395:(EMT). Although this is an important process for neural tube development and other fetal structures, it is also a necessary process for tumor metastasis as it can activate the TGF beta pathway in the event of an overexpression of GREM1. This has made GREM1 the proposed target for cancer therapeutics, however, more research is necessary before any major steps are taken.
391:(VEGF). Both molecules are capable of binding to the VEGF receptor to activate vascular differentiation and proliferation during development. In the absence of GREM1, it is possible to see unregulated bone growth as there is no inhibitory signal to regulate the bone morphogenetic proteins. Gremlin1 also plays a role in the
1217:
Jacqueline L. Norrie, Jordan P. Lewandowski, Cortney M. Bouldin, Smita
Amarnath, Qiang Li, Martha S. Vokes, Lauren I.R. Ehrlich, Brian D. Harfe, Steven A. Vokes. Dynamics of BMP signaling in limb bud mesenchyme and polydactyly (2014) Developmental Biology Volume 393, Issue 2, Pages
914:
Michos O, Gonçalves A, Lopez-Rios J, Tiecke E, Naillat F, Beier K, Galli A, Vainio S, Zeller R (2007). "Reduction of BMP4 activity by gremlin 1 enables ureteric bud outgrowth and GDNF/WNT11 feedback signalling during kidney branching morphogenesis".
468:. Conditional deletion of one copy of Grem1 does not produce an abnormal phenotype and deletion of both copies causes only a small difference in phenotype in one-month-old male mice, but this difference cannot be observed after 3 months of age.
366:
in limb buds and functions as part of a self-regulatory feedback signaling system, which is essential for normal limb bud development and digit formation. Inhibition of BMPs by Grem1 in limb buds allows the transcriptional up-regulation of the
1236:
Zuniga, A., Laurent, F., Lopez-Rios, J., Klasen, C., Matt, N. and Zeller, R. (2012). Conserved cis-regulatory regions in a large genomic landscape control SHH and BMP-regulated
Gremlin1 expression in mouse limb buds. BMC Dev. Biol. 12,
509:(CRMs) regulate when and where Grem1 is transcribed. It has been reported that a CRM acts as both a silencer and activator for Grem1 transcription in the mouse limb bud. There are additional CRMs that regulate Grem1 transcription.
341:
part of the DAN family and is a cysteine knot-secreted protein. Gremlin1 was first identified in differential screening as a transcriptional down-regulated gene in v-mos-transformed rat embryonic fibroblasts.
1863:
1227:
Li, Q., Lewandowski, J. P., Powell, M. B., Norrie, J. L., Cho, S. H. and Vokes, S. a (2014). A Gli silencer is required for robust repression of gremlin in the vertebrate limb bud. Development 141, 1906–14
779:
Bénazet JD, Bischofberger M, Tiecke E, Gonçalves A, Martin JF, Zuniga A, Naef F, Zeller R (2009). "A self-regulatory system of interlinked signaling feedback loops controls mouse limb patterning".
387:
While GREM1 functions as a BMP antagonist during limb bud formation, it also functions as a pro-angiogenic molecule. As stated above, GREM1 is a member of the cysteine-knot superfamily similar to
375:(SHH) ligands, which are part of the signaling system that controls progression of limb bud development. Grem1 regulation of BMP4 in mice embryos is also essential for kidney and lung branching
423:
and proliferation in some cancers. Grem1 expression is found in many cancers and is thought to play important roles in uterine cervix, lung, ovary, kidney, breast, colon, pancreas, and
1856:
574:
Stabile H, Mitola S, Moroni E, Belleri M, Nicoli S, Coltrini D, Peri F, Pessi A, Orsatti L, Talamo F, Castronovo V, Waltregny D, Cotelli F, Ribatti D, Presta M (2007).
1849:
830:
Khokha MK, Hsu D, Brunet LJ, Dionne MS, Harland RM (2003). "Gremlin is the BMP antagonist required for maintenance of Shh and Fgf signals during limb patterning".
1062:"Gremlin-1: An endogenous BMP antagonist induces epithelial-mesenchymal transition and interferes with redifferentiation in fetal RPE cells with repeated wounds"
198:
61:
1830:
1291:
1591:
1312:
1728:
1722:
1717:
1707:
1702:
1697:
1692:
1687:
1115:"Bone morphogenetic protein antagonist gremlin 1 is widely expressed by cancer-associated stromal cells and can promote tumor cell proliferation"
679:
Zúñiga A, Haramis AP, McMahon AP, Zeller R (1999). "Signal relay by BMP antagonism controls the SHH/FGF4 feedback loop in vertebrate limb buds".
958:
Shi W, Zhao J, Anderson KD, Warburton D (2001). "Gremlin negatively modulates BMP-4 induction of embryonic mouse lung branching morphogenesis".
3421:
441:, (whose binding site for Grem1 is between residues 61-80) and is seen as a potential therapeutic and diagnostic target against human cancers.
392:
3479:
875:"Gremlin-mediated BMP antagonism induces the epithelial-mesenchymal feedback signaling controlling metanephric kidney and limb organogenesis"
730:
Zuniga A, Michos O, Spitz F, Haramis AP, Panman L, Galli A, Vintersten K, Klasen C, Mansfield W, Kuc S, Duboule D, Dono R, Zeller R (2004).
