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The southern root-knot nematode (RKN, Meloidogyne incognita) is a major soil-inhabiting plant parasite that causes significant yield losses in cotton (Gossypium spp.). Progeny from crosses between cotton genotypes susceptible to RKN produced segregants in subsequent populations which were highly resistant to this parasite. A recombinant inbred line (RIL) population of 138 lines developed from a cross between Upland cotton TM-1 (G. hirsutum L.) and Pima 3–79 (G. barbadense L.), both susceptible to RKN, was used to identify quantitative trait loci (QTLs) determining responses to RKN in greenhouse infection assays with simple sequence repeat (SSR) markers. Compared to both parents, 53.6% and 52.1% of RILs showed less (P<0.05) root-galling index (GI) and had lower (P<0.05) nematode egg production (eggs per gram root, EGR). Highly resistant lines (transgressive segregants) were identified in this RIL population for GI and/or EGR in two greenhouse experiments. QTLs were identified using the single-marker analysis nonparametric mapping Kruskal-Wallis test. Four major QTLs located on chromosomes 3, 4, 11, and 17 were identified to account for 8.0 to 12.3% of the phenotypic variance (R2) in root-galling. Two major QTLs accounting for 9.7% and 10.6% of EGR variance were identified on chromosomes 14 and 23 (P<0.005), respectively. In addition, 19 putative QTLs (P<0.05) accounted for 4.5–7.7% of phenotypic variance (R2) in GI, and 15 QTLs accounted for 4.2–7.3% of phenotypic variance in EGR. In lines with alleles positive for resistance contributed by both parents in combinations of two to four QTLs, dramatic reductions of >50% in both GI and EGR were observed. The transgressive segregants with epistatic effects derived from susceptible parents indicate that high levels of nematode resistance in cotton may be attained by pyramiding positive alleles using a QTL mapping approach.
Citation: Wang C, Ulloa M, Mullens TR, Yu JZ, Roberts PA (2012) QTL Analysis for Transgressive Resistance to Root-Knot Nematode in Interspecific Cotton (Gossypium spp.) Progeny Derived from Susceptible Parents. PLoS ONE 7(4): e34874. https://doi.org/10.1371/journal.pone.0034874
Editor: Baohong Zhang, East Carolina University, United States of America
Received: September 7, 2011; Accepted: March 10, 2012; Published: April 13, 2012
This is an open-access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication.
Funding: Funding provided by Cotton Incorporated (grant #08-321) University of California Discovery Grant (Grant#06-10632). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
Competing interests: Cotton Incorporated through their research grants program designed to address cotton field production problems, funded a portion of the research in this study conducted in the Roberts lab, in a manner typical of other funding agency programs. No other form of support from this funder was provided. This does not alter the authors' adherence to all the PLoS ONE policies on sharing data and materials.
Host-plant resistance is highly effective in controlling crop loss from root-knot nematode (RKN) Meloidogyne incognita Kofoid and White (Chitwood) infection. Novel sources and enhanced levels of pathogen resistance are desirable for genetic improvement of crop plants. One source of enhanced resistance is that generated through transgressive segregation. Transgressive segregation is the phenomenon in which segregating hybrids exhibit phenotypes that are extreme or novel relative to the parental lines [1]–[2] and result from epistatic interactions between the genes assembled in novel combinations. Transgressive segregants for numerous traits have been reported, and can be derived from both intraspecific and interspecific crosses [2]–[4]. Breeding for optimal resistance must be based on selection of progeny with combinations of genes homozygous for resistance. However, a highly susceptible parent can contribute to nematode resistance via transgressive segregation [5]. These crosses can derive highly resistant lines, even when both parents have a susceptible phenotype. Such transgressive segregants can be used as improved resistance sources in crop breeding [5]–[6].
Transgressive segregation is one of the major selection sources in cotton (Gossypium hirsutum L.) for enhanced resistance to RKN, an important pest of cotton [7]. Wang et al. (2008) [5] reported that a segregant factor (RKN2) from one susceptible parent, G. barbadense L. Pima S-7 interacted with a major recessive gene rkn1 from G. hirsutum Acala NemX [8] to produce a highly resistant phenotype in the interspecific cross Pima S-7 x Acala NemX. We also observed that a susceptible genotype Acala SJ-2 contributed to the level of resistance in some F2:7 (NemX x SJ-2) RI homozygous resistant lines [9]. Transgressive segregation also has been observed in other pathogen-cotton systems, such as for Verticillium wilt resistance in an F2 (resistant Pima S-7 x susceptible Acala 44) [10] and in an F2:3 family (G. barbadense H7124 x G. hirsutum XLZ1) [11], for Fusarium wilt resistance in the F2 or F3 populations (resistant Pima S-7 x susceptible Acala NemX or Acala SJ-2) [12], and for bacterial blight resistance in the BC4F4 generation within an intraspecific cross of cotton [13]. Transgressive segregation for resistance also has been reported in other plant-pathogen systems, such as soybean - soybean cyst nematodes [14], soybean - Rhizoctonia root and hypocotyl rot [15], Arabidopsis – Leptosphaeria maculans, blackleg disease [16], wheat – stripe rust [17]–[20], barley – Pyrenophora teres [21] and barley - powdery mildew [22].
Individual cotton plants highly resistant to M. incognita (transgressive segregants) have been reported within Auburn 623 RNR and some N-lines [23]–[25]. Turcotte et al. (1963) [26] reported that two recessive genes determined M. incognita resistance based on susceptible F1 and segregating F2 in a study of crosses of root-knot-resistant G. darwinii x susceptible G. barbadense cv. Pima S-1 and Pima S-2 breeding stocks. Genetic and quantitative trait loci (QTL) analyses revealed at least one major recessive gene with strong additive effect in intraspecific crosses Acala NemX x Acala SJ-2 and Acala SJ-2 x Clevewilt and one major QTL with strong dominant effect in the interspecific cross Pima S-7 x Acala NemX [8]–[9], [12], [26]. A major telomeric segment on chromosome 11 harbors RKN resistance genes from different backgrounds [5], [8], [27]–[29].
Transgressive segregation appears to be quite common in allotetraploid cotton, although the genetic basis of the generated novel phenotypes is generally not known. For traits that involve complex interactions of multiple genes, a QTL mapping approach can be informative for studying inheritance and gene action [27]. In particular, detection of minor "modifier" genes that may otherwise escape detection can be improved by QTL analysis [30]–[31]. Here we report on QTL mapping analysis to characterize the transgressive segregation for RKN resistance in a recombinant inbred line (RIL) population developed from an interspecific cross of susceptible Upland TM-1 (G. hirsutum) x susceptible Pima 3–79 (G. barbadense) with the goal to improve introgression of novel combinations of RKN resistance QTLs into elite cotton cultivars using susceptible genotypes. The specific approach was to phenotype the RIL population to identify QTLs associated with RKN resistance, and to study epistatic effects among QTL regions associated with RKN resistance.
Materials and Methods
Plant materials and Nematode resistance screening
Original seed source of a recombinant inbred line (RIL) population developed from an interspecific cross between G. hirsutum Upland TM-1 and G. barbadense Pima 3–79 [32] was provided by Drs. RJ Kohel and JZ Yu of USDA-ARS in College Station, TX. G. hirsutum Acala NemX and Acala N901 were included as resistant controls and Acala SJ-2 and G. barbadense Pima S7 as susceptible controls. The population was used previously for mapping microsatellite or simple sequence repeat (SSR) markers [33]–[34]. One hundred thirty-eight recombinant inbred lines (RILs) [6], parents, and controls were evaluated for nematode response in infection assays under controlled conditions in a greenhouse at University of California, Riverside, CA in two experiments using previously published protocols [9]. Five replicate plants of each RIL were screened in each test. Each of the parents and resistant and susceptible controls were also represented by five replicate plants in each test, which were randomly positioned between the randomized RILs to confirm uniformity of test conditions. The five sets of replicates were arranged on greenhouse benches in a randomized complete block design. Briefly, three-week-old seedlings were inoculated with approximately 50,000 eggs of M. incognita race 3 (isolate Project 77, from a cotton field in California). Air temperatures in the greenhouse were maintained between 28 and 35°C during the day and 24°C at night. Individual plants were scored for phenotype 60 days after inoculation. A 0–10 root-gall rating scale [35] ranging from no galling (0) to severe galling reaction (10) was used to evaluate resistance reaction to nematodes (root galling index - GI) [9]. For experiment 1, GI was recorded, and assays of egg production per root system and per gram fresh root were made by extracting eggs from the roots in NaOCl [36] (nematode egg production - EGR). In experiment 2, only GI was recorded.
