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Population Structure Among African and Derived Populations of Drosophila simulans: Evidence for Ancient Subdivision and Recent Admixture
Martha T. Hamblin and Michel Veuille
Genetics September 1, 1999 vol. 153 no. 1 305-317
Martha T. Hamblin
Laboratoire d'Ecologie-EPHE, Université Pierre-et-Marie Curie, 75252 Paris Cedex 05, France
Michel Veuille
Previous studies based on allozyme variation have found little evidence for genetic differentiation in Drosophila simulans. On the basis of DNA sequence variation at two nuclear loci in four African populations of D. simulans, we show that there is significant structure to D. simulans populations within Africa. Variation at one of the loci, vermilion, appears to be neutral and supports an eastern African origin for European and American populations. Samples from the West Indies, Europe, and North America had a nucleotide diversity lower than that of African populations at vermilion and show nonequilibrium haplotype distributions at both vermilion and G6pd, consistent with a hypothesis of recent bottleneck and possibly also admixture in the history of these populations. Directional selection, previously documented at G6pd, appears to have occurred within the coalescence time of the species, obscuring deep population history.
THE closely related species Drosophila melanogaster and D. simulans are widely used in studies of evolution at the phenotypic, genetic, and molecular levels. In spite of many broad similarities, there are important differences between these species, the causes of which are not well understood (Parsons 1975). Both species evolved in tropical Africa and have spread worldwide as human commensals in historical times (Lachaiseet al. 1988), although D. melanogaster has apparently inhabited temperate regions for sufficient time to develop significant genetic differentiation between populations from different continents (Choudhary and Singh 1987; Begun and Aquadro 1993; Benassi and Veuille 1995). In contrast, little population structure has been detected in D. simulans as assessed by studies of phenotypic variation, allozymes, and mtDNA (Hyytiaet al. 1985). However, a recent study of microsatellite variation among one African and three non-African populations of D. simulans (Irvinet al. 1998) revealed more genetic differentiation between populations of D. simulans than had been observed using allozymes.
A discordance between DNA-level and protein-level evolution is also observed in comparisons between these species: while allozyme variation is slightly higher in D. melanogaster (Choudhary and Singh 1987), DNA sequence variation is two to three times higher in D. simulans (Aquadroet al. 1988; Moriyama and Powell 1996). Similarly, the ratio of replacement to silent DNA variation is significantly higher for D. melanogaster than D. simulans (Moriyama and Powell 1996). This genome-wide discrepancy between protein and (presumably largely neutral) DNA evolution implies that selection on protein variation has been different in these species. Patterns of codon usage also vary between the species, with a higher proportion of preferred codons having been fixed in the D. simulans lineage (Akashi 1996).
Several nonexclusive hypotheses have been proposed to explain these different patterns of molecular variation: D. simulans has a different "adaptive strategy" than D. melanogaster and/or has become a cosmopolitan species more recently than D. melanogaster (Choudhary and Singh 1987); D. simulans has a larger effective population size than D. melanogaster, leading to stronger effects of weak selection (Aquadroet al. 1988; Akashi 1996). Testing of these hypotheses should also consider possible differences in population history and structure between the species. Recent demographic events, in particular, are expected to affect genome-wide patterns of variation and may cause departures from the predictions of models assuming that populations are at equilibrium with respect to migration, mutation, and drift. Recovery from such events is expected to take on the order of 4Ne (where Ne is effective population size) generations, much longer than the few hundred years since the establishment of many non-African populations of D. melanogaster and D. simulans.
Progress in elucidating population history and structure for these species has been hampered by a dearth of studies of DNA sequence variation from true population samples representing geographically diverse populations, especially African populations that are potentially ancestral. This is particularly true for D. simulans, for which DNA sequence variation has been examined almost exclusively in North American and European population samples or in worldwide collections of alleles that are inappropriate for many population genetic tests. There is only one published report of DNA sequence variation in more than one population sample of D. simulans, a study of the vermilion locus by Begun and Aquadro (1995), which showed significant differentiation (FST = 0.25) at silent sites between a North American and Central African population. Furthermore, most of the sequence variants were not shared between populations, implying that the Central African population was not ancestral to the North American one. This, in turn, implies that African populations of D. simulans are genetically differentiated if one assumes that some other African population is ancestral.
The North American vermilion dataset showed a strong haplotype structure (i.e., alleles fell into very divergent classes for which few intermediate haplotypes were observed), a pattern that has also been observed at a number of other unlinked loci in North American samples of D. simulans alleles (Begun and Aquadro 1994; V. L. Bauer and C. F. Aquadro, personal communication). Such a pattern might result from mixing of alleles from genetically differentiated founder populations in the recent past. The presence of highly divergent, geographically restricted mitochondrial DNA (mtDNA) lineages in this species (Solignac and Monnerot 1986; Satta and Takahata 1990) lends plausibility to this hypothesis. If admixture has occurred, the observed level of variation might not be an accurate indicator of the long-term effective population size of the ancestral populations. Because differences in effective population size have frequently been invoked to explain differences in patterns of molecular variation between D. melanogaster and D. simulans from derived populations (Aquadroet al. 1988; Ohta 1992; Akashi 1996), the question of population admixture needs to be resolved.
The goal of this study was to use nuclear DNA sequence data to assess the level of population differentiation of African and non-African populations of D. simulans and to begin to reconstruct their evolutionary histories. In particular, we wanted to test the hypothesis that divergent alleles from derived populations reflect population structure in the ancestral populations from which they were founded. The possibility of population admixture has been raised before to explain divergent lineages in worldwide collections of alleles of D. simulans (e.g., Hassonet al. 1998), but this question can be directly addressed only by studying true population samples. We surveyed two unlinked regions on the X chromosome for which some sequence polymorphism data were already available: vermilion (Begun and Aquadro 1995) and glucose-6-phosphate dehydrogenase (G6pd; Eaneset al. 1996). For each locus, we sequenced a relatively short region (∼700 bp) containing a reasonable number of segregating sites and showing the pattern of divergent haplotypes described above.
Population samples
Population samples: Table 1 shows the dates and locations of the collections of D. simulans used in this study. Samples were obtained using attractive baits. Wild-collected flies were used to establish isofemale lines (Kenya, Tanzania, Antilles, Zimbabwe) or extract chromosomes using attached-X lines (Cameroon) or were frozen immediately in the laboratory (Italy). Flies from isofemale lines were frozen within 3 mo after trapping, except for Kenya (8 mo) and Antilles (1 yr). For vermilion, we included the sequences from Raleigh, North Carolina (United States), published by Begun and Aquadro (1995), because this sample comes from a geographic region not represented in our data.
DNA methods: DNA was prepared from single male flies by the method of Gloor et al. (1993). An 809-bp product was amplified from the vermilion locus using primers corresponding to bases 602–622 (forward) and 1410–1390 (reverse) of GenBank accession no. U27204. A 769-bp product was amplified from the G6pd locus using primers corresponding to bases 917–939 (forward) and 1685–1664 (reverse) of GenBank accession no. L13876. PCR products were separated on 1.2% agarose gels and the desired bands were cut out and purified using the QiaexII kit (Qiagen, Valencia, CA). DNAs were sequenced manually using the Thermosequenase kit (Amersham, Arlington Heights, IL). The PCR primers were used as sequencing primers, with the exception of the reverse sequencing primer for vermilion, which was an internal primer corresponding to bases 1376–1357. DNAs were sequenced on one strand except for a small area of overlap in the middle of the fragment. GenBank accession numbers for the vermilion sequences are AF149122–149191; accession numbers for the G6pd sequences are AF148146–148207.
Data analysis: The program DnaSP (Rozas and Rozas 1997) was used to obtain summary statistics of sequence polymorphism within populations, Tajima's D (Tajima 1989), and divergence between populations. The fixation index (FST) was calculated according to Hudson et al. (1992b) and the probability of panmixis was determined using the method of Hudson et al. (1992a).
Tests of haplotype number and haplotype diversity were conducted using the method of Depaulis and Veuille (1998). For G6pd, a recombination rate of Nr/bp = 0.005 was used; for vermilion, Nr/bp = 0.01. Note that for vermilion, estimates of r based on genetic data from D. melanogaster are 2 × 10i–6/bp (Searleset al. 1990) and that if N = 10–6, Nr = 2. (Rates of recombination in this region of the X chromosome are very similar in D. simulans and D. melanogaster; Sturtevant 1929.) Limitations of computational power prevented us from running the simulations under higher values of Nr; our P values therefore are conservative. Tests were one-tailed because the hypothesis predicted a departure in the direction of too few haplotypes.
—Polymorphism at vermilion in D. simulans. The numbering system is the same as in Begun and Aquadro (1995), based on the sequence of D. melanogaster as reported in GenBank accession no. M34147. cons, the most common base at each segregating site; i, intron; s, synonymous site. a The U.S. alleles are from Begun and Aquadro (1995).
DNA sequence variation: For estimation of historical patterns of gene flow between populations of D. simulans, we collected DNA sequence data from two unlinked loci on the X chromosome in samples from four continental African, one West Indian, and one European population. We determined the sequence of 728 bp of the vermilion gene in 10–13 individuals from each of the six populations. This 728-bp region spans three small introns, which total 186 bp in length. Haplotypes at vermilion, including the corresponding vermilion data for the United States (Begun and Aquadro 1995), are shown in Figure 1.
We also determined the sequence of 700 bp of the third exon of the G6pd gene in 10–12 individuals from each of the four African populations. Due to problems with amplification in the Italian lines, we have data from only five sequences from Italy, which we combined with the four sequences from France published by Eanes et al. (1996) to create a European sample. The four haplotypes from Italy are very similar to the three haplotypes from France, suggesting that the sample has not been biased by the amplification problems and that it is appropriate to combine the samples. Haplotypes at G6pd are shown in Figure 2.
Summary statistics for both loci are presented in Table 2. Note that these data are not appropriate for comparing variation between loci nor are they appropriate for making inferences about absolute effective population size, because the regions sequenced were not chosen at random; the goal was to understand population structure, not to estimate 4Neμ. Nevertheless, these statistics are appropriate for use in comparing variation between populations at a particular locus. At vermilion, the samples from African populations, particularly Tanzania and Kenya, are the most variable. This pattern is consistent with the hypothesis of Lachaise et al. (1988) that the ancestral population that became D. simulans inhabited coastal eastern Africa and/or the islands in the Indian Ocean. At G6pd, however, this pattern is not observed: the populations differ very little in the number of segregating sites, and Tanzania has the fewest. The ability to detect differences between populations is weaker at G6pd because there are about three times fewer polymorphic sites in this region, but other analyses suggest that the evolutionary history of the G6pd region has indeed been different from that of vermilion (see below).
