Experiment
Organisms on islands often undergo rapid morphological evolution, providing a platform for understanding mechanisms of phenotypic change. The authors used house mice living on the remote Gough Island - the largest wild house mice on record - to understand the genetics of rapid skeletal evolution in nature.
Compared to a mainland reference strain from the same subspecies (WSB/EiJ), the skeleton of Gough Island mice (GI/Wisc) is considerably larger, with notable expansions of the pelvis and limbs. The GI/Wisc skeleton also displays changes in shape, including elongations of the skull and the proximal vs. distal elements in the limbs.
Several partially inbred lines of GI/Wisc mice were created through full-sib mating for four filial generations, a procedure expected to reduce within-line heterozygosity by 60% (Silver 1995). To incorporate variation segregating among GI/Wisc mice, two partially inbred GI/Wisc lines were used for intercross experiments (denoted as crosses A and B). One pair of male and female siblings from each partially inbred line was crossed with WSB/EiJ to generate four independent F2 intercrosses. A total of 1374 F2 mice were generated: 497 from cross A (WSB/EiJ x GI/Wisc = 279 and GI/Wisc x WSB/EiJ = 218) and 877 from cross B (WSB/EiJ x GI/Wisc = 494 and GI/Wisc x WSB/EiJ = 383). From this F2 population, 827 mice were used for all skeletal phenotyping and analyses: 367 from cross A (WSB/EiJ x GI/Wisc = 206 and GI/Wisc x WSB/EiJ = 161) and 460 from cross B (WSB/EiJ x GI/Wisc = 252 and GI/Wisc x WSB/EiJ = 208).
All mice were weighed to the nearest milligram every week, beginning 1 week after birth and ending at 16 weeks. Dual energy X-ray absorptiometry (DXA) was used to measure bone morphology. Digital X-ray images (Carestream Health DXS Pro 4000) were collected for 43 F1 and 827 F2 individuals and mice from the four parental strains of the cross. X-ray images were taken at three postnatal time points (5, 10, 16 weeks of age) for each animal. A dorsal and lateral X-ray image was taken of each individual. Skeletal dimensions were measured from the X-ray images using Carestream Molecular Imaging Software (Carestream Health). Measurements were chosen to capture axes of known variation across laboratory mouse strains and across species. A total of 16 measurements were used for phenotyping, including lengths and diameters of long bones, pelvis, and skull.
All mice were genotyped using the Mega Mouse Universal Genotyping Array (MegaMUGA, Geneseek, Lincoln, NE). The MegaMUGA is an Illumina array platform containing ~77,800 markers. Most of these markers are SNPs, although some are structural variants and transgenic markers. The markers are densely and relatively evenly spaced at 33 kb across the genome, found across all autosomes, sex chromosomes, and the mitochondria. Following initial screening, the cleaned genomic data included the four GI/Wisc parents of the crosses, 70 F1 individuals, 1346 F2 individuals, and 33,191 markers. In all subsequent analyses, the authors focused on
a subset of 11,833 markers that were fixed in the four GI/Wisc parents and therefore segregated as in a standard F2 intercross
between inbred lines.
Single-trait QTL analysis was performed using HaleyKnott regression on a 0.5-cM grid across the genome, as implemented in R/qtl. Analysis was conducted separately for each of the 16 skeletal traits at 5, 10, and 16 weeks with sex, mother, and observer as additive covariates.
Additional single-trait scans were performed on F2s using two different methods to control for effects of body size on each skeletal trait: (1) by using relative skeletal sizes as the traits and (2) by treating body weight as an additive covariate in the model. Relative skeletal size was calculated by dividing each skeletal trait by the cube root of body weight for each individual (trait/body weight1/3) and is referred to as the shape ratio.
GI/Wisc mice have larger skeletons than WSB/EiJ mice across postnatal development, with all but one trait (height of the cranial vault (HCV) at 16 weeks) showing significant expansions at all three time points.
The QTL identified in this study exhibit modest, mostly additive effects, and GI/Wisc alleles are associated with larger skeletal size at most QTL. Many of these loci also colocalize with QTL for body weight. The high degree of QTL colocalization is consistent with an important contribution of pleiotropy to skeletal evolution.
A total of 208 QTL are identified across 16 skeletal traits and three time points (genome coordinates relative to GRCm38/mm10):
Skull length
Sklq1 (skull length QTL 1, 5 week) maps to Chr 1: 76 - 181.5 Mb with a peak LOD score of 4.8 at 159.6 Mb.
Sklq2 (skull length QTL 2, 5 week) maps to Chr 4: 36.7 - 129.1 Mb with a peak LOD score of 4.3 at 59.2 Mb.
Sklq3 (skull length QTL 3, 5 week) maps to Chr 6: 7.6 - 80.6 Mb with a peak LOD score of 4.8 at 75.5 Mb.
Sklq4 (skull length QTL 4, 5 week) maps to Chr 7: 65.6 - 126.9 Mb with a peak LOD score of 6.2 at 76.3 Mb.
Sklq5 (skull length QTL 5, 5 week) maps to Chr 10: 119.2 - 125 Mb with a peak LOD score of 22.4 at 123.4 Mb.