Marker and genetic linkage analysis
One thousand and fifty SSR markers with wide genome coverage [6], [8], [33]–[34], [37, CMD, www. cottonmarker.org] were tested on the RIL (Upland TM-1 x Pima 3–79) population for association with RKN resistance. PCR amplification of BNL, CIR, Gh, MUSB, MUCS, MUSS, and NAU cotton molecular markers was performed on a total volume of 15 µL containing 2 µL of DNA template (concentration 10 ng), following the protocol described by Ulloa et al. (2011) [6]. PCR products were separated for 4 to 5 hrs on a 3% super fine resolution (SFR™) agarose gel (Amresco, Solon, OH) containing 1X TBE at 90 volts, and were visualized by Alphaimager software (v. 5.5, Alpha Innotech Corporation, San Leandro, CA) after staining with ethidium bromide. Primer-pairs were scored if they resulted in discrete PCR banding patterns (amplicons) denoting a molecular marker. Informative bands were scored as present (+) or absent (−) for a dominant marker, and if alleles from both parents were identified, then the marker was scored as co-dominant. The genotypic ratio of 1∶1 was expected for both dominant and co-dominant markers.
The JoinMapR version 4.0 [38] computer program was used to test for Chi-square goodness-of fit for expected versus observed genotypic ratios, and to develop the linkage groups for chromosomes. Logarithm of odds (LOD) scores of 3 to 14 were examined for each population using the Kosambi map function, and a maximum distance of 40 cM was used to determine linkage between any two markers. Linkage analyses on the RILs used DNA isolated from Fusarium wilt-phenotyped plants [6], together with a previously developed genetic linkage map [33], [37]. Twenty-three linkage groups from the 24 cotton chromosomes were developed. The linkage groups/chromosomes were developed with LOD>6 to obtain strong linkage between two anchored markers. We selected this LOD score to represent specific chromosome regions of the cotton genome.
Data analysis
Phenotypic data were subjected to one-way analysis of variance (ANOVA). Fisher's Protected LSD test was used to compare the treatment means using SAS (SAS, ver. 9.1.3; SAS Institute, Cary, NC). Data for nematode egg production were transformed to Log10(x+1) for analysis (LogEGR). Root-galling index (GI), eggs per gram of root (nematode egg production - EGR), and LogEGR were used for quantitative trait loci (QTL) analyses.
QTL analysis
QTL analyses were conducted on GI, EGR, and LogEGR using MapQTL 5.0 [39]. Single-marker analysis was conducted by using nonparametric mapping [Kruskal-Wallis analysis (K*)] test equivalent of the one-way analysis of variance [39]. We used the nonparametric analysis, because in this test, no assumptions are being made for the probability distribution(s) of the quantitative trait, and even if the data are distributed normally, the nonparametric test is often as powerful as parametric methods. In addition, the nonparametric test uses all markers genotyped on the population regardless of their linkages (tests each locus separately without the use of the linkage map). Threshold value for a marker-QTL was determined at P<0.1. Significant QTLs were determined with a more stringent P<0.005 for Kruskal-Wallis [39]. In order to verify the cumulative effects of QTL on the GI and EGR nematode response phenotypes, RILs were classified according to the number of alleles contributing to the resistance (favorable or positive alleles) present at each of the QTLs. Then QTLs were pyramided by combining data with favorable alleles (+, ++, +++, or ++++) and null RIL-genotypes (−,−−,−−− or −−−−). Class-specific means of GI and LogEGR and standard error were calculated for each genotypic class.
Phenotype of parents
Susceptible TM-1, Pima 3–79, Pima S7 and Acala SJ-2 had higher (P<0.05) galling indices (GI) and supported greater (P<0.05) numbers of eggs per gram root (EGR) than the two resistant genotypes Acala N901 (GI, 2.7, and LogEGR, 1.15) and Acala NemX (GI, 3, and LogEGR, 1.52) (Figure 1). Susceptible Pima S7 (GI, 5.4 and LogEGR, 3.76) and TM-1 (GI, 5.8 and LogEGR, 3.97) had lower (P<0.05) GI than the susceptible Pima 3–79 (GI 7.4, and LogEGR, 4.46) and SJ-2 (GI, 7.3 and LogEGR 4.25) (Figure 1).
Data were collected 60 days after inoculation. The error bars show standard error.
Phenotyping RILs
The RILs were phenotyped in the two greenhouse experiments (I and II). Different severity levels of infection were observed between the two experiments. Experiment II had less infection than Experiment I. This difference may have been caused by different environmental conditions in the greenhouse related to the season of the year during which each test was performed. Therefore, the phenotypic data are shown below for Experiment I. Seventy-four of 138 lines had lower (P<0.05) GI than the parent TM-1 (5.8±0.25 SE) indicating 53.6% of the lines exhibited transgressive resistance (Figure 2A), and 72 lines had fewer (P<0.05) EGR (8398±2025SE) indicating 52.1% of the lines showed transgressive resistance (Figure 2B). These observations indicated that multiple genes from both parents are involved in determining the resistance phenotype in these transgressive segregants because both parents are in the typical range of the susceptible phenotype for both GI and EGR. A low correlation (R2 = 0.42) between GI and EGR in the RIL population indicated that different genes might control the GI and nematode reproduction responses.
Data were collected 60 days after inoculation. Galling Index: 0–10 scale; 0 no galling, and 10 severe galling. The arrows point to the phenotypic reaction score in Acala N901, Acala NemX, Acala SJ-2, Pima S-7, TM-1 and Pima 3–79.
QTL mapping and identification
Twenty-one linkage groups/chromosomes were identified and they were found to be involved in RKN transgressive resistance by QTL analyses in this study (Figure 3, Tables 1, S1, and S2). However, four highly significant (P<0.005) QTLs were revealed by the nonparametric mapping test analogous to one-way analysis of variance [39] on chromosomes 3 (Mi-GIh -C031, BNL3408), 4 (Mi-GIb -C041, MUSSS396_N-111), 11 (Mi-GIb -C111, BNL1231), and 17 (Mi-GIh -C171, MUSB0224_320) in Experiment I accounting for 12.3, 11.5, 10.1, and 7.95% of the phenotypic variance in GI, respectively (Table 1, Figure 3). Three of these QTLs were identified in both Experiment I and Experiment II even though weaker infection occurred in Experiment II (Figure 3), confirming that these QTLs contributed to transgressive segregation for resistance. Moreover, there were two significant (P<0.005) QTLs on chromosome 23 (Mi-EGRb -C231, BNL1672) and chromosome 14 (Mi-EGRh -C141, BNL3661) in Experiment I accounting for 10.6 and 9.7% of the phenotypic variance in nematode egg production, respectively (Table 1, Figure 3).
Single-marker analysis conducted using nonparametric mapping (Kruskal-Wallis analysis) test equivalent of the one-way analysis of variance (Van Ooijen 2004). Red bars - QTL influencing root-galling index phenotype; Green bars - QTL influencing egg production phenotype (For Experiment I).
Nineteen additional QTLs for GI and 15 for EGR were detected at P<0.05 (Table 1, Table S1). The map locations of some of the detected QTLs associated with resistance to RKN are shown in Figure 3. In total, eight pairs of homoeologous chromosomes (Chr 1–15, 3–17, 4–22, 5–19, 6–25, 8–24, 9–23 and 11–21) were involved in galling index and/or nematode production. Sixteen QTLs were involved in both GI and EGR (Figure 3, Table S2) in Experiment I. Two significant QTLs (Mi-GIh -C031 and Mi-GIh -C171) were associated with GI but not with EGR. However, four significant QTLs (Mi-GIb -C111, Mi-GIb -C041, Mi-EGRh -C141 and Mi-EGRb -C231) were involved in both GI and EGR (Figure 3, Table S2). Among 23 QTLs associated with GI, 10 had positive alleles from G. hirsutum TM-1 and 13 had positive alleles from G. barbadense Pima 3–79. Nine of 17 QTLs with positive alleles associated with EGR were from G. hirsutum TM-1 and 8 from G. barbadense Pima 3–79 (Table 1, Table S1).
