isolated both from apparently healthy animals and from cattle with a variety of clinical signs, including post-partum endometritis and abortion. BoHV-4 causes either a persistent or a latent infection in cells of the monocyte/macrophage lineage. Two groups of BoVH-4 strains have been defined based on their restriction patterns: the Movar-like strains (European prototype) and the DN 599-like strains (American prototype). The purpose of the present study was to genetically characterize wild type BoHV-4 strains isolated from vaginal discharges of aborted cows in Argentina. The virus was identified by isolation and nested PCR in all vaginal discharge samples from aborted cows, either as a sole agent or in association with other pathogens. Restriction enzyme profiling and phylogenetic analysis demonstrated that there is a high genetic variability among the studied field isolates. The existence of three groups of strains, which were designated as genotypes 1, 2 and 3, is described. Genotypes 1 and 2 possibly correspond to the Movar-like and DN 599-like groups, respectively, whereas Genotype 3 corresponds to a novel group. Two viral strains did not cluster into any of these three groups, indicating that other genotypes could be circulating in Argentina. These results suggest a complex epidemiological background for the Argentinean BoHV-4 strains, probably influenced by independent events of genetic drift. This hypothesis cannot be rejected based on the available data. However, there is no direct evidence supporting this possibility. Thus, it seems speculative to suggest that interspecific jumps are responsible for the observed phylogenetic grouping. On the other hand, our analyses suggest a geographical structure for the observed viral genotypes, since genotypes 1 and 2 included the European (Movar-like) and American (DN599-like) reference strains, respectively. Geographic dispersion is known to be a driver of herpes viruses diversification, and independent evolution in geographical isolated places ensures the emergence of particular mutations in each location due to genetic drift (Compans, 2007; Zong et al., 1999). Therefore, at this point, the genetic drift hypothesis is the one that requires less ad-hoc considerations and thus, to our understanding, is the one that fits to the findings from this study. The involvement of this genetic variability in the detection and pathogenesis of BoHV-4 remains to be investigated.

Genetic Analysis of Six Communities of Mbyá-Guaraní Inhabiting Northeastern Argentina by Means of Nuclear and Mitochondrial Polymorphic Markers

Autosomal STRs, Y-chromosome markers, and mitochondrial DNA sequences were investigated in six Mbyá-Guaraní villages (Fortín M'Bororé, Yryapu, Tabay, Kaaguy Poty, Jejy, and Yaboti), all of them settled within the province of Misiones, northeastern Argentina. One hundred twenty-one unrelated individuals were analyzed. The study involved typing fifteen autosomal STRs, nine Y-chromosome STRs, and four biallele loci in the nonrecombinant region of the Y chromosome, sequencing the mtDNA of hypervariable regions I and II, and detecting the 9-bp ins/del in region V of mtDNA. All autosomal STRs were in Hardy-Weinberg equilibrium. The four major native American mtDNA haplogroups were represented in the sample. Haplogroups A2 and D1 exhibited the highest frequencies (40.5% and 36.0%, respectively), and haplogroups B2 and C1 appeared to be less frequent (17.5% and 6.0%, respectively). The native American haplogroup Q1a3a was observed in a relevant proportion (88.8%). In addition, a nine-STR Y-chromosome haplo-type (DYS19*13, DYS389I*14, DYS389II*31, DYS390*24, DYS391*11, DYS392*14, DYS393*11, DYS385A*14, DYS385B*16) exhibited a frequency of more than 36%. Our results indicate that the analyzed Argentinean Guaraní individuals are genetically more closely related to Guaraní from Brazil [genetic distance (Δµ)(2) = 0.48] than to other related tribes that are geographically closer. Statistical approaches based on autosomal data do not support the hypothesis of genetic drift previously proposed; however, this apparent discrepancy might be due to the lack of sensitivity of the autosomal markers used here.

