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We combined the use of a congenic line and recombinant progeny testing (RPT) to characterize and fine map a previously identified region of distal mouse chromosome 2 (MMU2) harboring quantitative trait loci (QTL) with large effects on growth and fatness. The congenic line [M16i.B6-(D2Mit306-D2Mit52); MB2] was created using an inbred line (M16i) derived from a line that had undergone long-term selection for rapid weight gain (M16) as the recipient for an ∼38-cM region on MMU2 from the inbred line C57BL/6J. A large F2 cohort (1,200 mice) originating from a cross between MB2 and M16i was created, and 40 F2 males with defined recombinations within the QTL region were used to produce 665 segregating progeny. Linkage analysis of the F2 population detected QTL with very large effects on body weight, body fat, lean tissue mass, bone mineral density, and liver weight. Confidence intervals of the QTL were narrowed to regions of 1.5–4.5 cM. Analysis of progeny of the recombinant F2 males confirmed the existence of the QTL and further contributed to localization of their map positions. These efforts confirmed the presence of QTL with major effect on MMU2, narrowed the estimated region harboring the QTL from 38 to 12 cM, and further characterized phenotypic effects of the QTL, effectively culminating in a significantly decreased pool of positional candidate genes potentially representing these genes controlling predisposition to growth and fatness.
- fine mapping
- quantitative trait locus
- obesity
- congenic
- recombinant progeny testing
body weight, body composition, and energy stored in adipose tissue are determined by a complex combination of genetic and environmental factors (6, 44). Identification of genes underlying predisposition to these traits may help to elucidate the fundamental mechanisms of growth and obesity. In humans, such knowledge would be valuable in the development of diagnostic and therapeutic approaches to treat obesity and comorbidities. In food-producing animals, it could enable manipulation of growth and body composition through marker-assisted selection or by transgenic strategies.
Lines of mice that have undergone long-term selection provide a uniquely powerful model to dissect and understand the complex polygenic architecture of quantitative traits with relevance to biomedicine and animal agriculture. The M16 line was the result of 27 generations of selective breeding for rapid growth rate (15), resulting in correlated responses of polygenic obesity and symptoms of type II diabetes (2). Full-sib mating within M16 has led to a fully inbred derivative denoted as M16i (21).
One method for identifying genes that regulate complex traits is quantitative trait locus (QTL) mapping. Many successful efforts to map QTL for growth and obesity in mice (33) and humans (43) have been published, including large-scale studies employing the M16i line (37, 38). However, a deficiency of the QTL approach is that resolution of mapping includes broad genomic confidence intervals encompassing hundreds or even thousands of genes, rendering positional cloning an extremely difficult proposition.
Several strategies have been proposed to increase the physical resolution of QTL detection (1) including fine mapping, using interval-specific congenic strains (11, 12), genome-tagged mice (19, 20), and chromosome substitution strains of mice (42). Congenic strains differ from their founder strains only for the genomic region containing a specific, well-defined chromosomal interval. Other strategies such as recombinant progeny testing have been successfully incorporated into fine-mapping paradigms to increase resolution (8, 10, 18, 26, 27, 52). The basis of progeny testing is to identify recombinant individuals in the interval where the putative QTL is located, propagate progeny representing the alternative allelic forms in this region, and determine whether there is a phenotypic difference among allelic class means (26).
In our QTL studies using the M16i line, a common finding is that the distal region of chromosome 2 harbors one or more QTL with very large effects on traits related to growth, body weight, and fatness (33, 37, 38). To fine map these QTL, we developed a congenic line [M16i.B6-(D2Mit306-D2Mit52); MB2] using M16i as the recipient for a ∼38-cM region on mouse chromosome 2 (MMU2) from the inbred line C57BL/6J. The phenotypic characterization of the MB2 congenic line was described previously (21). Briefly, absence of M16i alleles in this 38-cM region on MMU2 resulted in a 60% decrease in adiposity and 15% decrease in body weight in MB2, confirming the existence of loci within this region that have large effects on these traits. Several other studies have found QTL in this region with effects on body weight and adiposity (34, 49), and recent reports have described characterization of congenic lines with a variety of donor and recipient backgrounds (13, 49) with relevancy to gene discovery on MMU2.
We now report on use of the MB2 congenic strain to fine map and better characterize the QTL on distal MMU2. Using a combination of a large F2 intercross between M16i and MB2, and extensive progeny testing of F2 males with defined recombinations within the QTL region, we have built a map of QTL in this region, estimated their locations within intervals of 1.4–4.5 cM, and further defined the phenotypic consequences of alternative QTL alleles. Results of this effort will be instrumental in facilitating positional cloning of the underlying polygenes in this region of the mouse genome that help regulate predisposition to growth and obesity.
Mouse lines.
Mouse lines used in these studies included the polygenically obese M16i line and the MB2 congenic line [M16i.B6-(D2Mit306-D2Mit52)] derived from the M16i line as recipient for a ∼38-cM region of MMU2 from the inbred line C57BL/6J (C57) obtained from the Jackson Laboratory (Bar Harbor, ME). The M16 line was derived from an outbred population (ICR) selected for rapid 3- to 6-wk weight gain for 27 generations (15). Subsequently, over 20 generations of full-sib mating were implemented to produce the inbred M16i line. Details of development of MB2 and phenotypic comparison between MB2 and M16i were previously reported (21).
Mouse care and maintenance.
Fraternal litter size was standardized 1 day after birth to 10 pups per litter. Pups were weaned at 3 wk and housed three to four per cage (by sex) with ad libitum access to feed (Teklad 8640 Rodent Chow) and water. Mice were reared at 22°C and 30% relative humidity and with a 12:12-h light-dark cycle, with lights on at 0700. All procedures and protocols were approved by the University of Nebraska Institutional Animal Care and Use Committee.
Experimental design.
The general experimental approach for fine mapping the MMU2 QTL involved creation of an F2 intercross population between M16i and MB2, followed by progeny testing of recombinant F2 males (Fig. 1). Work was divided into three specific phases: 1) initial selective genotyping of F2 with extreme fatness phenotypes to narrow the region containing the QTL, 2) QTL analysis of the entire F2 population to obtain additional mapping resolution, and 3) recombinant progeny testing of recombinant F2 males to evaluate specific and defined segments for the presence of QTL.
Creation of the M16i × MB2 F2 intercross population.
A large F2 population (n = 1,200 mice) was created over four consecutive time periods (replicates) from a cross between MB2 and M16i. Matings in periods 1 and 3 were repeated in periods 2 and 4, respectively. The resource population was created by reciprocal crossing of five males with five females representing each line to produce the F1 generation, and then intermating the resulting F1 males and F1 females to produce the F2 population.
Recombinant progeny testing.
Thirty-five males carrying a recombinant segment of chromosome 2 between markers D2Mit304 and D2Mit52 were used in the F2 population as sires for recombinant progeny testing (RPT). These males possessed one evident recombinant (based on marker genotyping) in at least one of seven possible marker-to-marker intervals of the congenic segment. The progeny test involved crossing each recombinant male to four C57 and two M16i females, generating 424 and 241 F3 mice, respectively.
Measurement of phenotypes.
The same phenotypic variables were measured in both the F2 and RPT populations. Body weight was measured at 3, 6, 9, 12, and 15 wk of age. At 15 wk, all mice were killed by cervical dislocation after 1.5 h of fasting. The head was removed and trunk blood was collected, with serum subsequently separated by centrifugation and stored at −80°C. The subcranial body was then scanned using dual-energy X-ray absorptiometry (DEXA; Lunar, Madison, WI) to measure total body fat (FAT), total lean tissue (LEAN), and overall bone mineral density (BMD). These DEXA traits were evaluated for replicates 3 and 4 of the F2 population and all RPT progeny. For all mice, the right-side epididymal (male) or perimetrial (female) fat pad (PAD) and liver (LIV) were dissected, weighed, snap frozen in liquid nitrogen, and stored at −80°C. Serum glucose was measured with the glucose oxidase method, using the SureStep Blood Glucose Monitoring System (LifeScan).
DNA lysate was prepared from toe clips (used to identify mice at 12 days of age) or from tail clip samples, following a general method described previously (35). Aliquots of 5 μl of DNA lysate were dispensed into 384-well plates, dried at room temperature, and stored at −40°C. Genotypes were determined by PCR using infrared dye labeling, followed by electrophoresis and analysis on the LI-COR Model 4200 IR2 system (LI-COR, Lincoln, NE). Gels were analyzed using Gene ImagIR analysis software (LI-COR) to determine allele sizes of each individual.
Genotyping was performed in three phases. Initially, selective genotyping was employed based on weight of epididymal and perimetrial PAD as percentages of body weight, selecting the highest and lowest (48 mice in each of the high and low groups, with approximately equal representation of each sex) from replicates 1 and 2. These 96 samples were scored for 16 markers covering the complete congenic segment of MMU2. Second, all F2 males from replicates 3 and 4 were genotyped for eight markers [between markers D2Mit304 and D2Mit52 (Table 1), as determined by analysis of the selective genotyping results]. This step identified males with specific recombinations to be used for RPT. Finally, all individuals in the F2 and RPT populations were subsequently genotyped for these eight markers.
Statistical analysis: selective genotyping.
Replicates 1 and 2 of the F2 population were used for these analyses. Deviation from Mendelian expectations (1:2:1) of individual markers within selectively genotyped mice was evaluated using two chi-square analyses. The first determined whether genotypic frequency for the given marker within the high or low selected phenotypic groups differed from the 1:2:1 Mendelian ratio expected for the null hypothesis of no linkage of the given marker to a hypothesized QTL in the region. The second analysis tested for marker allele frequency equivalence between the high and low selected phenotypic groups (High-Low), which is also expected under the null hypothesis of no linkage of the given marker to a QTL. Effects of individual marker genotypes were evaluated by analysis of variance, using the PROC MIXED procedure of SAS (39). Fixed effects included sex, marker genotype, and genotype-by-sex interaction, with dam as a random effect.
Statistical analysis: QTL detection.
A total of 18 traits were analyzed for the presence, most likely position, and effects of QTL in the complete F2 population (n = 1,200). Descriptive statistics and distributions for each trait were calculated using the MEANS and UNIVARIATE procedures of SAS (39). Various fixed effects were tested for each trait, including sex, replicate, family, and possible two-way interactions among these, using the GLM procedure of SAS (39). Traits displaying a significant departure from a normal distribution (PADP, FATP, LEANP; see results) were normalized, using a log transformation. Phenotypic correlations among the traits, adjusted for sex, were estimated using multivariate analysis of variance (PROC MANOVA; SAS).
To test for segregation distortion, marker genotype frequencies in the F2 were compared with the expected Mendelian ratio of 1:2:1 using a chi-square test. For all markers, the null hypothesis of expected Mendelian segregation was not rejected using a Bonferroni protected significance threshold of P > 0.006. This analysis also enabled evaluation of systematic genotyping errors, and none were found. Marker linkage maps were generated with MAPMAKER/EXP and marker distances calculated in this specific cross (Table 1) were used in the subsequent QTL analysis. Positions of markers and QTL (with their respective confidence intervals) were estimated in the physical map, using the Ensembl Mouse Genome Server.