Analysis of QTL combinations with positive alleles contributed by both parents revealed that some putative detected QTLs (P<0.05) dramatically decreased GI when combined with major (P<0.005) QTLs or other putative QTLs. A few examples of these QTL combinations are represented in Figure 4. Combinations of the QTLs for nematode root-galling index and egg production were clearly associated with nematode infection response in the RIL population. The lines with a single positive QTL allele (+) had significantly lower GI and EGR (P<0.05; Figure 4). However, the combination of two or more QTLs with positive effect alleles resulted in much greater suppression of galling index and egg production (in some combinations more than 50% reduction) than those with only a single positive QTL allele (Figure 4, Table 1), suggesting additive or epistatic effects in those RIL-genotypes carrying two or more favorable alleles. Even though each single putative QTL could not contribute to high resistance to nematodes, highly resistant lines were formed when a few putative QTLs were combined together, such as the 3-QTL combinations Mi-GI- C121xC141xC191 (GI, 3.33) and Mi-EGR-C112xC121xC191 (LogEGR, 3.38) (Figure 4, Table 1) compared with parents TM-1 (GI, 5.8; LogEGR 3.97), Pima 3–79 (GI, 7.4; LogEGR 4.46) and resistant Acala NemX (GI, 3.0; LogEGR 1.52). We found that the response of root-galling suppression differed depending on selected QTLs. The 3-QTL combination of Mi-GI-031xC121xC191 (GI, 2.48) conferred a lower (P<0.05) galling index than the combination of three significant (P<0.05) QTLs Mi-GI- C031xC041xC17 1 (GI, 4.14) or Mi-GI-C041xC111xC171 (GI, 4.14). The 4-QTL combination of Mi-GI-C031xC041xC111xC191 (GI, 1.47) conferred a lower (P<0.05) galling index than the combination of four significant (P<0.05) QTLs Mi-GI-C031xC041xC111xC171 (3.02) (Figure 4). Some combinations were observed to be better than others. For example, the combination of Mi-EGR-C112xC121xC141xC191 resulted in less (P<0.05) egg production (LogEGR, 2.89) than that conferred by the combination Mi-EGR-C111xC141xC191xC231 (LogEGR, 3.43). However, both combinations conferred lower eggs per root than the parents TM-1 (LogEGR, 3.97) and Pima 3–79 (LogEGR, 4.46). These epistatic effects in progeny derived from susceptible parents indicate that pyramiding of these QTLs presents a new level of nematode resistance in cotton.
Our QTL analyses of root-knot nematode resistance in a cotton RIL population developed from an interspecific cross demonstrates the extent to which transgressive segregation for nematode resistance can occur in interspecific progeny derived from plant types with standard susceptible phenotypes. Previously we documented transgressive segregation for resistance in progenies derived from interspecific parents in which one of the parents carried a known major gene for resistance, rkn1 on chromosome 11 [8]. In that case, the susceptible parent was found to contribute a gene, RKN2, that had an undetectable effect on nematode response phenotype in the susceptible parent Pima S-7. However, presence of both genes resulted in extremely high resistance beyond that determined by the major gene alone and indicated an epistatic interaction that strengthened the resistance phenotype. We had also discovered in an intraspecific G. hirsutum cross between an rkn1 parent (Acala NemX) and a susceptible genotype (Acala SJ-2) in which some progenies exceeded the resistance level of the resistant parent [9]. That work built on an earlier report by Shepherd (1974) [23] of transgressive segregants for nematode resistance in progenies developed from a cross of two partially resistant intraspecific G. hirsutum cotton lines (Clevewilt and Wild Mexico Jack Jones). In the latter case, recent analysis of derived lines indicated that a combination of two genes, one from each parent, residing on chromosomes 11 and 14, were responsible for the combined resistance phenotypes [40].
In the current study, even considering this range of documented transgressive segregation events for resistance in cotton, the extent of resistance expression in the progenies generated from susceptible parents was remarkable. We found that more than 50% of recombinant inbred lines were more resistant than either susceptible parent. Importantly, in some lines the levels of resistance measured by suppression of root-galling and (or) nematode egg production were higher than standard resistant control lines such as Acala NemX included in the phenotyping screens. The QTL analysis revealed 23 major and putative QTLs involved in suppression of root-galling and 17 for suppressing egg production, indicating high transgressive segregation ratios. Obviously, the combinations of positive allele QTLs existing in each parent are not effective in conferring nematode resistance. Rather, only combinations of certain positive alleles contributed from both parents provided the unique genotypes required for nematode resistance. Thus, a single QTL had low effect on galling index and egg production, but when combined in positive allele forms, higher resistance levels were conferred (Figure 4 and Table 1), including combinations involving some of the putative small-effect QTLs. The analysis of effect on phenotype of various combinations of up to several QTLs revealed that some combinations of four QTL could produce high levels of nematode resistance for both root-galling and egg production phenotypes. This is apparent from the comparisons of the phenotypic averages of lines with or without the positive alleles for the specific QTL combination (Figure 4). These small-set QTL combinations could provide the basis for effective plant selection targets for genetic improvement in cotton breeding programs.
QTL analysis indicated that there were eight homoeologous chromosomes involved in the root-galling index (GI) and/or nematode egg production (EGR) phenotypes. Chromosome 11 of cotton is rich in disease resistance genes, including resistance to root-knot nematode [8], [29], [40]–[43], reniform nematode [44]–[45], Fusarium wilt [6], Verticillium wilt [10] and black root rot [46]. The marker CIR316-191/196 on chromosome 11 was involved in both GI and EGR, supporting our previous finding with gene rkn1 that resistance genes could suppress both root-galling and egg production [8]–[9]. The marker CIR316-194/184 on chromosome 21 also contributed to decreasing root-galling indicating some homological gene region on the paired homoeologous chromosome 11. These two chromosomes (11 and 21) might share gene expression functions in cotton. Ulloa et al. (2010) [27] reported that minor genes on chromosome 21 associated with markers CIR316 and MUCS088 contributed to both egg production and root-galling phenotypes in the F2 population of Pima S-7 x Acala NemX. A minor gene on chromosome 21 was also identified to contribute to suppressing root-galling in the F2 and F7 populations of Acala NemX and Acala SJ-2 (NxS) and BC1F1 (Acala SJ-2 x F1 (NxS) (Wang and Roberts, unpublished data). The homoeologous chromosome pair 11/21 also may be involved in reniform nematode resistance based on genetic mapping studies [44]–[45], [47]–[48]. Depending on populations, the regions involved in both root-knot nematode and reniform nematode resistance are around 30 cM apart, based on linked markers [48]. The sequencing of markers in these regions associated with both chromosomes 11 and 21 is underway and should help in resolving the complexity of genome recombination between the pair of homoeologous chromosomes. In the current study marker BNL3661 on chromosome 14 contributed more to inhibiting egg production than to suppressing root-galling, with the positive allele derived from G. hirsutum TM-1. This result is consistent with the report that a gene in this region from G. hirsutum M240 contributed to reduced egg production [40]. The marker BNL3545 close to BNL3661 on chromosome 14 involved in reducing egg production was also identified with the F2 population of Acala NemX x Acala SJ-2 (Wang and Roberts, unpublished data). Our finding that 14 QTLs were involved in both GI and EGR suggested that the relative contributions of these genes to both root-galling and egg production phenotypes are complex. Sequencing these QTL regions and comparing those sequences with other plants would expedite molecular breeding in cotton and enhance understanding of cotton evolution. Cotton whole genome sequencing currently in progress (http://www.monsanto.com/newsviews/Pages/Monsanto-Illumina-Key-Milestone-Cotton-Genome-Sequencing.aspx) will provide critical information about the cotton genome, plant evolution and adaption as it relates to valuable processes including transgressive segregation.