—Polymorphism at G6pd in D. simulans. The numbering system is the same as that in Eanes et al. (1996). cons, the most common base at each segregating site. All sites are synonymous coding sites. a The alleles labeled "it" are from Italy. The alleles labeled "mt" are from France (Eaneset al. 1996).
Genetic differentiation between populations: We tested the null hypothesis that our population samples were drawn from a single panmictic population using the method of Hudson et al. (1992b). Estimates of FST were calculated according to Hudson et al. (1992a). In cases where the sample size differed between populations, the average pairwise difference was weighted by the sample size. The significance of FST was determined by resampling the data 1000 times. Populations were tested in all pairwise combinations, and data from the two loci were analyzed separately (Table 3). Because multiple nonindependent tests were performed, we have not chosen a significance threshold for this analysis. Instead, we make qualitative comparisons of P values.
At the vermilion locus, FST is essentially zero between the samples from Tanzania and Kenya, which were collected only ∼200 miles apart. The other distances, and most of the other FST's, are much larger: 13 of the 15 comparisons involving Antilles, Zimbabwe, and Cameroon have P values ≤0.01. P values for FST's involving Tanzania, the United States, and Italy are generally low (0.01 < P < 0.1) but probably not significant. Thus, our samples seem to fall into four genetically distinct groups: Zimbabwe, Cameroon, Lesser Antilles, and Tanzania/Kenya/Italy/United States. The Antilles sample, dominated by seven copies of a unique haplotype, is quite different from all other populations. Again, the relationships between populations at the G6pd locus are different from those at vermilion: Cameroon appears to be differentiated (P ≤ 0.005) from all other populations except Tanzania, but most other comparisons have much larger P values.
Summary statistics of population variation at vermilion and G6pd
Haplotype tests: Haplotypic diversity is quite high; at vermilion, there are 41 different haplotypes, of which 27 occur only once in the total of 82 sequences. At G6pd, there are 28 haplotypes, 16 of which are unique, in a total of 66 sequences. However, inspection of the data in Figures 1 and 2 reveals that all the samples that do not come from Africa contain multiple copies of a few highly divergent haplotypes: Italy/Europe and Antilles for both loci and the U.S. sample of vermilion. One prediction of a hypothesis of recent population admixture is that the number of haplotypes will be smaller than expected in a population at mutation-drift equilibrium, given the observed number of segregating sites, similar to the pattern produced by an old balanced polymorphism. We tested the observed number of haplotypes in each of our samples against the expectation of a neutral, equilibrium model, conditioned on the number of segregating sites (S) and the number of sequences surveyed (n) (Depaulis and Veuille 1998). Recombination was included in the model (see materials and methods), because both loci in our study experience significant amounts of recombination. All the samples from Italy/Europe, Antilles, and the United States show a significant departure from the expectation for at least one of the tests (Table 4), while none of the tests of the samples from Africa is significant. The P values for the vermilion sample from Zimbabwe, however, are much lower than those for the three other African populations.
Estimates of FST in D. simulans populations
The significance of the haplotype tests for the vermilion data may be somewhat overestimated, because we chose to survey the 5′ half of the gene because of its higher level of linkage disequilibrium. The entire region surveyed by Begun and Aquadro (1995) does not contain significantly too few haplotypes, although the haplotype diversity is still significantly lower than expected (Table 3). However, it should be noted that the rate of recombination used in the simulations was considerably lower than the rate of recombination estimated for the vermilion region (by ∼100-fold if we accept the estimate of 2 × 10–6/bp from Searleset al. 1990) due to limited computational ability. A more realistic recombination rate would have decreased the P value by an unknown amount. At G6pd, it is unlikely that including the entire gene would have changed the outcome of the tests, because there is little variation in the remaining sequence, and linkage disequilibrium was high throughout the gene (Eaneset al. 1996). Note also that the significant result for the European sample is not due to the combining of alleles from Italy and France; the divergent haplotypes are not associated with different locations.
Tests of haplotype number and haplotype diversity
Because the hypothesis of population admixture predicts a similar pattern throughout the genome, we conducted the same tests of haplotype number and diversity for all the other surveys of DNA polymorphism that we could find in the literature for population samples of D. simulans; unfortunately there are very few (Table 4). A four-cutter restriction site survey of the phosphogluconate dehydrogenase region (Pgd) from North Carolina (Begun and Aquadro 1994) gave very significant results; a sequence survey of the glucose dehydrogenase (Gld) region from the same population did not, although this sample did have a significantly positive Fu and Li's D statistic due to a complete lack of singletons (Hamblin and Aquadro 1996). Surveys of the yellow-achaete region from Europe and the United States (Martin-Camposet al. 1992; Begun and Aquadro 1991) as well as the supressor of forked region (Langleyet al. 1993) had too little variation to be informative.
Phylogeny of G6pd alleles: Because the FST analysis pools all alleles in a sample to produce one statistic of distance, it is a poor reflection of the complexity of these particular samples, and the relationships of these populations are problematic (see discussion). We therefore wanted to estimate phylogenies of alleles using parsimony to see where these divergent alleles arose with respect to the potentially ancestral African populations. Because both loci are from recombining regions of the nuclear genome, simple bifurcating phylogenies cannot be reconstructed.
For G6pd, it was possible to construct a network (Figure 3) using all 22 segregating sites and all but two haplotypes (Cameroon 3 and 12). We also included the alleles from the United States and Mexico surveyed by Eanes et al. (1996). There are two haplotypes (the consensus or "dot" and the Tanzania 9-type), separated by two steps, that are found in all three eastern African populations. Many of the Cameroon alleles are located between these two haplotypes in the middle of the network. Most of the Kenyan alleles are more closely related to the dot haplotype. There are two clusters of alleles from the derived populations at opposite ends of the network, separated by between five and nine steps. Both clusters are associated with alleles from Zimbabwe, so the network provides no evidence that they have different geographic origins.
Informative site analysis: Because of the large amount of recombination in the history of the vermilion alleles, we tried to construct phylogenies using short regions (Templetonet al. 1992). This approach was not successful, however; the regions for which it was possible to construct a network with little recombination had few informative sites, while even slightly larger regions produced extremely reticulate phylogenies (data not shown).
—Haplotype network of alleles at G6pd. Small black circles represent hypothetical intermediates not observed in our samples.
Instead, we did a site-by-site analysis of what we call "informative segregating sites," i.e., those sites that are segregating in at least two populations, but not in all of them. There were a total of 35 informative sites. For each pair of populations, the number of times they shared one of these informative sites was counted. The resulting counts were then divided by the average total number of segregating sites in the two populations, giving the fraction of the polymorphism shared by the two populations that is not present in all populations. The results are presented in Table 5. There are six comparisons in which >30% of the polymorphism is shared and informative. One of these, not surprisingly, is Kenya/ Tanzania. What is not obvious from the FST data is that Zimbabwe and Cameroon have the highest proportion of shared informative polymorphisms and that the other most similar pairs all involve comparisons among Italy, Antilles, the United States, and Zimbabwe.
At G6pd, there is also a high proportion of shared informative polymorphisms among Zimbabwe, Antilles, and Europe, none of which shares any informative sites with Tanzania. These associations can be seen in the network in Figure 3, where several derived haplotypes are shared among these populations.
This study, the first to compare African populations of D. simulans at the DNA sequence level, has revealed substantial genetic differentiation among those populations at the vermilion locus. These findings contrast with patterns of allozyme and mitochondrial DNA variation, which show little geographic structure. Haplotype structure in the non-African population samples at both vermilion and G6pd departs significantly from the expectation under a neutral, equilibrium model, suggesting a history of bottleneck (i.e., founder effect) and possible admixture in these recently established populations. At the vermilion locus, the three non-African population samples also have lower genetic diversity (average θ = 7.73 ± 0.692) than the four African samples (average θ = 12.03 ± 2.656), consistent with the inference of a founder effect.
Variation at the G6pd locus is not lower in the non-African samples, and there is much less evidence for population structure among all the samples. If both loci were neutral indicators of population sizes and structure, we would expect relative distances and diversity among populations at the two loci to be the same. While such a discrepancy might be due to chance, we believe that it is due primarily to past episode(s) of positive selection at G6pd in D. simulans as discussed in Eanes et al. (1996). In the following two sections, we present our case that the vermilion data are likely to reflect much more closely the population structure of this species in Africa.
Reduced coalescence time at G6pd: Of the 21 amino acid differences fixed between D. melanogaster and D. simulans at G6pd, 15 have become fixed in the D. simulans lineage, although only 1 has been fixed since the divergence of D. simulans and D. sechellia (Eaneset al. 1996). A significant McDonald-Kreitman test supports the interpretation that many of those amino acid differences have been fixed due to positive selection (Eaneset al. 1993). The coalescence time of D. simulans is very long and probably goes back before the simulans-mauritiana-sechellia split because D. simulans and D. mauritiana share polymorphisms (Kliman and Hey 1993). An episode of positive selection at a locus is expected to reduce the coalescence time and eliminate evidence of prior population history for that region. Using the method of Hudson et al. (1987), we found that the levels of polymorphism and divergence at G6pd in our samples were concordant with those at vermilion. One possible interpretation of this result is that the episodes of positive selection at G6pd happened sufficiently long ago that they have no detectable effect on present-day levels of polymorphism. However, the lack of a departure could also be due to the nonrandom nature of our sample: we deliberately collected sequence data from the more variable 3′ region of G6pd. The unsurveyed 5′ end of the gene was much less variable in the alleles surveyed by Eanes et al. (1996). While it therefore remains unproven that there is a significant hitchhiking effect at G6pd in D. simulans, this is clearly a strong possibility that must be kept in mind in the interpretation of these data. If the migration of D. simulans out of Africa has been very recent, however (i.e., more recent than selection at G6pd), our G6pd data are still useful for inferring the relationship between African and non-African populations (see below).
Shared informative sites
Variation at vermilion reflects population history: In contrast to G6pd, the vermilion locus appears to have been evolving under purifying selection since the D. melanogaster-D. simulans split. There is only one amino acid difference (a serine to threonine change) between the genes in the two species (Begun and Aquadro 1995). Rates of recombination in the vermilion region are high (2.2 × 10–6/bp/generation in D. melanogaster), so evolution at vermilion is substantially decoupled from evolution at linked loci. The D. simulans population samples surveyed by Begun and Aquadro (1995) showed no departure from neutrality in the original analyses or in a further analysis of these data using several heterogeneity tests (McDonald 1998). In our own data, Tajima's D's for several of the vermilion samples, although not significant, are large and negative. This is most likely due to recent population expansion, as the high levels of variation observed in most of these samples argue against other explanations such as a bottleneck or selective sweep. It therefore seems reasonable to use our vermilion data to make inferences about the history of African populations of D. simulans, keeping in mind that these inferences need to be confirmed by data from other, independent loci.