Sklq6 (skull length QTL 6, 5 week) maps to Chr 15: 6.1 - 100.9 Mb with a peak LOD score of 5.2 at 71.6 Mb.
Sklq7 (skull length QTL 7, 10 week) maps to Chr 4: 68.1 - 150.8 Mb with a peak LOD score of 8.1 at 93.6 Mb.
Sklq8 (skull length QTL 8, 10 week) maps to Chr 7: 68.2 - 124.6 Mb with a peak LOD score of 9 at 105.2 Mb.
Sklq9 (skull length QTL 9, 10 week) maps to Chr 10: 67.5 - 111.9 Mb with a peak LOD score of 14.2 at 110.4 Mb.
Sklq10 (skull length QTL 10, 10 week) maps to Chr 11: 3.3 - 63 Mb with a peak LOD score of 4.8 at 12.7 Mb.
Sklq11 (skull length QTL 11, 10 week) maps to Chr 15: 8.1 - 47.8 Mb with a peak LOD score of 10.3 at 30.6 Mb.
Sklq12 (skull length QTL 12, 16 week) maps to Chr 2: 24.7 - 55 Mb with a peak LOD score of 4.5 at 43 Mb.
Sklq13 (skull length QTL 13, 16 week) maps to Chr 4: 57.7 - 155.6 Mb with a peak LOD score of 4.7 at 127 Mb.
Sklq14 (skull length QTL 14, 16 week) maps to Chr 7: 68.2 - 125.4 Mb with a peak LOD score of 6.2 at 118.9 Mb.
Sklq15 (skull length QTL 15, 16 week) maps to Chr 10: 59.5 - 122.5 Mb with a peak LOD score of 9.4 at 67.1 Mb.
Sklq16 (skull length QTL 16, 16 week) maps to Chr 11: 3.3 - 72.8 Mb with a peak LOD score of 4.3 at 16.2 Mb.
Sklq17 (skull length QTL 17, 16 week) maps to Chr 15: 30.2 - 58.1 Mb with a peak LOD score of 7.3 at 46.5 Mb.
Skull width
Skwq1 (skull length QTL 1, 5 week) maps to Chr 3: 44 - 101.7 Mb with a peak LOD score of 8.9 at 58.3 Mb.
Skwq2 (skull length QTL 2, 5 week) maps to Chr 8: 120 - 131.2 Mb with a peak LOD score of 4.3 at 123.9 Mb.
Skwq3 (skull length QTL 3, 5 week) maps to Chr 10: 114.8 - 122.9 Mb with a peak LOD score of 18.8 at 121.2 Mb.
Skwq4 (skull length QTL 4, 5 week) maps to Chr 16: 8.6 - 91.1 Mb with a peak LOD score of 4.8 at 29.7 Mb.
Skwq5 (skull length QTL 5, 5 week) maps to Chr 17: 32.9 - 53.8 Mb with a peak LOD score of 6.5 at 48.1 Mb.
Skwq6 (skull length QTL 6, 10 week) maps to Chr 3: 52 - 102.4 Mb with a peak LOD score of 12.8 at 100.5 Mb.
Skwq7 (skull length QTL 7, 10 week) maps to Chr 10: 101.9 - 113.9 Mb with a peak LOD score of 20.3 at 107.4 Mb.
Skwq8 (skull length QTL 8, 10 week) maps to Chr 15: 6.1 - 68.4 Mb with a peak LOD score of 4.7 at 9.2 Mb.
Skwq9 (skull length QTL 9, 10 week) maps to Chr 17: 43.4 - 67.5 Mb with a peak LOD score of 6.2 at 47.4 Mb.
Skwq10 (skull length QTL 10, 16 week) maps to Chr 3: 44 - 65.4 Mb with a peak LOD score of 10.8 at 59.6 Mb.
Skwq11 (skull length QTL 11, 16 week) maps to Chr 10: 113.7 - 121.8 Mb with a peak LOD score of 19.6 at 121.2 Mb.
Skwq12 (skull length QTL 12, 16 week) maps to Chr 15: 6.1 - 50.9 Mb with a peak LOD score of 4.7 at 12.6 Mb.
Skwq13 (skull length QTL 13, 16 week) maps to Chr 17: 43.4 - 52.8 Mb with a peak LOD score of 6.5 at 48.1 Mb.
Skwq14 (skull length QTL 14, 16 week) maps to Chr 19: 3.2 - 35.9 Mb with a peak LOD score of 4.3 at 15.1 Mb.
Bi-zygomatic width
Bzwq1 (bi-zygomatic width QTL 1, 5 week) maps to Chr 3: 52.3 - 137.5 Mb with a peak LOD score of 4.6 at 128.9 Mb.
Bzwq2 (bi-zygomatic width QTL 2, 5 week) maps to Chr 7: 65.6 - 97.3 Mb with a peak LOD score of 6.5 at 81.3 Mb.
Bzwq3 (bi-zygomatic width QTL 3, 5 week) maps to Chr 10: 114.4 - 122.9 Mb with a peak LOD score of 20.5 at 119.5 Mb.
