Null Hypothesis Of Random Mating Research Articles

Genotyping by sequencing suggests overwintering of Peronospora destructor in southwestern Québec, Canada.

Several Peronospora species are carried by wind over short and long distances, from warmer climates where they survive on living plants to cooler climates. In eastern Canada, this annual flow of sporangia was thought to be the main source of Peronospora destructor responsible for onion downy mildew. However, the results of a recent study showed that the increasing frequency of onion downy mildew epidemics in eastern Canada is associated with warmer autumns, milder winters, and previous year disease severity, suggesting overwintering of the inoculum in an area where the pathogen is not known to be endogenous. In this study, genotyping by sequencing was used to investigate the population structure of P. destructor at the landscape scale. The study focused on a particular region of southwestern Québec—Les Jardins de Napierville—to determine if the populations were clonal and regionally differentiated. The data were characterized by a high level of linkage disequilibrium, characteristic of clonal organisms. Consequently, the null hypothesis of random mating was rejected when tested on predefined or nonpredefined populations, indicating that linkage disequilibrium was not a function of population structure and suggesting a mixed reproduction mode. Discriminant analysis of principal components performed with predefined population assignment allowed grouping P. destructor isolates by geographical regions, while analysis of molecular variance confirmed that this genetic differentiation was significant at the regional level. Without using a priori population assignment, isolates were clustered into four genetic clusters. These results represent a baseline estimate of the genetic diversity and population structure of P. destructor.

Genetic Structure of Rhizoctonia solani AG-2-2IIIB from Soybean in Illinois, Ohio, and Ontario

Rhizoctonia solani AG-2-2IIIB is an important seedling pathogen of soybean in North America and other soybean-growing regions around the world. There is no information regarding the population genetics of field populations of R. solani associated with soybean seedling disease. More specifically, information regarding genetic diversity, the mode of reproduction, and the evolutionary factors that shape different R. solani populations separated in time and space are lacking. We exploited genotyping by sequencing as a tool to assess the genetic structure of R. solani AG-2-2IIIB populations from Illinois, Ohio, and Ontario and investigate the reproductive mode of this subgroup. Our results revealed differences in genotypic diversity among three populations, with the Ontario population having greatest diversity. An overrepresentation of multilocus genotypes (MLGs) and a rejection of the null hypothesis of random mating in all three populations suggested clonality within each population. However, phylogenetic analysis revealed long terminal multifurcating branches for most members of the Ontario population, suggesting a mixed reproductive mode for this population. Analysis of molecular variance revealed low levels of population differentiation, and sharing of similar MLGs among populations highlights the role of genotype flow as an evolutionary force shaping population structure of this subgroup.

Genetic diversity evidence a mixed reproduction mode in Venturia oleaginea populations in Uruguay

Olive scab caused by Venturia oleaginea is one of the most important diseases of olive worldwide. The fungus infects young leaves and fruits by asexually produced conidia. The sexual stage and its importance in the epidemiology and genetics of the pathogen population is unknown. A collection of 52 fungal isolates was obtained from scabbed leaves collected in 16 orchards from three major regions in Uruguay. All isolates were identified as V. oleaginea by PCR with specific primers and sequence analysis of the TEF-1α gene region. Five colony morphotypes and different growing rates were observed on potato dextrose agar (PDA) medium. Universally primed polymerase chain reaction (UP-PCR) was successfully used for assessing the genetic diversity and population structure of the V. oleaginea isolates, whereas ISSR was unable to generate reproducible and polymorphic bands. Two populations genetically different were identified: [1] composed by 21 isolates mainly from Southern and South-Western regions; [2] composed by 31 isolates mainly from South-Eastern and Southern regions. Both populations showed a moderate gene diversity expressed as hNei = 0.163 and 0.212 for population [1] and [2], respectively. There was evidence of linkage disequilibrium in both populations, rejecting the null hypothesis of random mating as the predominant reproduction mode, a typical result of clonal populations. However, all isolates were genotypically different, an indirect evidence of recombination. We conclude that V. oleaginea populations in Uruguay present a mixed mode of reproduction, a characteristic of many pathogens that are difficult to control.

Population Biology and Genetic Variation of Spongospora subterranea f. sp. subterranea, the Causal Pathogen of Powdery Scab and Root Galls on Potatoes in South Africa.