Our finding that susceptible parents can produce highly resistant progenies has important potential for exploitation in plant breeding. While transgressive segregation for biotic and abiotic stress tolerance and resistance is not uncommon, we have found very few reports where such highly resistant novel genotypes have been identified. Traditionally, crosses are made between a known resistance donor parent and susceptible parent with favorable traits, or between two moderately resistant parents, and the most resistant progeny are selected for advancement. Wallwork and Johnson (1984) [49] reported in wheat that 10% (24/240) of lines in a F2 population and 21.2% of lines (50/235) in a F3 population showed more resistance to yellow rust than either susceptible parent. Welch and Rieseberg (2002) [50] and Lexer et al. [51] reported that progenies showed 5–14 times more salt tolerance than the parental species in wild sunflower. Both sensitivity to salt and corresponding QTLs involved in salt tolerance were mapped to the chromosome level and used to select salt tolerant genotypes [52]. Based on the current findings, we suggest that efforts to identify novel and valuable phenotypes for biotic and abiotic stress resistance traits among progenies developed from stress susceptible or sensitive parent combinations are worthy of increased attention in plant breeding programs. This work also adds further support to the importance of outcrossing events in natural populations which drive selective advantage, adaptation and speciation processes.
Supporting Information
Table S1.
QTLs associated by nonparametric mapping with root-galling and nematode egg production in TM-1 x Pima 3-79 RIL population.
Table S2.
QTLs associated by nonparametric mapping with both root-galling and nematode egg production with same maker locus in TM-1 x Pima 3-79 RIL population.
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Author Contributions
Conceived and designed the experiments: PAR CW MU. Performed the experiments: PAR CW MU TM. Analyzed the data: PAR CW MU TM JZY. Contributed reagents/materials/analysis tools: PAR CW MU TM. Wrote the paper: PAR CW MU.
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- 45. Dighe N, Robinson AF, Bell A, Menz M, Cantrell R, Stelly D (2009) Linkage mapping of resistance to reniform nematode in cotton (Gossypium hirsutum L.) following introgression from G. longicalyx (Hutch & Lee). Crop Sci 49: 1151–1164.N. DigheAF RobinsonA. BellM. MenzR. CantrellD. Stelly2009Linkage mapping of resistance to reniform nematode in cotton (Gossypium hirsutum L.) following introgression from G. longicalyx (Hutch & Lee).Crop Sci4911511164
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- 47. Romano GB, Sacks EJ, Stetina SR, Robinson AF, Fang DD, et al. (2009) Identification and genomic location of a reniform nematode (Rotylenchulus reniformis) resistance locus (Ren ari) introgressed from Gossypium aridum into upland cotton (G. hirsutum). Theor Appl Genet 120: 139–150.GB RomanoEJ SacksSR StetinaAF RobinsonDD Fang2009Identification and genomic location of a reniform nematode (Rotylenchulus reniformis) resistance locus (Ren ari) introgressed from Gossypium aridum into upland cotton (G. hirsutum).Theor Appl Genet120139150
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- 50. Welch ME, Rieseberg LH (2002) Habitat divergence between a homoploid hybrid sunflower species, Helianthus paradoxus (Asteraceae), and its progenitors. Am J Bot 89: 472–478.ME WelchLH Rieseberg2002Habitat divergence between a homoploid hybrid sunflower species, Helianthus paradoxus (Asteraceae), and its progenitors.Am J Bot89472478
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- 52. Lexer C, Welch ME, Durphy JL, Rieseberg LH (2003) Natural selection for salt tolerance quantitative trait loci (QTLs) in wild sunflower hybrids: implications for the origin of Helianthus paradoxus, a diploid hybrid species. Mol Ecol 12: 1225–1235.C. LexerME WelchJL DurphyLH Rieseberg2003Natural selection for salt tolerance quantitative trait loci (QTLs) in wild sunflower hybrids: implications for the origin of Helianthus paradoxus, a diploid hybrid species.Mol Ecol1212251235


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Explore Picnic Area, Park Service and more!
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Park Closes Roads and Trails due to Chimney 2 Fire Great Smoky Mountains National Park officials have closed Newfound Gap Road, Cherokee Orchard Road, Elkmont Road, and several trails due to the Chimney 2 Fire. The fire has grown to approximately 500 acres in size overnight due to extreme, sustained winds over 20 mph. National Park Service fire crews are focusing on suppression efforts to ensure public safety and protect park facilities at locations such as Chimneys Picnic Area. The fire…
At the over look before the chimney tops picnic area during tje fires
Park Fire Update Great Smoky Mountains National Park officials reported additional fire activity including the Park Headquarters area and a spot fire between Elkmont and Newfound Gap Road off of the Sugarland Mountain Trail approximately 1 mile south of the Husky Gap Trail intersection. The park has closed the Gatlinburg Bypass and Little River Road from Sugarlands Visitor Center to Metcalf Bottoms Picnic Area due to fire activity and downed trees. The park has evacuated employees from…
First responders with the Gatlinburg Fire Department
A man is trying to find his family during the tragic fires and evacuations in Sevier County.
Dollywood Foundation giving $1,000 per month for 6 months to families who lost everything in the wildfires.
Rev. Ed Taylor was found dead outside of his hime due to smoke inhalation, according to son who spoke with detectives.
Another chimney is all that's standing
More damage from the Gatlinburg fires
One of the trucks that was one of many burnt up in the Gatlinburg fires

subset: commoncrawl


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Art as a Coping Tool in Eating Disorder Recovery By Z Zoccolante Art can be an important tool in your eating disorder recovery. Eating Disorders thrive on rigidity. They tell us what to eat, what not to eat, how much to run, when we'll starve, purge, or hide away from the world. Eating disorders are built on control and structure. Art can be flexible and fluid. Art can be wild and spacious and free. Art can be what we want it...Read More
You Are the Constant in Eating Disorder Recovery By Z Zoccolante In life, and in your eating disorder recovery, you are the constant. Let's be honest, when we have an eating disorder, recovery feels like the most daunting path we face. There are reasons the eating disorder started; therefore, recovery is a process of exploring little black holes, and learning the skill sets we missed. Recovery can feel as though it depends on...Read More
How to Set a New Year's Resolution with an Eating Disorder By Z Zoccolante Listen to the audio of this post in the blue box below! *This post first appeared on HealthyPlace.com Each year, as the New Year rolls around, millions of us with eating disorders set our resolutions. One of the number one American resolutions is to exercise more, with gym memberships spiking in January and then waning soon after. Despite good intentions, many...Read More
The Link Between Substance Abuse And Eating Disorders Guest Post – By Chloe P. Substance abuse disorders can affect many age groups, but when they are combined with another disorder they can be deadly. Studies have found links between drug and alcohol abuse and eating disorders, depression, and PTSD, all of which are exacerbated by the interaction. In fact, research suggests that up to 50% of those living with an eating...Read More
Why You Don't Want Your Eating Disorder to Magically Disappear By Z Zoccolante (Listen to the audio of this post in the blue box below!) *This article first appeared on HeathyPlace.com You don't want your eating disorder to magically disappear. You may be thinking, "Um, yeah I do," but hear me out. In counseling, there's something known as the "miracle question." Often it sounds like, "If you woke up tomorrow and you no longer...Read More
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T2 Online Newsdesk
20 April 2017, 11:51 AM
A Yuvraj Singh over changed the tide of the game
Utathya Nag @utathya10
15 March 2017, 12:38 PM
The wicketkeeper-batsman recently surpassed Virat Kohli to become the fourth most successful batsman in T20Is
Utathya Nag @utathya10
14 March 2017, 01:02 PM
Ashwin replaced Bangladesh talisman Shakib Al Hasan to climb to the No.1 spot
Ayush Chatterjee @gbm_xii
27 October 2016, 12:30 PM
Captain Dhoni pulled of a brilliant run out to dismiss New Zealand's Ross Taylor but ultimately Team India were victims of their own poor batting...
