Table 2.
QTL region | Best candidate varianta | Number of traitsd | ||||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
N | BTA | From (bp) | To (bp) | # variants | #genes | bp | Variant ID | Rank | R2 | MAF | Effect | SE | − log10P | Gene | Functional annotation | Trait most strongly affected | CMP | Proteins | Fatty acids | Minerals |
1 | 1 | 144,389,419 | 144,398,814 | 6 | 1 | 144,395,375 | rs136069703 | 1 | 0.97 | 0.38 | 29.5 | 0.951 | 210.4 | SLC37A1 | Intron | P | 3 | 1 | 1 | 4 |
2 | 2 | 5,718,384 | 6,567,522 | 111 | 5 | 5,865,070 | rs136830094 | 1 | 0.91 | 0.22 | − 0.02 | 0.003 | 15.6 | INPP1 | Downstream | Lactose | 0 | 1 | 0 | 2 |
3 | 2 | 131,808,301 | 131,888,417 | 88 | 1 | 131,816,616 | rs133677653 | 1 | 0.91 | 0.49 | − 0.05 | 0.003 | 41.4 | ALPL | Intron | κ-CN | 1 | 3 | 0 | 1 |
4 | 3 | 7,442,755 | 8,149,715 | 56 | 5 | 7,935,102 | rs383033753 | 7 | 0.50 | 0.14 | − 0.65 | 0.096 | 10.7 | FCGR2B | Intron | CYDM | 2 | 0 | 1 | 0 |
5 | 3 | 15,514,034 | 15,928,379 | 401 | 7 | 15,525,599 | rs110073735 | 1 | 0.99 | 0.03 | − 2.78 | 0.410 | 10.9 | EFNA1 | Intron | Lactose | 1 | 0 | 0 | 3 |
6 | 3 | 34,235,208 | 34,355,357 | 35 | 4 | 34,327,146 | rs210558120 | 1 | 0.34 | 0.43 | 0.26 | 0.035 | 12.9 | KIAA1324 | Intron | aSC | 2 | 0 | 0 | 0 |
7 | 4 | 49,033,707 | 49,153,995 | 39 | 2 | 49,033,707 | rs380575157 | 1 | 0.47 | 0.20 | 0.02 | 0.001 | 30.7 | CBLL1 | Upstream | pH0_PCC | 2 | 3 | 0 | 2 |
8 | 4 | 75,743,094 | 79,803,738 | 59 | 5 | 77,825,429 | rs385069094 | 1 | 0.57 | 0.01 | − 2.34 | 0.360 | 10.1 | GCK | Intron | Mg | 0 | 1 | 0 | 3 |
9 | 4 | 92,588,016 | 92,966,245 | 227 | 7 | 92,624,543 | rs379514460 | 1 | 0.34 | 0.03 | − 29.0 | 4.226 | 11.2 | FSCN3 | Upstream | Ca | 0 | 0 | 0 | 1 |
10 | 5 | 29,947,476 | 31,423,430 | 315 | 25 | 29,947,476 | rs442522314 | 2 | 0.63 | 0.09 | − 2.26 | 0.383 | 8.4 | COX14 | Downstream | Lactose | 0 | 0 | 0 | 1 |
11 | 5 | 93,892,583 | 93,945,738 | 9 | 1 | 93,943,700 | rs210744452 | 1 | 0.77 | 0.05 | 0.11 | 0.010 | 27.7 | MGST1 | Intron | FC | 2 | 0 | 7 | 0 |
12 | 5 | 117,126,900 | 119,221,867 | 51 | 3 | 117,972,265 | rs525880746 | 1 | 0.79 | 0.04 | 42.4 | 2.186 | 83.0 | GRAMD4 | Upstream | Ca | 7 | 4 | 1 | 6 |
13 | 6 | 37,857,989 | 38,326,250 | 268 | 8 | 38,326,250 | rs382477515 | 1 | 0.37 | 0.42 | − 0.10 | 0.017 | 9.5 | IBSP | Upstream | β-CN | 0 | 1 | 0 | 0 |
14 | 6 | 46,555,489 | 47,082,793 | 89 | 3 | 46,876,802 | rs110408618 | 1 | 1.00 | 0.17 | 27.8 | 1.189 | 120.5 | SEL1L3 | Intron | K | 4 | 5 | 1 | 4 |