Spongospora subterranea f. sp. subterranea, causal agent of powdery scab and root galls of potatoes, occurs worldwide and is responsible for quality and yield losses in potato production in South Africa. Despite being one of the most important potato pathogens in South Africa, little information is available on the genetic structure and diversity of S. subterranea f. sp. subterranea, which could provide insight into the factors shaping its evolution and the role of inoculum sources in disease development. A total of 172 samples were collected from four potato growing regions in South Africa. An additional 27 samples obtained from Colombia were included for comparative purposes. The samples were screened against six informative microsatellite (simple-sequence repeat) markers. Of the 172 samples obtained from potato growing regions in South Africa, there were 75 multilocus genotypes (MLGs), only 16 of which were shared between potato growing regions, indicating substantial gene flow and countrywide dispersal of the pathogen. The presence of common MLGs among the root- and tuber-derived samples indicated a lack of specialization of S. subterranea f. sp. subterranea to either tuber or root infection. Nei's unbiased estimates of gene diversity for the clone-corrected data were low and ranged from 0.24 to 0.38. Analysis of molecular variance and discriminant analysis of principal components showed no population differentiation between different potato growing regions in South Africa and between root- and tuber-derived genotypes. The presence of MLGs, high considerable genotypic diversity, and failure to reject the null hypothesis of random mating in most populations are indicative of some kind of recombination, either sexual or asexual, in these S. subterranea f. sp. subterranea populations. Information from this study provides new insights into the genetic structure and diversity of S. subterranea f. sp. subterranea in South Africa. Continuous monitoring of the pathogen population dynamics will be helpful in implementing effective region-specific management strategies for the pathogen, especially in the development of resistant potato cultivars.

The Population Genetics of Alternaria tenuissima in Four Regions of China as Determined by Microsatellite Markers Obtained by Transcriptome Sequencing.

A total of 32,284 unigenes were obtained from the transcriptome of Alternaria tenuissima, a pathogenic fungus causing foliar disease in tomato, using next-generation sequencing (NGS) technology. In total, 24,670 unigenes were annotated using five databases, including NCBI non-redundant protein, Swiss-Prot, euKaryotic Orthologous Groups, Kyoto Encyclopedia of Genes and Genomes, and the Gene Ontology. A total of 1,140 simple sequence repeats were also identified for use as molecular markers. Sixteen of the simple sequence repeat loci were selected to study the population structure of A. tenuissima. A population genetic analysis of 191 A. tenuissima isolates, sampled from four geographic regions in China, indicated that A. tenuissima had a high level of genetic diversity, and that the selected simple sequence repeat markers could reliably capture the genetic variation. The null hypothesis of random mating was rejected for all four geographic regions in China. Isolation by distance was observed for the entire data set, but not within clusters, which is indicative of barriers to gene flow among geographic regions. The analyses of Bayesian and principal coordinates, however, did not separate four geographic regions into four separate genetic clusters. The different levels of historical migration rates suggest that isolation by distance did not represent a major biological obstacle to the spread of A. tenuissima. The potential epidemic spread of A. tenuissima in China may occur through the transport of plant products or other factors. The presented results provide a basis for a comprehensive understanding of the population genetics of A. tenuissima in China.

Population structure and reproductive mode of Didymella fabae in Syria

Didymella fabae is a highly destructive fungal pathogen of faba bean (Vicia faba) that represents a significant yield‐limiting biotic constraint in all locations where the crop is grown. However, nothing is known about the population structure of this pathogen anywhere in the world. Population genetic analyses employing 18 sequence‐tagged microsatellite (STMS) markers covering eight genetic linkage groups and a PCR‐based mating type marker were used to elucidate the genetic structure and reproductive mode of the pathogen in three populations in Syria. High gene diversity within populations but low genetic differentiation among populations was observed and the entire collection of isolates was assigned to a single genetic population using a Bayesian clustering algorithm. Independent analyses were performed based on four unlinked sets of STMS markers to infer reproductive mode. A simulation approach was used to estimate which of the repeated multilocus genotypes were probably the result of asexual reproduction and should be clone‐corrected. A 1:1 ratio of mating types could not be rejected in any clone‐corrected population, probably due to small sample sizes. Likewise, frequency of clones and sample size, but not marker linkage, had strong effects on multilocus gametic disequilibrium. The null hypothesis of random mating was rejected in the majority of populations for both non‐clone‐corrected and clone‐corrected samples and with four sets of unlinked markers indicating a predominance of asexual reproduction in these populations. This represents the first detailed screening of clonal and genetic composition of D. fabae populations in Syria.