Genetic Analysis of Six Communities of Mby�-Guaran� Inhabiting Northeastern Argentina by Means of Nuclear and Mitochondrial Polymorphic Markers

Autosomal STRs, Y-chromosome markers, and mitochondrial DNA sequences were investigated in six Mbyá-Guaraní villages (Fortín M'Bororé, Yryapu, Tabay, Kaaguy Poty, Jejy, and Yaboti), all of them settled within the province of Misiones, northeastern Argentina. One hundred twenty-one unrelated individuals were analyzed. The study involved typing fifteen autosomal STRs, nine Y-chromosome STRs, and four biallele loci in the nonrecombinant region of the Y chromosome, sequencing the mtDNA of hypervariable regions I and II, and detecting the 9-bp ins/del in region V of mtDNA. All autosomal STRs were in Hardy-Weinberg equilibrium. The four major native American mtDNA haplogroups were represented in the sample. Haplogroups A2 and D1 exhibited the highest frequencies (40.5% and 36.0%, respectively), and haplogroups B2 and C1 appeared to be less frequent (17.5% and 6.0%, respectively). The native American haplogroup Q1a3a was observed in a relevant proportion (88.8%). In addition, a nine-STR Y-chromosome haplotype (DYS19*13, DYS389I*14, DYS389II*31, DYS390*24, DYS391*11, DYS392*14, DYS393*11, DYS385A*14, DYS385B*16) exhibited a frequency of more than 36%. Our results indicate that the analyzed Argentinean Guaraní individuals are genetically more closely related to Guaraní from Brazil [genetic distance (Δμ)2 = 0.48] than to other related tribes that are geographically closer. Statistical approaches based on autosomal data do not support the hypothesis of genetic drift previously proposed; however, this apparent discrepancy might be due to the lack of sensitivity of the autosomal markers used here.

Sex differences in fat storage, fat metabolism, and the health risks from obesity: possible evolutionary origins

Human beings are susceptible to sustained weight gain in the modern environment. Although both men and women can get fat, they get fat in different ways, and suffer different consequences. We review differences between men and women in the incidence of obesity, fat deposition patterns, fat metabolism, and the health consequences of obesity, and examine potential evolutionary explanations for these differences. Women generally have a larger proportion of body mass as fat, and are more likely to deposit fat subcutaneously and on their lower extremities; men are more likely to deposit fat in the abdominal region. Excess adipose tissue in the abdominal region, especially visceral fat, is associated with more health risks. Women have higher rates of reuptake of NEFA into adipose tissue; however, they also have higher rates of fat oxidation during prolonged exercise. Oestrogen appears to underlie many of these differences. Women bear higher nutrient costs during reproduction. Fat and fertility are linked in women, through leptin. Low leptin levels reduce fertility. Ovarian function of adult women is associated with their fatness at birth. In our evolutionary past food insecurity was a frequent occurrence. Women would have benefited from an increased ability to store fat in easily metabolisable depots. We suggest that the pattern of central obesity, more commonly seen in men, is not adaptive, but rather reflects the genetic drift hypothesis of human susceptibility to obesity. Female obesity, with excess adiposity in the lower extremities, reflects an exaggeration of an adaptation for female reproductive success.

The geometry of the wing of Aedes ( Stegomyia) aegypti in isofemale lines through successive generations

Under a common laboratory environment, three isofemale lines of Aedes (Stegomyia) aegypti were used to score metric properties (size and shape) of the wings during 10 generations. Since the number of generations was much higher than the number of founders in each line, genetic drift was expected to occur. Size tended to slightly increase with time, but its variation among the successive generations did not show any detectable information specific to each line. Shape could discriminate among lines in females. Males of lines A and B were not discriminated before generation 8, after which they became completely separated. For each line at each generation, the variance of size and the metric disparity index (as an estimate of shape variance) were higher in females than in males. From one generation to another, the within line shape variance decreased in both sexes, while shape similarity progressively and consistently decreased between males of lines A and B. At each generation, in both sexes, shape variance of the pooled lines was higher than that of each line separately, a pattern not observed for size variance. In conclusion, shape, as a metric property, was closer to a genetic character than size: (i) it showed modifications compatible with the hypothesis of genetic drift and (ii) its variation was related to the complexity of the sample.