15 | 6 | 87,125,482 | 87,961,577 | 143 | 7 | 87,392,899 | rs382350292 | 3 | 0.53 | 0.23 | 0.03 | 0.002 | 76.4 | CSN3 | Downstream | K10/RSC | 8 | 8 | 0 | 1 |
16 | 6 | 108,967,622 | 109,145,154 | 91 | 5 | 109,026,822 | rs208213463 | 1 | 0.93 | 0.47 | − 8.42 | 0.904 | 19.9 | GAK | Intron | Ca | 2 | 2 | 0 | 2 |
17 | 7 | 958,741 | 1,146,962 | 359 | 4 | 975,515 | rs383246531 | 1 | 0.93 | 0.37 | − 1.56 | 0.180 | 17.4 | 47695b | Intron | Lactose | 0 | 0 | 0 | 2 |
18 | 7 | 41,576,519 | 46,646,915 | 399 | 23 | 46,452,656 | rs209828204 | 6 | 0.87 | 0.07 | 1.36 | 0.127 | 26.0 | FSTL4 | Downstream | aPCC | 2 | 2 | 0 | 1 |
19 | 9 | 102,731,669 | 103,130,410 | 207 | 3 | 102,885,120 | rs134445867 | 15 | 0.90 | 0.03 | − 0.14 | 0.023 | 9.3 | MPC1 | Intron | C4_C10 | 0 | 0 | 1 | 0 |
20 | 10 | 1,984,741 | 2,326,212 | 86 | 1 | 2,096,282 | rs385793060 | 67 | 0.99 | 0.35 | − 2.04 | 0.380 | 7.1 | 47622b | Upstream | Na | 0 | 0 | 0 | 1 |
21 | 10 | 48,359,318 | 50,266,445 | 213 | 1 | 49,459,919 | rs109896326 | 1 | 1.00 | 0.14 | 0.30 | 0.046 | 9.8 | RORA | Intron | UNSAT | 0 | 0 | 4 | 0 |
22 | 10 | 99,714,774 | 99,843,583 | 20 | 0 | 99,714,774 | rs440530756 | 1 | 0.22 | 0.02 | − 0.18 | 0.028 | 9.8 | – | Intergenic | PUNSAT | 0 | 0 | 2 | 0 |
23 | 11 | 9,023,594 | 9,684,851 | 11 | 3 | 9,684,851 | rs384459785 | 8 | 0.62 | 0.07 | − 2.82 | 0.385 | 12.6 | POLE4 | Intron | CYFAT-PROT | 3 | 0 | 3 | 0 |
24 | 11 | 14,152,677 | 15,493,112 | 132 | 6 | 14,284,886 | rs384594145 | 48 | 0.90 | 0.10 | − 0.01 | 0.001 | 18.7 | XDH | Upstream | pH0_PCC | 1 | 0 | 0 | 1 |
25 | 11 | 86,907,041 | 86,916,295 | 9 | 1 | 86,912,990 | rs481567394 | 1 | 0.21 | 0.05 | 0.02 | 0.003 | 16.7 | ATP6V1C2 | Intron | pH0_PCC | 1 | 0 | 0 | 0 |
26 | 11 | 103,273,963 | 103,322,890 | 214 | 4 | 103,301,982 | rs109907194 | 48 | 0.89 | 0.46 | − 1.09 | 0.017 | 559.7 | PAEP | Intron | ΣWP | 8 | 7 | 2 | 3 |
27 | 12 | 68,616,690 | 77,578,414 | 337 | 7 | 70,162,028 | rs721489054 | 22 | 0.83 | 0.17 | 0.22 | 0.019 | 29.0 | ABCC4 | Intron | C14:0 | 0 | 0 | 7 | 2 |
28 | 13 | 20,094,707 | 22,437,171 | 291 | 4 | 21,053,894 | rs378591536 | 7 | 0.34 | 0.07 | − 0.24 | 0.034 | 12.1 | 23216b | Intron | C14:0 | 0 | 0 | 1 | 0 |
29 | 13 | 45,394,264 | 48,611,254 | 75 | 4 | 46,734,011 | rs379821485 | 18 | 0.76 | 0.15 | − 0.01 | 0.001 | 10.6 | RF0026 | Downstream | pH0_PCC | 1 | 0 | 1 | 0 |
30 | 13 | 52,289,279 | 55,114,121 | 80 | 9 | 54,938,610 | rs110422533 | 2 | 1.00 | 0.39 | − 6.60 | 1.007 | 10.3 | GID8 | Synonymous | K | 0 | 0 | 2 | 1 |