3484:
1544:
388:
1394:
222:
85:
1636:
732:"Mouse limb deformity mutations disrupt a global control region within the large regulatory landscape required for Gremlin expression"
490:. Overexpression of grem1 in stromal and osteoblastic cells in addition to the inhibition of BMP, grem 1 inhibits activation of Wnt/
1284:
210:
73:
3340:
2766:
2410:
2106:
1961:
1776:
1499:
1334:
203:
66:
486:. In addition, overexpression of Grem1 in the mouse limb bud inhibits BMP signaling which can lead to digit loss as well as
3172:
3008:
2873:
2756:
2658:
2584:
2396:
2287:
1951:
1786:
1459:
1452:
1447:
1442:
1437:
3298:
3208:
3144:
2964:
2841:
2729:
2540:
2186:
1886:
1872:
1432:
1427:
1422:
1417:
1412:
1407:
630:"The bone morphogenetic protein antagonist gremlin 1 is overexpressed in human cancers and interacts with YWHAH protein"
359:
1277:
351:
3140:
2960:
2837:
2536:
2182:
2075:
1936:
1738:
1402:
413:
355:
326:
322:
1003:"Insights into the Role of Gremlin-1, a Bone Morphogenic Protein Antagonist, in Cancer Initiation and Progression"
3284:
2177:
2065:
1521:
1304:
368:
1619:
1564:
1548:
1263:
1253:
479:
215:
78:
506:
495:
131:
1126:
788:
688:
268:
3431:
1671:
1356:
416:
2625:
2236:
2139:
1259:
1249:
3381:
3256:
2019:
1878:
1746:
1535:
1177:
Gazzerro E, Smerdel-Ramoya A, Zanotti S, Stadmeyer L, Durant D, Economides AN, Canalis E (2007).
983:
940:
855:
812:
712:
3213:
3056:
2895:
2610:
2814:
2715:
2503:
2498:
2468:
2463:
2363:
2483:
1946:
1326:
1200:
1154:
1091:
1073:
1042:
1024:
975:
932:
896:
847:
804:
761:
704:
661:
607:
553:
2705:
2630:
2453:
2353:
2241:
2144:
1994:
1179:"Conditional deletion of gremlin causes a transient increase in bone formation and bone mass"
3426:
3416:
2508:
2458:
2149:
1999:
1791:
1781:
1540:
1322:
1190:
1144:
1134:
1081:
1032:
1014:
967:
924:
886:
839:
796:
751:
743:
696:
651:
641:
597:
587:
543:
318:
263:
126:
2809:
2620:
2231:
2134:
2129:
3237:
3167:
3162:
3080:
3003:
2998:
2868:
2863:
2686:
2579:
2574:
2334:
2221:
2216:
2101:
1771:
1110:
483:
435:
420:
292:
155:
135:
1269:
272:
1130:
792:
692:
1841:
1675:
1300:
1149:
1114:
1086:
1061:
1037:
1002:
656:
629:
409:
372:
756:
731:
3473:
3391:
1984:
971:
376:
987:
859:
816:
3411:
2794:
2789:
2478:
2438:
2433:
2009:
944:
716:
472:
452:
of human lung tissue, which implies a role for Grem1 in the development of cancer.
445:
431:
338:
491:
475:
and a lesser known function later in adulthood. Overexpression of Grem1 decreases
337:
Gremlin1, previously known as Drm, is a highly conserved 20.7-kDa, 184 amino acid
1109:
Sneddon JB, Zhen HH, Montgomery K, van de Rijn M, Tward AD, West R, Gladstone H,
1019:
592:
576:"Bone morphogenic protein antagonist Drm/gremlin is a novel proangiogenic factor"
575:
532:"Skeletal overexpression of gremlin impairs bone formation and causes osteopenia"
239:
102:
3401:
3099:
2799:
2443:
1801:
1761:
487:
449:
329:
and is implicated in disorders of increased bone formation and several cancers.
302:
165:
3441:
3352:
3348:
3344:
3134:
2954:
2778:
2774:
2770:
2473:
2422:
2418:
2414:
2281:
2118:
2114:
2110:
2059:
1973:
1969:
1965:
1925:
1657:
1349:
1344:
1339:
530:
Gazzerro E, Pereira RC, Jorgetti V, Olson S, Economides AN, Canalis E (2005).
476:
465:
246:
109:
1077:
1028:
460:
Deletion of Grem1 in mice after birth increased bone formation and increased
3334:
3330:
3321:
3231:
3227:
3198:
3130:
3126:
3074:
3070:
3046:
2950:
2946:
2913:
2909:
2751:
2710:
2493:
2391:
2358:
2328:
2324:
2305:
2277:
2273:
2070:
2055:
2051:
1931:
1921:
1917:
1494:
1139:
800:
464:
bone volume, whereas overexpression causes inhibition of bone formation and
461:
427:
3307:
2738:
1204:
1195:
1178:
1158:
1095:
1046:
979:
936:
900:
851:
808:
765:
708:
665:
646:
628:
Namkoong H, Shin SM, Kim HK, Ha SA, Cho GW, Hur SY, Kim TE, Kim JW (2006).