Ayush Chatterjee @gbm_xii
23 September 2016, 05:52 PM
After the hosts were dismissed for 318 early on in Day Two, Kiwi skipper Kane Williamson and Tom Latham provided stability to take the Blackcaps...
Prabhjeet Singh Sethi @SinghOfGoodTMZ
22 September 2016, 05:29 PM
With an economical spell of 3/67, Trent Boult choked the flow of runs for India. Cheteshwar Pujara and Murali Vijay failed to convert their stable...
Prabhjeet Singh Sethi @SinghOfGoodTMZ
21 September 2016, 05:16 PM
At Green Park Kanpur, Ravichandran Ashwin will lead the Indian bowling attack and Kiwi skipper Kane Williamson will be the star batting attraction...
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21 September 2016, 03:23 PM
Did you know Rahul Dravid is the most successful batsman in Test matches between the two nations? Here are some more interesting facts for you
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Williamson's world class batting, Ish Sodhi's sharp deliveries that can disturb the Indian batting line-up and Ross Taylor's vast experience. Here...
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United States
Metal Injection
All Types
Metal Injection
yesterday at 22:56.
Southern rock legend Gregg Allman of Allman Brothers Band has passed away. How many bands today wouldn't exist because of this legend.
Southern Rock Legend Gregg Allman Dead At 69; Tributes Pour In. - Metal Injection
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491 likes - 28 comments
KyLo-Ren Spike Forgony
today at 00:38
Rob Zombie for one. Dude is heavily influenced by old school southern rock
Mack Johnson
yesterday at 23:06
First Chuck Berry, then Chris Cornell, and now Gregg Allman! This is 2016 all over, again!
Florio Daniel Robert
yesterday at 23:42
First harmonized guitar leads and killer solos I heard were from Allman Brothers. RIP.
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Metal Injection
yesterday at 19:16.
Broken Hope bring the brutality. What do you think of their new music video?
Consume Gore With BROKEN HOPE's Video For "The Carrion Eaters" - Metal Injection
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64 likes - 5 comments
Metal Injection
yesterday at 18:02.
Take a bow, Wacken Open Air – this is so awesome.
Germany's Wacken Open Air 2017 Will Have Its Own Beer Pipeline, Because Of Course It Will! - Metal Injection
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1659 likes - 126 comments
Anthony Russo II
yesterday at 18:08
"Both fresh and waste water will run through these pipes too. Of course we will control the hygiene of the tubes all the time."
Wait. What? Did I read this right?
Steve Knight
yesterday at 18:07
Depends on what kind of beer you put in there. If it's something like Budweiser, that may as well be a sewer pipeline.
Stanley E. Supranowicz Jr.
yesterday at 18:59
Because beer is such an integral part of any great music festival. God forbid you go to something without drinking yourself stupid.
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Metal Injection
yesterday at 17:45.
New Incantation officially announced!
INCANTATION To Release Profane Nexus In August - Metal Injection
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107 likes - 4 comments
Micah Stackhouse
yesterday at 17:55
"Incantation is all geared up to bring forth its next sprawling void of black metal". I think there's something wrong with that statement...
Ben Johnson
yesterday at 18:17
Incantation are not black metal and don't have a facebook page. WTF is guy talking about?
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Metal Injection
yesterday at 16:53.
Everybody from Metallica to Foo Fighters to Linkin Park to actors like Brad Pitt and Christian Bale came out to pay their respects to Soundgarden's Chris Cornell at a memorial service.
METALLICA, LINKIN PARK, PEARL JAM Members Pay Tribute To Chris Cornell At Memorial Service (Photos + Video) - Metal Injection
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7232 likes - 99 comments
Alexandra Díaz
yesterday at 17:12
His death made me realized that i need and asked for help.😔
Chris Munoz
yesterday at 16:59
Rock in peace
Dave Johnson
yesterday at 16:54
Linkin Park punched a Black Hole Sun into someone's face, in tribute. And cause they brought up the new album.
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Metal Injection
yesterday at 15:25.
Remember when Danzig said he didn't want to do any more Misfits shows? Well now Danzig is open to more Misfits shows, while taking a jab at other metal bands…
Now DANZIG Says He'd Do More Shows With MISFITS, Slams Metal Bands Who Can't Play Live - Metal Injection
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1017 likes - 139 comments
Josh Taber
yesterday at 15:31
I love the misfits (every era) but if you can play guitar hero on medium.. you play just about any, if not all misfits songs live.
Sheikh Yasser Jamil
yesterday at 16:01
I wish when people made comments like this would at least name the band or the era the band is from. Big 4 still kick ass, Iron Maiden and the rest are killing it, Newer bands like Gojira are phenomenal.
Who are you talking about specifically?!
Blaine Haniff
yesterday at 15:44
What metal bands do this exactly? I know some use backing tracks for keyboards/absent mates, but the whole show? Pretty sure no one I listen to or even know of does that.
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Metal Injection
05/26/2017 at 21:45.
Soundgarden tributes, System of a Down news, Lamb of God thoughts, and much more in the 10 trending stories of the week!
A Drunk Black Metal Band, Tributes To SOUNDGARDEN's Chris Cornell, and 8 Other Trending Stories Of The Week - Metal Injection
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363 likes - 3 comments
Matt Grabon
05/26/2017 at 23:16
I still can't get over that Megadeth cover.
Chris Munoz
yesterday at 02:38
Still cant process cornells death
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Metal Injection
05/26/2017 at 21:25.
We're looking for production interns to join the team. See the post for details on what we're looking for.
METAL INJECTION Is Looking For Production Interns! - Metal Injection
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67 likes - 18 comments
Marc Jarrell
yesterday at 04:11
This would be something from a dream to work and do something that involves one of your passions and one of your lucky to possibly get paid would be unimaginable but ultimately would be amazing haha #keeping it real
Lance Marshall
05/26/2017 at 21:36
If you need someone to go round and round with Phil labonte some more I'm your guy lol
Eric Fogle
05/26/2017 at 21:39
Does it require one to be sober at work ... Cuuuuuase that's kinda a deal breaker ...
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Metal Injection
05/26/2017 at 21:00.
Enter this week's caption contest for your chance to win something from the giveaway pile!
CAPTION CONTEST: Enter Your Funniest Caption & You Could Win A Free CD From Our Giveaway Pile - Metal Injection
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59 likes - 27 comments
Lee Boughton
yesterday at 01:39
When you get an alien so stoned he thinks the closet is the bathroom and he waits in line so as not to draw attention to himself.
Muhammed MacTavish Behlol
yesterday at 16:01
When you earn your friends respect after telling mom that you need money to buy books, but bought metal tees.
Aaron Adair
yesterday at 02:45
Back in black dog. Stairway to hell. For those about to ramble on.
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Metal Injection
05/26/2017 at 20:30.
Our Funeral Doom Friday column investigates who should be on the Mount Rushmore of doom. You can help us decide!
Funeral Doom Friday: Who Is On The Mount Rushmore of Funeral Doom? - Metal Injection
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55 likes - 13 comments
David Mackowiak Jr.
05/26/2017 at 21:30
Esoteric, for sure. One of the biggest, most distinct acts in the genre. All the prog, psych and whatever-the-hell else they casually combine still astounds me, all these years later.
James Howlett
05/26/2017 at 21:36
Definitely Ahab if they still count. Last few releases weren't really funeral doom but they still rule.
Derek Ayers
05/26/2017 at 23:39
The last Ahab release was garbage, not even funeral doom! Esoteric are the true kings of funeral doom
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Metal Injection
05/26/2017 at 20:00.
We chat with Horisont about their amazing new album, and also post some killer live footage on the site!
HORISONT 'About Time' Interview - Metal Injection
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20 likes - 1 comments
Tim Petersön
yesterday at 02:57
Pontus is such a sick drummer. Love these guys.
Metal Injection
05/26/2017 at 19:30.
The Night Flight Orchestra's new album is the 80's infused party you want to be at! Here's our full review.