Relationships among African populations: At vermilion, Tanzania and Kenya have the highest levels of polymorphism and haplotype diversity and are not significantly differentiated from each other. Collectively, they contain 18 polymorphisms that are not observed in any other population and have the lowest FST's in comparisons with the non-African populations. The other two African populations have lower levels of polymorphism and are genetically differentiated from Tanzania/Kenya and from each other. The accumulation of genetic differences between distant populations suggests that these populations have not been very recently established. These observations are consistent with the biogeographic hypothesis of Lachaise et al. (1988), which proposes that early populations of D. simulans were restricted to coastal eastern Africa and islands in the Indian Ocean until the end of the Pleistocene (Lachaiseet al. 1988), at which time they expanded into central Africa. The time of the putative expansion corresponds to ∼0.1 Ne generations ago (assuming Ne ≥ 106 and 10 generations/year), far less than the time required to reach mutation-drift equilibrium.
The higher level of genetic variation in the Tanzania/ Kenya samples suggests that the population in this region may be older and/or larger than the others. This interpretation, however, assumes that the observed differences in levels of polymorphism reflect real differences in long-term effective population size. Several issues need to be considered here; first is the variance associated with estimates of 4Neμ. While the stochastic variance of independent samples of a single, nonrecombining locus is very large, note that substantial recombination at vermilion results in estimates of 4Neμ that represent several partially independent evolutionary trajectories, substantially reducing the variance (Pluzhnikov and Donnelly 1996). Even if we assume free recombination, however, none of our estimates of 4Neμ is significantly different from any other.
Perhaps more importantly, the populations surveyed in this study share a recent common ancestry in a species with a very long coalescence time (4Ne generations, where Ne ≥ 106). Many alleles within these populations are likely to share a most recent common ancestor with an allele from another population, and much of the genealogy of these alleles probably took place prior to the divergences of these populations from the ancestral population(s). Wakely (1996) has shown that the variance of pairwise differences within and between samples decreases with the decrease in time of population splitting (T). At very small T, the variance converges on the value expected in a single, randomly mating population. In other words, the variance of θ among recently separated populations approaches the sampling variance of a single population.
Another issue is that, in populations recently descended from a common ancestral population, estimates of 4Neμ will predominately reflect ancestral population size and contain little information about the sizes of the descendant populations. This is particularly true if these populations have experienced rapid expansion, which slows down genetic drift (Nichols and Beaumont 1996). Instead, the observed differences in levels of polymorphism among these population samples may reflect differences in the amount of genetic variation sampled at foundation, which may in turn reflect proximity to an ancestral population. Interestingly, the African samples show a significant relationship between levels of genetic diversity and distance from Tanzania (r2 = 0.948, P = 0.026). This strong pattern in the data suggests that the observed differences in variation are not simply random fluctuations about a common value of 4Neμ. When the distance from either Zimbabwe or Cameroon is used as the independent variable, the P values are 0.910 and 0.199, respectively. When the non-African samples were included in the analysis, their much lower variation and long distances result in a significant regression regardless of which African location was the focus, showing only that variation in Africa is higher than variation out of Africa.
Inferences of gene flow based on shared informative sites: As discussed above, biogeographical analysis suggests that continental African populations of D. simulans may have undergone an expansion on the order of 0.1Ne generations ago. Nichols and Beaumont (1996) have shown that exponential growth following foundation has the effect of reducing genetic drift: distributions of genetic variation become "frozen in place" and reflect the foundation event and early migration events, when the population is still small, much more than they reflect subsequent gene flow. This means that our estimates of FST also reflect early events and may be misleading when used to make inferences about recent gene flow.
To try to separate foundation events from subsequent gene flow, we identified a subset of polymorphic sites that are segregating in more than one sample but not in all samples as informative segregating sites. We have not yet explored the expected properties of informative sites, so this analysis is only qualitative and suggestive, but it reveals some interesting properties of the dataset that are not revealed in any of the other analyses and may reflect underlying processes different than the FST analysis.
The FST statistics indicate rather similar levels of differentiation among all the African populations except between Kenya and Tanzania, presumably because all these populations share an ancestral set of polymorphisms that have been sorting for similar amounts of time. For example, FST at vermilion between Zimbabwe and Tanzania is 0.122 (P = 0.007) and FST at vermilion between Zimbabwe and Cameroon is 0.159 (P = 0.014). Yet Zimbabwe and Tanzania share only ∼14% of their informative sites, while Zimbabwe and Cameroon share ∼40%, the same proportion as Kenya and Tanzania (Table 5). The high proportion of informative sites shared by Zimbabwe and Cameroon suggests that gene flow between these populations has been significant. Similarly, FST is very high between Antilles and the United States (0.238; P < 0.000), but shared informative sites are high (30%).
Conversely, FST's at vermilion are much smaller in comparisons between Tanzania, Italy, and the United States than in comparisons between Zimbabwe, Italy, and the United States. However, there are more derived polymorphisms shared among Italy, the United States, and Zimbabwe than among Italy, the United States, and Tanzania (Table 5). This suggests that Zimbabwe is as likely as Tanzania to be ancestral to these recently established populations or that alleles from another population not sampled in this study have migrated differentially.
As we would expect, at G6pd, where much of the shared ancestral polymorphism appears to have been eliminated during episodes of directional selection, the FST and shared informative sites analyses are much more concordant. Both indicate a closer relationship among Zimbabwe, Europe, and Antilles than among Tanzania, Europe, and Antilles, in agreement with the informative sites analysis at vermilion.
Interaction of selection, migration, and recombination at G6pd: If we did not have independent evidence of positive selection at G6pd (Eaneset al. 1996), the discordance between geographic patterns of variation at G6pd and vermilion would make interpretation of our data for both loci problematic. In contrast to the distribution of variation at vermilion, the G6pd alleles from Tanzania have the least variation of all five samples (Table 2). At G6pd, we see three private polymorphisms each in Cameroon and Zimbabwe, one in Kenya, and none in Tanzania. At vermilion, where there is no evidence of selection, Tanzania and Kenya each have five private polymorphisms, while Zimbabwe and Cameroon each have only one.
Positive selection at the G6pd locus has most likely reduced the coalescence time for this region such that it no longer reflects the early population history of D. simulans in Africa. In this case, the pattern observed at G6pd reflects the impact of directional selection on a recombining locus in a geographically structured species. A selectively favored mutation will go to fixation most quickly within the population in which it arises, in a process that may approximate a simple selective sweep model. Migration to distant populations will be slower and may allow opportunities for recombination during the fixation process, such that more variation may be preserved in those distant populations. Many different patterns of variation across populations could result from such a process, depending on the strength of selection, rate of recombination, rate of migration, and effective population size. The fact that variation at G6pd is lowest in Tanzania and is not reduced in the derived populations is one such outcome and is not inconsistent with the vermilion data, given that there is strong independent evidence for directional selection at this locus.
The hypothesis of population admixture: Our data indicate that populations of D. simulans in Europe and America are young and far from equilibrium. At vermilion, Italy, Antilles, and the United States have about half as much variation as Tanzania and Kenya. The five samples of D. simulans from non-African populations all show a deficiency in haplotype number and/or diversity. None of the eight samples from Africa shows such a departure (although the P values for the Zimbabwe samples are low). At vermilion, the exact significance values of these haplotype tests are somewhat compromised by the fact that the region surveyed was not chosen at random (see results), but the difference in results between African and non-African populations is nevertheless real. Unlike the FST analysis, both vermilion and G6pd give the same results in these tests, suggesting that the phenomenon responsible for these unusual haplotype structures is more recent than any episode of selection at G6pd. The fact that we observe the same significant haplotype structure at three out of four unlinked loci (Table 4) also suggests that this is a population-level phenomenon rather than multiple instances of diversifying selection.
The unusual haplotype structure and reduced variation provide strong evidence for a founder event in non-African populations as was inferred from the microsatellite survey of Irvin et al. (1998). The additional question of admixture is not resolved by our data. However, there are haplotypes observed at high frequencies in the non-African samples that are not present in any of the 46 alleles surveyed from the African populations. For example, the Italy 3 haplotype is present in 10 out of 36 non-African alleles, while the Antilles 3 haplotype is present seven times. These haplotypes, along with the U.S. 14 type, harbor five polymorphisms (at positions 1214, 1225, 1454, 1461, and 1654) that are not segregating in the 46 African alleles. These "non-African" variants may have come from some divergent African population not included in our survey. The most likely sources of divergent alleles are the subpopulations associated with the mitochondrial races siI, siII, and siIII (Baba-Aïssa and Solignac 1984; Solignac and Monnerot 1986), which have distinct lineages over 1 million years old (Satta and Takahata 1990). In worldwide continental populations of D. simulans, there is no evidence for admixture based on mtDNA haplotypes, which are uniformly of the siII type; this may be a consequence of natural selection against the siI and siIII mtDNAs in these populations (Randet al. 1994; Ballardet al. 1996). Although mtDNAs show evidence of admixture in only a few isolated populations (e.g., Madagascar, the Seychelles), worldwide patterns of neutral nuclear DNA variation may reveal the contributions from these ancient lineages in a way that allozyme variation does not.
Evidence of admixture should be reflected in patterns of haplotype structure throughout the genome, although this evidence will decay at different rates at different loci due to differences in recombination. Selection in a new environment could also eliminate evidence of admixture. At the two loci we have studied, as well as at Pgd, haplotype number and/or diversity are inconsistent with the equilibrium expectation; data from the Gld locus do not show such a departure. Clearly, more data from other loci are needed to test this hypothesis.
Conclusions and implications for population genetic studies:
Populations of D. simulans in continental Africa are genetically differentiated and therefore not very young, although they are probably still not at equilibrium.
Non-African populations of D. simulans are very young, far from equilibrium, and have experienced a bottleneck during their foundation.
Compelling, although not conclusive, evidence suggests that there has been admixture among genetically differentiated lineages in population(s) ancestral to European and American populations.