611:
557:
3458:
548:
531:
251:
114:
3396:
3386:
3326:
3223:
3122:
3066:
2942:
2905:
2320:
2269:
2047:
1913:
1756:
1751:
1001:
Elemam NM, Malek AI, Mahmoud EE, El-Huneidi W, Talaat IM (2022-01-28).
747:
602:
424:
928:
891:
874:
873:
Michos O, Panman L, Vintersten K, Beier K, Zeller R, Zuniga A (2004).
3447:
3245:
3241:
3109:
3088:
3084:
2924:
2694:
2690:
2488:
2342:
2338:
1814:
1713:
1683:
1629:
1624:
1601:
1596:
1586:
1579:
1574:
234:
97:
17:
2523:
2164:
843:
3376:
3308:
3266:
3203:
3051:
3040:
3036:
2992:
2988:
2984:
2857:
2825:
2824:
2760:
2739:
2676:
2670:
2604:
2568:
2564:
2560:
2404:
2400:
2378:
2310:
2299:
2210:
2206:
2095:
2091:
1940:
1809:
1641:
1569:
1514:
1509:
1504:
1383:
1376:
1371:
1366:
700:
438:
3192:
3188:
3184:
3180:
3176:
3156:
3152:
3148:
3032:
3028:
3024:
3020:
3016:
3012:
2980:
2976:
2972:
2968:
2889:
2885:
2881:
2877:
2853:
2849:
2845:
2666:
2662:
2640:
2600:
2596:
2592:
2588:
2556:
2552:
2548:
2544:
2518:
2373:
2295:
2291:
2251:
2202:
2198:
2194:
2190:
2159:
2087:
2083:
2079:
2029:
1955:
1895:
1489:
1484:
1479:
1474:
1469:
1464:
1361:
363:
350:
Gremlin1 (Grem1) is known for its antagonistic interaction with
227:
90:
1845:
1273:
3267:
412:
of cancer and non-cancer tissues suggest that grem1 and other
180:
gremlin 2, cysteine knot superfamily, homolog (Xenopus laevis)
35:
gremlin 1, cysteine knot superfamily, homolog (Xenopus laevis)
494:
signaling activity. The interaction between Grem1 and the
1060:
Li D, Yuan D, Shen H, Mao X, Yuan S, Liu Q (2019-10-21).
482:
or the inhibition of bone formation and the ability for
3271:
1873:
2641:
2519:
2374:
2252:
2160:
2030:
1900:
1896:
3366:
3306:
3297:
3265:
3108:
2923:
2823:
2737:
2728:
2639:
2517:
2372:
2250:
2158:
2028:
1894:
1885:
1823:
1800:
1737:
1670:
1650:
1612:
1557:
1534:
1392:
1320:
1311:
298:
288:
283:
262:
257:
245:
233:
221:
209:
197:
189:
184:
179:
161:
151:
146:
125:
120:
108:
96:
84:
72:
60:
52:
44:
39:
34:
29:
Inhibitor protein in the TGF beta signaling pathway
2645:
2256:
2034:
3110:
2925:
1172:
1170:
1168:
2929:
623:
621:
525:
523:
521:
569:
567:
434:(between residues 1 to 67) interacts with the
1857:
1285:
8:
3303:
2734:
1891:
1864:
1850:
1842:
1554:
1317:
1292:
1278:
1270:
1113:, Morganroth GS, Oro AE, Brown PO (2006).
280:
143:
1262:at the U.S. National Library of Medicine
1252:at the U.S. National Library of Medicine
1194:
1148:
1138:
1085:
1036:
1018:
890:
755:
655:
645:
601:
591:
547:
444:Grem1 also plays a BMP-dependent role in
419:are important in the survival of cancer
517:
960:Am. J. Physiol. Lung Cell Mol. Physiol
176:
31:
7:
3374:Endogenous antagonists/inhibitors:
389:vascular endothelial growth factor
25:
471:Grem1 plays an important role in
393:epithelial-mesenchymal transition
972:10.1152/ajplung.2001.280.5.L1030
1313:TGF beta superfamily of ligands
430:. More specifically, the Grem1
358:. Grem1 inhibits predominantly
2150:ML-347 (LDN-193719, VU0469381)
2000:ML-347 (LDN-193719, VU0469381)
1460:Growth differentiation factors
1:
3480:Genes on human chromosome 15
1119:Proc. Natl. Acad. Sci. U.S.A
1020:10.3390/biomedicines10020301
593:10.1182/blood-2006-06-032276
3485:Genes on human chromosome 1
2489:RepSox (E-616452, SJN-2511)
1403:Bone morphogenetic proteins
352:bone morphogenetic proteins
3501:
502:Transcriptional regulation
356:TGF beta signaling pathway
323:TGF beta signaling pathway
2479:Galunisertib (LY-2157299)
498:is not fully understood.