Album Review THE NIGHT FLIGHT ORCHESTRA Amber Galactic - Metal Injection
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53 likes - 1 comments
Eric May
yesterday at 02:14
One of the best albums of the last decade for me. :)
Metal Injection
05/26/2017 at 19:00.
Looking for a new band name? A research scientist used a neural network to generate unused metal band names that are up for grabs!
Here's What Happens When A Machine Learns Metal Band Names And Then Makes Up Its Own - Metal Injection
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484 likes - 53 comments
Andrés Zúñiga
yesterday at 00:50
I'm gonna be honest. Vultrum sounds like something I would listen. Folk/black from Germany? Not bad at all
Austin Chu
yesterday at 07:19
Executioner- death, thrash
Exorcise- black, doom
Ungod- black
Dunkleosteun (as in the giant, shark-eating prehistoric fish)- death, thrash
Lizardforce (Dragonforce cover band)- power
Trumpium (political themed Trivium parody band)- death, thrash
Tesla Coil (Lacuna Coil cover band)- gothic
Dethwish (Nightwish cover band)- symphonic, folk
Those are just a few that popped up in my head immediately. Feel free to take if you need ideas... :)
Eric Evans
05/26/2017 at 20:22
Ive come up with couple names that I thought would make killer band names. Death Camp Revival is one, tho it may stir up some controversy.
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Metal Injection
05/26/2017 at 18:30.
Seattle based members in 7 Horns 7 Eyes, Arch Enemy, and Ghost Ship Octavius pay a very metal tribute to Soundgarden's Chris Cornell!
Seattle Based Members Of 7 HORNS 7 EYES, ARCH ENEMY, and GHOST SHIP OCTAVIUS Come Together To Record An Amazing Cover Of SOUNDGARDEN's "Black Hole Sun" - Metal Injection
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1240 likes - 34 comments
Shane Krajewski
05/26/2017 at 19:51
That was an awesome cover. Glad to hear it was sung by someone with some chops and they didn't go overboard on the music. Fantastic stuff. RIP Chris. 😔🤘
Fabrizio Di Gioia
05/26/2017 at 21:34
Very good performance! They maintain the heart of song untouched, but the customizations sound all good! Much better than those who forget the lyrics or sing them out of tune too!
Darryl Jeffrey
yesterday at 05:04
I actually think this is a lower key vocal version of the actual a Capella Black Hole Sun played over their new music. So no wonder it sounds like Chris. Correct me if I'm wrong.
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Metal Injection
05/26/2017 at 18:03.
Here's video of Tool performing "Eulogy" for the first time in 16 years!
TOOL Performed "Eulogy" Live For The First Time In 16 Years - Metal Injection
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4897 likes - 313 comments
Amanda Scott
05/26/2017 at 19:19
My bf and I were there last night, as part of my 30th birthday present! It was an amazing show, and to hear both Eulogy and Aenima (My fave Tool song) was such a treat! I've seen them 3 times in the last 11 years they've toured the East Coast, and if I never get to see Tool live again, I can die a happy woman.
Eam Gerard
05/26/2017 at 18:34
I was there and knew it was a treat but didn't realize it's been that long since they played it. What a show, they sounded incredible.
Brandon Fitzgerald
05/26/2017 at 18:48
Fuck yes! When I saw them, this is THE song I wanted to hear...And didn't :( they had a boring as fuck hour long intermission of laser lights and alien noises, instead of playing more songs. Needless to say, I thought they sucked live for that reason
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Metal Injection
05/26/2017 at 17:30.
The new Queens of the Stone Age album sounds promising!
QUEENS OF THE STONE AGE Frontman Says New Album Is Uptempo, Won't Reveal Release Date - Metal Injection
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781 likes - 40 comments
Darrell Schuldiner
yesterday at 02:55
Why is this even on metal injection??? In real metal news, Kalmah is about to start recording their next album. Why don't you give us some details on that instead metal injection?
Michael Garner
05/26/2017 at 20:23
Would be nice if Josh Homme went back to doing his stuff from Kyuss. That's some pretty damn good Stoner Rock.
Anthony Thomas
yesterday at 16:17
At least use a recent picture, Joey Castillo haven't been in Queens for at least 3 years lol
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Metal Injection
05/26/2017 at 17:00.
Matt Pike (High On Fire, Sleep) jamming with kids might be the cutest thing you see all day!
SLEEP's Matt Pike Jammed With Kids From School Of Rock Portland, And It Ruled - Metal Injection
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557 likes - 25 comments
Lyall Storey
05/26/2017 at 18:52
I love matt pike, but he doesn't strike me as the kind of guy I'd like to have around children
Matt Phillips
yesterday at 12:39
Who's the old dude with a shirt on? I thought Matt Pike was in this vid?
Tom Sekerak
05/26/2017 at 23:37
Its good to see musicians jamming with kids. Keep laying the foundations for future music.
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Metal Injection
05/26/2017 at 16:30.
Uh Oh, The Faceless have canceled another tour! What's going on here?
THE FACELESS Cancels Tech Fest 2017 Appearance, Entire European Tour - Metal Injection
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282 likes - 89 comments
Jon Milles
05/26/2017 at 16:40
Why worry about the faceless when you can listen to ENTHEOS, Alluvial, John Frum, INFERI, Good Tiger, Black Crown Initiate and all of the other amazing bands with former faceless members?
Jesse Gray
05/26/2017 at 16:36
I don't know why they don't just call it a day? It's been a good run... I just I think it's time for everyone to move on both fans and band members included =/
Chris Aguilar
05/26/2017 at 17:15
Instead of promoting/posting about band that obviously doesn't give a shit music, touring, fans etc....how about you take a second and check out a band that does.
Machines of Man
Releasing debut album "Dreamstates" 6/27/17 - mixed and mastered by Jamie King (Between the Buried and Me, The Contortionist). Indiegogo campaign up to raise money for tour! Link below!
[ Igg.me Link ]
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Metal Injection
05/26/2017 at 16:03.
Danzig, Dream Evil, Alestorm and more have albums out today! Which are you picking up?
THE WEEKLY INJECTION: New Releases From SOLSTAFIR, ALESTORM, and More Out Today - 5/23 - Metal Injection
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185 likes - 19 comments
Andrea Allensworth
05/26/2017 at 18:28
Lol @ Fucked with an Anchor, I made sure to see and dedicate that gem by Alestorm to my best friend.
Fernando Sierra
yesterday at 15:01
Been waiting for Dream Evil Long time! Specially since Mark u Black is back from the dead ! ;) \m/ \m/
Diego Thomsen
05/26/2017 at 20:22
Man, the new Cydemind is incredibly good. Worth the wait.
Prog metal lovers will surely love it.
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Metal Injection
05/26/2017 at 15:30.
Born of Osiris are in pre-production for the next album, and teased some very heavy music!
BORN OF OSIRIS Is Demoing A New Album, And It Sounds Heavy - Metal Injection
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1611 likes - 114 comments
Remus Stingaciu
05/26/2017 at 17:15
Sounds very promising. Just hope they drop their failed attempts at djent. Soul Sphere was a 50/50 deal, with killer tracks and boring tracks. Maybe their re-recording of New Reign inspired them to do more of that.
Ashe Frankie LeBlanc
05/26/2017 at 22:40
I've loved BOS for a long time and I also liked soul sphere a lot (a rare combination, I know)...so I fully expect their next album to be just as bad ass as everything they put out 🤘🏻
Josh McGaughey
05/26/2017 at 16:52
bring back the sound of the new reign and/or the discovery pls, every album after has become far too repetitive and predictable, you guys used to be one of my fav bands now your just kind of alright...