Genomic regions involved in response to grain yield selection at high and low nitrogen fertilization in maize

In order to validate the role of genomic regions involved in nitrogen use efficiency and detected in a population of recombinant inbred lines (RIL), we have applied from the same population a recurrent selection for adaptation to low N-input (N0) and to high N-input (N1). Variation of allele frequency at neutral marker during the two cycles of recurrent selection may provide information about markers linked to QTLs. Significant temporal variation of allele frequency was investigated using the test of Waples, which tests the hypothesis of genetic drift versus selection. Most genomic regions (12/19) responding to selection were detected for selection at high N-input and only two were common to selection at high and low N-inputs. This was consistent with the greater grain yield response to selection observed for the population selected under high N-input compared with the population selected under low N-input, when they were evaluated at high N-fertilization. In contrast, when they were evaluated at low N-input both types of selection gave similar yield. As was expected, in the first cycle we observed selection of markers linked to grain yield QTLs. In the course of the second cycle three situations were observed: the confirmation of most regions already selected in C1 including all C1 regions overlapping with grain yield QTLs; the non-confirmation of some C1 regions (2/9); and the identification of new genomic zones (10/17). The detected marker-QTL associations revealed the consistency of the involvement of some traits, such as root architecture and glutamine synthetase activity, which would be of major importance for grain yield setting whatever the nitrogen fertilization.

Spatial variation in phenotypic selection on floral characteristics in an epiphytic orchid

Spatial variation in twelve floral characters was examined in an epiphytic orchidLepanthes rupestris to evaluate the strength and direction of phenotypic selection in seven riparian populations along two river basins in the Caribbean National Forest “El Yunque” for a range of 18–34 months. We evaluated selection on floral characters based on male (pollinaria removal) and female fitness (fruit set). Simple linear and quadratic regressions were used to evaluate the strength of directional, disruptive and stabilizing selections. Univariate and multivariate analyses were used to estimate the total strength of the selection acting on a character. Phenotypic selection was inconsistent among characters and populations. Few of the characters appeared to be under selection and none of them was found to be consistent throughout all populations. Inconsistency in selection coefficients among populations could suggest that selection is spatially variable. We only noted one character (column length) which had some consistency in differential selection coefficients among populations. Previous studies have shown that effective population sizes inL. rupestris are small and the observed “fitness differences” among populations could as easily be explained as stochastic events at play. We argue that the observed “fitness differences” in most characters and inconsistency among populations are likely from stochastic noise and not phenotypic selection. Consequently, we propose that random selection on character state support the hypothesis of genetic drift in small orchid populations.

GENETIC AND GEOGRAPHIC DIFFERENTIATION IN THE RIO NEGRO TUCO-TUCO (CTENOMYS RIONEGRENSIS): INFERRING THE ROLES OF MIGRATION AND DRIFT FROM MULTIPLE GENETIC MARKERS

Among tuco-tucos, Ctenomys rionegrensis is especially amenable to the study of the forces driving population differentiation because of the restricted geographic range it occupies in Uruguay. Within this limited area, the Rio Negro tuco-tuco is limited to sandy soils. It nonetheless exhibits remarkable variation in pelage color, including melanic, agouti, and dark-backed individuals. Two hypotheses have been put forth to explain this pattern: (1) local differentiation and fixation of alternative pelage types by genetic drift under limited gene flow; or (2) fixation by natural selection that may take place even in the presence of gene flow. A previous allozyme study rejected the genetic drift hypothesis on the basis of high inferred levels of migration. New estimates of gene flow from microsatellites and mitochondrial cytochrome b sequences were obtained for C. rionegrensis populations to further test these hypotheses. Much lower levels of gene flow were estimated with these more sensitive markers. Microsatellite-based estimates of gene flow are close to zero and may come closest to estimating current levels of migration. A lack of equilibrium between migration and genetic drift is also strongly suggested by the absence of an isolation-by-distance pattern found in all three genetic datasets. The microsatellite genotype data show that the species is strongly structured geographically, with subpopulations constituting distinct genetic entities. If current levels of gene flow are very low, as indicated by the new data, the local fixation of alternative alleles, including those responsible for pelage color polymorphism, is possible by drift alone. A scenario is thus proposed in which the species expanded in the recent past from a more restricted geographic range and has subsequently differentiated in near isolation, with genetic drift possibly playing a primary role in overall genetic differentiation. The local fixation of pelage color types could also be due to drift, but selection on this trait cannot be ruled out without direct analysis.