369:fibroblast growth factors
279:
142:
3427:Thrombospondin 1 (THBS1)
3209:Osteogenin (BMP3, BMP3A)
1620:Activin type 2 receptors
1565:Activin type 1 receptors
1264:Medical Subject Headings
1254:Medical Subject Headings
325:. It primarily inhibits
1140:10.1073/pnas.0606857103
801:10.1126/science.1168755
2626:Dorsomorphin (BML-275)
2237:Dorsomorphin (BML-275)
2140:Dorsomorphin (BML-275)
1522:Anti-müllerian hormone
1305:TGFβ signaling pathway
1196:10.1074/jbc.M701317200
647:10.1186/1471-2407-6-74
507:Cis-regulatory modules
496:Wnt signaling pathway
399:Clinical significance
1876:receptor superfamily
1260:GREM2+protein,+human
1250:GREM1+protein,+human
549:10.1210/en.2004-0766
3444:(against myostatin)
1357:Activin and inhibin
1131:2006PNAS..10314842S
793:2009Sci...323.1050B
693:1999Natur.401..598Z
371:(FGFs) 4 and 8 and
3450:(against endoglin)
2807:Kinase inhibitors:
2703:Kinase inhibitors:
2618:Kinase inhibitors:
2451:Kinase inhibitors:
2351:Kinase inhibitors:
2229:Kinase inhibitors:
2127:Kinase inhibitors:
1992:Kinase inhibitors:
1536:TGF beta receptors
748:10.1101/gad.299904
327:bone morphogenesis
3467:
3466:
3362:
3361:
3293:
3292:
2724:
2723:
2017:Other inhibitors:
1839:
1838:
1739:Ligand inhibitors
1666:
1665:
1530:
1529:
929:10.1242/dev.02861
892:10.1242/dev.01251
687:(6753): 598–602.
383:Fetal Development
312:
311:
308:
307:
175:
174:
171:
170:
16:(Redirected from
3492:
3310:
3304:
3273:
3269:
3254:Decoy receptors:
3199:Myostatin (GDF8)
3112:
3097:Decoy receptors:
3047:Myostatin (GDF8)
2931:
2927:
2827:
2741:
2735:
2647:
2643:
2525:
2521:
2380:
2376:
2306:Myostatin (GDF8)
2258:
2254:
2166:
2162:
2036:
2032:
2007:Decoy receptors:
1902:
1898:
1892:
1875:
1866:
1859:
1852:
1843:
1555:
1318:
1294:
1287:
1280:
1271:
1238:
1234:
1228:
1225:
1219:
1215:
1209:
1208:
1198:
1189:(43): 31549–57.
1174:
1163:
1162:
1152:
1142:
1106:
1100:
1099:
1089:
1066:Molecular Vision
1057:
1051:
1050:
1040:
1022:
998:
992:
991:
955:
949:
948:
923:(13): 2397–405.
911:
905:
904:
894:
870:
864:
863:
827:
821:
820:
787:(5917): 1050–3.
776:
770:
769:
759:
727:
721:
720:
676:
670:
669:
659:
649:
625:
616:
615:
605:
595:
571:
562:
561:
551:
527:
473:bone development
281:
177:
144:
32:
21:
3500:
3499:
3495:
3494:
3493:
3491:
3490:
3489:
3470:
3469:
3468:
3463:
3382:Cerberus (CER1)
3358:
3289:
3261:
3168:Eptotermin alfa
3163:Dibotermin alfa
3104:
3004:Eptotermin alfa
2999:Dibotermin alfa
2919:
2869:Eptotermin alfa
2864:Dibotermin alfa
2819:
2720:
2635:
2580:Eptotermin alfa
2575:Dibotermin alfa
2513:
2368:
2246:
2222:Eptotermin alfa
2217:Dibotermin alfa
2154:
2102:Eptotermin alfa
2024:
1881:
1870:
1840:
1835:
1819:
1796:
1733:
1662:
1646:
1608:
1538:
1526:
1388:
1335:TGF beta family
1307:
1298:
1246:
1241:
1235:
1231:
1226:
1222:
1216:
1212:
1176:
1175:
1166:
1125:(40): 14842–7.
1108:
1107:
1103:
1059:
1058:
1054:
1000:
999:
995:
957:
956:
952:
913:
912:
908:
885:(14): 3401–10.
872:
871:
867:
829:
828:
824:
778:
777:
773:
742:(13): 1553–64.