Residuals from the fixed effects models were utilized with QTL Cartographer (v.1.15; Ref. 4) to first perform simple interval mapping (SIM) analysis (23). To obtain better resolution and precision of QTL location and effect, a composite interval mapping (CIM) analysis (53, 54) was performed. A forward-backward stepwise regression procedure (4) with a 0.01 threshold for the addition and elimination of new markers was used to select background factors for CIM, and a 0.5-cM window size was adopted. Experiment-wise significance thresholds were established with at least 1,000 permutations (4, 9). Additive (a) and dominance (d) effects of each QTL were estimated. Effects of QTL were estimated as the percentage of residual phenotypic variance explained by the QTL, and confidence intervals (CI) for each QTL were determined using a 1-LOD drop from the QTL peak (23).
Statistical analysis: RPT.
To further fine map QTL, two approaches were used: the recombinant ancestral haplotype analysis (17) and the contrast mapping method (48).
Phenotypic averages of haplotypes or offspring groups were calculated and contrasted to derive a specific comparison for each marker interval (48). The expectation is that the interval-specific contrast will be maximal in the true interval; therefore, the maximum P value (more significant) observed among the observed contrasts is assumed to contain the QTL. Data were analyzed using the GLM procedure of SAS (39) with the fixed effect of haplotype, sex, and sire. Haplotype effects were analyzed by defining contrasts to test specific allele substitution effects to determine the QTL effect at each specific interval.
Phenotypic characterization of the M16i × MB2 F2 population.
Means, standard deviations, and ranges for all traits averaged across males and females in the M16i × MB2 F2 population are presented in Table 2. These traits include body weight at 6 (WT6), 9 (WT9), 12 (WT12), and 15 wk (WT15); body weight gain between 3 and 6 (GAIN3–6), 6 and 9 (GAIN6–9), 9 and 12 (GAIN9–12), and 12 and 15 wk (GAIN12–15); liver weight (LIV); liver weight expressed as percent kill weight (LIVP); epididymal (males) or perimetrial (females) fat pad weights (PAD); fat pad weight expressed as percent kill weight (PADP); total fat weight (FAT); total fat weight expressed as percent kill weight (FATP); total lean weight (LEAN); total lean weight expressed as percent kill weight (LEANP); BMD; and blood glucose. A very large variation in phenotypes was created in the F2 population, with mice ranging from 34.47 to 76.60 g (mean 56.16, standard deviation 7.45) for WT15, from 2.56 to 20.66 g (mean 9.12, standard deviation 2.18) for FAT, and from 5.88 to 36.51% (mean 17.05, standard deviation 3.98) for FATP.
Phenotypic correlations between WT15, body composition traits and glucose in the F2 population are presented in Table 3. As expected, WT15 is highly correlated with PAD, LIV, FAT, LEAN, and BMD. However, these correlations were much smaller in magnitude when composition traits were expressed as a percentage of body weight. Both LIV and LIVP are positively correlated with LEAN and LEANP and negatively correlated with all fat traits. The high correlation between PAD and FAT (r = 0.832) demonstrates that PAD is a relatively good predictor of FAT in this population. All fat traits had high negative correlations with lean traits. Correlations between weight and composition traits with glucose were low (0.1–0.2) but significantly (P < 0.01) positive.
Selective genotyping of the M16i × MB2 F2 population.
Observed segregation ratios for 16 markers covering the congenic region (61–99 cM) of MMU2 in mice with the highest and lowest phenotypes for PADP in the M16i × MB2 F2 population did not deviate from expected (1:2:1) in the high group, but differences were observed in the low group and between the two groups (High-Low) for markers located between 71 and 99 cM (Table 4). Eight markers mapping between 73 and 99 cM on chromosome 2 showed potential linkage (P < 0.05) with putative QTL regulating adiposity (Table 4). These eight markers were thus genotyped in the entire F2 population, and all subsequent description and discussion of results (including all figures) relate to this reduced region (D2Mit304 to D2Mit52) and not the full congenic region (D2Mit306 to D2Mit52).
QTL analysis.
Positions of markers based on linkage analysis in the F2 population and comparison with those found in the Mouse Genome Database (MGD; http://www.informatics.jax.org/) and with the physical map from the Ensembl Mouse Genome Server (MGSC; http://www.ensembl.org/Mus_musculus/, accessed October 2004, build v.25.33a.1) are presented in Table 1. In general, our linkage map agreed with the linkage map from the MGD; the largest discrepancy is for the interval between D2Mit106 and D2Mit411 (8.4 vs. 2.0 cM). A discrepancy between our linkage map and the physical map from the MGSC was observed for markers D2Mit411 and D2Mit343. On the basis of the MGSC, marker D2Mit411 is located at 160.2 Mb, distal to marker D2Mit285, and marker D2Mit343 is located at 169.9 Mb, distal to marker D2Mit52. Clearly, knowing the true locations of the markers used in this study is critical in resolving QTL positions based on fine mapping. These discrepancies will likely not be adequately clarified until additional sequencing is completed across several mouse strains and a consensus physical map is generated with very high confidence. Because the linkage map generated in this study is based on a very large number of meioses, we consider this order and spacing of markers to be suitable for use in interpreting fine-mapping data within this population.
SIM analysis detected loci influencing WT9, WT12, WT15, GAIN6–9, PAD, PADP, LIV, LIVP, FAT, FATP, LEANP, and BMD (data not shown). The experiment-wise 5% LOD for SIM, based on permutation analysis of all traits, ranged between 2.8 and 3.5. For simplicity, the average of 3.15 was used as the significant threshold for all traits.
The location of, peak LOD score for, and effect of QTL, using CIM, are presented in Table 5. The 5% experiment-wise significant threshold (LOD) determined from 1,000-permutation testing ranged from 3.0 to 4.5 across the traits measured (displayed as horizontal dotted lines in Figs. 2–6). Most of the QTL map to positions that are similar to those determined in earlier studies; thus new symbols representing these QTL are not provided, and reference should be made to the QTL symbols proposed by Rocha et al. (37, 38).
Figure 2 graphically displays CIM results for WT9, WT12, WT15, and GAIN6–9. A single QTL flanked by markers D2Mit194 and D2Mit343, but more closely linked to D2Mit194, appears to explain the observed variation in these four weight traits. The CI for this QTL is between 136.5 and 147.5 Mb (78.1–82.4 cM). The proportion of residual phenotypic variance explained by this body weight QTL ranged from 2.5 to 3.8%. A QTL detected for BMD and another for LEANP mapped to slightly different positions but still close to marker D2Mit194 (Fig. 3). The most likely positions for each of these QTL were essentially within each of their respective CIs (Table 5), which might imply a single pleiotropic QTL. The QTL for BMD and that for LEANP explained 8.5 and 11.7% of the phenotypic variance for each trait, respectively.
For traits representing measures of adiposity, evidence for up to three different QTL was detected (Figs. 4 and 5). A QTL influencing fat traits unadjusted for body weight (PAD and FAT) is located near marker D2Mit194, with a CI between 142.4 and 153.3 Mb (80.4–84.2 cM) that partially overlaps with the CI for body weight QTL. Effects of this QTL on PAD and FAT explain 13.3 and 18.4% of their residual phenotypic variances, respectively. Very strong evidence for a QTL influencing fat traits adjusted for body weight (PADP and FATP) was detected between markers D2Mit285 and D2Mit263, with a CI between 157.3 and 161.3 Mb (88.4–91.6 cM) This QTL explains 8.7 and 10.8% of the phenotypic variation of PADP and FATP, respectively. A third QTL for PADP was detected between D2Mit343 and D2Mit285, with a CI between 154.0 and 156.9 Mb (84.6–86.4 cM). This QTL explains 8.3% of the phenotypic variance in PADP.
Figure 6 shows a QTL influencing LIV and LIVP mapping close to D2Mit106, in an interval between D2Mit304 and D2Mit106. This QTL accounts for ∼4.0% of the residual phenotypic variance for LIV, with a CI between 128.5 and 134.5 Mb (73.0–76.6 cM).
The mode of gene action for all QTL detected in this study was almost entirely additive in nature (Table 5). Estimates of additive effects for each of the detected QTL demonstrate that the C57 allele always decreases body weight, fat, and BMD, while increasing lean tissue, relative to the M16i allele.
Progeny were grouped into distinct paternal multi-marker haplotypes, with haplotypes differing as to the location of single recombination events. Details of each haplotype class, genotype, and markers present in each interval are presented in Table 6.
The P value from an interval-specific contrast was used as the criterion for determining QTL effect. Table 6 shows the defined contrasts and the P values for the parental haplotypes. The maximum P value observed between contrasts is assumed to represent the interval harboring the QTL. Recombinant progeny with haplotype MH2 (interval 75.6–77.6 cM) were significantly different from those with MH4 (interval 81.4–84 cM) for WT12 (Fig. 7). Furthermore, the reciprocal comparison (haplotypes HM4 vs. HM2) was also significantly different for WT9 and WT12. In both groups, mice with homozygous M16 genotypes at D2Mit411 and at D2Mit194 were heavier than mice that had heterozygous genotypes at those markers. The contrast between recombinants from M16i dams showed significant phenotypic effects upon allele substitution in two adjacent marker intervals (D2Mit106–D2Mit411 and D2Mit194–D2Mit343); however, no mice with the intermediate haplotype HM3 or MH3 were obtained in this study. As a result, a precise determination of which of the two intervals is more likely to harbor the QTL is not possible from the current data.
As was seen for body weight, mice with recombinations in interval 2 (75.6–77.6 cM) had significantly different body composition phenotypes vis a vis those with recombinations in interval 4 (81.4–84 cM). Mice with haplotype HM4 (interval 4) possessed more PAD, FAT, and FATP (Fig. 7) than those with HM2 (interval 2). The interval-specific contrast was maximized in terms of P value within the marker interval D2Mit343–D2Mit285 (Table 6). Haplotype MH5 (interval 84.2–86.0 cM) was different from MH6 (interval 86.0–92 cM) in terms of PAD, PADP, FAT, and FATP phenotypes. Progeny with two copies of the M16i allele at marker D2Mit285 had more PAD and FAT than those with one copy. When comparing recombinants from the C57 background, the interval from 81.4 to 84.2 cM also appeared to harbor a QTL. The haplotype with recombinants in the interval 81.4–84 cM (HC4) had lower PAD, PADP, FAT, and FATP than the haplotype with recombinants in the interval 84.2–86.0 cM (HC5) (Fig. 7). No significant contrast differences were detected for LIV or BMD (data not shown).
Interestingly, nearly all contrasts found to be significant (P < 0.01) within recombinant haplotype classes were found in the subset of data generated by M16i dams. The parallel contrasts in the subset of data generated by the C57 dams for the most part approached significance, so it is likely that this observation reflects differences in magnitude or scale of QTL effects in the two genetic backgrounds.
Whole genome scans have proven to be extremely useful for coarse mapping of the chromosomal locations of QTL. However, proceeding from initial QTL mapping to eventual cloning of the underlying responsible gene(s) remains a very difficult and elusive task. Further mapping using congenic lines and subsequently subcongenic lines (or recombinant progeny testing) offers a means for more precise QTL localization, and establishes definitive limits for a QTL-containing region (31). The combination of interval-specific congenic strains and recombinant progeny testing has led to successful reduction of QTL intervals, and positional identification of at least one confirmed QTL with very large effect has resulted from such efforts (3).
Using the M16i model of polygenic obesity, Pomp (33) reported two QTL (Pfat1, Pfat2) in an interval of ∼12 cM on MMU2 affecting both fat and body weight. Rocha et al. (37, 38) also reported QTL for body weight and fat-related traits in a second cross utilizing M16i, providing significantly more details on location, effects, and gene action of the QTL. Many other studies (7, 16, 25, 29, 47, 50), using a variety of different mouse strains, have also reported QTL for body weight and/or fatness within or near this same interval, supporting the significance of distal MMU2 for the genetic regulation of growth and body composition. This region of MMU2 is uniquely conserved relative to HSA20 (55), which harbors QTL for obesity and non-insulin-dependent diabetes mellitus phenotypes based on multiple reports.