Estimates of nucleotide diversity in non-African populations do not appear to be inflated but are nonetheless unlikely to reflect long-term effective population size due to the combined effects of bottleneck and possible admixture. Ratios of silent to replacement variation might also be affected by a bottleneck if frequency distributions of those variants are different (i.e., if one or both are not neutral). For these reasons, non-African population samples of D. simulans are unsuitable for testing models that assume mutation-drift equilibrium. African populations of both D. melanogaster and D. simulans are probably more suitable for testing population genetic models but may also violate assumptions of equilibrium. In addition, comparisons between the species will need to be put in the context of their evolutionary histories, which may be quite different.
We thank Frantz Depaulis for providing the computer programs for the population subdivision test and the haplotype tests and for helpful discussions; Chip Aquadro for providing office space during preparation of the manuscript; the Aquadro lab group and two reviewers for comments on the manuscript; and Walt Eanes for editorial assistance. M.T.H. was supported by a Chateaubriand Fellowship from the French Embassy to the United States. M.V.'s research is supported by CNRS-Unité Mixte de Recherche 7625.
Communicating editor: W. F. Eanes
Received November 11, 1998.
Copyright © 1999 by the Genetics Society of America
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I wrote this piece in 2006 upon my return from my first post-war visit to Lebanon. Starting with the 2006 war I developed what you can safely call a Lebanon obsession. It was apparent in everything I thought or spoke about. Until then I still hadn't known much about the country beyond the books I read or the things I heard. I was specifically interested in what many of us in the Arab world call Lebanon's "culture of resistance," which took many forms throughout its turbulent modern history. The closer I got to the people the more specific the obsession became. I became more obsessed with Hizbullah and its people, that revolutionary Shiism that is the eternal fuel that prepares its people to sacrifice, any sacrifice. As intimidating as that may be to some, my humble experience has taught me what I will try to reveal in my chronicle: they are very common people whose experience is not at all different to that of any people subjected to war and invasion. They just want to live free, even if they have to die for it. And so are most of the Lebanese.
My feelings were mixed. I could not decide why I was so happy as I looked out the plane window and began to see the beautiful mountains as they hugged the Mediterranean coast. All those buildings and homes clustered next to each other on mountains and hills, forming Beirut, one of the Arab world's most beloved cities.
I was headed to Lebanon on November 16, 2006, to participate in a conference that gathered hundreds of activists and media professionals from all over the world who came together in support of the resistance of Lebanon in all its forms.
When I made the decision to revisit Lebanon after the war, I wanted to make sure I remained as detached and professional as I could, not just driven by emotion. I struggled with this equilibrium for a long time. I doubted myself all along.
But I couldn't contain myself as Beirut became more and more visible through the window. This was a reunion with a city I had seen and experienced before only once but had read so much about. It was also a first encounter with a wounded city that only recently came out of a vicious war that took more than 1400 civilian lives. I didn't know what the buildings would look like, and most importantly, I didn't know how the people would receive an Egyptian, a citizen of a country whose government didn't do much beyond standing by and watching as destruction was eating its way throughout the country. Would they have changed? Would they have overcome the pain already, ready to recall it to a stranger like me, trying to see in reality what the war I was following minute by minute has brought Lebanon to?
After more than 30 years of covering the Middle East, not even British journalist Robert Fisk could hide his emotions when writing about Lebanon. In his famous book Pity the Nation, he lamented a country he saw tearing apart for long years, losing much of its beauty to conflicts often meant to serve interests of outside powers.
But Lebanon's tragedies have given it a special enduring character. Suffering has given it a spirit of perseverance that finds breath even through smallest holes of hope. After the recent July 2006 war, I discovered that death and destruction have bred life in an ironic twist of fate, showing only the proud side of a nation quickly picking up the pieces, never looking back except to find its way forward.
It was Ali*, an acquaintance of a friend of mine I'd heard so much about and learned about his ground stories of the war. He was rushing the wounded fighters and literally sweeping pools of blood along the streets of Bint Jbeil — the southern Lebanese town that is considered by Hizbullah the "capital of the resistance," and that witnessed the fiercest fighting during the war.
I used to listen to his stories through my friend and follow them so closely during the war that I had a vision in my head of how he must have looked like. And I wasn't mistaken.
I was so overwhelmed by the man's kindness that half of my talk was thanks and praises. Such is the way with the Lebanese no matter what their background: They smother you with hospitality and kindness and render you absolutely speechless. In this trip I realized that the war hasn't changed them one bit.
With each street we passed through, my heart embraced this courageous city. I could not help stare at each corner, afraid I would miss an inch. The airport road was a celebration of Hizbullah's victory in the war. The streets were filled with gigantic posters of the Secretary General Sayyid Hassan Nasrallah, each poster quoted a different statement he gave in one of his speeches.
More than any other Arab city, Beirut's streets speak of its people and their character. They speak of the nation's sorrows and triumphs. You don't need to attend lectures or read essays on the nature and spirit of Lebanon. Just walk down one of its streets.
Ali and his wife invited me for dinner. I dove into some of the best Lebanese dishes as I heard unspeakable stories of how they were separated when the war erupted. Sometimes what you hear is too devastating that any words of consolation can appear trivial. So I focused on my food as I listened to how Ali's wife and the children were caught in Bint Jbeil in the South, unable to reach him in Beirut. They remained hiding for 10 consecutive days with barely any food. She told me of how they would spend each night in a different hiding place, and how miraculously each place was bombed only after they had left it.
Shortly afterwards we were joined by a young man, a close relative of Ahmad Qassir** He was introduced to me in two small sentences: "Akram* is a graphic designer. He is with the resistance." And I was left with that.
So much has happened in each Arab country that many of us now feel separated by ages of identity formation, hardship, and struggle. As an Egyptian born in the 70s, I grew up in the "Camp David era," where Egypt supposedly brought its long years of confrontation and war with Israel to an end. I grew up hearing about Palestinian suffering and watching the Lebanese lying in pools of blood on the television screen as I sat comfortably in the safety of my home. I had no idea that with years of suffering generations are brought up with a cause, with a clear mission that shapes everything a person says and does. It is so entrenched in a person's making that it becomes natural, expected.
So Ali introduced Akram in those two sentences so casually to me and went on to discuss the next topic, leaving me still trying to grasp the magnitude of those two sentences and what they implied about Akram's life and upbringing. I was looking at a 23-year-old professional young man who almost literally designs graphics for publications during the day and gets trained by Hizbullah at night.
Remembering the information and accounts I heard about Hizbullah resistance in Lebanon, it is not uncommon to run into fighters everyday as you talk to a shopkeeper, a taxi driver, or any other Lebanese citizen. And Akram was no exception.
Akram listened more than he spoke and he occasionally gave me a warm welcoming smile. As I turned his smile back I couldn't help thinking about many youth back at home and how they felt lost and frustrated, unable — and in many cases unwilling — to channel the energy they have in anything fruitful. Yes. We've been surpassed by so much. They've certainly come a long way over here. They lived danger and faced it, they knew the true size of it, they grasped the true meaning of life and held on to it dearly and survived, while we remained in our shells, looking at the outside world through our television screens, fearing what we chose to keep unknown.
Many Arabs hold a stereotype of Lebanon being the land of blond women wearing bikinis. I remember when I was going on my first visit to Lebanon how I received a few comments on how I was heading to the "Paris of the Middle East." I had known some people from Lebanon who did not relate, even remotely, to this stereotype. The Lebanese in my mind have always been good-natured people who appreciated a joke even in the darkest of times. They always rose from the ruins of conflict and war to go about their daily lives, seeing beauty in everything, appreciating what they had, and always looking at the full side of the glass. The Sunnis and Shiites that I met were committed Muslims with a clear mission guiding them in almost everything they said and did. Life to them is a means to an end so alive in their minds, so real that nothing could distract them from it.
As the night was over and they drove me to the hotel, I stored the memory as I tried to focus on my duties the next day, the first day in the conference where I was supposed to give a presentation on IslamOnline.net's coverage of the Lebanon war.
*I changed all the names of persons used in this article for their privacy.
**Ahmad Qassir is the first fighter who carried out a martyrdom operation against the Israeli occupation in 1982, killing 17 Israeli soldiers. Since then, that day has been celebrated by Hizbullah each year as Yom Al Shahid (The Day of the Martyr).
This entry was posted on November 17, 2009, 12:13 am and is filed under Uncategorized. You can follow any responses to this entry through RSS 2.0. You can leave a response, or trackback from your own site.
It's good to know about how life is goes on in Lebanon, cause, for me, it's unimaginable and may be complicated to understand how the Lebanese can go along with this chaotic life politically, economically, socially….. But what really caught my eyes, is the gigantic poster of Sayyid Nasrallah everywhere… I guess it's trouble, cause it magnifies his power and his image more than what really is… like what happens with all the Arab leaders..
But I love the rest and looking forward the part 2.
Thank you so much for your comment. The truth is that Lebanon is quite unique in the Arab world, there's just no state quite like it. I suppose the cultural and political diversity, as much as it has the ability to fragment the society, it has also enrich it in many ways.
As for the big poster you see of Sayyid Nasrallah, you see it only in areas that are predominantly Shiite or somehow fall within the realm of Hizbullah. If you go to other places in Western Beirut, for example, you start seeing posters of Rafik Hariri and his son Saad, who is now the prime minister. And if you move to certain predominantly Christian areas (not all Christian areas since they too have their differences) you start seeing posters of Pierre Jmeyil, etc. It's many worlds living together. So you can't quite compare Beirut to any Arab city.
Thanks, Miri, for your comment. It never fails to mesmerize me how war often brought out the best in many people and allowed them to experience a true liberty in thought and in spirit, something they would not otherwise have experienced. It's always when a person see death, faces it, or even challenge it that they begin to understand the true value of their lives and what they stand for. Of the people that I have spoken to or the examples of the people you mention, I feel that they all have one thing that binds them: it's this clear vision that allows them to see things with their true colors, and maybe call them by their names if they have to.
lol!!!! i also experienced a lebanon obsession after i got back to nyc in dec 2006. for 12 days in december i went to the rallies against the western backed sinora gov't .
i couldn't stop thinking about my time there. it got to the point that i when i was day dreaming about lebanon my friends would say, "your thinking about lebanon again arn't you?" of course they were right!
please let me know when the next conference is, i would love to go!