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Featured Videos
In recognition of Memorial Day, WWE and its Superstars thank and remember the heroes who made the ultimate sacrifice for freedom.05/25/2017 - 20:00
Bayley tries out a new Seth Rollins-inspired hairdo and talks Kendo sticks with Tommy Dreamer. Cathy Kelley celebrates Red Nose Day, looks at what The Architect had to say and more.05/25/2017 - 15:45
In celebration of the 40th anniversary of the original "Star Wars" film, Zack Ryder checks out Hasbro's line of highly detailed Black Series action figures, which are contained in throwback Kenner packaging.05/25/2017 - 12:00
The Internet can't stop arguing about rompers for men, but would Superstars like The Miz, Dolph Ziggler and Enzo Amore wear one05/24/2017 - 14:45
Karl Anderson throws out the first pitch at Great American Ball Park before the Cincinnati Reds take on the Cleveland Indians.05/24/2017 - 17:00
The WWE Champion spared no expense in celebrating his improbable victory. See slow-motion highlights of Jinder Mahal's Punjabi Celebration from SmackDown LIVE.05/24/2017 - 00:00
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The Higher Education Ministry denied news reports that some 2,000 professors submitted a petition against a Higher Education bill.
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Sarah Michelle Gellar Spends the Night in the Emergency Room With Her 5-Year-Old Son
Wochit Headline News - Friday, 13 January 2017
Sarah Michelle Gellar may be an experienced mother of two, but she still faced a mommy first Thursday night when she had to pay a visit to the emergency room. "Well I made it 7 years before I had to take one of my children to the ER (I think that's a pretty good stretch)," she told fans Friday morning on social media. The star described how she had to bring her youngest child, 5-year-old son Rocky James Prinze, to the hospital because he was battling a virus and was having trouble breathing. "It was a long night and apparently a nasty virus, but thankfully Rocky was released," she explained. "I'm not going to lie, watching your child not be able to breathe, is the most helpless I have ever felt." As Rocky recuperated, he stayed entertained with the help of an iPhone. "Thank goodness for #iphones and #inspectorgadget when your little one is sick in the ER," the actress tweeted. While her little one has since returned home, the devoted mom expressed special gratitude for his caretakers and noted the silver lining of the entire experience. "Thank you to all the great doctors and nurses (and staff) who work tirelessly 24 hours a day to help. And thank you to his big sister, Charlotte who would not leave his side," she wrote on Instagram. "(The best moment was when I was filling out his paperwork, and I was asked if Rocky was single- laughter is definitely a great medicine)."
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Ice pops
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Creamy, dreamy, Oreo cookies blended with healthy coconut milk for a fabulously refreshing popsicle!
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Just give in to Big Brother. There is no other way.
Just give in to Big Brother. There is no other way.
Edible star glitter! I will always be blown away by the variety of food decorations that look inedible but are in fact safe! Also, the little fondant mug on top of this cupcake is filled with butterbeer so you get a little sip before you eat the creamy goodness below. Genius.
He's just popping up to say hello, no need to be alarmed.
Each mustache has its own personality, just like every morning has its mood. Make sure they match up when you're drinking your coffee.
Finally, a mug that looks like duct tape but doesn't get all gummy and nasty when it gets warm.


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We stopped here while staying at Hunting Island. The food was really good and they have the very best egg rolls that I can remember having!
1 - 10 of 14 reviews
We stopped here while staying at Hunting Island. The food was really good and they have the very best egg rolls that I can remember having!
We ordered Chinese to take home to share with our family. I have eaten.Chinese from New York to Miami, and to Seattle This was only fair at best. I ordered shrimp with cashews and the cashews were crammed on top of the order and the...More
I am hard to please when it comes to Chinese food: I have lived in and travelled extensively through China. I was married to a Chinese woman was a great cook, too. So I generally stay away from small strip mall Chinese restaurants as my...More
if you need something fast, good tasting, and filling for a good value for the price, get some good Chinese food at Peking Gourmet. They are nice, get your order the first time over the phone, and ready for pickup in less than 15 mins...More
Peking Gourmet always has delicious, fast, and always fresh food. The servings are more than generous and extremely well-priced. Love this place! Try the pineapple chicken, the lemon chicken, the hot and sour soup, and everything else on the menu. We love this place!
Good Chinese food and a big variety of things to buy (and eat). I enjoy looking at the large menu and ordering several things in smaller size to have a mixture of things to eat. Eating there is OK, but I would rather take it...More
First visit here, was glad to see the number of customers. Should have noted they were all waiting. Counter woman was pleasant and smiling, but was highly distracted by trying to conduct business while arguing loudly with her daughter on speaker phone. The stir fry...More
We stopped here on vacation and while this has the typical hole in the wall Chinese restaurant look, the food was very good and much better than others I have been to. We ended up ordering at the counter and stayed to eat there. We...More
I've eaten food from this restaurant many times and the food has been consistently delicious but I insist that they go easy on the oil. There is a large selection of menu items which are moderately priced and the food is prepared to order.
We wanted something other than fast food, so we looked for Chinese close by our hotel. Peking Gourmet came up on the map and we decided to give it a try. It looks just like all the Chinese restaurants in strip malls around our home...More


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Foster J. Sayers
|Foster Joseph Sayers
April 27, 1924
Centre County, Pennsylvania
|November 12, 1944
(aged 20)
Thioville, France
|Place of burial
|Howard, Pennsylvania
|United States of America
|United States Army
|Years of service
|1943 - 1944
|Private First Class
|357th Infantry, 90th Infantry Division
|World War II
|Medal of Honor
Bronze Star
Purple Heart
Foster Joseph Sayers (April 27, 1924–November 12, 1944) was a 20-year-old infantryman from Centre County. He joined the Army from Howard, Pennsylvania in March 1943.[1]
He received the Medal of Honor for acts of bravery near Thionville, France on November 12, 1944. Foster Joseph Sayers Reservoir at Bald Eagle State Park near his hometown of Howard, Pennsylvania is named in his honor.
He is buried in Schenck's Cemetery in Howard, Pennsylvania.[2]
Medal of Honor citation[edit]
Private First Class, U.S. Army, Company L, 357th Infantry, 90thInfantry Division. Entered service at: Howard, Pa. Birth: Marsh Creek, Pa. Place and date: Near Thionville, France, November 12, 1944. Killed in action. G.O. No.: 89, October 19, 1945.
He displayed conspicuous gallantry above and beyond the call of duty in combat on 12 November 1944, near Thionville, France. During an attack on strong hostile forces entrenched on a hill he fearlessly ran up the steep approach toward his objective and set up his machinegun 20 yards from the enemy. Realizing it would be necessary to attract full attention of the dug-in Germans while his company crossed an open area and flanked the enemy, he picked up his gun, charged through withering machinegun and rifle fire to the very edge of the emplacement, and there killed 12 German soldiers with devastating close-range fire. He took up a position behind a log and engaged the hostile infantry from the flank in an heroic attempt to distract their attention while his comrades attained their objective at the crest of the hill. He was killed by the very heavy concentration of return fire; but his fearless assault enabled his company to sweep the hill with minimum of casualties, killing or capturing every enemy soldier on it. Pfc. Sayers' indomitable fighting spirit, aggressiveness, and supreme devotion to duty live on as an example of the highest traditions of the military service.[3]
See also[edit]
- ^ WWII Army Enlistment Records
- ^ "Foster J. Sayers". Claim to Fame: Medal of Honor recipients. Find a Grave. Retrieved 2007-11-13.
- ^ "Foster Joseph Sayers - Congressional Medal of Honor". United States Congress. Retrieved 2006-12-19.
- "Foster J. Sayers". Claim to Fame: Medal of Honor recipients. Find a Grave. Retrieved 2007-11-13.
- "Foster Joseph Sayers - Congressional Medal of Honor". United States Congress. Retrieved 2006-12-19.


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When visiting Burton, Red Maple Inn is a great choice to consider. Guests can visit the fitness center for a workout, and free perks include WiFi and continental breakfast daily between 7 AM and 10 AM. Rooms offer jetted tubs, flat-screen TVs, and DVD players.


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Isakson, Chambliss appear ready to OK Syria military action
Both of Georgia's Republican U.S. Senators released statements over the weekend in which they appear ready to vote in favor of the U.S. taking military action against Syria, reports WABE.
The statements followed President Barack Obama's speech Saturday regarding his plan to seek authorization from Congress to use military action against Syria. Congress returns to Washington, D.C., on Sept. 9 from recess.
U.S. Senator Johnny Isakson said he supports the use of military action. U.S. Senator Saxby Chambliss said he believes the evidence is clear Obama's 'red-line' was crossed long ago, WABE reports.