Genetic Structure ofColletotrichum gloeosporioidessensu lato Isolates Infecting Papaya Inferred by Multilocus ISSR Markers

Colletotrichum gloeosporioides sensu lato is widely distributed throughout temperate and tropical regions and causes anthracnose disease in numerous plant species. Development of effective disease management strategies is dependent on, among other factors, an understanding of pathogen genetic diversity and population stratification at the intraspecific level. For 132 isolates of C. gloeosporioides sensu lato collected from papaya in Trinidad, inter-simple-sequence repeat-polymerase chain reaction (ISSR-PCR) generated 121 polymorphic loci from five ISSR primers selected from an initial screen of 22 ISSR primers. The mean percentage of polymorphic loci was 99.18%. Bayesian cluster analysis inferred three genetic subpopulations, where group 1 consisted exclusively of isolates collected in the southern part of Trinidad whereas groups 2 and 3, although genetically distinct, were mixtures of isolates collected from both the northern and southern parts of Trinidad. Principal coordinates analysis and unweighted pair-group method with arithmetic mean phylogeny were concordant with Bayesian cluster analysis and supported subdivision into the three subpopulations. Overall, the total mean gene diversity was 0.279, the mean within-population gene diversity was 0.2161, and genetic differentiation for the Trinidad population was 0.225. Regionally, northern isolates had a lower gene diversity compared with southern isolates. Nei's gene diversity was highest for group 1 (h = 0.231), followed by group 2 (h = 0.215) and group 3 (h = 0.202). Genotypic diversity was at or near maximum for all three subpopulations after clone correction. Pairwise estimates of differentiation indicated high and significant genetic differentiation among the inferred subpopulations (Weir's θ of 0.212 to 0.325). Pairwise comparisons among subpopulations suggested restricted gene flow between groups 1 and 2 and groups 1 and 3 but not between groups 2 and 3. The null hypothesis of random mating was rejected for all three inferred subpopulations. These results suggest that pathogen biology and epidemiology as well as certain evolutionary factors may play an important role in population substructuring of C. gloeosporioides sensu lato isolates infecting papaya in Trinidad.

Variability of the Ramularia collo‐cygni Population in Central Europe

AbstractThe genetic structure of the fungal barley pathogen Ramularia collo‐cygni (Rcc) population in Central Europe involving the isolates from the Czech Republic, the Slovak Republic, Germany and Swiss was determined using amplified fragment length polymorphism (AFLP) analysis. One hundred and eighty‐four markers were chosen to determine genetic and genotypic diversity and to test the hypothesis of random mating and population differentiation of Rcc isolates. Among the 337 isolates collected, the overall gene diversity was moderate ( = 0.216). The level of multilocus genotypic diversity was higher within populations than among them. All individuals had unique multilocus genotypes. Genetic differentiation was significant among populations in localities, but at a moderate level (θ = 0.12; P < 0.001), suggesting that gene flow is occurring among populations. The isolates from all twelve clusters produced by Structure were found in all local populations, although at different frequencies. Therefore, the inferred clusters did not represent geographical populations. Although the null hypothesis of random mating in Rcc populations was rejected, the high level of genotypic diversity suggests that the Rcc population structure appears to be generated by a mixed reproductive system including both asexual and sexual reproduction, along with a rather high migration rate.