In a previous study (21), we described development and characterization of the MB2 congenic line [M16i.B6-(D2Mit306-D2Mit52)], consisting of the M16i line as host harboring ∼38 cM of distal MMU2 from C57BL/6J. The replacement of M16i alleles with C57 alleles in the congenic region on MMU2 resulted in a decreased adiposity of 60% and a decreased body weight of 15%, as well as decreased levels of plasma leptin, insulin, and glucose. In this study, we used a cross between M16i and MB2 to further confirm the presence in this region of M16 alleles with large effects on growth and fatness, and to fine map their locations.
Initially, selective genotyping based on percentage epididymal/perimetrial fat (PADP) in the F2 population allowed the chromosomal interval harboring QTL to be reduced from 38 to 26 cM. Genotypic frequencies deviating from Mendelian expectation, which suggests linkage to a QTL, were observed only within the low selected group. This result strongly suggests a strong relationship of the low fat phenotype with the absence of the M16 allele in the distal MMU2. There may be more genetic heterogeneity within the high fat extreme relative to the low fat extreme, which has been observed previously in a different mouse model of polygenic obesity (40). Using the full F2 population, we next delineated QTL effects to smaller, marker-defined regions, effectively reducing the QTL intervals for body weight, total lean, total fat, and BMD to regions of 1.5–4.5 cM. This also allowed us to definitively determine that the large QTL effects on MMU2 observed in previous crosses using M16i (37, 38), and confirmed in the characterization studies of the MB2 line (21), are actually the result of multiple linked genes as previously suspected (13, 33, 37, 38) for this region. The general finding that an initially strong QTL peak is represented by the presence of more than one underlying gene confirms previous theory (33) and supports similar recent evidence from other regions (45).
The close proximity of a significant QTL [likely representing Epfq1 and Scfq1, from Rocha et al. (37, 38)] associated with fat, lean, and BMD to a QTL affecting body weight [likely representing W10q1, from Rocha et al. (37)] suggests strong pleiotropic effects of a single underlying gene. The strong correlation structure between these traits supports this theory. Further evaluation such as multiple-trait QTL analysis may help test this hypothesis of the presence of a single QTL with pleiotropic effect on body weight and body composition, but definitive proof will require identification of the underlying gene and its function.
Epididymal (or perimetrial) fat is a reliable predictor of total body fat (r = 0.82). The expectation in this study was to find the same QTL for both traits. However, we found one QTL influencing PADP but not (simultaneously) influencing percentage of total body fat. Mehrabian et al. (29) found differential genetic effects on measures of regional body fat. The proximal (Mob7) and central (Mob6) QTL on MMU2 influenced subcutaneous fat, whereas a more distal QTL (Mob5) affected percent total body fat. A similar pattern was found in this study. Our distal QTL mostly affected total body fat, whereas the proximal QTL influenced percentage of epididymal/perimetrial fat.
As suggested above, the fact that the QTL affecting fat-related traits (PAD and FAT) are located at the same marker interval as the QTL for body weight (D2Mit194–D2Mit343) is probably due to variation in body weight. However, the second peak located between D2Mit343 and D2Mit263 [likely representing Epfq2, from Rocha et al. (37, 38)] appears to influence adiposity independently of body weight. Together, these two QTL regions explain ∼30% of the variation in the F2 population, which is in line with expectations based on the heritability of body fat levels.
A QTL was found to affect liver weight within the interval D2Mit304–D2Mit106 (3.2 cM), similar to what was found (Lvrq1; Ref. 37) using the M16i × L6 cross. Moody et al. (30) reported seven QTL affecting liver weight, each accounting for 2–5% of residual variance, but none of these QTL mapped to MMU2. Leamy et al. (24), using a backcross between Cast/Ei and M16i, reported a QTL for liver weight close to marker D2Mit164, which is 2 cM distal to that found in this study. Most likely, the QTL from the present study and from Leamy et al. represent the same underlying gene.
BMD is a predictor of skeletal fragility underlying osteoporosis (36). Masinde et al. (28) reported nine QTL on chromosomes 1, 2, 4, 9, 11, 14, and 15 for BMD in a mouse intercross. One of the two QTL on chromosome 2 reported by Masinde et al. coincides with the QTL reported in this study, also supporting additional previous findings (5, 22).
The recombinant progeny testing element of this study involved identifying recombinants in the congenic interval, propagating these recombinants into progeny, and then performing contrast analysis to determine the existence of phenotypic differences among marker (or haplotype) class means (8, 26). Because the alleles at the QTL segregate, marker allelic means should give rise to phenotypic contrasts that will be large or small depending on where the QTL lies with respect to a marker-interval recombination point. For all traits except BMD, analysis of the segregation of parental recombinant haplotypes confirmed the existence of QTL but did not further increase mapping resolution relative to the large F2 intercross. Considered independently, both methods provided similar results and were equally valuable in this fine-mapping effort, but their benefits were not additive or synergistic. The absence of recombinant progeny in interval 3 impeded making gains in mapping resolution. The distance between markers D2Mit194 and D2Mit343 is 2.8 cM, so that additional recombinants within that interval would help reduce the size of the QTL region. Further genotyping within the RPT population will be useful in this regard and may thus provide benefits beyond those obtained from the F2 mapping. Denser genotyping will also help resolve the actual placement of recombinations in haplotype classes where there are two or more males, which would aid in achieving finer mapping of the QTL.
Several potential candidate genes that may underlie these fine-mapped QTL can be identified based on their map positions and their associations with known biological mechanisms of growth and/or fatness. There are also still many other genes in this region with functions that are as yet unknown. And furthermore, the nature of a QTL may be regulatory and may not represent genetic variation within primary genes of a pathway. Microarray analysis can be an important tool in candidate gene selection, especially if the array included essentially all transcripts residing in the QTL interval. However, utility of expression analysis in this regard assumes existence of detectable differences in steady-state mRNA levels at the temporal and spatial coordinates selected for study. A previous transcriptional profiling revealed differential expression of several genes in liver and adipose tissue between the M16i and MB2 lines (21), several of which map within the CIs of the QTL for liver, body weight, lean, and fat (Table 7). On the basis of this differential expression and on their map position, these genes become viable positional candidates for the QTL fine mapped in the present study, although none of them maps in close proximity to any of the QTL for body weight. In particular, somatostatin receptor 4 (Smstr4, located at 148 Mb based on the MGSC) maps directly on top of the peak for a QTL impacting fat and lean tissue mass. Additional examples of differentially expressed genes mapping in close proximity to QTL detected in this study include the acetyl-CoA synthetase-like (Acas2), MAPK phosphatase (Dusp15), and growth differentiation factor-5 (Gdf-5) loci (Table 7). Strategies for determining that a gene underlies a QTL effect have been recently summarized (1).
While detailed description of every candidate gene is not feasible within the context of this discussion, it is worth noting that the diminutive locus (dm), discovered on the basis of a spontaneous mouse mutant that was shorter than average with low body weight (45), maps to 80 cM on MMU2, which coincides with one of the fine-mapped intervals harboring a body weight and fatness QTL in this study. It is thus possible that this QTL represents a hypomorph of the dm spontaneous mutation, the underlying gene for which has yet to be identified.
By identifying expression QTL that coincided with a QTL peak for, and had correlations with, subcutaneous fat mass, Schadt et al. (40, 41) proposed the NM_025575 and NM_15731 genes as candidates for the distal MMU2 QTL for dietary-induced obesity, previously described (25). These candidate genes are located at ∼165 Mb based on the MGSC, thus falling outside the CIs for any of the QTL localized using the M16i selection line of mice.
The QTL characterized in this study are all harbored in the region of 130–160 Mb, based on the MGSC. The adiposity QTL recently fine mapped (49) using the UW-H3b we Pax1un at/Sn (B10.UW) congenic strain is found in this same general region of MMU2. In addition, the B6.S-D2Mit194-D2Mit311 congenic strain was recently used to isolate a QTL for percent body lipid within the same interval (13). In each of these cases, a C57-derived strain was used as recipient in the congenic line, and loss of C57 alleles led to a decrease in adiposity. This is in direct contrast to our work, where C57 was the donor strain for the congenic line, and where inclusion of C57 alleles dramatically decreased body weight and fatness. When the MB2 line was originally developed, a reciprocal congenic line (M16i as donor and C57 as recipient) was also initiated but did not reach fruition due to reproductive problems. The selection of C57 germplasm as a suitable contrast for M16 was based on two criteria. First, C57 is a widely used inbred line in biomedical (and obesity) research, and second, C57 mice have been shown to be relatively lean when fed a diet that was not high fat in nature (51). The opposite effects of C57 alleles in the present study relative to previous studies (13, 49) may be attributable to our use of the polygenically obese M16i strain as recipient in the MB2 congenic line, or other genetic background interactions.
Extensive sequencing of genes within this region has not been carried out within the M16i line. Such information would be extremely valuable to identify possible polymorphisms between M16i and C57BL/6J. It would also provide increased power to the method of identifying candidate genes based on the pattern of polymorphisms across different strains (32).
In summary, identification of genes underlying quantitative traits is a major challenge. Congenic strains and recombinant progeny testing, among other resources and strategies, are valuable tools for fine mapping of QTL. The polygenic obese M16 selection line is an excellent resource population to initiate positional cloning of novel genes with effects on growth and fatness that have been previously localized to distal MMU2, affecting growth at different ages, fat, lean, and BMD. In this study, the presence of several QTL on distal MMU2 affecting fatness and growth was confirmed and their locations mapped to intervals of 1.5–4.5 cM. Although recombinant progeny testing validated the existence and location of the QTL for most of the traits, little gain in resolution was obtained relative to mapping in the F2 population. However, further addition of markers within the significant marker intervals will likely increase the mapping precision of the QTL using RPT. Several potential candidate genes were identified based on their map position relative to the QTL and their association with growth and fat metabolism.
N. Jerez-Timuare was supported by fellowships from La Universidad del Zulia of Venezuela. This research is a contribution of the University of Nebraska Agricultural Research Division (Lincoln, NE; Journal Series No. 14686) and the North Carolina Agricultural Research Service and was supported in part by funds provided through the Hatch Act. This research was also partially based on work supported by the National Science Foundation under grant no. 0091900 (Nebraska EPSCOR Infrastructure Improvement Grant).
We are appreciative of the efforts of Sara Olberding, Jeryl Hauptman, Mark Allan, and Stephanie Wesolowski in assisting with mouse husbandry and phenotypic data collection. We thank Dr. John Rocha and Dr. James Specht for assistance with QTL analysis. This manuscript was significantly improved on the basis of insightful revisions suggested by two anonymous reviewers.
Present address of N. Jerez-Timuare: La Universidad del Zulia, Facultad de Agronomía, Zulia, Venezuela.
Article published online before print. See web site for date of publication (http://physiolgenomics.physiology.org).
Address for reprint requests and other correspondence: D. Pomp, Dept. of Animal Science, Univ. of Nebraska, Lincoln, NE 68583-0908 (e-mail: dpomp{at}unl.edu).
- Copyright © 2005 the American Physiological Society


