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The evolution of programmed death in a spatially structured population.

Recent studies have identified a number of genes that regulate life span. When these genes are inactivated, organisms live longer, but suffer no obvious adverse effects. These results have generated considerable debate, largely because current evolutionary theory is unable to explain them. In this article, I report results from an individual-based spatial model in which a programmed age of death is allowed to evolve. In a freely mixing population with global dispersal, evolution selects for individuals with ever-increasing life span. However, in a spatially structured population with localized dispersal, a programmed age of death evolves. The exact age of death that evolves depends critically on the scale of dispersal. Within this model, individuals are genetically programmed to die, even though they are still able to reproduce. These results suggest that death can be adaptive and offer an explanation for the evolution of "death genes."

Cell Death↗

On the determinants of population structure in antigenically diverse pathogens.

Many pathogens exhibit antigenic diversity and elicit strain-specific immune responses. This potential for cross-immunity structure in the host resource motivates the development of mathematical models, stressing competition for susceptible hosts in driving pathogen population dynamics and genetics. Here we establish that certain model formulations exhibit characteristics of prototype pattern-forming systems, with pathogen population structure emerging as three possible patterns: (i) incidence is steady and homogeneous; (ii) incidence is steady but heterogeneous; and (iii) incidence shows oscillatory dynamics, with travelling waves in strain-space. Results are robust to strain number, but sensitive to the mechanism of cumulative immunity.

Animals↗

Reduction of structured population models to threshold-type delay equations and functional differential equations: a case study.

It is shown by way of a simple example that certain structured population models lead naturally to differential delay equations of the threshold type and that these equations can be transformed in a natural way to functional differential equations. The model examined can be viewed as a model of competition between adults and juveniles of a single population. The results indicate the possibility that this competition leads to instability.

Age Factors↗

Allelic diversity and population structure in Oenococcus oeni as determined from sequence analysis of housekeeping genes.

Oenococcus oeni is the organism of choice for promoting malolactic fermentation in wine. The population biology of O. oeni is poorly understood and remains unclear. For a better understanding of the mode of genetic variation within this species, we investigated by using multilocus sequence typing (MLST) with the gyrB, pgm, ddl, recP, and mleA genes the genetic diversity and genetic relationships among 18 O. oeni strains isolated in various years from wines of the United States, France, Germany, Spain, and Italy. These strains have also been characterized by ribotyping and restriction fragment length polymorphism (RFLP) analysis of the PCR-amplified 16S-23S rRNA gene intergenic spacer region (ISR). Ribotyping grouped the strains into two groups; however, the RFLP analysis of the ISRs showed no differences in the strains analyzed. In contrast, MLST in oenococci had a good discriminatory ability, and we have found a higher genetic diversity than indicated by ribotyping analysis. All sequence types were represented by a single strain, and all the strains could be distinguished from each other because they had unique combinations of alleles. Strains assumed to be identical showed the same sequence type. Phylogenetic analyses indicated a panmictic population structure in O. oeni. Sequences were analyzed for evidence of recombination by split decomposition analysis and analysis of clustered polymorphisms. All results indicated that recombination plays a major role in creating the genetic heterogeneity of O. oeni. A low standardized index of association value indicated that the O. oeni genes analyzed are close to linkage equilibrium. This study constitutes the first step in the development of an MLST method for O. oeni and the first example of the application of MLST to a nonpathogenic food production bacteria.

Alleles↗

Using reproductive value to estimate key parameters in density-independent age-structured populations.

The dynamics of reproductive value are used to provide a simple derivation of Tuljapurkar's approximation for the long-run growth rate and environmental variance of lnN, in a density-independent age-structured population in a random environment. With no environmental autocorrelation, the dynamics of total population size, N, generally shows time lags and autocorrelation caused by life history, which may strongly bias estimates of environmental variance obtained by ignoring age structure. In contrast, the total reproductive value, V, is Markovian and obeys a first-order autoregressive process. This suggests a simple method for estimating the environmental variance, and avoiding potentially large bias due to age-structure fluctuations, by converting a multivariate time series of age structure to a univariate time series of lnV. We illustrate the method by estimating the long-run growth rate and the environmental variance in an exponentially growing population of Bighorn Sheep.

Age Distribution↗

Genetic population structure of marine viral haemorrhagic septicaemia virus (VHSV).