Isakson added that it's appropriate to seek authorization from Congress, but that he would have preferred a vote be called immediately rather than waiting until Sept. 9.
Chambliss said while he appreciates that Obama sought Congressional approval, he should have already presented Congress with a strategy and objectives for military action, including what impact such action will have on U.S. allies and enemies in the region.
WABE adds that on Friday, U.S. Representative David Scott, a Democrat who represents parts of Atlanta, urged the President to consult with Congress before taking action in Syria. Scott told the station Friday that U.S. action in Syria doesn't seem warranted. Click here to read more of that interview and for a link to hear the broadcast.
Click here to read more of WABE's report, which includes the full statements released by Isakson and Chambliss.


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Karen Shepard releases statement in her bid to unseat Chris Winters on the Williamstown Planning Board
|Karen Shepard, third from right, with her husband, Jim, left, and two children
- VIDEO: Planning Board candidate forum
- VIDEO: Meet the candidates on WilliNet
- Shepard's personal blog
- Be the kind of person who
wants to listen and learn.
- Have to have common sense; you have to have the energy to try to make things happen, and, most importantly, it seems to me, you have to be able to put yourself in other people's shoes.
- Listen to what everyone thinks they want and need, and then have open, honest, and civil discussions, and figure out first what needs to be done and second how to roll up your sleeves and do it.
- The Planning Board Should Represent the Town. The Planning Board should represent and support the town's vision for the future of our community; individual agendas should not outweigh public vision.
- The Town's Vision is an Achievable Vision. Economic development, encouraging growth, and the protection of open space can work with not against each other.
- We Need Flexible Residential Areas. Increasing the flexibility of residential areas through high density, mixed-income, multi-family, and affordable housing, and modified zoning in concentrated areas will promote a vibrant, populated residential center that will energize not just the tourist trade, but the local residents' day-to-day life, and will also care for our growing elderly population and attract younger residents to town.
- Concentrated Development is Good Development. Concentrated development reduces sprawl, allowing for the protection of our valued natural resources in larger parcels on the outskirts of town. We need zoning that encourages sustainable high-density, mixed-use development in certain districts.
- Protecting Open Space is Pro-development. It increases the value of abutting lots; it attracts tourists; protecting open space on the outskirts of town encourages development at the center.
- An Educated Town is a More Prosperous Town. Did you know, for example, that
although our town is 83% open space, only 29% of that is protected, the
smallest percentage among eight Northern Berkshire towns? Or that when you apply absolute and
partial constraints to the Limited Industrial and Planned Business districts,
the amount of land available to develop more or less disappears? The more we know, the more we can
- We Need Change on the Planning Board. A town is only as strong as the number of people involved in the way it's run. Too often the same handful of people fill the same local government positions. The incumbent has been on the Planning Board for ten years. Someone who isn't as familiar with "the way things have always been done" can bring fresh eyes to old problems.


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Automotive Business Strategy and Innovation
Future of Sports Utility Vehicles in the United States, 2016-2022
Socio-Economic and Political Influence Projected to Drive SUV Segment Market Share to 42% in the US by 2022Release Date : 18-May-2017 Region : North AmericaSUVs in the US have been growing steadily since 2012 and have over taken passenger cars as the largest selling passenger vehicle segment in US in 2016. This recent spike in SUV sales has encouraged auto OEMs to capitalize on the demand and increase their SUV model line-up. Over 20 new SUV name plate...USD 1,500
Women as Vehicle Customers—Voice of North America Consumer Findings, 2016
40% will Consider Digital Retail Platforms for their Next Vehicle Purchase (2020)Release Date : 12-May-2017 Region : North AmericaThe percentage of women holding driving licenses in North America is greater than that of men. However, women are unhappy with the entire process of car buying. Hence, understanding women car purchase preferences is essential for OEMs to ensure that the process is amicable. With the growth of digita...
African Commercial Vehicles Market, Forecast to 2025
Despite Slowdown, the Market is Expected to Grow at a Robust CAGR of 6.1% until 2025, Driven by Increasing Infrastructure DevelopmentsRelease Date : 05-May-2017 Region : AfricaThe African commercial vehicles (CV) market, despite the slowdown due to low oil and commodity prices, is expected to grow at a compound annual growth rate (CAGR) of 6.1%, largely driven by increasing infrastructural developments and rising urbanisation. South Africa is the largest contributor to th...
Global Urban Rail Vehicle Database, 2016
Interactive Dashboard for Rail OEMs, Rail Operators, and Vendors with Details of Networks and Vehicles of Global Urban Rail Transit SystemsRelease Date : 05-May-2017 Region : GlobalThis research service covers all types of urban rail transit vehicles and rolling stock around the world. The vehicles are listed by region, sub-region, country, city, transit agency, transit classification (Metro, light rail, automated people mover, monorail, streetcar, heritage streetcar, automate...
Global Bus Rapid Transit System (BRT) Database, 2016
Interactive Dashboard for Bus OEMs, Transit Operators and Vendors with Details of Networks and Buses of Global BRT SystemsRelease Date : 04-May-2017 Region : GlobalThe Global Bus Rapid Transit System Database, 2016 is an interactive dashboard for bus original equipment manufacturers (OEMs), transit operators, and vendors with details of networks and buses of global bus rapid transit (BRT) systems. The base year is 2016 (2014/2015 data has been provided where t...
BMW M Division—Dedicated Driving Dynamics
Pushing the Limits of Luxury and Performance for 45 YearsRelease Date : 21-Apr-2017 Region : EuropeBMW prides itself in creating best-in-class performance luxury vehicles in terms of driving dynamics. BMW M takes this a step further by providing higher power and torque output powertrains and handling capabilities that enhance overall vehicle performance. The division has 3 distinct strategies to ...USD 1,500
Analysis of Top 30 Features of Mass Market and Luxury OEMs in North America, 2016–2025
Driver-centric Technologies Growing at 15%, Creating Opportunities for OEMs and Tier I Suppliers AlikeRelease Date : 11-Apr-2017 Region : North AmericaGovernment mandates on safety and emission regulations are driving auto Original Equipment Manufacturers (OEMs) to offer advanced technology features in cars. Advancement in the field of Artificial Intelligence (AI), in-vehicle connectivity, and electric vehicle technology has enabled OEMs and suppl...
Game-changing Automotive Policy Developments in Africa, 2016
Automotive Investment Policy Review for Africa and its Impetus to OEMsRelease Date : 31-Mar-2017 Region : AfricaGame-changing Automotive Policy Developments in Africa Automotive Investment Policy Review for Africa and its Impetus to OEMs As Africa speeds up its efforts to industrialise, the automotive sector has attracted attention from a number of governments on the continent. With over 1 billion inhabita...USD 3,000
Vehicle Platforms and Fuel Strategies of Car Manufacturers in India
15 Modular Platforms Anticipated to Account for 74.8% of the Total Market Production by FY 2023Release Date : 30-Mar-2017 Region : South Asia, Middle East & North AfricaThis research service focuses on various modular platform and fuel strategies of car manufacturers including competitive analysis, technology proliferation, and key manufacturer profiling. Car manufacturers in India are currently investing in modular platform development to minimize costs, enhance c...
Outlook of Select ASEAN Automotive Markets, Forecast to 2017
Sales Will Reach 3.25 Million Units in 2017, with Preference towards Crossover and Entry-level VehiclesRelease Date : 22-Mar-2017 Region : Asia PacificThis study aims to provide a detailed overview and analysis of the trends in the Association of Southeast Asian Nations (ASEAN) automotive market, with a focus on trends in the 3 key markets of Malaysia, Indonesia, and Thailand. Research Scope • This ASEAN automotive outlook includes key market tr...


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The women of Oak Hill are involved with ongoing ministries throughout the church body including weekly Bible Studies, Sunday School, local and international missions, Worship & Arts, and much more. Please join us as we serve along side each other, pray for each other, and encourage each other on the journey!
Tuesday Morning and Evening Bible Studies will kick-off again in August! Check out the latest activities and friends on facebook.