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text:
Interview for RollingStone • Tuesday, June 7, 2016
Paul McCartney Talks Summer Tour, New 'Pure McCartney' Comp
Press interview • Interview of Paul McCartney
Interview by:
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Timeline More from year 2016
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Previous interview 2016 • Paul McCartney interview for paulmccartney.com
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Songs mentioned in this interview
Ob-La-Di, Ob-La-Da
Officially appears on The Beatles (Mono)
Temporary Secretary
Officially appears on McCartney II
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The interview below has been reproduced from this page . This interview remains the property of the respective copyright owner, and no implication of ownership by us is intended or should be inferred. Any copyright owner who wants something removed should contact us and we will do so immediately.
"It's like the Bob Dylan thing, isn't it? The 'Neverending Tour,'" Paul McCartney says with casual good cheer, chatting over the phone between show dates. This is how he likes the road these days, a few months on, a few months off. Last year was his 27-date Out There Tour, and now he's traveling the U.S. for One on One, playing marathon sets of solo songs and Beatles classics.
"You're putting a few new numbers in, changing the presentation a little bit, just so anyone who saw that tour and wants to come to this one isn't bored," he says. "We just switch it up and make some changes. Then you're allowed to call it something else."
McCartney and his band will be on the road through the summer and early fall, closing with two Saturdays in Indio, California, on October 8th and 15th as part of Desert Trip, the classic-rock cousin to Coachella. His current touring pattern was initially forced on him during a custody battle over his youngest daughter, Beatrice Milly, requiring him to be close to home. He could only schedule brief runs of concerts on the road.
"That actually turned out to be great because it meant that you'd get this time off," he says, "which would then leave you kind of hungry to get back onstage – instead of never knowing quite where you were, whether you were in Des Moines or Detroit."
He began the tour April 13th in Fresno, California, on a redesigned stage set with a familiar sound: the opening clang from "A Hard Day's Night," a song he hadn't performed in 50 years. "That chord is pretty iconic," he says. "I suggested that to the band and we all got sort of goosebump-y."
The rarities reach back to "In Spite of All the Danger," an adoring ballad from when the Beatles were still known as the Quarrymen. During sets of nearly three hours, McCartney mingles the Beatles' generational anthem "Let It Be" with the explosive Wings hits "Live and Let Die" and "Jet," as well as an emotional reading of "Here Today" – his intimate remembrance of John Lennon – and the early solo hit "Maybe I'm Amazed," now the opening track of his new four-CD retrospective, Pure McCartney.
There's also the warm effervescence of the Beatles' "And I Love Her" and complex psychedelia of "Being for the Benefit of Mr. Kite!" (from 1967's Sgt. Pepper's Lonely Hearts Club Band), a song McCartney once vowed was too difficult to attempt live. "I'll just hear a song, and I'll go, 'Oh, that was a great song,'" McCartney says. "It's really down to what excites us. If it excites us, it will probably excite an audience – fingers crossed."
The songs are brought to life by McCartney's band of the past decade: keyboardist Paul "Wix" Wickens, bassist-guitarist Brian Ray, guitarist Rusty Anderson and drummer Abe Laboriel Jr. "We like playing together. We can talk about stuff. We can throw ideas around. I think we make a pretty good noise," McCartney says. "That's what you want in a band. You want to have friendship, people you get on with. You want to respect them and their playing, and you want to be able to say, 'Hey, look, I think you could do that better,' and be able to communicate."
For his One on One dates, McCartney is including an acoustic version of "FourFiveSeconds," last year's multi-platinum collaboration with Rihanna and Kanye West. The former Beatle plays it in the original key, which West later adjusted for Rihanna's voice.
"I like the grooves that I originally did, and that Kanye must have liked. It was good to take it back to where it started," he says. "The younger people are the ones who really know it. I must admit, I think, 'Oh, my God, all the older people who are all going to be singing "Ob-La-Di, Ob-La-Da" are not going to know what this is.' They know more than you think. In a show that spans many years, it's nice to have something recent."
More obscure is "Temporary Secretary," a funky, bouncy, New Wave electronic track from 1980 rediscovered in recent years by DJs in Brighton, England, and across Europe. Word got to McCartney that the song was being played in clubs. "I started to hear rumors that this guy is going crazy over 'Temporary Secretary,'" he says. "I'd always enjoyed the record, which I made when I was sort of experimenting with synthesizers and sequencers when they first came out on the record I made, McCartney II. I'd pretty much forgotten it."
It was Wix who had to figure out how to program the 36-year-old electronic song, originally recorded with an old Arp sequencer. "I like that things I thought were good at the time, that maybe didn't get that noticed, then come around on their second lap," McCartney says. "Suddenly it resurfaces. That's something I love about music. It's very rewarding for me."
Responding to the song's unexpected rebirth, he performed "Temporary Secretary" live for the first time ever a year ago at London's O2 Arena. "The message is very all-time," he says. "For me, it was a little bit tongue-in-cheek, a little bit saucy: 'Send me along a girl, just for a little while/When I send her back, make sure she stays on the right track.' It's kind of silly, but it has a sort of meaning, and it's a meaning that people still understand."
His balance of Beatles and post-Beatles songs has evolved over the years. In the early days of Wings, he refused to perform anything from his former band. After the international success of Band on the Run in 1973, he loosened up the set list to include Beatles tracks, but still makes a point of spotlighting new music.
"Every band you talk to knows, if you play the old hits, people love it, and all the phones light up so it becomes like a stellar galaxy," he says. "When you play the new stuff, it goes like a black hole. I sometimes bust the audience for that: 'We know which ones you like. We can tell.' But we enjoy doing it.
"Even though there's a lot of Beatles fans and Wings fans in the audience, there's still a lot of people who want to hear something different or something new or a little bit of a deep cut. I like to keep them happy too."
He suggested a series of concerts filled with the deepest cuts. "You'd have to call the tour 'You're Not Going to Like This One,'" McCartney jokes. "It would be a different vibe altogether."
The full 67-track version of his new collection, Pure McCartney (also available as a shorter two-disc set), is a balance of post-Beatles hits and rarities, set for a June 10th release. He chose to open with the original studio version of "Maybe I'm Amazed," a love song to his first wife, Linda Eastman, from his 1970 solo debut. In 1977, a live recording of the song was a Top 10 U.S. single.
"I do like the studio version. It embodied that album I made, which was the first thing I did right after the Beatles. It has good associations for me," says McCartney, noting that he heard the track was even a favorite of Cabaret singer-actress Liza Minnelli. "It kind of surprised me. I would have expected more of a show song, but that's cool. This one's for Liza."
The Pure collection was suggested by a young woman in McCartney's office who craved something for a long car trip. "I developed this playlist of stuff you would like to hear on a long journey," he says. "There was not too much deep thinking. It was stuff that we think is good, but there's not a massive theme to it."
Even as McCartney has been revisiting his discography for the new compilation, road life still inspires. Late during that first set in Fresno, McCartney began reading aloud from the signs fans brought to the gig. Most were song lyrics and notes of devotion: "I Love Paul," "Macca-nificent!" and "It's my 64th concert since '64. Autograph?"
Little kids bounced with signs the entire night, and even fans well past 60 danced and sang with the energy of early Beatlemania. After performing "Yesterday," McCartney pulled to the stage a family who carried signs that noted a mom had named her son "Jude," after the Beatles anthem "Hey Jude." One of the girls in the family promised to name her firstborn daughter "Penny Lane." There were hugs and autographs, and genuine affection between the former Beatle and his fans.
"With the Beatles, we thought we'd last 10 years," McCartney says later. "To realize that there's still large numbers of people who were into what we did then, and are now into what I do now, it becomes like a family thing. They're your musical family. It's like long-lost cousins. I enjoy that."