text:
KN v Secretary of State for Work and Pensions (ESA): [2016] UKUT 521 (AAC)
- From:
- Ministry of Justice, HM Courts & Tribunals Service, and Upper Tribunal (Administrative Appeals Chamber)
- Published:
- 12 December 2016
- Decision date:
- 16 November 2016
- Categories:
- Employment and support allowance and Tribunal procedure and practice
- Sub-categories:
- Employment and support allowance - Regulation 35, Tribunal procedure and practice - evidence, and Tribunal procedure and practice - statements of reasons
- Judges:
- Gray, P
Upper Tribunal Administrative Appeals Chamber decision by Judge Gray on 16 November 2016.
Read the full decision in CE/1665/2016.
Judicial Summary
Tribunal practice and procedure: statement of reasons and dealing with the evidence ESA: regulation 35 and the presence of a third party If the FTT is considering whether the presence of a third party will reduce any risk in relation to work-related activities or travelling to them there must be findings of fact what their role would need to be, and the reasonableness of them being available. Where relevant consideration will need to be given to whether anxiety in relation to whether a person will be available of itself creates a substantial risk to health. Where the tribunal says that it relies upon the HCP report but comes to a different conclusion as to functional ability than that in the report there is more need to explain the basis of the reliance than where the conclusions of the report are adopted. The lack of a complaint by an appellant about the conduct of the examination or the accuracy of the report to the organisation providing the HCP report should not be used as a significant credibility pointer in relation to whether the appellant's evidence can be accepted as to any limitations of the report.
Document information
Published: 12 December 2016
Decision date: 16 November 2016
From: Ministry of Justice HM Courts & Tribunals Service Upper Tribunal (Administrative Appeals Chamber)
Categories: Employment and support allowance Tribunal procedure and practice
Sub-categories: Employment and support allowance - Regulation 35 Tribunal procedure and practice - evidence Tribunal procedure and practice - statements of reasons
Judges: Gray, P