Distinguishing Galactomyces citri-aurantii from G. geotrichum and Characterizing Population Structure of the Two Postharvest Sour Rot Pathogens of Fruit Crops in California

A growth assay in lemon juice and polymerase chain reaction amplifications using newly designed species-specific primers from endopolygalacturonase and β-tubulin genes rapidly differentiated isolates of the morphologically similar fruit sour rot pathogens Galactomyces citriaurantii and G. geotrichum. Isolates of both species were collected from agricultural soils and decaying fruit at locations within and outside California, including worldwide locations, and were used in population genetic studies based on amplified fragment length polymorphic (AFLP) DNA markers. For all four geographically defined subpopulations (three counties of California and locations outside California) among 97 isolates of G. citri-aurantii and for the two subpopulations (origin within or outside California) among 35 isolates of G. geotrichum, the proportion of polymorphic loci and haplotypic diversity was high. In total, 82 unique haplotypes were identified for G. citri-aurantii for the four subpopulations and, of these, 80 haplotypes were unique among subpopulations. For G. geotrichum, 25 unique haplotypes were identified among the two subpopulations and no haplotype was shared. Indices of genetic differences (F(ST)) between subpopulations within each species were all low (e.g., 0.038 for G. geotrichum and 0.085 to 0.226 for G. citriaurantii), indicating a low level of genetic differentiation. Following clone correction, mating type segregation ratios for G. citri-aurantii did not significantly (P > 0.1) deviate from a 1:1 ratio for all four subpopulations or the entire population. Tests of the index of association (I(A)) and parsimony tree-length permutation tests (PTLPT) supported a random mating structure for clone-corrected data for the Kern, Tulare, and Ventura County subpopulations and the null hypothesis of random mating could not be rejected. Additionally, PTLPT also supported random mating for the "outside of California" population. For G. geotrichum, random mating was only tested using I(A) and PTLPT and the null hypothesis of random mating was not rejected (P > 0.05) using clone-corrected data. Further evidence that sexual recombination likely occurs in both species of Galactomyces was the lack of grouping consistency in the unweighted pair-group method with arithmetic mean clustering of AFLP data. A high confidence based on bootstrap values was obtained for only a few of the nodes in each of the two trees. A mixed reproduction system with an out-crossing sexual mating system and a prolific asexual phase is proposed for both species.

No evidence for inbreeding avoidance through active mate choice in red-billed gulls

Except for cooperative breeders, most studies on wild birds have failed to find evidence for inbreeding avoidance via kin discriminative mate choice. This, together with evidence for kin avoidance through dispersal, has led to the general view that dispersal is often a sufficient inbreeding avoidance mechanism and active discrimination through mate choice is unnecessary. Yet, the study of inbreeding avoidance in the wild is difficult and long-term studies of pedigreed wild populations can provide important insights. We studied the occurrence of inbreeding avoidance in a highly philopatric red-billed gull (Larus novaehollandiae scopulinus) population subject to an individual-based field study since 1958 in Kaikoura, New Zealand. Despite a wealth of breeding and pedigree data, we did not observe a single inbred pair. This observation was a small but significant deviation from the expectation under the null hypothesis of random mating when we looked at annual breeding attempts, suggesting inbreeding avoidance. However, the difference disappeared when we examined pair bonds rather than annual breeding attempts. Our results are consistent with the expectation that close inbreeding occurs rarely in large random-mating populations. They also demonstrate how mating systems, in this case long-term monogamous pair bonds with sex differences both in the age of first breeding and breeding dispersal at natal subcolonies, can reduce the likelihood of inbreeding.

Clonality, genetic diversity and support for the diversifying selection hypothesis in natural populations of a flower-living yeast

Vast amounts of effort have been devoted to investigate patterns of genetic diversity and structuring in plants and animals, but similar information is scarce for organisms of other kingdoms. The study of the genetic structure of natural populations of wild yeasts can provide insights into the ecological and genetic correlates of clonality, and into the generality of recent hypotheses postulating that microbial populations lack the potential for genetic divergence and allopatric speciation. Ninety-one isolates of the flower-living yeast Metschnikowia gruessii from southeastern Spain were DNA fingerprinted using amplified fragment length polymorphism (AFLP) markers. Genetic diversity and structuring was investigated with band-based methods and model- and nonmodel-based clustering. Linkage disequilibrium tests were used to assess reproduction mode. Microsite-dependent, diversifying selection was tested by comparing genetic characteristics of isolates from bumble bee vectors and different floral microsites. AFLP polymorphism (91%) and genotypic diversity were very high. Genetic diversity was spatially structured, as shown by amova (Φ(st) = 0.155) and clustering. The null hypothesis of random mating was rejected, clonality seeming the prevailing reproductive mode in the populations studied. Genetic diversity of isolates declined from bumble bee mouthparts to floral microsites, and frequency of five AFLP markers varied significantly across floral microsites, thus supporting the hypothesis of diversifying selection on clonal lineages. Wild populations of clonal fungal microbes can exhibit levels of genetic diversity and spatial structuring that are not singularly different from those shown by sexually reproducing plants or animals. Microsite-dependent, divergent selection can maintain high local and regional genetic diversity in microbial populations despite extensive clonality.