729:
728:
724:
678:
677:
673:
627:
626:
619:
573:
572:
565:
529:
528:
519:
515:
504:
484:bone remodeling
480:differentiation
458:
436:binding protein
406:
401:
385:
348:
335:
30:
23:
22:
15:
12:
11:
5:
3498:
3496:
3488:
3487:
3482:
3472:
3471:
3465:
3464:
3462:
3461:
3452:
3451:
3445:
3435:
3434:
3429:
3424:
3419:
3414:
3409:
3404:
3399:
3394:
3389:
3384:
3379:
3370:
3368:
3364:
3363:
3360:
3359:
3357:
3356:
3338:
3324:
3315:
3313:
3301:
3295:
3294:
3291:
3290:
3288:
3287:
3278:
3276:
3263:
3262:
3260:
3259:
3250:
3249:
3235:
3217:
3216:
3211:
3206:
3201:
3196:
3170:
3165:
3160:
3138:
3116:
3114:
3106:
3105:
3103:
3102:
3093:
3092:
3078:
3060:
3059:
3054:
3049:
3044:
3006:
3001:
2996:
2958:
2936:
2934:
2921:
2920:
2918:
2917:
2899:
2898:
2893:
2871:
2866:
2861:
2831:
2829:
2821:
2820:
2818:
2817:
2812:
2803:
2802:
2797:
2792:
2783:
2782:
2764:
2754:
2745:
2743:
2732:
2726:
2725:
2722:
2721:
2719:
2718:
2713:
2708:
2699:
2698:
2680:
2679:
2674:
2652:
2650:
2637:
2636:
2634:
2633:
2628:
2623:
2614:
2613:
2608:
2582:
2577:
2572:
2530:
2528:
2515:
2514:
2512:
2511:
2506:
2501:
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2408:
2394:
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2370:
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2285:
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2239:
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2225:
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2219:
2214:
2180:
2171:
2169:
2156:
2155:
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2152:
2147:
2142:
2137:
2132:
2123:
2122:
2104:
2099:
2073:
2068:
2063:
2041:
2039:
2026:
2025:
2023:
2022:
2013:
2012:
2003:
2002:
1997:
1988:
1987:
1978:
1977:
1959:
1949:
1944:
1934:
1929:
1907:
1905:
1889:
1883:
1882:
1871:
1869:
1868:
1861:
1854:
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1528:
1527:
1525:
1524:
1519:
1518:
1517:
1512:
1507:
1502:
1497:
1495:Myostatin/GDF8
1492:
1487:
1482:
1477:
1472:
1467:
1457:
1456:
1455:
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1445:
1440:
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1369:
1364:
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1342:
1331:
1329:
1315:
1309:
1308:
1301:Cell signaling
1299:
1297:
1296:
1289:
1282:
1274:
1268:
1267:
1257:
1245:
1244:External links
1242:
1240:
1239:
1229:
1220:
1210:
1164:
1101:
1052:
993:
966:(5): L1030–9.
950:
906:
865:
844:10.1038/ng1178
822:
771:
722:
671:
617:
586:(5): 1834–40.
563:
516:
514:
511:
503:
500:
457:
454:
405:
402:
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397:
384:
381:
373:sonic hedgehog
354:(BMPs) in the
347:
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3:
2:
3497:
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3460:
3457:
3454:
3453:
3449:
3446:
3443:
3440:
3437:
3436:
3433:
3432:Tomoregulin 1
3430:
3428:
3425:
3423:
3420:
3418:
3415:
3413:
3410:
3408:
3407:Gremlin (Drm)
3405:
3403:
3400:
3398:
3395:
3393:
3390:
3388:
3385:
3383:
3380:
3378:
3375:
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2128:
2125:
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2097:
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2049:
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2027:
2021:
2018:
2015:
2014:
2011:
2008:
2005:
2004:
2001:
1998:
1996:
1993:
1990:
1989:
1986:
1985:Ascrinvacumab
1983:
1980:
1979:
1975:
1971:
1967:
1963:
1960:
1957:
1953:
1950:
1948:
1945:
1942:
1938:
1935:
1933:
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1598:
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1501:
1498:
1496:
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1478:
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1473:
1471:
1468:
1466:
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1458:
1454:
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1426:
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1404:
1401:
1400:
1398:
1396:
1391:
1385:
1382:
1378:
1375:
1373:
1370:
1368:
1365:
1363:
1360:
1359:
1358:
1355:
1351:
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1341:
1338:
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1336:
1333:
1332:
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1328:
1324:
1319:
1316:
1314:
1310:
1306:
1302:
1295:
1290:
1288:
1283:
1281:
1276:
1275:
1272:
1265:
1261:
1258:
1255:
1251:
1248:
1247:
1243:
1233:
1230:
1224:
1221:
1214:
1211:
1206:
1202:
1197:
1192:
1188:
1184:
1183:J. Biol. Chem
1180:
1173:
1171:
1169:
1165:
1160:
1156:
1151:
1146:
1141:
1136:
1132:
1128:
1124:
1120:
1116:
1112:
1105:
1102:
1097:
1093:
1088:
1083:
1079:
1075:
1071:
1067:
1063:
1056:
1053:
1048:
1044:
1039:
1034:
1030:
1026:
1021:
1016:
1012:
1008:
1004:
997:
994:
989:
985:
981:
977:
973:
969:
965:
961:
954:
951:
946:
942:
938:
934:
930:
926:
922:
918:
910:
907:
902:
898:
893:
888:
884:
880:
876:
869:
866:
861:
857:
853:
849:
845:
841:
837:
833:
826:
823:
818:
814:
810:
806:
802:
798:
794:
790:
786:
782:
775:
772:
767:
763:
758:
753:
749:
745:
741:
737:
733:
726:
723:
718:
714:
710:
706:
702:
701:10.1038/44157
698:
694:
690:
686:
682:
675:
672:
667:
663:
658:
653:
648:
643:
639:
635:
631:
624:
622:
618:
613:
609:
604:
599:
594:
589:
585:
581:
577:
570:
568:
564:
559:
555:
550:
545:
542:(2): 655–65.