subset: commoncrawl


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A Masters degree is an academic degree awarded to individuals who successfully denote a higher level of expertise. There are two main types of Masters - taught and research.
Finance is the study of money management, and includes everything from personal savings management to accounting to investment banking. A finance student may choose to specialize in a subfield like retirement planning or business investments, or might keep their education general with a survey of all areas of finance.
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OCEANWEST will be built in multiple phases, as many as 12 or 15, over the next 10 years. Projected completion is 2017, depending on the real estate market. The following areas are currently offered for sale.
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text:
So I had the opportunity to see some of the techniques that were in the video about Whole Brain Learning. See the post.
And to be completely fair I didn't hate them in action.
Here's some of the techniques that I watched in person that worked well with middle schoolers.
+Ritualized Call and Response to gain attention
"Claaaaassssss….. Yeeees."
It worked because teacher had complete control and attention from students without raising her voice; and the kids returned to focusing very quickly.
+Students teaching each other
In this case however, the students were paired up to explaining the parts of a cell. In each pair one person explained (in their own words) about the cell and the other student was listening to their explanation.
It worked because the students had to give explanations out loud multiple times and in their own words. They weren't talking over their partners. And it gave an opportunity for the teacher to check in with all students to figure out who was struggling without drawing attention to them.


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text:
He said the company will aim to create several other forums "where people can feel comfortable to speak freely". In his manifesto, Damore explained why there are not more women in tech jobs and called for more ideological diversity in Google's workplace.
After the Galaxy Note 7 mishap past year that resulted in its official discontinuation, it is the Galaxy Note 8 that needs to be an attractive package to improve Samsung's image in the market. Reportedly another more powerful "special edition" variant too might launch in the future. Even Blass, a trusted industry tipster who often publishes leaks that end all leaks for good, has just uploaded a render of the Galaxy Note 8 .
Thanks to the innumerable leaks and rumors , we already have a fair idea about what the Samsung Galaxy Note 8 will bring to the table. Thanks to this additional layer of security around it, the Samsung Galaxy S8 Active is about 1.5 times thicker than the regular Samsung Galaxy S8 .
F-150 Police Responder joins the broadest lineup of law enforcement vehicles in the industry. The new F-150 will have upgraded brake calipers and pads, upgraded front stabilizer bar, steel wheels, skid plates and plenty of other goodies that can happily take a constant beating.
Trump was Macron's guest of honour at national day celebrations in the capital as this year also marked the centenary of the United States entering World War OneI. Over their two day meeting, Mr Trump and Mr Macron vowed to continue cooperative over security and counter-terrorism forces. Trump came to France beset by allegations of Russian interference in the 2016 US election, with emails released on Tuesday suggesting his eldest son welcomed an offer of Russian help against his ...
It seems as though cheaters love a Steam Sale as much as the rest of us. This looks like it could be the case, since nearly every day after the mass ban, the numbers of people removed from the service has been less than 1,000 a day - normally, it's around 3,000 a day.
Low-key meetings between the prime minister and members of his legal team arrived at the decision to challenge the JIT report - in the top court as well as in public - sources in the ruling camp said. The report alleged that there was "irregular movement of huge amounts in shape of loans and gifts" to Sharif's family and Pakistan Based companies from the companies based in Saudi Arabia , UAE and the UK.
While Instagram aped the " Stories " feature straight off Snapchat , the feature has grown in popularity, with many Insta users posting Stories regularly. A sticker of the original story will only be visible to the user so that they know to which story the reply was made. Users can tap on the button to reply with a photo or a video.
The average salary for men in the White House is around $104,000, while for women it's around $83,000, according to CNN's look at the White House's annual report to Congress. Bush's administration, showing Trump performing far worse than the 43 President. At $187,100, senior policy adviser Mark House is the top earning White House staffer.
After the firing, a large contingent of police, CRPF and army personnel had arrived in the area - a highly fortified zone - a few hundred metres away from the Indian Army's 15 Corps headquarters. "Restrictions under Sec 144 imposed from Ram Munshi Bagh to Sempora (around Pantha Chowk attack site) with immediate effect", Srinagar's Deputy Commissioner Farooq Ahmad Lone told ANI.
YET another new smartphone brand, Nubia , has entered the already competitive Malaysian market. The smartphone features a metal body with a layer of curved glass on top. Further Sony Cmos sensor brings an f/2.2 aperture, and supports a number of features including Beauty mode, HDR 4.0 and anti-shake. In particular, 3 GB RAM , HD screen with a diagonal of 5.5 inches and a matrix IPS, which is installed on top of the protective glass, and the slot under microSD memory cards up to 128 ...
More details in the full report linked below. In the recent months, Apple has been in conversations with health IT industry groups that can help them achieve their goals. "At any given time, only about 10 to 15 per cent of patients care about this stuff", said Micky Tripathi, president and CEO of the MA eHealth Collaborative and a health IT expert.
Once you select the video or photo a simple swipe will showcase the filters for you to choose from. The update also introduces the ability to add filters directly to the GIFs, photos, and videos shared within the WhatsApp camera. Now, when you receive multiple photos from the same sender in a row, they will be grouped together as an album . For now all these features are limited to the iOS platform, but are soon expected to be made available on Android OS as well; however, not ...
Digital preorders are now available for Sonic Mania on Steam and Xbox One , and will be for PlayStation as well later today. The retro-inspired platformer will launch for Switch and other platforms on August 15, for US$19.99/C$25.99.
The Sun recounts the great lengths-literally-Harry went to to bring Markle to the reception: it was a 100-mile round-trip drive for him to pick her up. Shortly after the photo was taken he was seen holding up three fingers, so it seems he was actually counting. Well, if things seem to be going great for Harry , apparently we can not say the same thing about William .
BTG plc is engaged in the business of healthcare, focusing on Interventional Medicine therapies for liver cancer, emphysema and vascular disorders, specialty pharmaceuticals for acute care uses, and a licensing business. The stock of BTG plc (LON:BTG) earned "Neutral" rating by JP Morgan on Tuesday, January 17. Finally, Canaccord Genuity reiterated a "hold" rating and issued a GBX 160 ($2.07) price objective on shares of ITE Group plc in a report on Friday, January 27th.
This year alone, Netflix plans to spend $6 billion to produce more than 1,000 hours of original content. The Australian reported Netflix has been applying price increases to its plans for some Australian customers, but only on weekends.
If we take the AirPods rumour, for example, Apple now has a supply issue meaning AirPods continue to face a six-week shipping estimate on Apple's website; if the company bundled these it could cause further delays of iPhone 8 shipments.
While still far from mass adoption, virtual assistants are becoming more widely used by Americans, according to eMarketer's first forecast on users of digital assistants, such as Siri, and voice-enabled speakers, like Amazon Echo . "I think voice assistants are incredibly powerful, their intelligence is going to grow, they're gonna do more for us, but the role of the screen is gonna remain very important to all of this", he said.
The nostalgia is strong with this one. In our own little celebration of Star Wars Day , we've searched through Twitter for some of the most enjoyable, adorable, and down right amusing tweets for you. And lastly, Star Wars: The Last Jedi will mark the last time the late, and great, Carrie Fisher will be seen on the big screen after her sad and unexpected passing a year ago.
It also will run the new Windows 10S operating system, software the company presented Tuesday that will be faster and offer better security than previous versions. Unlike other Surface models, it does not run the traditional version of Windows. Every license of Minecraft for Education will be offered for free with the Windows 10 education PCs and will be primarily targeted towards schools.
Apple released its quarterly reports on Tuesday, affirming most analysts' projections on the financial status of the Cupertino-based tech giant at this point in the year. Apple's contribution amounts to a tiny fraction of its $257 billion in cash. In the second quarter, Apple reported results that were slightly under what Wall Street expected.
As reports suggested, we're expecting to see Microsoft's own answer to Chromebooks and even Google's Chromebook Pixel. The device offers 14.5 hours of battery life and it is available in four colors: platinum, burgundy, cobalt blue, and graphite gold.
Canaccord Genuity reissued a "buy" rating and issued a $43.00 target price on shares of Intel in a report on Thursday, January 19th. Wells Fargo & Co reissued a "top pick" rating and set a $45.00 price target on shares of Intel in a research note on Tuesday, January 3rd.
Now, however, Google is reportedly set to solve this issue with a new upgrade to its Chrome mobile app which it says can make copying and pasting easier than ever. Aside from these new features, the developers have also added a new gesture. Apple introduced a similar feature in iOS 10 that used Siri's smarts to automatically generate contextual suggestions like calendar events, contacts, and locations in the default keyboard.