Local scale population genetic structure of Entomophthora muscae epidemics

Abstract Isolates of the entomopathogenic fungus Entomophthora muscae were obtained from houseflies sampled at five stables on Zealand during an epidemic in fall 2005. DNA fingerprints were generated from single conidia from 40 E. muscae isolates using the PCR-based method of Inter-Simple Sequence Repeats (ISSR). This resulted in fingerprint patterns consisting of about 50 fragments, of which, 14 were polymorphic. From the polymorphic loci we estimated the reproductive mode, genetic differentiation and gene diversity of E. muscae populations using the statistics of index of association, Weir & Cockerhams theta and Nei's analysis of gene diversity in subdivided populations. Our results revealed no significant differences in allele frequencies among the five populations. The index of association test rejected the null hypothesis of random mating, but the test for paired locus compatibility showed weak or no linkage between loci indicating recombination or homoplasy in the dataset. From this study we cannot exclude the possibility that the population genetic structure underlying E. muscae epidemics could be panmictic consisting of several lineages with a high level of reciprocal migration.

Population Structure of Seedborne Phaeosphaeria nodorum on New York Wheat

ABSTRACT Population genetic and epidemiological studies have resulted in different hypotheses about the predominant source of primary inoculum in the Phaeosphaeria nodorum-wheat pathosystem (i.e., sexually derived, windborne ascospores versus asexual or seedborne inoculum). We examined the genetic structure of seedborne populations of P. nodorum as a further step toward evaluating the hypothesis that seedborne inoculum is an important contributor to foliar epidemics in New York's rotational wheat fields. In all, 330 seedborne isolates from seven field populations were genotyped at 155 amplified fragment length polymorphism loci. Seedborne populations possessed high levels of genotypic diversity, with virtually every isolate (326/330) having a unique haplotype. As in previous population genetic studies of P. nodorum, we found low levels of gametic disequilibrium, although we could reject the null hypothesis of random mating with the index of association test for two populations. Thus, genotypically diverse and seemingly panmictic populations of P. nodorum that have been observed in wheat foliage could be derived from seedborne primary inoculum. Although sexual reproduction and recombination may contribute to the diversity of foliar populations of P. nodorum, population genetic data do not rule out seed as a source of primary inoculum. Further experimentation will be needed to determine definitively the relative importance of windborne ascospores and seed-borne asexual inoculum in epidemics of Stagonospora nodorum blotch in New York.

Genetic structure of populations of the red sea urchin, Strongylocentrotus franciscanus.

Population subdivision was evaluated in the red sea urchin, Strongylocentrotus franciscanus, using DNA sequence data from 134 adult individuals collected in 1995 and 1996. On average 22 individuals were sequenced from six geographic locations between Alaska and Baja California (N=134), nearly the full extent of the species range. DNA sequence data was obtained from direct sequencing of a 273 base pair region of the bindin gene, which encodes a sperm fertilization protein. Results indicate that bindin is sufficiently polymorphic to serve as a genetic marker. We identified 14 unique alleles present in the entire range sampled with a maximum of eight alleles at a specific site. Mean pairwise comparison of the 14 unique alleles indicates moderate sequence diversity (p-distance=1.06). Although there is conflicting evidence to suggest that Alaska populations may deviate from the Hardy-Weinberg expectations, analysis of bindin genotype frequencies indicate that it is not possible to reject the null hypothesis of random mating throughout the species range. The results of a chi-square test with pooling conform to Hardy-Weinberg expectations for all populations (P>0.05) except for the Alaska population (P=0.037). Inbreeding coefficients are consistent with this result and suggest that for the bindin locus, there is high gene flow. These results are compared with previously published results of genetic substructuring in sea urchins to examine relationships among population structure, dispersal potential and biogeography.