Bzwq4 (bi-zygomatic width QTL 4, 5 week) maps to Chr 15: 6.1 - 72.6 Mb with a peak LOD score of 5.4 at 11.3 Mb.
Bzwq5 (bi-zygomatic width QTL 5, 5 week) maps to Chr 16: 37.5 - 84.9 Mb with a peak LOD score of 6.1 at 76.3 Mb.
Bzwq6 (bi-zygomatic width QTL 6, 10 week) maps to Chr 3: 100.7 - 137.5 Mb with a peak LOD score of 10.9 at 133.9 Mb.
Bzwq7 (bi-zygomatic width QTL 7, 10 week) maps to Chr 7: 68.2 - 105.8 Mb with a peak LOD score of 6.7 at 81.3 Mb.
Bzwq8 (bi-zygomatic width QTL 8, 10 week) maps to Chr 10: 45 - 116.1 Mb with a peak LOD score of 10.6 at 110.3 Mb.
Bzwq9 (bi-zygomatic width QTL 9, 10 week) maps to Chr 12: 16.1 - 47.7 Mb with a peak LOD score of 4.4 at 27.7 Mb.
Bzwq10 (bi-zygomatic width QTL 10, 10 week) maps to Chr 15: 6.1 - 71.6 Mb with a peak LOD score of 4.5 at 24.9 Mb.
Bzwq11 (bi-zygomatic width QTL 11, 16 week) maps to Chr 3: 125.2 - 137.5 Mb with a peak LOD score of 6.6 at 133.9 Mb.
Bzwq12 (bi-zygomatic width QTL 12, 16 week) maps to Chr 6: 3.4 - 139.1 Mb with a peak LOD score of 4.3 at 15.4 Mb.
Bzwq13 (bi-zygomatic width QTL 13, 16 week) maps to Chr 7: 62.1 - 121.8 Mb with a peak LOD score of 5.4 at 81.3 Mb.
Bzwq14 (bi-zygomatic width QTL 14, 16 week) maps to Chr 10: 45.6 - 120.8 Mb with a peak LOD score of 10.1 at 113.5 Mb.
Bzwq15 (bi-zygomatic width QTL 15, 16 week) maps to Chr 12: 3.6 - 29.7 Mb with a peak LOD score of 5.6 at 27.7 Mb.
Bzwq16 (bi-zygomatic width QTL 16, 16 week) maps to Chr 15: 6.1 - 38 Mb with a peak LOD score of 4.7 at 24.9 Mb.
Zygomatic length
Zlq1 (zygomatic length QTL 1, 5 week) maps to Chr 4: 53.7 - 103.4 Mb with a peak LOD score of 9.3 at 93.9 Mb.
Zlq2 (zygomatic length QTL 2, 5 week) maps to Chr 10: 113.7 - 125.9 Mb with a peak LOD score of 6.4 at 123.8 Mb.
Zlq3 (zygomatic length QTL 3, 10 week) maps to Chr 3: 3.2 - 142.7 Mb with a peak LOD score of 5.3 at 121.2 Mb.
Zlq4 (zygomatic length QTL 4, 10 week) maps to Chr 4: 85.2 - 105.1 Mb with a peak LOD score of 9 at 93.9 Mb.
Zlq5 (zygomatic length QTL 5, 10 week) maps to Chr 10: 68 - 125 Mb with a peak LOD score of 7.7 at 115.6 Mb.
Zlq6 (zygomatic length QTL 6, 10 week) maps to Chr 15: 6.8 - 45.1 Mb with a peak LOD score of 6.1 at 30.6 Mb.
Zlq7 (zygomatic length QTL 7, 10 week) maps to Chr 16: 4 - 64.8 Mb with a peak LOD score of 4.9 at 48.2 Mb.
Zlq8 (zygomatic length QTL 8, 16 week) maps to Chr 4: 74.8 - 108.5 Mb with a peak LOD score of 9.2 at 88.2 Mb.
Zlq9 (zygomatic length QTL 9, 16 week) maps to Chr 10: 64.1 - 116.1 Mb with a peak LOD score of 9.6 at 76.7 Mb.
Zlq10 (zygomatic length QTL 10, 16 week) maps to Chr 15: 20.6 - 50.9 Mb with a peak LOD score of 4.4 at 34.3 Mb.
Cranial vault height
Cvht1 (cranial vault height 1, 5 week) maps to Chr 10: 24.8 - 121.2 Mb with a peak LOD score of 5.1 at 31.3 Mb.
Cvht2 (cranial vault height 2, 5 week) maps to Chr 15: 28.8 - 102.1 Mb with a peak LOD score of 4.8 at 41.5 Mb.
Cvht3 (cranial vault height 3, 5 week) maps to Chr 17: 46.2 - 82.1 Mb with a peak LOD score of 5.3 at 60.3 Mb.
Cvht4 (cranial vault height 4, 10 week) maps to Chr 3: 16 - 109.8 Mb with a peak LOD score of 6.6 at 21.2 Mb.
Cvht5 (cranial vault height 5, 10 week) maps to Chr 10: 45.7 - 69.3 Mb with a peak LOD score of 8.6 at 63.2 Mb.