text:
Israel News
Middle East
American Politics
Edition française
Green Israel
price tag attack on Mughayyir mosque DO NOT USE.
(photo credit:ISSAM RIMAWI /Flash 90)
Three IDF soldiers were arrested on Tuesday for alleged involvement in "price tag" attacks which occurred in the West Bank recently, the IDF Spokesman's office said in a statement.The soldiers were transferred to the custody of military police for interrogation.
By Gil Stern Stern HOFFMAN
By Gil Stern Stern HOFFMAN
Sponsored Content
By YONAH JEREMY BOB
By HERB KEINON
Please insert a valid email address


text:
District Attorney Lynne Abraham told City Council today that the local gun control legislation signed into law last week by Mayor Nutter is unconstitutional.
Asked during a budget hearing if she would prosecute people arrested under the laws, Abraham said, "nope!"
Abraham said after the hearing that state law does not permit Philadelphia to enact local gun laws.
"Until or unless the Supreme Court tells me otherwise, I'm obligated to follow the law the Supreme Court has enunciated," Abraham said.
The state Supreme Court in a 1996 ruling declared that only the General Assembly has the power to regulate guns, tossing out the city's last effort at gun control.
But Nutter last week signed a package of five gun bills passed unanimously by City Council. The laws limit handgun purchases to one a month, make it a crime not to report lost or stolen guns in 48 hours, allow police to confiscate guns from people considered a danger to themselves or others and ban semiautomatic weapons with clips that hold more than 10 rounds.
Nutter said he planned to start enforcing the laws immediately, except for the one-gun-a-month bill, which will take six months.
Abraham was skeptical about the enforcement plans.
"If there are wholesale arrests which turn out to be illegal, this city is going to get the pants sued off them," Abraham said.
Nutter wants to sit down with Abraham to discuss the laws, which he says are legitimate. She says she's happy to talk.


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text:
|{all images courtesy of the lovely Jen Lauren Grant of Birds of a Feather}
Gosh, it was fun. A little DIY and a lot of Pinterest inspiration really came together with the help of our amazing vendors, for which I am forever grateful. Wish we could do it all again!
photography: Jen Lauren Grant // florals: Milieu Florals {by Hannah Koo} // venue: Spanish Hills Country Club // cake: La Starr & Co // hair and makeup: Susie Chhuor // DJ: DJ Jeff of VOX DJs // cermony harpist: Vonette Yanaginuma // videography: Skipping Stones Productions


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Apple may be planning 10.5-inch iPad Pro in 2017
New middle option between 9.7-inch and 12.9-inch models
Over the weekend, highly reputable Apple analyst Ming-Chi Kuo released a new research report claiming that the company may be planning to release a "middle-sized" 10.5-inch iPad Pro sometime in 2017 to accompany the existing 9.7-inch and 12.9-inch models.
Intel cheers the USB Type-C
Backing Apple
While users and the rest of the world think Apple's move to kill off the headphone jack is a bad idea, Intel is cheering a general move to a more mobile USB Type-C.
Encryption fault in Apple's imessage
Flaws in the method
Cryptography researchers at John Hopkins University have found another flaw in the encryption used by Apple's iMessage and while Jobs' Mob might have fixed it, the researchers say that the company's encryption methodology is borked.
Apple jumps on the AR bandwagon
Claims it is working on a new AI platform
Fruity cargo cult Apple is trying to convince the world it is "coming up with something new" by talking a lot about Artificial Reality.
Apple's tenth-gen flagship iPhone to feature Smart Connector
Simultaneous release of standard and "S" models possible
A week ago, we reported that alleged prototypes of Apple's upcoming tenth-generation smartphones were released in a YouTube video comparing dimensions, features and aesthetics between three new devices.
Microsoft plans three new Surfaces
Expect more expensive Apple clones to follow
Software king of the world Microsoft is expected to release three new flavours of Surface three in one desktops later this year.
Apple shamed into upgrading its Mac Pro range
Can't take the jibes from Fudzilla
For a while now we have been mocking the fruity cargo cult Apple for being unable to upgrade its Macbook Pro. Now it seems that Jobs Mob has suddenly woken up to the fact that you can't claim you are cutting edge while peddling four-year-old stock.
New Apple Watch will be buried in iPhone 7 release
Nothing to see here, move on please
A Chinese analyst claims that Apple will release the "new and improved" iWatch when all the coverage is going to be focused on the iPhone 7.
Apple's TV programming hits new low
Front people named as secret minutes revealed
The fruity cargo cult, Apple's mission to lower the standards of the world hit a new "high" today after it announced which celebs were going to host their new self-reverential app developer "reality" show. Gwyneth Paltrow, will.i.am have been named as the celebrities which will add weight to "Planet of the Apps." It seems such a strange decision that we asked one of our Apple deep throats to find us the minutes of the meeting where it was made. We provide it verbatim:
London's coppers stuck in Windows XP time warp
27,000 PCs still run the ancient software
More than 27,000 PCs in use by London's Metropolitan Police are running Windows XP despite an IT modernisation programme and fears that it makes them vulnerable to hackers.