Reproductive Success and Assortative Mating amongPomoxisSpecies and Their Hybrids

We investigated the patterns of reproductive isolation and gene flow between crappie species Pomoxis spp. by conducting a series of controlled breeding experiments using both pure parental species and various forms of interspecific hybrids (i.e., F1, F2) as parents. To assess relative reproductive success of mating types, progeny from each of three different experimental treatments were assayed for genotypic composition at four diagnostic allozyme loci. In the first experimental treatment, we confirmed the viability and fertility of F1 hybrids and observed that genotypic and allelic frequencies among the F2 progeny departed from predicted Mendelian genotypic proportions. In the second experimental treatment, we confirmed that F3 hybrids can arise from intermating F2 hybrids. However, the small number of progeny recovered from experimental ponds suggests that either prezygotic or postzygotic barriers to gene flow had limited reproductive success. In the third experimental treatment, we evaluated the relative reproductive success of the parental species and both reciprocal F1 hybrids. We rejected the null hypothesis of random mating from equal mixtures of parental and hybrid brood based on a maximum likelihood mixture model estimate of resulting progeny. Within each replicate, one or both parental species were most successful, although F2 hybrid progeny also occurred more often than expected; first generation hybrids and backcrosses occurred less frequently than expected. These and previously reported observations concerning the viability of hybrids and backcrosses indicate that the two crappie species can interbreed but that they mate assortatively. Because reproductive isolation is imperfect, we caution against intentional mixing or creation of zones of secondary contact between these species.

DNA fingerprinting and analysis of population structure in the chestnut blight fungus, Cryphonectria parasitica.

We analyzed DNA fingerprints in the chestnut blight fungus, Cryphonectria parasitica, for stability, inheritance, linkage and variability in a natural population. DNA fingerprints resulting from hybridization with a dispersed moderately repetitive DNA sequence of C. parasitica in plasmid pMS5.1 hybridized to 6-17 restriction fragments per individual isolate. In a laboratory cross and from progeny from a single perithecium collected from a field population, the presence/absence of 11 fragments in the laboratory cross and 12 fragments in the field progeny set segregated in 1:1 ratios. Two fragments in each progeny set cosegregated; no other linkage was detected among the segregating fragments. Mutations, identified by missing bands, were detected for only one fragment in which 4 of 43 progeny lacked a band present in both parents; no novel fragments were detected in any progeny. All other fragments appeared to be stably inherited. Hybridization patterns did not change during vegetative growth or sporulation. However, fingerprint patterns of single conidial isolates of strains EP155 and EP67 were found to be heterogenous due to mutations that occurred during culturing in the laboratory since these strains were first isolated in 1976-1977. In a population sample of 39 C. parasitica isolates, we found 33 different fingerprint patterns with pMS5.1. Most isolates differed from all other isolates by the presence or absence of several fragments. Six fingerprint patterns each occurred twice. Isolates with identical fingerprints occurred in cankers on the same chestnut stems three times; isolates within the other three pairs were isolated from cankers more than 5 m apart. The null hypothesis of random mating in this population could not be rejected if the six putative clones were removed from the analysis. Thus, a rough estimate of the clonal fraction of this population is 6 in 39 isolates (15.4%).

Allozyme Variation in Mojave Populations of the Desert Tortoise, Gopherus agassizi

Whole blood samples were obtained from a total of 146 desert tortoises from western (Kramer Hills region) and eastern (Chemehuevi Valley) parts of the Mojave Desert in San Bernardino County, California. These samples were examined electrophoretically for allozyme expression at 23 loci, eight of which were polymorphic. Heterozygosity levels and other measures of genetic variability were comparable for the eastern and western Mojave samples. Intrapopulation chi-square tests did not falsify our null hypothesis of random mating at each location. Genetic distance measures and interpopulation FsT values suggested that genetic differentiation of eastern and western Mojave Desert tortoises was minimal. However, a contingency chi-square test and multivariate analyses demonstrated significant differences between these geographic samples, falsifying the hypothesis that these Mojave samples are part of a single randomly breeding population. From the perspective of allozyme characteristics, we have no reservations regarding relocation of tortoises between the Kramer and Chemehuevi sites. Allozyme characters of transplanted tortoises should be documented for historical record.