The nucleotide sequences of a specific region of the nucleoprotein gene were compared in order to investigate the genetic population structure of marine viral haemorrhagic septicaemia virus (VHSV). Analysis of the sequence from 128 isolates of diverse geographic and host origin renders this the most comprehensive molecular epidemiological study of marine VHSV conducted to date. Phylogenetic analysis of nucleoprotein gene sequences confirmed the existence of the 4 major genotypes previously identified based on N- and subsequent G-gene based analyses. The range of Genotype I included subgroups of isolates associated with rainbow trout aquaculture (Genotype Ia) and those from the Baltic marine environment (Genotype Ib) to emphasise the relatively close genetic relationship between these isolates. The existence of an additional genotype circulating within the Baltic Sea (Genotype II) was also confirmed. Genotype III included marine isolates from around the British Isles in addition to those associated with turbot mariculture, highlighting a continued risk to the development of this industry. Genotype IV consisted of isolates from the marine environment in North America. Taken together, these findings suggest a marine origin of VHSV in rainbow trout aquaculture. The implications of these findings with respect to the future control of VHSV are discussed. The capacity for molecular phylogenetic analysis to resolve complex epidemiological problems is also demonstrated and its likely future importance to disease management issues highlighted.

Animals↗

Distinct population structure in a phenotypically homogeneous rock-dwelling cichlid fish from Lake Tanganyika.

Several lineages of cichlid fishes in the East African Great Lakes display stunning levels of morphological diversification. The rapid evolution of rock-dwelling polygynous mouthbrooders in Lake Malawi, for example, was in part ascribed to their allopatric distribution on disjunct stretches of rocky coast, where even short habitat discontinuities reduce gene flow effectively. However, as seen in other cichlids, ecological barriers do not always prevent gene flow, whereas genetic structure can develop along continuous habitat, and morphological diversification does not necessarily accompany genetic differentiation. The present study investigates the population structure of Variabilichromis moorii, a monogamous substrate-brooding lamprologine of rocky coasts in Lake Tanganyika, which occurs over about 1000 km of shoreline almost without phenotypic variation. Phylogeographic analyses of mitochondrial DNA sequences indicated that dispersal is infrequent and generally occurs between adjacent locations only. Exceptions to this pattern are closely related haplotypes from certain locations on opposite lakeshores, a phenomenon which has been observed in other species and is thought to reflect lake crossing along an underwater ridge in times of low water level. Genetic population differentiation, estimated from mitochondrial DNA and microsatellite data in six adjacent populations, was equally high across localities separated by sandy shores and along uninterrupted stretches of rocky shore. Our results suggest that ecological barriers are not required to induce philopatric behavior in Variabilichromis, and that morphological stasis persists in the face of high levels of neutral genetic differentiation.

Alleles↗

Population structure of brush-tailed rock-wallaby (Petrogale penicillata) colonies inferred from analysis of faecal DNA.

Genetic data obtained using faecal DNA were used to elucidate the population structure of four brush-tailed rock-wallaby (Petrogale penicillata) colonies located in Wollemi National Park, New South Wales. The results suggested that the four sampled colonies are genetically differentiated and do not form a panmictic unit. Based on assignment tests, approximately 5% of sampled individuals were inferred to be dispersers and both male and female migrants were detected. Multilocus spatial autocorrelation analyses provided evidence for increased philopatry among females compared to males within the largest colony in the valley. Females in close spatial proximity were more genetically similar than expected under a random distribution of females, and females separated by more than 400 m were less genetically similar than expected. In contrast, there was no evidence of a significant clustering of related males. This suggests that within-colony dispersal is male biased. We also investigated the best strategies for conserving genetic diversity in this population. All of the four sampled colonies were found to contain distinct components of the genetic diversity of the Wolgan Valley P. penicillata population and loss of any colony is likely to result in the loss of unique alleles. Conservation and management plans should take into account that these colonies represent genetically differentiated discrete subpopulations. This approach is also the best strategy for maintaining the genetic diversity of the populations in this valley.

Analysis of Variance↗

A novel mitochondrial intergenic spacer reflecting population structure of Pacific oyster.

Nucleotide sequence divergence in a novel major mitochondrial DNA intergenic spacer (IGS) of Pacific oyster Crassostrea gigas was analyzed for 29 cultured individuals within the Goseong population (Korea). A total of 7 variable sites were detected within the IGS, and the relative frequency of nucleotide alteration was determined to be 1.16%;. All alterations were due to a single nucleotide substitution, and 5 transitions and 2 transversions were observed. Among 29 specimens, only 8 haplotypes could be identified, and 6 of the haplotypes were unique to particular specimens. Pairwise genetic diversity of all 8 haplotypes was calculated to be 0.412+/-0.134 from multiple sequence substitutions based on the two-parameter model. The phylogenetic tree obtained for these haplotypes according to the neighbor-joining method illustrated a single cluster of linkages, which comprised 5 haplotypes associated with 23 specimens, while the other 3 haplotypes associated with 6 specimens were scattered. The results indicate that the IGS is higher polymorphic and thus more suitable as a genetic marker for population structure analysis of Pacific oyster than the mtDNA coding regions, such as cytochrome c oxidase I and 16S ribosomal RNA genes.