text:
Lord of Temptation(Lost Lords of Pembrook,Book 2)(85) by Lorraine Heath
He wept.
For the boy he'd been.
For the man he was.
For the lord he wished he might have become.
And he screamed out because in the end, his uncle had won. He'd destroyed Lord Tristan Easton. And Captain Crimson Jack didn't know how to find him again.
Chapter 26
In two hours Anne would be married, yet as she stood in front of the cheval glass in her gown of satin, lace, and tiny beaded pearls, she felt no measure of excitement. She liked Chetwyn. She surely did. Marriage to him would be proper. She would be proper. She adjusted the veil that fell from a wreath of orange blossoms and wished she'd chosen some other sort of blossom because oranges always reminded her of Tristan. And she didn't want to think of him today. She didn't want to think of him ever again.
She was wrapping about her finger the strip of leather that he had once used to bind his hair, to bind hers. She needed to toss it away, but she knew, instead, she would return it to her jewelry box before she left for the church.
"Don't you look lovely, my lady," Martha said. "Lord Chetwyn is such a fortunate man."
"It is I who am fortunate." The words were the proper thing to say, so why were her eyes burning? "I think you'll be happy in Chetwyn's household."
Anne turned to find her maid's brow furrowed so deeply that she was surprised the woman didn't yelp in pain. "Ew?"
Martha released a deep sigh. "I was going to tell you after the wedding—"
"Tell me what?"
She smiled brightly. "Mr. Peterson has asked me to marry him. I've said yes."
Anne took Martha's hands. "Oh, that's wonderful. Congratulations. Although I don't understand why that should make you dread Chetwyn's household."
"Oh, I don't dread his house, but telling you, m'lady, that I won't be going. I'm giving my notice."
Releasing her hold on Martha, Anne scoffed. "That's a silly thing to do. It'll be years before he returns—"
"No, he came back last night. Said he missed me too much and had the captain turn the ship about."
Anne's heart slammed against her ribs. "They're in port?"
Martha nodded. "Yes, miss."
Anne's gaze shot to the window. What was she expecting for God's sake? To see Tristan clambering into her bedchamber?
"But they're setting sail again this afternoon," Martha continued. "Just not with Mr. Peterson. He's given up the sea. He's going to work in a shipping office or some such. He's saved his money so we can purchase a home. I don't have to work any more."
"Oh, Martha, I'm so happy for you."
"I'm happy for myself." Her smile grew. "I never thought to find love. He's a good man."
"I've no doubt of that."
A brisk knock sounded on her door. Martha hurried over to open it. Stiff and clearly unhappy, Jameson stood beside Chetwyn. "Leave us, Martha," her brother ordered.
Martha gave Anne a quick look before scampering into the hallway.
"Chetwyn wishes to speak with you before the nuptials. Highly unusual, but I've granted him permission. However, the door is to remain op—"
Chetwyn stepped into the bedchamber and slammed the door shut in her brother's face. Anne pressed a hand to her mouth to stifle her laughter. She could only imagine Jameson's startled expression. She'd never seen Chetwyn so forceful. It was a bit disconcerting to realize that it excited her to see him this way.
He strode to the fireplace, raised his arm, pressed it against the mantel, and stared into the cold empty hearth. "Do wish I'd stopped by your father's study for a bit of spirits."
"I have some brandy."
Looking over his shoulder at her, he grinned. "Do you?"
"Yes, would you like some?"
He shook his head. "No, I suppose not. You should know, Anne, that I will treat you kindly."
"I never doubted that."
"You will never want for anything. I am convinced and believe with all my heart that I can provide you with a satisfactory life. But I daresay that I believe you deserve more."
"I don't understand."
"I think Lord Tristan is a rotten bastard," he continued. "But be that as it may, I've seen the way he looks at you and more, I've seen the way you look at him."
"How is that, my lord?" she dared to ask.
"As though you are the only two people who exist in the world." He faced her completely. "Do you love me, Anne?"
She dreaded answering him. She didn't want to hurt him but she couldn't begin today with a lie.
"I don't love you either," he said as though she had responded. "I asked you to marry me because of Walter's letter. I've come to the unfortunate conclusion at a rather inconvenient time that it's not enough upon which to base a marriage."
"Walter's letter?"
He reached into a pocket inside his jacket and removed a yellowed crumpled piece of paper. "He was ill when he wrote it. I suspect he knew he would die. He asked me to see that you were happy, and I thought that I could ensure that best if you were my wife. I thought I owed him that at least. I pushed him into joining a regiment, into making his own way. Our coffers are thin, you see, and I didn't want to give him an allowance. Then we declared war on Russia and I told him to sell his commission. Marriage to you would bring him a dowry; he could make do with that. But he didn't want to be seen as a coward. It's my fault he's dead."
Copyright © 2015 by Read Best Books Free Online


text:
- Description
Quick Overview
Perfect for securing your hair or accenting a beautiful updo, these pretty pearl hair pins will add that last, elegant touch to your look.
Delicate faux pearls adorn a darling set of bobby pins perfect for securing your hair in place or accenting an elegant updo.
Having launched in 2010, Toniq accessory is a brand synonymous with young and fashion forward accessories. Over the past 5 years, Toniq has grown exponentially and is currently available at over 70 stores across the country. Each collection consists of over 300 styles.
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|iPhone apps
Mark of Mafia: Master Edition
Updated VERSION 7.01
====The BEST online Massive Multi Player Crime Game for the iPhone and Touch keeps getting better!====
Start as a rookie and progress to be a MADE MAN by outsmarting, out-buying, and out-GUNNING other real players in REAL TIME and AROUND THE WORLD!
Immerse yourself in the criminal underbelly of The City as you progress to become the Don of your Mafia family -
Download MARK OF MAFIA now - a life of crime and power can be YOURS!
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Rated 4/5 based on 3 customer reviews.
5/5 stars
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5/5 stars
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CV Reference Number: 430
|CV Headline
|Santos Martin Moreta United Kingdom 29/10/2015 Dear Sir/Madam I am writing to you to enquire whether there is an opportunity for me to join your team as a Technical Agricultural Engineer, I would appreciate very much if you would take a few minutes of your time to consider my application. I am currently working as a Demi-Chef, where my main responsibilities include the smooth running and operations of the kitchen. This means, that I have developed an organisational skillset that can be transferred to different business sectors. I have worked in the service industry ever since I came to the UK and I have learnt the importance of delivering first class service to customers. This means I have learnt to deal with people at all levels and have been able to gain a set of skills where an eye for detail was of utmost importance to deliver the high standards of service. You will also be able to see from my attached CV, that I have a degree in Technical Agricultural Engineering, which I completed at the University of Salamanca in Spain, with a focus on mixed farming. In addition, I completed a course in AutoCad to an advanced level, which means I have the skills to design agricultural machinery components and equipment. I believe that a creative and innovative approach to the industry is essential nowadays in order to achieve success. Ideally, I would like to work in an environment that enables me to put into practice my existing skills as well as to develop new ones. I would thus be grateful if you would consider my application and enclose my CV for your kind consideration. I would be pleased to discuss my application with you in more detail in an interview. I look forward to hearing from you. Yours faithfully, Santos Martin Moreta
|Relevant Qualifications
|Other Qualifications
|Other Professional Qualifications
|Farming Experience
|Agricultural Engineering, Beef/Stock, Mixed, Pig
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|Type of Work Wanted
|Agricultural Engineer, Arable, Beef, Diary, Horticulture, Mixed, Organic, Sheep
|Driving Licence
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Argentinian coach hails Messi as world's best
Normally a reserved man who avoids hype, Argentina coach Alejandro Sabella isn't shy about declaring Lionel Messi to be the best player in the world.
Comparing the Argentina captain with Arjen Robben of the Netherlands and Brazil's Neymar, Sabella today held up Messi as the undisputed No 1.
''Robben is a great player. He is very important for Holland. Like Neymar is for Brazil and Messi for Argentina,'' Sabella said. ''Each of them is important for their teams but Messi is the best of them all.''
After a disappointing World Cup four years ago, Messi has carried Argentina into the semifinals in Brazil, scoring four goals - the same tally Neymar reached before injury ruled him out of the tournament. Robben, who faces Argentina in the second semifinal tomorrow, has scored three.
Though he's been named world player of the year four times for his accomplishments with Barcelona, Messi hasn't had the same kind of success with Argentina, which hasn't won a major title since the 1993 Copa America.
This is the first time since 1990 that Argentina has reached a World Cup semifinal, which Sabella said was a victory in itself.
''After 24 years Argentina is among the four best teams in the world,'' Sabella said. ''I think this group of players has already won. Let's see if we can win even more.''
If they win the semifinal, Argentina won't have to worry about rivals Brazil in the final. Sabella was just as stunned as everyone else to see Brazil lose 7-1 to Germany in the first semifinal, a game Argentina's players watched the end of on TV at Itaquerao Stadium, even delaying their training session for around 20 minutes.
Sabella said the heavy defeat for Brazil was clearly ''not normal.''
''But in football you have to expect these things,'' Sabella said. ''It's football. It's the most illogical of sports. Things happen that you cannot foresee.''


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Just a quick public post to say a HUGE thank you for recommending my illustrations yesterday…thank you Ali Crooke, Mel HP and Anneliese B for your kindness, and believing in my creativity – working on my own, I can sometimes sink into my own thoughts and often wonder if what I'm doing is the right thing. Clearly I am – you're my version of an appraisal, which is the only thing I miss from good ol corporate days!
Lizzie x


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Discussion in 'Items For Sale or Trade' started by er4smiths, Sep 7, 2012.
If anyone is interested. $250 - price includes shipping.
Check EBAY again - there are 2 auctions.
I've got a loose never Transformed Takara version.
p.s. I apologize publicly for seeming like I'm trying to scalp. I hate scalpers.
I just checked ebay and thought that was the going rate.
I saw Takara versions for $400 and the ToysRUs auctions were at $150 with days left in the auction and shipping was like $50
Will ya take 125 and or a trade ?
Thread cleaned up. Good luck on your sales.
EDIT: Removed, was able to get one within my budget - good luck!
Separate names with a comma.