541:
537:
536:Endocrinology
533:
526:
524:
522:
518:
512:
510:
508:
501:
499:
497:
493:
489:
485:
481:
478:
474:
469:
467:
463:
455:
453:
451:
447:
442:
440:
437:
433:
429:
426:
422:
418:
415:
411:
403:
398:
396:
394:
390:
382:
380:
378:
377:morphogenesis
374:
370:
365:
361:
357:
353:
345:
343:
340:
332:
330:
328:
324:
320:
316:
304:
301:
297:
294:
291:
287:
282:
278:
275:
274:
270:
267:
265:
261:
256:
253:
250:
248:
244:
241:
238:
236:
232:
229:
226:
224:
220:
217:
214:
212:
208:
205:
202:
200:
196:
192:
188:
183:
178:
167:
164:
160:
157:
154:
150:
145:
141:
138:
137:
133:
130:
128:
124:
119:
116:
113:
111:
107:
104:
101:
99:
95:
92:
89:
87:
83:
80:
77:
75:
71:
68:
65:
63:
59:
55:
51:
47:
43:
38:
33:
27:
19:
3455:
3438:
3406:
3373:
3318:
3281:
3253:
3221:Antagonists:
3220:
3119:
3096:
3064:Antagonists:
3063:
2939:
2903:Antagonists:
2902:
2834:
2806:
2795:Lerdelimumab
2790:Fresolimumab
2786:
2748:
2702:
2684:Antagonists:
2683:
2655:
2617:
2533:
2450:
2439:Lerdelimumab
2434:Fresolimumab
2430:
2388:
2350:
2318:Antagonists:
2317:
2266:
2228:
2174:
2126:
2044:
2016:
2010:Dalantercept
2006:
1991:
1981:
1910:
1766:
1232:
1223:
1213:
1186:
1182:
1122:
1118:
1104:
1069:
1065:
1055:
1010:
1007:Biomedicines
1006:
996:
963:
959:
953:
920:
916:
909:
882:
878:
868:
838:(3): 303–7.
835:
831:
825:
784:
780:
774:
739:
735:
725:
684:
680:
674:
637:
633:
583:
579:
539:
535:
505:
470:
459:
446:angiogenesis
443:
432:binding site
407:
386:
349:
339:glycoprotein
336:
314:
313:
271:
134:
53:Alt. symbols
26:
3439:Antibodies:
3402:Follistatin
3257:Ramatercept
3100:Sotatercept
2800:Metelimumab
2787:Antibodies:
2444:Metelimumab
2431:Antibodies:
2020:Disitertide
1982:Antibodies:
1802:Coreceptors
1762:Follistatin
1672:Transducers
1072:: 625–635.
917:Development
879:Development
603:11379/29323
488:polydactyly
450:endothelium
417:antagonists
410:microarrays
293:Swiss-model
185:Identifiers
156:Swiss-model
40:Identifiers
3474:Categories
3442:Stamulumab
3392:DAN (PARN)
3311:(β-glycan)
3214:Radotermin
3057:Radotermin
3037:11 (BMP11)
2993:15 (GDF9B)
2896:Radotermin
2761:10 (BMP3B)
2671:11 (BMP11)
2611:Radotermin
2569:15 (GDF9B)
2474:LY-2109761
2405:11 (BMP11)
2401:10 (BMP3B)
2300:11 (BMP11)
1879:modulators
1658:betaglycan
1393:Ligand of
1321:Ligand of
1013:(2): 301.
832:Nat. Genet
634:BMC Cancer
513:References
477:osteoblast
466:osteopenia
462:trabecular
428:carcinomas
408:Data from
289:Structures
284:Search for
258:Other data
152:Structures
147:Search for
121:Other data
3322:Avotermin
3285:AMH (MIS)
3282:Agonists:
3193:7 (BMP12)
3189:6 (BMP13)
3185:5 (BMP14)
3120:Agonists:
3029:7 (BMP12)
3025:6 (BMP13)
3021:5 (BMP14)
2940:Agonists:
2890:7 (BMP12)
2886:6 (BMP13)
2882:5 (BMP14)
2835:Agonists:
2815:LY-364947
2752:Avotermin
2749:Agonists:
2716:SB-505124
2711:SB-431542
2656:Agonists:
2597:7 (BMP12)
2593:6 (BMP13)
2589:5 (BMP14)
2534:Agonists:
2504:SB-525334
2499:SB-505124
2494:SB-431542
2469:LY-364947
2464:GW-788388
2392:Avotermin
2389:Agonists:
2364:SB-505124
2359:SB-431542
2267:Agonists:
2178:AMH (MIS)
2175:Agonists:
2071:Avotermin
2066:AMH (MIS)
2045:Agonists:
1932:Avotermin