redpajama_set_name: RedPajamaCommonCrawl


text:
class SlackService < Service
  prop_accessor :webhook, :username, :channel
  validates :webhook, presence: true, if: :activated?

  def title
    'Slack'
  end

  def description
    'A team communication tool for the 21st century'
  end

  def to_param
    'slack'
  end

  def fields
    [
      { type: 'text', name: 'webhook',
        placeholder: 'https://hooks.slack.com/services/...' },
      { type: 'text', name: 'username', placeholder: 'username' },
      { type: 'text', name: 'channel', placeholder: '#channel' }
    ]
  end

  def supported_events
    %w(push issue merge_request note tag_push)
  end

  def execute(data)
    return unless supported_events.include?(data[:object_kind])
    return unless webhook.present?

    object_kind = data[:object_kind]

    data = data.merge(
      project_url: project_url,
      project_name: project_name
    )

    # WebHook events often have an 'update' event that follows a 'open' or
    # 'close' action. Ignore update events for now to prevent duplicate
    # messages from arriving.

    message = \
      case object_kind
      when "push", "tag_push"
        PushMessage.new(data)
      when "issue"
        IssueMessage.new(data) unless is_update?(data)
      when "merge_request"
        MergeMessage.new(data) unless is_update?(data)
      when "note"
        NoteMessage.new(data)
      end

    opt = {}
    opt[:channel] = channel if channel
    opt[:username] = username if username

    if message
      notifier = Slack::Notifier.new(webhook, opt)
      notifier.ping(message.pretext, attachments: message.attachments)
    end
  end

  private

  def project_name
    project.name_with_namespace.gsub(/\s/, '')
  end

  def project_url
    project.web_url
  end

  def is_update?(data)
    data[:object_attributes][:action] == 'update'
  end
end

require "slack_service/issue_message"
require "slack_service/push_message"
require "slack_service/merge_message"
require "slack_service/note_message"

redpajama_set_name: RedPajamaGithub


Dreams are the outcome of certain mental states, conscious and subconscious , caused by various positive and negative effects of heavenly bodies. Dream is as real as this world is when we are awake. Whenever we sleep, we die and as our sould leaves our body. Many of the times the dreams that we see are not just dreams but also real experience of the soul, messages from our protectors, omens, warnings and much more.
Thus, interpretation of dreams can point towards the mysteries of future. However, one must keep certain guidelines in mind in order to interpret the dreams and deduce their effects. the dreams that occur during the period beginning from 3 am till dawn yield their results within 7 days, the dreams of midnight give their result in a month and the dreams that come before midnight show their effect in a year. The dreams that come during daytime do not yield any result at all. If one sees several dreams in a night, only the last one happens to be effective.
Dreaming is a part of everyone's life and the fact goes that everyone dreams for as long as they live. Many even say that unborn babies dream in the womb of their mothers. Dreams are meant for the reality which is yet to happen and is a thing that takes place in the mind when the entire body has gone to rest. Several also opine that dreams are the gateway to an inner world, a world which is just as real as the one outside. Dream analysis and interpretation informs us in a visual language about reserved experiences and other process of the unconscious. The analysis of dream images and activities, however strange and even meaningless they might seem, depict symbolic structures. If you want a proper analysis and interpretation of your dreams then there is no better place than Mystik Astrology, where our panditji will assess you and provide you with the best possible outcomes. Remember, astrology is one of the best mediums to analyze and interpret your dreams.