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San Francisco City College health benefits
Think about what career is best suited for you, and where you want to study. Request more information now.
Think about what career is best suited for you, and where you want to study. Request more information now.
Think about what career is best suited for you, and where you want to study. Request more information now.
Information about San Francisco City College health benefits, and registering for elective seminars. Which college degrees give you the best chances of finding employment after graduation? Qualifications for nursing programs vary widely. Learn about the various medical specializations available within technical training programs. Concentrating on San Francisco City College health benefits, for example, you may gain an advantage in corporate job recruiting on-campus. Check with the health department for course registration deadlines, and healthcare program requirements. Please use the form above to request admissions info for San Francisco City College.
Preventive care helps doctors catch health problems early, with a better chance of successful treatment. Blood pressure, diabetes, and cholesterol tests, cancer screenings, vaccines, and child screening may be covered without any out-of-pocket cost. It is your responsibility to understand the risks and benefits of your treatment, so educate yourself before undergoing any serious treatment plan, such as a surgical operation. Also, protect your health information by obtaining your own copy of your medical records.
Read your health plan's summary of benefits to learn what is covered and what is excluded. Health insurance only pays for care that is medically necessary, which is defined in your health insurance policy. Your insurance plan may not pay for experimental procedures or advanced medical testing, but if you have a serious illness, you can ask for an independent medical review. You do not need pre-approval for emergency care, such as admittance to the emergency room, or ambulance service from the scene of an accident. Beyond major requirements within the health department, you may consider Valley City State University surgical assistant program as a complementary elective to your primary course of study.
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no subjectDate: Friday, May 12th, 2017 11:37 am (UTC)
From: miss_s_b
Recording. We went to the tour a couple of years ago in Manchester. Sandi was presenting. We saved all the kazoos and made a kazooman centipede when we got home :)
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I've never really been the one to talk about marriages and settling down seriously. I just feel like I'm too young to even be thinking about it. Be that as it may, it'd be a lie for me to say that I'd never pictured myself being someone's wife. Cooking meals for my future husband is my marriage wet dream. The thought of going grocery shopping and making the food in the kitchen, with my husband anticipating the meal just excites me. Imagine the yes-this-is-what-I'm-talking-about smile curving on his face after the first bite. What a total bliss that would be. Anyway, this is definitely one of the many things I'd look forward to if I were to settle down. Apart from the passionate lovemaking, of course.
Category Archives: Uncategorized
Sorry. I have been feeling such way to everyone the whole 22 years of my existence. Sparing people's feelings is one of my favorite pastimes. Taking other people's feelings into account when saying something has become a habit of mine. For once I just want to be spared. I want to be able to show my true feelings. I want to be allowed to be upset, mad, angry, and the list goes on. I want people to take my feelings into account. I want myself as well as my feelings to be considered. I want to be given the benefit of the doubt. I want to be understood. I want to be told "I know it hurts". Is it too much to ask for? Gosh, I am only after all, human just like the rest of you.
Life is a Constant Dysmenorrhea
It has been ages since I last poured my soul out in words. A lot has happened over the past months. I found myself completely trapped in a Hotel California like slump for months which I received tremendous loathes and mockeries instead of support and encouragement from those who I consider or considered to be my people. I was perpetually depressed and hating on myself for feeling so useless. I sought after shoulders to cry on but as usual, everyone's life was far more interesting than mine for anyone to take a minute and listen to my never ending crises. No matter how hard I've tried, it will never be enough. My decisions in life will always be according to my family's convenience. My problems will always sound petty because everything is a bloody competition to everyone. Living is indeed, a formidable thing to do. Particularly when you're not a heartless opportunist who constantly takes pleasure in utilizing other people's abilities to satisfy your needs. Which is why I have decided to be more selective in doing favours for anyone and not end up like my generous father who perpetually finds himself mercilessly used by those who know him, from now on. I realized that I should've stuck to writing in journals rather than venting out to people who could give two shits about me and my so called petty problems. At the end of the day, it's every man for himself. Innit?
Cursed is what I am.
If only you'd know how pathological you are.
For years I have been imprisoned by the thought of you.
And for years you have failed to acknowledge that.
Being free is what I desire.
Free from the curse of you.
An Overdue Apology
I wish those who met and got to know me before could've met and got to know me now instead. I don't really appreciate the younger version of myself. I was naive and rather a tad bit ignorant if you ask me. Thus, I apologize on behalf of my kid self. Nevertheless, everyone should be given a room to make a fool out of themselves. How are we to learn if we never flawed? I guess my frontal lobe was still developing, hence my semi underdeveloped personality. Now that I am older, I try to keep an open mind and not be prejudicial to those whom I am engaged in a conversation with. Be that as it may, I refuse to give in to the stereotypical bunch who keep putting their ginormous schnozzles into other people's business and never fail to find ways to make themselves feel like they're above everyone. Yeah, these people have got to go. Period.


text:
10 Reasons for a Cultural Boycott of Israel wants to
Tell musicians: Don't entertain apartheid Israel!
We have 10 Reasons to Boycott Israel, read more
We have 10 Reasons to Boycott Israel, read more
Please express support and sympathy for the families who lost loved ones in the Sandy Hook shooting, and those injured.
This Facebook page is an absolute disgrace. This page is doing nothing but causing those who have/had loved ones in the
WTF the government think they can just keep kicking people while they are down. It has gotten worse and I didn't think


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City oF Shiraz » Attractions » Historical Sites » The Ancient Site of Pasargadae
Shiraz Historical Sites
The Ancient Site of Pasargadae
مجموعه باستانی پاسارگاد
The World Heritage Site of Pasargadae is a collection of ancient structures remained from the Achaemenid era located in the city of Pasargadae in Fars province.
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Please only ask questions about Iranian culture and Iran travel tips. Please ask in English and be brief. We reserve the right to not publish inappropriate questions.
Your question will be translated to Persian and posted in our Iranian forum . Once an answer is received from a real Iranian, it will be translated to English and posted here.
Photos by UntoldIran's members


text: Sorting type: Most visited Most visited Alphabetical | The different Ethnic Groups on the Wiki Subcategories Arabic-Yemeni Ethnicity Caucasian-American Ethnicity Chinese-Japanese Ethnicity Egyptian-American Ethnicity Hispanic-American Ethnicity Indian Ethnicity Retrieved from "http://nikaraworld.wikia.com/wiki/Category:Ethnicitys?oldid=2961" Ad blocker interference detected! Wikia is a free-to-use site that makes money from advertising. We have a modified experience for viewers using ad blockers Wikia is not accessible if you've made further modifications. Remove the custom ad blocker rule(s) and the page will load as expected. Categories: List Pages Protected Pages


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If there's one rule to follow when it comes to eating healthy, it's "the fresher, the better." It's no secret that we want to eat healthier, while still enjoying the food we eat. Unfortunately, there are hundreds, if not thousands of marketing ploys used to try to get us to buy food that seems healthy, but [...]
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Apple's ARKitadmin
Apple announced a tool called ARKit, which has been designed to provide advanced Augmented Reality (AR) features and capabilities on iOS. Here are 11 facts to help everyone understand why this release is so important and exciting for customers. https://appreal-vr.com/blog/11-conclusions-from-wwdc-2017-ios-arkit/
How Many Attorneys are in the USAadmin
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$15 million in funding in Victorian budget for Emergency Services
Victoria's state budget will include a major shakeup of how the State Emergency Service is funded.
Currently Councils and the Victorian Government jointly fund the SES, but since the government introduced rate-capping many councils have raised issues with the funding arrangement.
Today's state budget will include $15 million to support the SES.
'Sea of surpluses' in Victorian budget. Congratulations Premier, well done. #springst #auspol https://t.co/OF1QrEYeBB— the man from (@UNCLE80075178) May 1, 2017
The money will cover the operational costs of SES units and assist with long term leases for stations located on council land.
Meanwhile, Health and Education are expected to be among the big winners when the budget is delivered after midday.
$174 million will go towards an elective surgery blitz, helping 200,000 Victorians get surgery sooner.
Victorian budget tonight up stamp duty -up land tax why would you invest or live here.ps I still feel very unsafe— Alan Schwarz (@moneysport1) May 1, 2017


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Home » Worksheet » Browse 30 Images »
Rocks and minerals geology products on pinterest free worksheets transparencies reproducible worksheets. 1000 images about rocks and minerals on pinterest natural resources activities student. Activities the ojays and rock on pinterest free rocks minerals worksheets third grade science greatschools. Rocks and minerals worksheets rock on pinterest. Worksheets rocks and minerals delwfg com learning about layers of minerals.