1911:Agonists:
1078:1090-0535
1029:2227-9059
736:Genes Dev
492:β-catenin
333:Structure
319:inhibitor
240:NM_022469
199:NCBI gene
103:NM_013372
62:NCBI gene
3459:Endoglin
3367:Unsorted
3319:Ligands:
3299:Type III
2878:2 (BMP9)
2484:R-268712
1956:2 (BMP9)
1947:Cetermin
1747:Cerberus
1205:17785465
1159:17003113
1111:Chang HY
1096:31700227
1047:35203511
988:16350339
980:11290528
937:17522159
901:15201225
860:19761724
852:12808456
817:25309127
809:19229034
766:15198975
709:10524628
666:16545136
612:17077323
558:15539560
346:Function
303:InterPro
166:InterPro
56:CKTSF1B1
3456:Others:
3397:Decorin
3387:Chordin
3327:Inhibin
3224:Inhibin
3123:Activin
3067:Inhibin
2943:Activin
2906:Inhibin
2730:Type II
2706:A 83-01
2631:K-02288
2454:A 83-01
2354:A 83-01
2321:Inhibin
2270:Activin
2242:K-02288
2145:K-02288
2048:Activin
1995:K-02288
1914:Activin
1767:Gremlin
1757:Decorin
1752:Chordin
1651:TGFBR3:
1613:TGFBR2:
1558:TGFBR1:
1541:Activin
1218:270–281
1150:1578503
1127:Bibcode
1087:6817737
1038:8869528
945:7867216
789:Bibcode
781:Science
717:4372393
689:Bibcode
657:1459871
425:sarcoma
321:in the
315:Gremlin
299:Domains
247:UniProt
162:Domains
132:Chr. 15
110:UniProt
3448:TRC105
3417:Noggin
3309:TGFβR3
3111:ACVR2B
2926:ACVR2A
2740:TGFβR2
2646:ACVR1C
2524:BMPR1B
2509:SD-208
2459:D-4476
2379:TGFβR1
2257:ACVR1B
2165:BMPR1A
2035:ACVR1A
1901:ACVRL1
1887:Type I
1815:Cripto
1782:Noggin
1714:I-SMAD
1684:R-SMAD
1630:ACVR2B
1625:ACVR2A
1602:BMPR1B
1597:BMPR1A
1587:ACVRL1
1580:ACVR1C
1575:ACVR1B
1549:family
1350:TGF-β3
1345:TGF-β2
1340:TGF-β1
1266:(MeSH)
1256:(MeSH)
1203:
1157:
1147:
1094:
1084:
1076:
1045:
1035:
1027:
986:
978:
943:
935:
899:
858:
850:
815:
807:
764:
757:443518
754:
715:
707:
681:Nature
664:
654:
640:: 74.
610:
556:
421:stroma
404:Cancer
317:is an
269:Chr. 1
252:Q9H772
235:RefSeq
228:608832
190:Symbol
136:q11-13
115:O60565
98:RefSeq
91:603054
45:Symbol
3412:LTBP1
3377:BAMBI
3268:AMHR2
3238:Lefty
3204:Nodal
3081:Lefty
3052:Nodal
2930:ACVR2
2826:BMPR2
2810:DMH-2
2687:Lefty
2677:Nodal
2621:DMH-2
2335:Lefty
2311:Nodal
2232:DMH-2
2135:DMH-2
2130:DMH-1
1824:Other
1810:BAMBI
1792:THBS1
1777:LTBP1
1772:Lefty
1729:SMAD4
1723:SMAD7
1718:SMAD6
1708:SMAD9
1703:SMAD5
1698:SMAD3
1693:SMAD2
1688:SMAD1
1642:BMPR2
1637:AMHR2
1592:BMPR1
1570:ACVR1
1515:GDF15
1510:GDF11
1505:GDF10
1453:BMP15
1448:BMP10
1443:BMP8b
1438:BMP8a
1384:Nodal
1377:INHBC
1372:INHBB
1367:INHBA
1327:TGFBR
984:S2CID
941:S2CID
856:S2CID
813:S2CID
713:S2CID
580:Blood
439:YWHAH
264:Locus
216:17655
204:64388
193:GREM2
127:Locus
67:26585
48:GREM1
18:GREM1
3422:TGIF
3341:TGFβ
3272:AMHR
2767:TGFβ
2642:ALK7
2520:ALK6
2411:TGFβ
2375:ALK5
2253:ALK4
2161:ALK3
2107:TGFβ
2031:ALK2
1962:TGFβ
1897:ALK1
1874:TGFβ
1831:SARA
1787:PARN
1676:SMAD
1500:GDF9
1490:GDF7
1485:GDF6
1480:GDF5
1475:GDF3
1470:GDF2
1465:GDF1
1433:BMP7
1428:BMP6
1423:BMP5
1418:BMP4
1413:BMP3
1408:BMP2
1395:BMPR
1362:INHA
1323:ACVR
1201:PMID
1155:PMID
1092:PMID
1074:ISSN
1043:PMID
1025:ISSN
976:PMID
933:PMID
897:PMID
848:PMID
805:PMID
762:PMID
705:PMID
662:PMID
608:PMID
554:PMID
456:Bone
364:BMP4
362:and
360:BMP2
223:OMIM
211:HGNC
86:OMIM