text:
Keeneland
For the town, see Keeneland, Kentucky.
Keeneland Association, Inc. is an equine business based in Lexington, Kentucky. It includes two distinct divisions: the Keeneland Race Course, a Thoroughbred racing facility, and Keeneland Sales, a horse auction complex.[2][3][4] It is also known for its reference library.[5]
Lexington, Kentucky, United States
38°2′44″N 84°36′38″W / 38.04556°N 84.61056°W / 38.04556; -84.61056Coordinates: 38°2′44″N 84°36′38″W / 38.04556°N 84.61056°W / 38.04556; -84.61056
Keeneland Association Inc.
Date opened
Flat / Thoroughbred
Notable races
Ashland Stakes (G1)
Turf Mile Stakes (G1)
Breeders' Futurity Stakes (G1)
Blue Grass Stakes (G2)
U.S. National Register of Historic Places
U.S. National Historic Landmark
A view of Keeneland's grandstand at dawn, taken from the last turn leading into the home stretch
NRHP reference No.
Added to NRHP
Designated NHL
In 2009, the Horseplayers Association of North America introduced a rating system for 65 Thoroughbred racetracks in North America. Keeneland was ranked #1 of the top ten tracks. It is listed on the National Register of Historic Places and designated a National Historic Landmark in 1986.
2 Keeneland Sales
3 Layout
4 TV personalities
5 Keeneland Mark of Distinction
6 Racing
HistoryEdit
Keeneland originated as a nonprofit racing–auction entity on 147 acres (0.59 km2) of farmland west of Lexington, which had been owned by the son of James R. Keene, Jack Keene, a driving force behind the building of the facility.[6] It has used proceeds from races and its auctions to further the thoroughbred industry as well as to contribute to the surrounding community.
The start of the 2016 Appalachian Stakes, one of many graded races run during Keeneland's spring meet every year.
Keeneland Race Course has conducted live race meets in April and October since 1936. The 15-day spring meet is one of the richest in North America, with fifteen graded stakes races featuring the Blue Grass Stakes, a prep race for the Kentucky Derby.[7][8][9] The 17-day fall meet features seventeen graded stakes races, six of which are Grade One events used as Breeders' Cup preps.[10][11]
Keeneland takes pride in maintaining racing traditions; it was the last track in North America to broadcast race calls over its public-address system, not doing so until 1997. Most of the racing scenes of the 2003 movie Seabiscuit were shot at Keeneland, in part because of the track's "retro feel".[12] Keeneland was also used in the 2005 movie Dreamer[13] and the 2010 movie Secretariat for several key scenes, including the running of the Belmont Stakes where the horse completes the Triple Crown.[14]
Nonetheless, Keeneland has adopted several innovations. In 1984 in preparation for a visit by Queen Elizabeth II, it built a trackside Winner's Circle and created the Queen Elizabeth II Challenge Cup Stakes. In 1985, it installed a turf course over which the Challenge Cup, as well as a number of other turf races, is now run.[15] It reshaped the main track and replaced the dirt surface with the proprietary Polytrack surface over the summer of 2006 in time for its fall race meeting.[16] The track was restored to a dirt racing surface during the summer of 2014.[17]
Keeneland was designated a National Historic Landmark in 1986,[18] for its role in the growth of the horseracing sport and its revitalization in Lexington.[19]
Keeneland hosted the Breeders' Cup for the first time in 2015. The Breeders' Cup Classic was won by Triple Crown winner American Pharoah by six and a half lengths. He became the first to win the unofficial Grand Slam of horse racing; winning the Kentucky Derby, Preakness Stakes, Belmont Stakes and Breeders' Cup Classic. Many horse industry personnel were skeptical of Keeneland as a suitable venue because the track and town were too small to host such a large event. However it was a huge success and even had a Thoroughbred Daily News writer report "I was wrong...it was spectacular" and how he "couldn't be more impressed".[20] In August 2018 Keeneland was named as the host of the 2020 Breeders' Cup.[21]
In 2020 due to the COVID-19 pandemic in the United States, Keeneland cancelled their Spring meeting which was to be held from 2 April 2020 to 24 April 2020[22] and moved their meet to a shortened five day period from 8 July 2020 to 12 July 2020.[23] Five stakes race were not scheduled: Bewitch Stakes, Ben Ali Stakes, Commonwealth Stakes, Doubledogdare Stakes and Lexington Stakes. The Breeders' Cup was won by Authentic, which had won the Kentucky Derby.
Keeneland SalesEdit
Main article: Keeneland Sales
Keeneland is the world's largest Thoroughbred auction house, conducting three sales annually: The September Yearling Sale, November Breeding Stock Sale, and January Horses of All Ages Sale.[24] Horses sold at Keeneland sales include 82 horses that won 88 Breeders' Cup World Championship races; 19 Kentucky Derby winners; 21 Preakness winners; 18 Belmont winners; 11 recipients of the Eclipse Award as Horse of the Year; and five Epsom Derby winners. Graduates of Keeneland sales.[25] The Keeneland Team travels to over 25 countries and invests in over $700,000 annually towards international market development to deliver the world's deepest buying bench.[26] The auctions have sold horses to owners worldwide that have won large-scale races. Nine of the 14 highest ranked horses in the Kentucky Oaks were sold in sales through Keeneland.[27]
LayoutEdit
The track has a 1+1⁄16 miles (1,700 m) dirt oval and a seven and one-half furlong (0.875 miles (1,408 m)) turf oval. The turf course uses two configurations: the Keeneland Course setup has a temporary rail set 15 feet (4.6 m) out, while the Haggin Course has no temporary rail.[28]
TV personalitiesEdit
Mike Battaglia (1974–present)
Katie Mikolay Gensler (2013–present)
Keeneland Mark of DistinctionEdit
Outside view of Keeneland
Since 1968 Keeneland has honored the contributions of individuals to both Keeneland itself and the Thoroughbred industry with the Mark of Distinction. Receipents of the honor are given a button that bears the colors of their racing or farm silks. Each spring and fall, members of the Keeneland Club wear the button.[29]
RacingEdit
Keeneland has two racing seasons: a Spring Meeting in April and a Fall Meeting in October. The following stakes races have been run at Keeneland, some with changing names and sponsorships over the years.
Grade 1 :
Alcibiades Stakes
Ashland Stakes
Breeders' Futurity Stakes
First Lady Stakes
Jenny Wiley Stakes
Keeneland Turf Mile Stakes
Madison Stakes
Maker's 46 Mile Stakes
Queen Elizabeth II Challenge Cup Stakes
Spinster Stakes
Appalachian Stakes
Elkhorn Stakes
Fayette Stakes
Jessamine Stakes
Marathon Stakes
Phoenix Stakes
Raven Run Stakes
Shakertown Stakes
Thoroughbred Club of America Stakes
Woodford Stakes
Beaumont Stakes
Ben Ali Stakes
Bewitch Stakes
Bourbon Stakes
Bryan Station Stakes
Franklin County Stakes
Commonwealth Stakes
Doubledogdare Stakes
Dowager Stakes
Lexington Stakes
Sycamore Stakes
Transylvania Stakes
Valley View Stakes
Non-graded stakes races
Giant's Causeway Stakes
A race at Keeneland
^ "NPS Focus". National Register of Historic Places. National Park Service. Retrieved July 1, 2010.
^ "Keeneland Sales". Keeneland Association, Inc. 2019-06-28. Retrieved 2019-06-28.
^ Keeneland Magazine - Blood-Horse Publications designation approved by Keeneland Association, Inc.
^ "Keeneland Sales portal". Keeneland Association, Inc. 2019-06-28. Retrieved 2019-06-28.
^ "The Keeneland Library: An uncommon collection | Topics: Keeneland | Thoroughbred Racing Commentary". Thoroughbred Racing Commentary. Retrieved 2016-08-01.
^ "Keeneland History". keeneland.com.
^ "Keeneland Spring Produces Strong Attendance, Wagering". BloodHorse.com. Retrieved 25 June 2017.
^ "Keeneland Spring Meet boasts increase in purses, 15 graded stakes races". Lexington Herald-Leader. Retrieved 25 June 2017.
^ "Toyota Blue Grass Website". keeneland.com.
^ "Keeneland Announces $5.325 Million Stakes Schedule". BloodHorse.com. Retrieved 25 June 2017.
^ "Breeders' Cup Challenge". breederscup.com.
^ Copley, Rich. "Flashback: When 'Seabiscuit' filmed at Keeneland | Copious Notes". Copious Notes. Retrieved 25 June 2017.
^ "Keeneland Part 1". WikiLex: A History Hub. Lexington History Museum. Retrieved 15 March 2018.
^ "'Secretariat' is infused with Kentucky spirit, director says". kentucky. Retrieved 25 June 2017.
^ "A Winner's Circle Fit for a Queen". keeneland.com. Retrieved 25 June 2017.
^ "Rogers Beasley Keeneland Director of Racing". Bloodhorse.com. 2006-09-27. Archived from the original on 2009-04-19. Retrieved 2009-10-24.
^ "Keeneland Dirt Track Debut". Courier-Journal.com. The Courier-Journal.
^ "National Register Information System". National Register of Historic Places. National Park Service. March 13, 2009.
^ "NHL nomination for Keeneland". National Park Service. Retrieved 2020-01-15.
^ One For the Ages by Barry Weisbord
^ "Future Host Sites 2019-2021". Breeders' Cup. 17 August 2018.
^ BloodHorse Staff (16 March 2020). "Keeneland Cancels 2020 Spring Meet". BloodHorse.com. Retrieved 22 May 2020.
^ King, Byron (22 May 2020). "Keeneland Applies to Run Summer Meet July 8–12". BloodHorse.com. Retrieved 22 May 2020.
^ "About Keeneland Sales". keeneland.com. Keeneland Sales.
^ "Keeneland Classic Winners". Keeneland.com. Keeneland.
^ "Sales | Keeneland". www.keeneland.com. Retrieved 2018-04-19.
^ "Keeneland Sales Alumni Capture Final Derby, Oaks Preps | Keeneland". www.keeneland.com. Retrieved 2018-04-19.
^ "Keeneland Track Configuration". keeneland.com. Keeneland Racing.
^ "Mark of Distinction" (PDF). Keeneland. 2013-03-20. Retrieved 2021-01-06.
Wikimedia Commons has media related to Keeneland.
Classic Race winners sold at Keeneland Auction
Keeneland's First Meet, 1936
Retrieved from "https://en.wikipedia.org/w/index.php?title=Keeneland&oldid=1105443766"
This page was last edited on 20 August 2022, at 09:03 (UTC).