Genetic trends in a population evolving antibiotic resistance.

The evolution of antibiotic resistance provides a well-documented, rapid, and recent example of a selection driven process that has occurred in many bacterial species. An exhaustive collection of Moraxella catarrhalis that spans a transition to chromosomally encoded penicillin resistance was used to analyze genetic changes accompanying the transition. The population was characterized by high haplotypic diversity with 148 distinct haplotypes among 372 isolates tested at three genomic regions. The power of a temporally stratified sample from a single population was highlighted by the finding of high genetic diversity throughout the transition to resistance, population numbers that remained high over time, and no evidence of departures from neutrality in the allele frequency spectra throughout the transition. The direct temporal analysis documented the persistence, antibiotic status, and haplotypic identity of strains undergoing apparent clonal expansions. Several haplotypes that were beta-lactamase nonproducers in early samples converted to producers in later years. Maintenance of genetic diversity and haplotype conversions from sensitive to resistant supported the hypothesis that penicillin resistance determinants spread to a diverse array of strains via horizontal exchange. Genetic differentiation between sample years, estimated by F(ST), was increasing at a rate that could cause complete haplotype turnover in less than 150 years. Widespread linkage disequilibrium among sites within one locus (copB) suggested recent mutation followed by clonal expansion. Nonrandom associations between haplotypes and resistance phenotypes provided further evidence of clonal expansion for some haplotypes. Nevertheless, the population structure was far from clonal as evidenced by a relatively low frequency of disequilibria both within sites at a second locus (M46) as well as between loci. The haplotype-antibiotic resistance association that was accompanied by gradual haplotype turnover is consistent with a hypothesis of genetic drift at marker loci with directional selection at the resistance locus.

Evaluation of ECG-synchronized technique on a half-second multi-slice CT

Under a common laboratory environment, three isofemale lines of Aedes (Stegomyia) aegypti were used to score metric properties (size and shape) of the wings during 10 generations. Since the number of generations was much higher than the number of founders in each line, genetic drift was expected to occur.Size tended to slightly increase with time, but its variation among the successive generations did not show any detectable information specific to each line. Shape could discriminate among lines in females. Males of lines A and B were not discriminated before generation 8, after which they became completely separated.For each line at each generation, the variance of size and the metric disparity index (as an estimate of shape variance) were higher in females than in males. From one generation to another, the within line shape variance decreased in both sexes, while shape similarity progressively and consistently decreased between males of lines A and B. At each generation, in both sexes, shape variance of the pooled lines was higher than that of each line separately, a pattern not observed for size variance.In conclusion, shape, as a metric property, was closer to a genetic character than size: (i) it showed modifications compatible with the hypothesis of genetic drift and (ii) its variation was related to the complexity of the sample.