Cvht6 (cranial vault height 6, 10 week) maps to Chr 15: 45.7 - 69.3 Mb with a peak LOD score of 6.7 at 94.9 Mb.
Cvht7 (cranial vault height 7, 16 week) maps to Chr 3: 28.9 - 119.1 Mb with a peak LOD score of 4.7 at 59.6 Mb.
Cvht8 (cranial vault height 8, 16 week) maps to Chr 10: 23 - 91.2 Mb with a peak LOD score of 5 at 57.1 Mb.
Cvht9 (cranial vault height 9, 16 week) maps to Chr 15: 42.3 - 95.6 Mb with a peak LOD score of 4.7 at 71.8 Mb.
Humerus length
Huml1 (humerus length 1, 5 week) maps to Chr 7: 30.5 - 109.4 Mb with a peak LOD score of 4.7 at 95.4 Mb.
Huml2 (humerus length 2, 5 week) maps to Chr 10: 24.7 - 41.7 Mb with a peak LOD score of 4.9 at 36.6 Mb.
Huml3 (humerus length 3, 10 week) maps to Chr 4: 75.7 - 100.3 Mb with a peak LOD score of 4.3 at 94.4 Mb.
Huml4 (humerus length 4, 10 week) maps to Chr 11: 11.6 - 68.6 Mb with a peak LOD score of 4.6 at 63.6 Mb.
Huml5 (humerus length 5, 16 week) maps to Chr 10: 46.4 - 69.4 Mb with a peak LOD score of 5.2 at 58.6 Mb.
Humerus midshaft diameter
Humsd1 (humerus midshaft diameter 1, 5 week) maps to Chr 15: 48.5 - 101.8 Mb with a peak LOD score of 6.4 at 99.5 Mb.
Humsd2 (humerus midshaft diameter 2, 10 week) maps to Chr 5: 69.9 - 132.5 Mb with a peak LOD score of 5.1 at 127.5 Mb.
Humsd3 (humerus midshaft diameter 3, 10 week) maps to Chr 10: 22.5 - 51.2 Mb with a peak LOD score of 5 at 31.6 Mb.
Humsd4 (humerus midshaft diameter 4, 10 week) maps to Chr 13: 47.7 - 73 Mb with a peak LOD score of 4.9 at 58.7 Mb.
Humsd5 (humerus midshaft diameter 5, 10 week) maps to Chr 15: 48.5 - 98.5 Mb with a peak LOD score of 6.3 at 54.9 Mb.
Humsd6 (humerus midshaft diameter 6, 16 week) maps to Chr 7: 79 - 137.3 Mb with a peak LOD score of 5.1 at 135.1 Mb.
Humsd7 (humerus midshaft diameter 7, 16 week) maps to Chr 13: 3.9 - 75.9 Mb with a peak LOD score of 4.8 at 58.7 Mb.
Humsd8 (humerus midshaft diameter 8, 16 week) maps to Chr 15: 29.3 - 76.2 Mb with a peak LOD score of 5.5 at 53.2 Mb.
Ulna length
Ulnl1 (ulna length 1, 5 week) maps to Chr 1: 67.3 - 129.8 Mb with a peak LOD score of 4.4 at 78.1 Mb
Ulnl2 (ulna length 2, 5 week) maps to Chr 4: 57.7 - 94.9 Mb with a peak LOD score of 9.2 at 86.1 Mb.
Ulnl3 (ulna length 3, 5 week) maps to Chr 7: 73.5 - 143.6 Mb with a peak LOD score of 4.2 at 131.4 Mb.
Ulnl4 (ulna length 4, 5 week) maps to Chr 15: 56.9 - 87.8 Mb with a peak LOD score of 5.1 at 75.6 Mb.
Ulnl5 (ulna length 5, 10 week) maps to Chr 1: 128.8 - 181.5 Mb with a peak LOD score of 5.2 at 145.9 Mb.
Ulnl6 (ulna length 6, 10 week) maps to Chr 4: 66.8 - 98.7 Mb with a peak LOD score of 11.7 at 75.6 Mb.
Ulnl7 (ulna length 7, 10 week) maps to Chr 7: 66.8 - 98.7 Mb with a peak LOD score of 4.6 at 133.3 Mb.
Ulnl8 (ulna length 8, 10 week) maps to Chr 15: 7.2 - 55.4 Mb with a peak LOD score of 4.6 at 30.6 Mb.
Ulnl9 (ulna length 9, 16 week) maps to Chr 4: 65.4 - 119 Mb with a peak LOD score of 10.3 at 104.1 Mb.
Ulnl10 (ulna length 10, 16 week) maps to Chr 7: 81.2 - 127.1 Mb with a peak LOD score of 7.3 at 87.9 Mb.
Ulnl11 (ulna length 11, 16 week) maps to Chr 15: 6.1 - 76.5 Mb with a peak LOD score of 5 at 11.2 Mb.
Pelvic girdle length
Pglq1 (pelvic girdle length QTL 1, 5 week) maps to Chr 1: 76 - 128.6 Mb with a peak LOD score of 11 at 80.4 Mb.
Pglq2 (pelvic girdle length QTL 2, 5 week) maps to Chr 4: 69.2 - 131 Mb with a peak LOD score of 6.6 at 94.1 Mb.