Animals↗

Population structure in the Connecticut Valley: II. A comparison of multidimensional scaling solutions of migration matrices and isonymy.

In a previous paper (Swedlund et al., 1984) we have described the population structure of the historical Connecticut River Valley of Massachusetts in terms of matrimonial migration matrices. Using procedures described by Morton (1973), Harpending and Jenkins (1974), Jorde (1980), and others the exchanges between subdivisions which make up the matrices are made column stochastic and analyzed to predict genetic kinship. Subsequently the kinship estimates within and between subdivisions can be interpreted as genetic covariance and compared to the actual geographic distances between the respective subdivisions using a principal components analysis. In the present paper we extend these results by applying nonmetric multidimensional scaling to the migration matrices, and to isonymy matrices based on the same communities. We demonstrate that the multidimensional scaling configurations of marital migration represent the actual geographic relationships between the communities quite effectively for this particular case study from historical Massachusetts. Moreover, we argue that while these migration data may provide good estimates of social and genetic exchange between the subdivisions, surname analysis may also be informative of processes not revealed in the migration matrices alone.

Demography↗

Population structure in the Mediterranean basin: a Y chromosome perspective.

The Mediterranean region has been characterised by a number of pre-historical and historical demographic events whose legacy on the current genetic landscape is still a matter of debate. In order to investigate the degree of population structure across the Mediterranean, we have investigated Y chromosome variation in a large dataset of Mediterranean populations, 11 of which are first described here. Our analyses identify four main clusters in the Mediterranean that can be labelled as North Africa, Arab, Central-East and West Mediterranean. In particular, Near Eastern samples tend to separate according to the presence of Arab Y chromosome lineages, suggesting that the Arab expansion played a major role in shaping the current genetic structuring within the Fertile Crescent.

Chromosomes, Human, Y↗

Microsatellite analysis of the population structure of a vancouver island sockeye salmon (Oncorhynchus nerka) stock complex using nondenaturing gel electrophoresis

Variation at four microsatellite loci, Omy77, Ots100, Ots3, and Ots103, was characterized for three populations of sockeye salmon (Oncorhynchus nerka) found within 50 km of each other in the Barkley Sound region of Southwest Vancouver Island, British Columbia. We used a simple polyacrylamide gel electrophoresis technique to analyze microsatellite alleles amplified from total genomic DNA extracts of liver and archived scales. Classes from several years were examined for each population, with little significant interannual variability detected. Each microsatellite locus was highly polymorphic with 10 alleles at Omy77, 12 alleles at Ots100, 16 alleles at Ots3, and 17 alleles at Ots103; average observed heterozygosities were also high at 75%, 76%, 69%, and 85% respectively. Fst values and pairwise tests indicated that the populations had significantly different allele frequencies, suggesting that the three populations are reproductively isolated. Our results suggested that microsatellite loci can be analyzed with a simple technique to reveal population structure of sockeye salmon on a small geographic scale.

Journal Article↗

Genetic population structure of Italy. II. Physical and cultural barriers to gene flow.

Three approaches were employed to evaluate the relative importance of geographic and linguistic factors in maintaining genetic differentiation of Italian populations as shown by blood groups and erythrocyte and serum markers. Genetic distances are closer to linguistic than to geographic distances. Gene-frequency change across 12 linguistic boundaries is significantly more rapid than at random locations. The zones of sharp genetic variation correspond to physical barriers to gene flow and to boundaries between dialect families, which overlap widely. However, two linguistically differentiated populations appear genetically differentiated despite the absence of physical obstacles to gene flow around them. The Po River is associated with abrupt genetic change only in the area where it corresponds to a dialect boundary. At most loci the genetic population structure seems affected by linguistic rather than geographic factors; exceptions are the systems that were subject to malarial selection in geographically close but linguistically heterogeneous localities. Gene flow appears to homogenize gene frequencies within regions corresponding to dialect families but not between them, leading to the patchy distributions of allele frequencies that were detected in an earlier study.

Alleles↗

Macrogeographic population structure in a parasitic nematode with avian hosts.