Child custody is one of the most contentious areas of family law. Most parents want custody and possession of, and access to, their children and are willing to do just about anything to get it. One of the experienced child custody lawyers from Orsinger, Nelson, Downing & Anderson can help get the results you need in court by skillfully addressing all areas of concern.
When custody issues arise, the courts will always keep the best interests of the child first and foremost in their decision-making. This will determine everything from visitation rights, the child's living arrangements, and who is named as a conservator of the child.
Texas courts designate "conservators" as those charged with caring for a child and making decisions on their behalf. This role very often falls to the child's parents. In fact, most judges favor placing a child with their natural mother or father before considering other options. Of course, there are exceptions to this. Sometimes the natural parents of the child are addicted to drugs or alcohol, or have a history of violent behavior.
In some instances a conservator can be a grandparent or some other relative. Grandparent custody is more common than some might think, and uncles, aunts, cousins, and even siblings are also eligible for custody (conservatorship) of a child.
In Texas, it's presumed that both parents are to be designated as "joint managing conservators." However, the needle concerning that presumption may move based on the evidence presented to the court. What this means is that courts in Texas view each parent as having important input to decisions relating to the child.
Regarding "the best interests of the child," as identified above, the court may limit the rights of a joint managing conservator, such that one conservator's rights are more significant than the other. For example, the court may simply find that one parent is especially attuned to the educational needs of the child, and award that parent the exclusive right to educational decision-making. Another example is that one parent may be a medical doctor. That parent may, by way of their professional training and experience, demonstrate that it would be in the child's best interests if they had a superior right to make decisions with regard to the child's health care treatment.
The biggest issue related to conservatorship tends to be when a conservator is determined by the court to be the "joint managing conservator with the exclusive right to designate the primary residence of the child" (i.e., which parent is designated as the child's home base). This may play a factor in possession (or visitation) of the child, but it won't necessarily determine what that possession will look like.
In framing the court's perspective as to what the "child's best interests" actually are, the testimony and evidence presented by the parents are obviously very significant. The parents are often the most significant component of the court's consideration with respect to the child and how the court would seek to allocate important rights and duties to each parent, such as who gets the right to make educational decisions, who gets the right to make important health care and psychological/psychiatric decisions, and who gets to designate the primary residence of the child.
The evidence and testimony provided to the court by the parents is critical because it is the parents who truly are at the "front lines" with regard to their child in most circumstances. If both parents, for instance, wake the child up, get them ready for school, take them to school, take them to their activities, help them with their schoolwork, and keep a close eye on their child's health care, the court may have little difficulty in appointing both parents as joint managing conservators with rights and duties that may be close to equal.
What happens if there's a case where things are more contested or where one parent is without question more responsible and substantially more attuned to the child's needs and care? This may also be a "simple" case for the court, insofar as some of the facts speak for themselves. For example, if dad doesn't know the name of the homeroom teacher, never attended a single parent-teacher conference, never has once spoken with a pediatrician, doesn't participate in any activities with the child, and instead spends his time closely attached to a bottle of gin, the court may likely be very comfortable giving that parent little control over the child's life and limiting that parent's rights and duties by giving mom superior decision-making power.
When things are very contested and it may not be so clear to the court who's in a better position to have such control over decisions (or how they should be made with respect to the child, i.e., exclusive to one, jointly, or independently), you will likely need to expand your supporting information sources. Witnesses to the lives of the family become more important. Thus, as you proceed with litigation, it's important to have a broad awareness of who else is playing an important role in the child's life.
Collateral witnesses (people apart from the parents or conservators) can offer insight on how a child is doing or functioning when the child is not with both parents. Alternatively, a collateral witness – a teacher, for example – may describe how one parent is particularly "good," and another may be relatively "bad" from their perspective. To the court, these collateral witnesses may present a fuller picture as to what is going on in a child's life and what would be in the child's best interests in regard to an outcome in the case. However, a word of caution: the friend, the teacher, the coach, whatever third party there may be that may have special knowledge of the child, may not be as assertive with their position when brought to court as they were on the phone with you the day before. The confines of the courtroom can be very intimidating, especially to people who are not directly involved in a custody case.
Another complication exists with bringing in multiple witnesses to support your client's position. Court can be a cumbersome and a time-consuming process: things get delayed, reset for another day, or they can go very long. It can be particularly burdensome for parties outside of the dispute. It can be expensive for the witnesses too. That third-party witness may have to take off work. They may have to travel. Also, they truly may not like the discomfort that comes with live testimony, especially cross-examination. They may distance themselves from you and the child as a ramification of their good intention of support.
In order to avoid this possible negative impact on your relationships, and still present critical evidence, the best option for supportive testimony may be a mental health professional.
There are several reasons why. But do not go through this process without the counsel of an experienced custody dispute attorney.
Many mental health professionals are educated and trained to deal with the issues that are involved in family law custody litigation. But first, what do we mean (or courts mean) when we say "mental health professional"? This designation covers a broad range of professionals, including psychologists, psychiatrists, licensed professional counselors, licensed professional social workers, parenting facilitators, and custody evaluators. These are individuals who are licensed by the state and who may even follow particular roles defined by statute in the Texas Family Code. However, not all mental health professionals have similar credentials. For example, a psychiatrist is a medical doctor while a psychologist with a Ph.D. is not.
One of the most important things that a mental health expert may be able to do, in addition to meeting with the child and maybe both parties, is contact and discuss things relevant to the case and the child with all the collateral witnesses. The mental health professional can often convey the information that they have learned, whether from counseling sessions, testing or from their discussions with collateral witnesses, to the court.
Because of their training and education, a mental health professional involved in the custody case will likely have an opinion to share with the court. However, an opinion as to an ultimate custody decision is only appropriate when the mental health professional has met all the parties involved: the respective parents and the child. Nevertheless, even if a mental health professional cannot opine as to an ultimate custody decision, they can likely add very important color to the court's consideration of what is in the child's best interests.
A contested child custody case isn't typically a matter fought out between the parties in isolation. It will often involve a lot of components such as collateral witnesses and mental health professionals. It will sometimes entail a group effort for the court to define what is in the child's best interest. It's important to work with your child custody attorney to identify your interests, your strengths and weaknesses, and ways to either accentuate and enhance them for presentation or mitigate the problems presented.
It's also important for you and your custody lawyers to work together to identify those other people who share a role in the upbringing of your child. Consider discussing the benefits or consequences of retaining a mental health expert or having one appointed, as it can provide important supporting information to the court related to the child's best interests.
It's important to recognize that when things are contested in a custody case, organizing and dealing with these myriad parts and people can take time.
You're talking about a matter that has significance to you and your relationship with your child and your child's development for years to come. Once a judgment is finalized, it is difficult to modify a child custody order. So in that regard, be patient and work with your attorney and any mental health professional involved. In the end, you (and your relationship with your child) will likely benefit greatly.
In Texas, the issue of custody can be decided by a jury if one side requests it.


text:
Q: 86 it ! using 86 as a verb
Possible Duplicate:
What does the term "86'd" relate to?

I found that you can say "86 something" as a verb when we want to "cancel" something...
Is it used for everything??
For example, if I'm in a restaurant and changed my mind about one dish, is it correct to say "Hey, 86 it"?
Note: I'm not asking about the meaning of the verb to 86 something, but rather whether the verb can be used generally or only in specific contexts.

A: Wikipedia link.
Speaking as someone who spent almost a decade working in restaurants and bars, I've never heard a customer use the term. It's jargon that restaurant employees would use speaking to each other.
In my experience it's used either to indicate you're refusing service to a particular customer, or that the establishment has either run out of an item or removed it from the menu. Even if you ordered a burger then changed your mind, I wouldn't tell the cook to "86 that burger". A much more likely scenario is I put your burger order in and the cook comes back and says that "86 on burgers" because they've run out of ground beef.

redpajama_set_name: RedPajamaStackExchange


A young girl named Avery landed a big bass this week using a tiny Barbie pole. Her reaction is priceless. I'm sure she will never forget that experience. Our hats are off to you Avery.
This girl is our hero.
The video is guaranteed to put a smile on your face. Watch it below.
We are huge advocates of getting kids fishing. We love to see that she's fishing and making incredible memories like this one.