31 | 13 | 64,648,620 | 64,870,118 | 19 | 3 | 64,812,464 | rs43717459 | 1 | 0.33 | 0.10 | − 0.24 | 0.027 | 18.9 | ACSS2 | Intron | C4–C10 | 0 | 0 | 1 | 0 |
32 | 13 | 79,225,285 | 79,326,265 | 6 | 1 | 79,326,265 | rs109862148 | 3 | 0.38 | 0.07 | − 0.04 | 0.007 | 8.9 | FAM65C | Downstream | αs2-CN | 0 | 1 | 0 | 0 |
33 | 14 | 1,622,956 | 1,881,116 | 27 | 9 | 1,629,753 | rs109035586 | 1 | 0.36 | 0.33 | 0.17 | 0.007 | 122.7 | GPT | Upstream | FC | 6 | 6 | 10 | 4 |
34 | 15 | 39,854,757 | 40,283,508 | 140 | 2 | 39,885,845 | rs134953698 | 2 | 0.79 | 0.18 | − 0.04 | 0.005 | 10.4 | ARNTL | Intron | PUNSAT | 0 | 0 | 1 | 0 |
35 | 15 | 52,993,384 | 54,612,060 | 341 | 10 | 53,943,342 | rs381948106 | 1 | 0.47 | 0.04 | 17.5 | 2.747 | 9.7 | RAB6A | Intron | P | 0 | 0 | 2 | 2 |
36 | 16 | 1,607,369 | 3,050,452 | 10 | 0 | 1,609,129 | rs42450079 | 1 | 0.54 | 0.28 | − 0.04 | 0.005 | 17.0 | – | Intergenic | κ-CN | 2 | 6 | 0 | 0 |
37 | 16 | 60,539,357 | 63,891,341 | 178 | 3 | 60,646,127 | rs137615589 | 72 | 0.75 | 0.14 | 0.02 | 0.004 | 9.8 | 38238b | Upstream | aSC | 2 | 1 | 0 | 0 |
38 | 16 | 67,700,538 | 67,811,269 | 167 | 2 | 67,758,163 | rs42465711 | 1 | 1.00 | 0.4 | 0.31 | 0.044 | 11.9 | SWT1 | Intron | CYDM | 2 | 1 | 2 | 1 |
39 | 16 | 70,170,086 | 71,493,899 | 59 | 1 | 71,432,479 | rs109766366 | 5 | 0.79 | 0.43 | − 0.23 | 0.032 | 12.3 | PROX1 | Intron | UNSAT | 0 | 0 | 4 | 2 |
40 | 17 | 29,348,215 | 30,211,790 | 203 | 9 | 29,938,428 | rs207509104 | 21 | 0.55 | 0.36 | 0.10 | 0.015 | 10.3 | LARP1B | Synonymous | C4–C10 | 0 | 0 | 1 | 0 |
41 | 17 | 52,753,338 | 53,240,467 | 283 | 6 | 53,072,959 | rs448501071 | 5 | 1.00 | 0.06 | − 0.32 | 0.019 | 62.4 | BRI3BP | Intron | C4–C10 | 2 | 3 | 2 | 1 |
42 | 18 | 10,566,605 | 11,091,131 | 68 | 5 | 11,002,789 | rs41867427 | 2 | 0.61 | 0.06 | 14.3 | 2.071 | 11.3 | CRISPLD2 | Intron | aSC | 2 | 0 | 0 | 1 |
43 | 19 | 51,304,834 | 51,538,272 | 67 | 2 | 51,383,847 | rs136067046 | 1 | 0.83 | 0.32 | 0.20 | 0.013 | 49.4 | FASN | Upstream | C14:0 | 0 | 0 | 9 | 0 |
44 | 19 | 55,229,384 | 57,240,571 | 267 | 10 | 57,151,350 | rs42848485 | 1 | 0.63 | 0.46 | 9.07 | 1.079 | 16.4 | FADS6 | Intron | Ca | 2 | 1 | 1 | 1 |
45 | 19 | 60,407,923 | 62,177,206 | 142 | 0 | 61,135,270 | rs41923848 | 1 | 0.91 | 0.13 | − 2.28 | 0.210 | 26.6 | – | Intergenic | Lactose | 0 | 3 | 0 | 3 |
46 | 20 | 58,245,970 | 58,457,768 | 87 | 1 | 58,446,058 | rs137085630 | 22 | 0.99 | 0.06 | − 1.01 | 0.036 | 176.5 | ANKH | Intron | Citrate | 5 | 4 | 2 | 5 |