Much remains to be discovered about the population genetic structure of parasites, despite the importance of such knowledge to understanding the processes involved in the spread of drug resistance through populations. Here we present a study of population genetic diversity in Trichostrongylus tenuis, an avian parasitic nematode infecting both poultry and game birds, where anthelmintic use is common. We examined diversity of nicotinamide dehydrogenase subunit 4 (nad4) mtDNA sequences within and between seven locations: five in the UK (red grouse hosts), one in Iceland (domestic goose) and one in Norway (willow grouse). Within-UK comparisons showed high nucleotide diversity (pi=0.015, n=23) but no structure between locations (Phi(ST)=0.022, P=0.27), with over 97% of variation distributed within-hosts. The highest diversity was found in Iceland (pi=0.043, n=4), and the lowest in Norway (pi=0.003, n=4). Differentiation between countries was considerable (Phi(CT)=0.44, P<0.05), in spite of the potential mixing effects of gene flow via migrating wild hosts and the poultry trade. However, significant pairwise F(ST) values were found only between Norway and the other locations. Phylogenetic analysis provided statistical support for a separate clade for Norwegian samples only, with unresolved diversity leading to a star-shaped relationship between Icelandic and UK haplotypes. These results suggest that Norwegian T. tenuis are isolated, but that there is some connectivity between UK and Icelandic populations. Although anthelmintic resistance has not yet been reported for T. tenuis, the population structure is such that emerging resistance has the potential to spread by gene flow over a large geographic scale.

Animals↗

Abundance and population structure of ammonia-oxidizing bacteria that inhabit canal sediments receiving effluents from municipal wastewater treatment plants.

A polyphasic, culture-independent study was conducted to investigate the abundance and population structure of ammonia-oxidizing bacteria (AOB) in canal sediments receiving wastewater discharge. The abundance of AOB ranged from 0.2 to 1.9% and 1.6 to 5.7% of the total bacterial fraction by real-time PCR and immunofluorescence staining, respectively. Clone analysis and restriction endonuclease analysis revealed that the AOB communities influenced by the wastewater discharge were dominated by Nitrosomonas, were similar to each other, and were less diverse than the communities outside of the immediate discharge zone.

Ammonia↗

Impact of population structure, effective bottleneck time, and allele frequency on linkage disequilibrium maps.

Genetic maps in linkage disequilibrium (LD) units play the same role for association mapping as maps in centimorgans provide at much lower resolution for linkage mapping. Association mapping of genes determining disease susceptibility and other phenotypes is based on the theory of LD, here applied to relations with three phenomena. To test the theory, markers at high density along a 10-Mb continuous segment of chromosome 20q were studied in African-American, Asian, and Caucasian samples. Population structure, whether created by pooling samples from divergent populations or by the mating pattern in a mixed population, is accurately bioassayed from genotype frequencies. The effective bottleneck time for Eurasians is substantially less than for migration out of Africa, reflecting later bottlenecks. The classical dependence of allele frequency on mutation age does not hold for the generally shorter time span of inbreeding and LD. Limitation of the classical theory to mutation age justifies the assumption of constant time in a LD map, except for alleles that were rare at the effective bottleneck time or have arisen since. This assumption is derived from the Malecot model and verified in all samples. Tested measures of relative efficiency, support intervals, and localization error determine the operating characteristics of LD maps that are applicable to every sexually reproducing species, with implications for association mapping, high-resolution linkage maps, evolutionary inference, and identification of recombinogenic sequences.

Alleles↗

Spatial population structure in a patchily distributed beetle.

The dynamics and evolution of populations will critically depend on their spatial structure. Hence, a recent emphasis on one particular type of structure--the metapopulation concept of Levins--can only be justified by empirical assessment of spatial population structures in a wide range of organisms. This paper focuses on Aphodius fossor, a dung beetle specialized on cattle pastures. An agricultural database was used to locate nearly 50 000 local populations of A. fossor in Finland. Several independent methods were then used to quantify key processes in this vast population system. Allozyme markers and mitochondrial DNA (mtDNA) sequences were applied to examine genetic differentiation of local populations and to derive indirect estimates of gene flow. These estimates were compared to values expected on the basis of direct observations of dispersing individuals and assessments of local effective population size. Molecular markers revealed striking genetic homogeneity in A. fossor. Differentiation was only evident in mtDNA haplotype frequencies between the isolated Aland islands and the Finnish mainland. Thus, indirect estimates of gene flow agreed with direct observations that local effective population size in A. fossor is large (hundreds of individuals), and that in each generation, a substantial fraction (approximately one-fifth) of the individuals move between populations. Large local population size, extreme haplotype diversity and a high regional incidence of A. fossor all testify against recurrent population turnover. Taken together, these results provide strong evidence that the whole mainland population of A. fossor is better described as one large 'patchy population', with substantial movement between relatively persistent local populations, than as a classical metapopulation.

Animals↗