Pglq3 (pelvic girdle length QTL 3, 5 week) maps to Chr 7: 81.6 - 124.6 Mb with a peak LOD score of 9.2 at 118.6 Mb.
Pglq4 (pelvic girdle length QTL 4, 5 week) maps to Chr 10: 119.2 - 125 Mb with a peak LOD score of 12.9 at 119.5 Mb.
Pglq5 (pelvic girdle length QTL 5, 5 week) maps to Chr 11: 3.3 - 19.8 Mb with a peak LOD score of 4.4 at 12.2 Mb.
Pglq6 (pelvic girdle length QTL 6, 10 week) maps to Chr 1: 76 - 148.9 Mb with a peak LOD score of 4.5 at 82.8 Mb.
Pglq7 (pelvic girdle length QTL 7, 10 week) maps to Chr 4: 88.2 - 108.5 Mb with a peak LOD score of 7.5 at 82.8 Mb.
Pglq8 (pelvic girdle length QTL 8, 10 week) maps to Chr 7: 88.2 - 108.5 Mb with a peak LOD score of 9.1 at 124.1 Mb.
Pglq9 (pelvic girdle length QTL 9, 10 week) maps to Chr 10: 33.9 - 71 Mb with a peak LOD score of 6.8 at 65.8 Mb.
Pglq10 (pelvic girdle length QTL 10, 10 week) maps to Chr 11: 3.3 - 18.5 Mb with a peak LOD score of 4.5 at 16.2 Mb.
Pglq11 (pelvic girdle length QTL 11, 10 week) maps to Chr 15: 8.1 - 38.9 Mb with a peak LOD score of 4.4 at 30.7 Mb.
Pglq12 (pelvic girdle length QTL 12, 16 week) maps to Chr 4: 83.2 - 108.5 Mb with a peak LOD score of 10.9 at 93.9 Mb.
Pglq13 (pelvic girdle length QTL 13, 16 week) maps to Chr 7: 73.4 - 126.1 Mb with a peak LOD score of 7 at 87.9 Mb.
Pglq14 (pelvic girdle length QTL 14, 16 week) maps to Chr 8: 73.4 - 126.1 Mb with a peak LOD score of 5 at 31.2 Mb.
Pglq15 (pelvic girdle length QTL 15, 16 week) maps to Chr 10: 39.3 - 129.8 Mb with a peak LOD score of 5.1 at 65.8 Mb.
Pglq16 (pelvic girdle length QTL 16, 16 week) maps to Chr 11: 3.3 - 18.2 Mb with a peak LOD score of 5.6 at 12.2 Mb.
Sacral vertebrae length
Scvln2 (sacral vertebrae length 2, 5 week) maps to Chr 1: 94.2 - 148.9 Mb with a peak LOD score of 8.3 at 135.5 Mb.
Scvln3 (sacral vertebrae length 2, 5 week) maps to Chr 4: 55.6 - 129.1 Mb with a peak LOD score of 5.4 at 94.1 Mb.
Scvln4 (sacral vertebrae length 2, 5 week) maps to Chr 7: 70.3 - 120.9 Mb with a peak LOD score of 7.3 at 97 Mb.
Scvln5 (sacral vertebrae length 2, 5 week) maps to Chr 10: 119.2 - 125 Mb with a peak LOD score of 11.5 at 97 Mb.
Scvln6 (sacral vertebrae length 2, 5 week) maps to Chr 11: 3.3 - 44.2 Mb with a peak LOD score of 5.1 at 16.2 Mb.
Scvln7 (sacral vertebrae length 2, 5 week) maps to Chr 16: 3.5 - 40 Mb with a peak LOD score of 5.9 at 4.8 Mb.
Scvln8 (sacral vertebrae length 2, 10 week) maps to Chr 1: 94.2 - 172.2 Mb with a peak LOD score of 5.7 at 137.3 Mb.
Scvln9 (sacral vertebrae length 2, 10 week) maps to Chr 4: 68.1 - 107.3 Mb with a peak LOD score of 10.6 at 94.1 Mb.
Scvln10 (sacral vertebrae length 2, 10 week) maps to Chr 7: 81.8 - 126.9 Mb with a peak LOD score of 6.1 at 97 Mb.
Scvln11 (sacral vertebrae length 2, 10 week) maps to Chr 10: 110.2 - 125.6 Mb with a peak LOD score of 6.6 at 121.2 Mb.
Scvln12 (sacral vertebrae length 2, 10 week) maps to Chr 11: 3.3 - 34.6 Mb with a peak LOD score of 5 at 12.2 Mb.
Scvln13 (sacral vertebrae length 2, 16 week) maps to Chr 1: 117.6 - 180 Mb with a peak LOD score of 4.7 at 162.2 Mb.
Scvln14 (sacral vertebrae length 2, 16 week) maps to Chr 4: 68.1 - 130.5 Mb with a peak LOD score of 7.2 at 94 Mb.
Scvln15 (sacral vertebrae length 2, 16 week) maps to Chr 7: 62.1 - 124.1 Mb with a peak LOD score of 4.6 at 87.9 Mb.