47 | 21 | 40,120,343 | 44,138,058 | 63 | 2 | 41,638,428 | rs137153434 | 25 | 0.21 | 0.12 | 0.07 | 0.009 | 12.7 | G2E3 | Upstream | PC | 3 | 1 | 0 | 1 |
48 | 22 | 32,877,755 | 33,466,544 | 46 | 2 | 32,877,755 | rs208141216 | 10 | 0.30 | 0.08 | 0.08 | 0.007 | 25.7 | FAM19A4 | Intron | PC | 8 | 1 | 1 | 2 |
49 | 22 | 55,186,094 | 55,273,619 | 72 | 1 | 55,254,221 | rs43597796 | 1 | 0.93 | 0.49 | 0.00 | 0.001 | 12.6 | ATP2B2 | Intron | pH0_PCC | 1 | 1 | 0 | 0 |
50 | 22 | 61,257,725 | 61,312,492 | 18 | 1 | 61,284,069 | rs109001472 | 1 | 1.00 | 0.47 | − 0.09 | 0.008 | 28.3 | KLF15 | Intron | C4–C10 | 0 | 0 | 2 | 0 |
51 | 24 | 50,420,365 | 50,550,020 | 114 | 2 | 50,465,348 | rs383068825 | 4 | 0.53 | 0.34 | − 10.7 | 1.282 | 16.0 | SKA1 | Downstream | K | 1 | 1 | 0 | 2 |
52 | 24 | 58,744,952 | 58,825,217 | 23 | 3 | 58,817,202 | rs208779762 | 1 | 0.93 | 0.43 | − 6.57 | 0.807 | 15.4 | LMAN1 | Upstream | P | 1 | 1 | 0 | 1 |
53 | 25 | 2,994,081 | 3,261,509 | 28 | 3 | 3,241,838 | rs137696417 | 16 | 0.61 | 0.18 | − 0.03 | 0.007 | 6.5 | ADCY9 | Downstream | κ-CN | 0 | 1 | 0 | 0 |
54 | 25 | 25,642,563 | 29,605,418 | 82 | 11 | 26,498,356 | rs137150057 | 1 | 1.00 | 0.34 | − 0.11 | 0.020 | 11.3 | FAM57B | 5′UTR | C18:0 | 1 | 1 | 6 | 0 |
55 | 26 | 20,727,700 | 21,427,109 | 61 | 4 | 21149234 | rs136334180 | 11 | 1.00 | 0.31 | 0.39 | 0.035 | 27.6 | SCD | Upstream | UNSAT | 0 | 0 | 11 | 0 |
56 | 26 | 32,773,808 | 33,925,908 | 95 | 2 | 33,233,277 | rs385554497 | 36 | 0.27 | 0.02 | 1.55 | 0.270 | 8.1 | ACSL5 | intron | aPCC | 1 | 0 | 0 | 0 |
57 | 27 | 36,165,492 | 36,235,730 | 11 | 1 | 36,212,352 | rs208675276 | 1 | 0.54 | 0.41 | 0.65 | 0.040 | 60.4 | GPAT4c | 5′ UTR | C16:0 | 3 | 0 | 7 | 1 |
58 | 28 | 6,008,464 | 7,038,810 | 141 | 3 | 6,027,037 | rs382911338 | 1 | 0.37 | 0.18 | − 0.01 | 0.001 | 17.1 | PCNX2 | Intron | pH0_PCC | 1 | 2 | 0 | 1 |
59 | 29 | 9,253,006 | 9,622,389 | 145 | 3 | 9,343,362 | rs133715120 | 6 | 0.61 | 0.33 | − 2.76 | 0.264 | 24.9 | EED | Intron | Lactose | 0 | 3 | 1 | 4 |
aWhen a gene was present in the confidence interval of the QTL, the best candidate variant was the genic variant with the most significant effects (intergenic variants were discarded)
b47695, 47622, 23216, and 38238 for ENSBTAG00000047695, ENSBTAG00000047622, ENSBTAG00000023216, and ENSBTAG00000038238, respectively
cAlso named AGPAT6
dNumber of milk cheese-making (CMP), protein, fatty acid, and mineral composition traits with significant effects