79:2001
74:HGNC
3173:GDF
3141:BMP
3009:GDF
2961:BMP
2874:GDF
2838:BMP
2757:GDF
2659:GDF
2585:GDF
2537:BMP
2397:GDF
2288:GDF
2183:BMP
2076:BMP
1952:GDF
1937:BMP
1545:BMP
1325:or
1191:doi
1187:282
1145:PMC
1135:doi
1123:103
1082:PMC
1033:PMC
1015:doi
968:doi
964:280
925:doi
921:134
887:doi
883:131
840:doi
797:doi
785:323
752:PMC
744:doi
697:doi
685:401
652:PMC
642:doi
598:hdl
588:doi
584:109
544:doi
540:146
448:on
414:BMP
273:q43
3476::
3351:,
3347:,
3333:,
3244:,
3230:,
3191:,
3187:,
3183:,
3179:,
3155:,
3151:,
3147:,
3135:AB
3133:,
3129:,
3087:,
3073:,
3041:15
3039:,
3035:,
3031:,
3027:,
3023:,
3019:,
3015:,
2991:,
2989:8B
2987:,
2985:8A
2983:,
2979:,
2975:,
2971:,
2967:,
2955:AB
2953:,
2949:,
2912:,
2888:,
2884:,
2880:,
2858:10
2856:,
2852:,
2848:,
2844:,
2777:,
2773:,
2693:,
2669:,
2665:,
2605:15
2603:,
2599:,
2595:,
2591:,
2567:,
2565:8B
2563:,
2561:8A
2559:,
2555:,
2551:,
2547:,
2543:,
2421:,
2417:,
2403:,
2341:,
2327:,
2298:,
2294:,
2282:AB
2280:,
2276:,
2211:8B
2209:,
2207:8A
2205:,
2201:,
2197:,
2193:,
2189:,
2117:,
2113:,
2096:8B
2094:,
2092:8A
2090:,
2086:,
2082:,
2060:AB
2058:,
2054:,
1972:,
1968:,
1941:10
1926:AB
1924:,
1920:,
1547:,
1543:,
1303::
1237:23
1199:.
1185:.
1181:.
1167:^
1153:.
1143:.
1133:.
1121:.
1117:.
1090:.
1080:.
1070:25
1068:.
1064:.
1041:.
1031:.
1023:.
1011:10
1009:.
1005:.
982:.
974:.
962:.
939:.
931:.
919:.
895:.
881:.
877:.
854:.
846:.
836:34
834:.
811:.
803:.
795:.
783:.
760:.
750:.
740:18
738:.
734:.
711:.
703:.
695:.
683:.
660:.
650:.
636:.
632:.
620:^
606:.
596:.
582:.
578:.
566:^
552:.
538:.
534:.
520:^
379:.
3355:)
3353:3
3349:2
3345:1
3343:(
3337:)
3335:B
3331:A
3329:(
3274:)
3270:(
3248:)
3246:2
3242:1
3240:(
3234:)
3232:B
3228:A
3226:(
3195:)
3181:3
3177:1
3175:(
3159:)
3157:7
3153:6
3149:4
3145:2
3143:(
3137:)
3131:B
3127:A
3125:(
3091:)
3089:2
3085:1
3083:(
3077:)
3075:B
3071:A
3069:(
3043:)
3033:9
3017:3
3013:1
3011:(
2995:)
2981:7
2977:6
2973:5
2969:4
2965:2
2963:(
2957:)
2951:B
2947:A
2945:(
2932:)
2928:(
2916:)
2914:B
2910:A
2908:(
2892:)
2876:(
2860:)
2854:7
2850:6
2846:4
2842:2
2840:(
2781:)
2779:3
2775:2
2771:1
2769:(
2763:)
2759:(
2697:)
2695:2
2691:1
2689:(
2673:)
2667:3
2663:1
2661:(
2648:)
2644:(
2607:)
2601:9
2587:(
2571:)
2557:7
2553:6
2549:5
2545:4
2541:2
2539:(
2526:)
2522:(
2425:)
2423:3
2419:2
2415:1
2413:(
2407:)
2399:(
2381:)
2377:(
2345:)
2343:2
2339:1
2337:(
2331:)
2329:B
2325:A
2323:(
2302:)
2296:3
2292:1
2290:(
2284:)
2278:B
2274:A
2272:(
2259:)
2255:(
2213:)
2203:7
2199:6
2195:5
2191:4
2187:2
2185:(
2167:)
2163:(
2121:)
2119:3
2115:2
2111:1
2109:(
2098:)
2088:7
2084:6
2080:5
2078:(
2062:)
2056:B
2052:A
2050:(
2037:)
2033:(
1976:)
1974:3
1970:2
1966:1
1964:(
1958:)
1954:(
1943:)
1939:(
1928:)
1922:B
1918:A
1916:(
1903:)
1899:(
1865:e
1858:t
1851:v
1725:)
1716:(
1710:)
1686:(
1674:/
1551:)
1539:(
1293:e
1286:t
1279:v
1207:.
1193::
1161:.
1137::
1129::
1098:.
1049:.
1017::
990:.
970::
947:.
927::
903:.
889::
862:.
842::
819:.
799::
791::
768:.
746::
719:.
699::
691::
668:.
644::
638:6
614:.
600::
590::
560:.
546::
20:)
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