GENETIC TRENDS IN A POPULATION EVOLVING ANTIBIOTIC RESISTANCE

The evolution of antibiotic resistance provides a well-documented, rapid, and recent example of a selection driven process that has occurred in many bacterial species. An exhaustive collection of Moraxella catarrhalis that spans a transition to chromosomally encoded penicillin resistance was used to analyze genetic changes accompanying the transition. The population was characterized by high haplotypic diversity with 148 distinct haplotypes among 372 isolates tested at three genomic regions. The power of a temporally stratified sample from a single population was highlighted by the finding of high genetic diversity throughout the transition to resistance, population numbers that remained high over time, and no evidence of departures from neutrality in the allele frequency spectra throughout the transition. The direct temporal analysis documented the persistence, antibiotic status, and haplotypic identity of strains undergoing apparent clonal expansions. Several haplotypes that were β-lactamase nonproducers in early samples converted to producers in later years. Maintenance of genetic diversity and haplotype conversions from sensitive to resistant supported the hypothesis that penicillin resistance determinants spread to a diverse array of strains via horizontal exchange. Genetic differentiation between sample years, estimated by FST, was increasing at a rate that could cause complete haplotype turnover in less than 150 years. Widespread linkage disequilibrium among sites within one locus (copB) suggested recent mutation followed by clonal expansion. Nonrandom associations between haplotypes and resistance phenotypes provided further evidence of clonal expansion for some haplotypes. Nevertheless, the population structure was far from clonal as evidenced by a relatively low frequency of disequilibria both within sites at a second locus (M46) as well as between loci. The haplotype-antibiotic resistance association that was accompanied by gradual haplotype turnover is consistent with a hypothesis of genetic drift at marker loci with directional selection at the resistance locus.Corresponding Editor: D. Roff

VARIATION IN GENETIC ARCHITECTURE OF CALLING SONG AMONG POPULATIONS OF ALLONEMOBIUS SOCIUS, A. FASCIATUS, AND A HYBRID POPULATION: DRIFT OR SELECTION?

Predictions using quantitative genetic models generally assume that the variance-covariance matrices remain constant over time. This assumption is based on the supposition that selection is generally weak and hence variation lost through selection can be replaced by new mutations. Whether this is generally true can only be ascertained from empirical studies. Ideally for such a study we should be able to make a prediction concerning the relative strength of selection versus genetic drift. If the latter force is prevalent then the variance-covariances matrices should be proportional to each other. Previous studies have indicated that females in the two sibling cricket species Allonemobius socius and A. fasciatus do not discriminate between males of the two species by their calling song. Therefore, differences between the calling song of the two males most likely result from drift rather than sexual selection. We test this hypothesis by comparing the genetic architecture of calling song of three populations of A. fasciatus with two populations of A. socius. We found no differences among populations within species, but significant differences in the G (genetic) and P (phenotypic) matrices between species, with the matrices being proportional as predicted under the hypothesis of genetic drift. Because of the proportional change in the (co)variances no differences between species are evident in the heritabilities or genetic correlations. Comparison of the two species with a hybrid population from a zone of overlap showed highly significant nonproportional variation in genetic architecture. This variation is consistent with a general mixture of two separate genomes or selection. Qualitative conclusions reached using the phenotypic matrices are the same as those reached using the genetic matrices supporting the hypothesis that the former may be used as surrogate measures of the latter.

The genetic structure of natural populations of Drosophila melanogaster. XXII. Comparative study of DNA polymorphisms in northern and southern natural populations.

Restriction map variation in four gene regions (Adh, Amy, Pu and Gpdh) was surveyed for 86 second chromosomes from northern (Aomori) and southern (Ogasawara) Japanese populations of Drosophila melanogaster (43 chromosomes from each population). The regions examined cover a total of 62 kilobases. Estimates of nucleotide diversity (pi) were approximately constant across the gene regions and populations examined. The distribution of restriction site polymorphisms was compatible with the expectation from the neutral mutation-random genetic drift hypothesis, but insertion/deletion polymorphisms were not consistent with it. While the two populations shared a majority of restriction site polymorphisms, frequencies of individual restriction site variants were significantly different between the two populations at 7 out of 35 segregating sites. In addition, an insertion in the Amy region was found in 15 chromosomes from the Ogasawara sample but absent in the Aomori sample. A considerable difference was observed in the number of rare insertions and deletions between the two populations. The numbers of aberrations uniquely represented were 16 in the Ogasawara sample and only 3 in the Aomori sample. These findings suggest that the two populations were differentiated from each other to some degree by means of random genetic drift and/or other factors.