Scvln16 (sacral vertebrae length 2, 16 week) maps to Chr 8: 89 - 130.8 Mb with a peak LOD score of 4.2 at 128.5 Mb.
Scvln17 (sacral vertebrae length 2, 16 week) maps to Chr 10: 118.6 - 125 Mb with a peak LOD score of 6.9 at 119.5 Mb.
Scvln18 (sacral vertebrae length 2, 16 week) maps to Chr 11: 3.3 - 60.8 Mb with a peak LOD score of 6 at 16.2 Mb.
Scvln19 (sacral vertebrae length 2, 16 week) maps to Chr 12: 62.1 - 99.5 Mb with a peak LOD score of 4.5 at 70.1 Mb.
Scvln20 (sacral vertebrae length 2, 16 week) maps to Chr 15: 26.6 - 66.5 Mb with a peak LOD score of 4.3 at 30.7 Mb.
Femur length
Feml5 (femur length 5, 5 week) maps to Chr 1: 122.1 - 165 Mb with a peak LOD score of 5.8 at 145.4 Mb.
Feml6 (femur length 6, 5 week) maps to Chr 4: 87.6 - 103.4 Mb with a peak LOD score of 10.5 at 94.1 Mb.
Feml7 (femur length 7, 5 week) maps to Chr 7: 86.5 - 99.3 Mb with a peak LOD score of 15.2 at 90.8 Mb.
Feml8 (femur length 8, 5 week) maps to Chr 10: 119.2 - 125.9 Mb with a peak LOD score of 6.1 at 123.4 Mb.
Feml9 (femur length 9, 5 week) maps to Chr 11: 7.1 - 16.8 Mb with a peak LOD score of 5.1 at 12.9 Mb.
Feml10 (femur length 10, 5 week) maps to Chr 12: 68.3 - 113.4 Mb with a peak LOD score of 4.8 at 92.6 Mb.
Feml11 (femur length 11, 5 week) maps to Chr 15: 6.1 - 76 Mb with a peak LOD score of 6.6 at 72.3 Mb.
Feml12 (femur length 12, 5 week) maps to Chr 16: 3.5 - 64.8 Mb with a peak LOD score of 4.5 at 4.9 Mb.
Feml13 (femur length 13, 10 week) maps to Chr 2: 102.8 - 160.1 Mb with a peak LOD score of 4.4 at 149.7 Mb.
Feml14 (femur length 14, 10 week) maps to Chr 4: 85 - 107.3 Mb with a peak LOD score of 17.7 at 98.7 Mb.
Feml15 (femur length 15, 10 week) maps to Chr 7: 87.6 - 126.9 Mb with a peak LOD score of 14 at 125.2 Mb.
Feml16 (femur length 16, 10 week) maps to Chr 11: 87.6 - 126.9 Mb with a peak LOD score of 4.7 at 12.7 Mb.
Feml17 (femur length 17, 10 week) maps to Chr 15: 7.6 - 55.4 Mb with a peak LOD score of 9.5 at 44.3 Mb.
Feml18 (femur length 18, 16 week) maps to Chr 2: 94.4 - 154.4 Mb with a peak LOD score of 6.1 at 117.9 Mb.
Feml19 (femur length 19, 16 week) maps to Chr 4: 91 - 108.6 Mb with a peak LOD score of 15.8 at 104.1 Mb.
Feml20 (femur length 20, 16 week) maps to Chr 7: 85.4 - 124.6 Mb with a peak LOD score of 14.7 at 91.2 Mb.
Feml21 (femur length 21, 16 week) maps to Chr 11: 11.8 - 66.1 Mb with a peak LOD score of 4.8 at 51.8 Mb.
Feml22 (femur length 22, 16 week) maps to Chr 15: 7.6 - 59.4 Mb with a peak LOD score of 5.7 at 46.5 Mb.
Femur midshaft diameter
Femd1 (femur midshaft diameter 1, 5 week) maps to Chr 4: 124.1 - 144.6 Mb with a peak LOD score of 5.3 at 133.1 Mb.
Femd2 (femur midshaft diameter 2, 5 week) maps to Chr 8: 4.9 - 34.6 Mb with a peak LOD score of 5.3 at 29.1 Mb.
Femd3 (femur midshaft diameter 3, 5 week) maps to Chr 11: 81 - 102 Mb with a peak LOD score of 7 at 85 Mb.
Femd4 (femur midshaft diameter 4, 5 week) maps to Chr 14: 8.7 - 47 Mb with a peak LOD score of 5.6 at 32.6 Mb.
Femd5 (femur midshaft diameter 5, 10 week) maps to Chr 7: 53.6 - 143.6 Mb with a peak LOD score of 4.5 at 63.1 Mb.
Femd6 (femur midshaft diameter 6, 10 week) maps to Chr 8: 37.4 - 109.6 Mb with a peak LOD score of 5.3 at 60.1 Mb.
Femd7 (femur midshaft diameter 7, 10 week) maps to Chr 12: 9.5 - 84.1 Mb with a peak LOD score of 4.5 at 44.9 Mb.
Femd8 (femur midshaft diameter 8, 10 week) maps to Chr 14: 8.7 - 46.5 Mb with a peak LOD score of 4.4 at 10.8 Mb.
Femd9 (femur midshaft diameter 9, 10 week) maps to Chr X: 33.9 - 56.8 Mb with a peak LOD score of 6.1 at 48.1 Mb.
Femd10 (femur midshaft diameter 10, 16 week) maps to Chr 2: 124.6 - 168.4 Mb with a peak LOD score of 4.9 161 Mb.
Femd11 (femur midshaft diameter 11, 16 week) maps to Chr 7: 57.3 - 134.4 Mb with a peak LOD score of 4.3 at 71 Mb.
Femd12 (femur midshaft diameter 12, 16 week) maps to Chr X: 35.4 - 57.8 Mb with a peak LOD score of 8 at 48.1 Mb.
Tibia length
Tibl1 (tibia length 1, 5 week) maps to Chr 1: 95.6 - 162.2 Mb with a peak LOD score of 8.5 at 145.4 Mb.
Tibl2 (tibia length 2, 5 week) maps to Chr 4: 91 - 111.4 Mb with a peak LOD score of 12.7 at 106.9 Mb.
Tibl3 (tibia length 3, 5 week) maps to Chr 7: 75.2 - 122.6 Mb with a peak LOD score of 10.9 at 98.1 Mb.
Tibl4 (tibia length 4, 5 week) maps to Chr 10: 119.2 - 125.6 Mb with a peak LOD score of 9.6 at 119.5 Mb.
Tibl5 (tibia length 5, 5 week) maps to Chr 11: 68.3 - 102 Mb with a peak LOD score of 4.3 at 82.1 Mb.
Tibl6 (tibia length 6, 5 week) maps to Chr 12: 60.6 - 92.7 Mb with a peak LOD score of 5.7 at 76.4 Mb.
Tibl7 (tibia length 7, 5 week) maps to Chr 14: 30.6 - 48.6 Mb with a peak LOD score of 7.5 at 45.6 Mb.
Tibl8 (tibia length 8, 5 week) maps to Chr 15: 6.9 - 84.7 Mb with a peak LOD score of 4.6 at 74.2 Mb.
Tibl9 (tibia length 9, 5 week) maps to Chr 16: 3.5 - 51.6 Mb with a peak LOD score of 5.4 at 4.9 Mb.
Tibl10 (tibia length 10, 10 week) maps to Chr 1: 121.8 - 165 Mb with a peak LOD score of 6.5 at 145.9 Mb.
Tibl11 (tibia length 11, 10 week) maps to Chr 4: 88.6 - 108.5 Mb with a peak LOD score of 17.1 at 94.1 Mb.
Tibl12 (tibia length 12, 10 week) maps to Chr 7: 87.6 - 126.9 Mb with a peak LOD score of 10.2 at 125.2 Mb.
Tibl13 (tibia length 13, 10 week) maps to Chr 10: 19 - 129.8 Mb with a peak LOD score of 4.6 at 45.7 Mb.
Tibl14 (tibia length 14, 10 week) maps to Chr 12: 68.3 - 98.8 Mb with a peak LOD score of 4.7 at 79.8 Mb.
Tibl15 (tibia length 15, 10 week) maps to Chr 14: 30.6 - 87.7 Mb with a peak LOD score of 5 at 71.6 Mb.
Tibl16 (tibia length 16, 10 week) maps to Chr 15: 6.9 - 76 Mb with a peak LOD score of 6.2 at 30.6 Mb.
Tibl17 (tibia length 17, 16 week) maps to Chr 4: 88.6 - 108.6 Mb with a peak LOD score of 16.2 at 93.9 Mb.
Tibl18 (tibia length 18, 16 week) maps to Chr 7: 82.2 - 126.9 Mb with a peak LOD score of 5.1 at 125.2 Mb.
Tibl19 (tibia length 19, 16 week) maps to Chr 10: 39.3 - 129.8 Mb with a peak LOD score of 5.4 at 123.4 Mb.
Tibl20 (tibia length 20, 16 week) maps to Chr 12: 59.1 - 93.9 Mb with a peak LOD score of 5 at 70.8 Mb.
Tibl21 (tibia length 21, 16 week) maps to Chr 14: 34.4 - 52.7 Mb with a peak LOD score of 4.6 at 46.2 Mb.
Tibia midshaft diameter
Tibmd1 (tibia midshaft diameter 1, 5 week) maps to Chr 11: 62.7 - 109.5 Mb with a peak LOD score of 5.1 at 66.1 Mb.
Tibmd2 (tibia midshaft diameter 2, 5 week) maps to Chr 15: 38 - 86.8 Mb with a peak LOD score of 4.6 at 75.6 Mb.
Tibmd3 (tibia midshaft diameter 3, 10 week) maps to Chr 3: 96.7 - 148.1 Mb with a peak LOD score of 5.2 at 115.5 Mb.
Tibmd4 (tibia midshaft diameter 4, 10 week) maps to Chr 11: 94.3 - 119 Mb with a peak LOD score of 5.7 at 109 Mb.
Tibmd5 (tibia midshaft diameter 5, 10 week) maps to Chr 15: 6.1 - 86.8 Mb with a peak LOD score of 5.3 at 70.6 Mb.
Tibmd6 (tibia midshaft diameter 6, 16 week) maps to Chr 3: 101.4 - 150.1 Mb with a peak LOD score of 4.9 at 116.4 Mb.
Tibmd7 (tibia midshaft diameter 7, 16 week) maps to Chr 6: 17.7 - 53.5 Mb with a peak LOD score of 4.4 at 35.7 Mb.
Proximal tibia width
Tibw1 (tibia width 1, proximal, 5 week) maps to Chr 10: 102.9 - 123.4 Mb with a peak LOD score of 7.6 at 116.7 Mb.
Tibw2 (tibia width 2, proximal, 10 week) maps to Chr 4: 43 - 108.5 Mb with a peak LOD score of 5.5 at 62.3 Mb.
Tibw3 (tibia width 3, proximal, 10 week) maps to Chr 10: 119 - 125.6 Mb with a peak LOD score of 10.7 at 119.6 Mb.
Tibw4 (tibia width 4, proximal, 10 week) maps to Chr 15: 6.1 - 78.8 Mb with a peak LOD score of 4.5 at 30.6 Mb.
Tibw5 (tibia width 5, proximal, 10 week) maps to Chr 18: 50.3 - 62.5 Mb with a peak LOD score of 5.8 at 58.6 Mb.
Tibw6 (tibia width 6, proximal, 16 week) maps to Chr 5: 72.7 - 140 Mb with a peak LOD score of 4.9 at 129.3 Mb.
Tibw7 (tibia width 7, proximal, 16 week) maps to Chr 10: 108.8 - 125.6 Mb with a peak LOD score of 8.6 at 119.3 Mb.
Tibw8 (tibia width 8, proximal, 16 week) maps to Chr 15: 6.9 - 76 Mb with a peak LOD score of 4.3 at 50.9 Mb.
Tibw9 (tibia width 9, proximal, 16 week) maps to Chr 18: 3.2 - 60.8 Mb with a peak LOD score of 4.4 at 21 Mb.
Metatarsal-calcaneal length
Metcl1 (metatarsal-calcaneal length 1, 5 week) maps to Chr 1: 121.8 - 139.8 Mb with a peak LOD score of 8.3 at 128.4 Mb.
Metcl2 (metatarsal-calcaneal length 2, 5 week) maps to Chr 4: 88.2 - 118.7 Mb with a peak LOD score of 17.2 at 106.9 Mb.
Metcl3 (metatarsal-calcaneal length 3, 5 week) maps to Chr 7: 68.2 - 119.9 Mb with a peak LOD score of 6.1 at 86.2 Mb.
Metcl4 (metatarsal-calcaneal length 4, 5 week) maps to Chr 10: 118.3 - 125.6 Mb with a peak LOD score of 10.1 at 119.4 Mb.
Metcl5 (metatarsal-calcaneal length 5, 5 week) maps to Chr 11: 4.7 - 46.4 Mb with a peak LOD score of 4.9 at 12.2 Mb.
Metcl6 (metatarsal-calcaneal length 6, 5 week) maps to Chr 14: 30.6 - 52.7 Mb with a peak LOD score of 5.7 at 45.6 Mb.
Metcl7 (metatarsal-calcaneal length 7, 10 week) maps to Chr 1: 121.8 - 154 Mb with a peak LOD score of 7.6 at 137.9 Mb.
Metcl8 (metatarsal-calcaneal length 8, 10 week) maps to Chr 4: 87.6 - 107.3 Mb with a peak LOD score of 12.9 at 88.6 Mb.
Metcl9 (metatarsal-calcaneal length 9, 10 week) maps to Chr 7: 75.2 - 122.6 Mb with a peak LOD score of 6.2 at 90.6 Mb.
Metcl10 (metatarsal-calcaneal length 10, 10 week) maps to Chr 10: 119.2 - 125 Mb with a peak LOD score of 10.7 at 122.9 Mb.
Metcl11 (metatarsal-calcaneal length 11, 10 week) maps to Chr 11: 8.9 - 61.5 Mb with a peak LOD score of 4.4 at 122.9 Mb.
Metcl12 (metatarsal-calcaneal length 12, 10 week) maps to Chr 14: 33.5 - 59.2 Mb with a peak LOD score of 5.4 at 45.6 Mb.
Metcl13 (metatarsal-calcaneal length 13, 16 week) maps to Chr 1: 121.8 - 153.9 Mb with a peak LOD score of 6.7 139.4 Mb.
Metcl14 (metatarsal-calcaneal length 14, 16 week) maps to Chr 4: 87.6 - 108.5 Mb with a peak LOD score of 13.9 at 94.1 Mb.
Metcl15 (metatarsal-calcaneal length 15, 16 week) maps to Chr 10: 119.2 - 125 Mb with a peak LOD score of 10.7 at 122.5 Mb.
Metcl16 (metatarsal-calcaneal length 16, 16 week) maps to Chr 11: 5.3 - 35.2 Mb with a peak LOD score of 5.2 at 12.2 Mb.
Analysis Tools