435:, which later will form the mesenchyme in the head. The somitomeres organize into somites which grow in pairs. In the fourth week, the somites lose their organization and cover the notochord and spinal cord to form the backbone. In the fifth week, there are 4 occipital somites, 8 cervical, 12 thoracic, 5 lumbar, 5 sacral and 8 to 10 coccygeal that will form the axial skeleton. Somitic derivatives are determined by local signaling between adjacent embryonic tissues, in particular the neural tube, notochord, surface ectoderm and the somitic compartments themselves. The correct specification of the deriving tissues, skeletal, cartilage, endothelia and connective tissue is achieved by a sequence of morphogenic changes of the paraxial mesoderm, leading to the three transitory somitic compartments: dermomyotome, myotome and sclerotome. These structures are specified from dorsal to ventral and from medial to lateral. Each somite will form its own sclerotome that will differentiate into the tendon cartilage and bone component. Its myotome will form the muscle component and the dermatome that will form the dermis of the back. The myotome and dermatome have a nerve component.
416:. The prechordal cells migrate to the midline to form the notochordal plate. The chordamesoderm is the central region of trunk mesoderm. This forms the notochord, which induces the formation of the neural tube, and establishes the anterior-posterior body axis. The notochord extends beneath the neural tube from the head to the tail. The mesoderm moves to the midline until it covers the notochord. When the mesoderm cells proliferate, they form the paraxial mesoderm. In each side, the mesoderm remains thin, and is known as the lateral plate. The intermediate mesoderm lies between the paraxial mesoderm and the lateral plate. Between days 13 and 15, the proliferation of extraembryonic mesoderm, primitive streak, and embryonic mesoderm take place. The notochord process occurs between days 15 and 17. Eventually, the development of the notochord canal and the axial canal takes place between days 17 and 19, when the first three somites are formed.
479:
131:
25:
444:
tube activate the protein SHH, which helps the somite to form its sclerotome. The cells of the sclerotome express the protein PAX1 that induces the cartilage and bone formation. The neural tube activates the protein WNT1 that expresses PAX 2 so the somite creates the myotome and dermatome. Finally, the neural tube also secretes neurotrophin 3, so that the somite creates the dermis. Boundaries for each somite are regulated by
359:. The intermediate mesoderm connects the paraxial mesoderm with the lateral plate. Eventually it differentiates into urogenital structures that consist of the kidneys, gonads, their associated ducts, and the adrenal cortex. The lateral plate mesoderm gives rise to the heart, blood vessels, and blood cells of the circulatory system, as well as to the mesodermal components of the limbs.
461:
decisions. Human embryonic stem cells for example have the potential to produce all of the cells in the body and they are able to self-renew indefinitely so they can be used for a large-scale production of therapeutic cell lines. They are also able to remodel and contract collagen and were induced to express muscle actin. This shows that these cells are multipotent cells.
495:
covers the yolk sac. The two layers cover the intraembryonic cavity. The parietal layer, together with overlying ectoderm, forms the lateral body wall folds. The visceral layer forms the walls of the gut tube. Mesoderm cells of the parietal layer form the mesothelial membranes or serous membranes, which line the peritoneal, pleural, and pericardial cavities.
143:
412:
the hypoblast and create the endoderm, and other cells migrate between the endoderm and the epiblast to create the mesoderm. The remaining cells form the ectoderm. After that, the epiblast and the hypoblast establish contact with the extraembryonic mesoderm until they cover the yolk sac and amnion. They move onto either side of the
443:
Surrounding structures such as the notochord, neural tube, epidermis and lateral plate mesoderm send signals for somite differentiation
Notochord protein accumulates in presomitic mesoderm destined to form the next somite and then decreases as that somite is established. The notochord and the neural
469:
The intermediate mesoderm connects the paraxial mesoderm with the lateral plate mesoderm, and differentiates into urogenital structures. In upper thoracic and cervical regions, this forms the nephrotomes. In caudal regions, it forms the nephrogenic cord. It also helps to develop the excretory units
494:
splits into the parietal (somatic) and visceral (splanchnic) layers. The formation of these layers starts with the appearance of intercellular cavities. The somatic layer depends upon a continuous layer with mesoderm that covers the amnion. The splanchnic layer depends upon a continuous layer that
460:
and zebrafish have analyzed the factors of this development and how they interact in signaling and transcription. However, there are still some doubts in how the prospective mesodermal cells integrate the various signals they receive and how they regulate their morphogenic behaviours and cell-fate
411:
on the surface of the epiblast. The cells of the layers move between the epiblast and the hypoblast, and begin to spread laterally and cranially. The cells of the epiblast move toward the primitive streak and slip beneath it, in a process called "invagination". Some of the migrating cells displace
294:
is protected from degradation by GSK-3. Beta-catenin acts as a co-factor that alters the activity of the transcription factor tcf-3 from repressing to activating, which initiates the synthesis of gene products critical for mesoderm differentiation and gastrulation. Furthermore, mesoderm has the
430:
During the third week, the paraxial mesoderm is organized into segments. If they appear in the cephalic region and grow with cephalocaudal direction, they are called somitomeres. If they appear in the cephalic region but establish contact with the neural plate, they are known as
362:
Some of the mesoderm derivatives include the muscle (smooth, cardiac, and skeletal), the muscles of the tongue (occipital somites), the pharyngeal arches muscle (muscles of mastication, muscles of facial expressions), connective tissue, the dermis and subcutaneous layer of the
452:
in vertebrates. The bilaterally symmetric body plan of vertebrate embryos is obvious in somites and their derivates, such as the vertebral column. Therefore, asymmetric somite formation correlates with a left-right desynchronization of the segmentation oscillations.
351:(cartilage and bone), and dermatomes (subcutaneous tissue of the skin). Signals for somite differentiation are derived from surroundings structures, including the notochord,
188:
519:
448:
and a combination of FGF8 and WNT3a. So retinoic acid is an endogenous signal that maintains the bilateral synchrony of mesoderm segmentation and controls
1079:
677:
1130:
862:
1100:
1063:
1040:
1017:
990:
967:
622:
569:
1158:
89:
108:
61:
68:
1485:
46:
407:
creates a mesodermal layer between the endoderm and the ectoderm. This process begins with the formation of a
1116:
718:
Yusuf, Faisal (2006). "The eventful somite: Patterning, fate determination and cell division in the somite".
183:
75:
1443:
1388:
1167:
881:"Human embryonic stem cell-derived mesoderm-like epithelium transitions to mesenchymal progenitor cells"
1136:
287:
814:
Vermot, J.; Gallego Llamas, J.; Fraulob, V.; Niederreither, K.; Chambon, P.; Dollé, P. (April 2005).
57:
42:
1448:
1360:
1245:
1151:
171:
1212:
279:
35:
1433:
1332:
491:
328:
195:
1051:
1001:
978:
606:
1495:
1438:
1428:
1272:
1028:
951:
815:
747:
324:
308:
272:
225:
879:
Boyd, N.L.; Robbins KR, K.R.; Dhara SK, S.K.; West FD, F.D.; Stice SL., S.L. (August 2009).
1480:
830:
561:
554:
8:
1490:
1182:
1144:
260:
834:
1073:
1006:
905:
880:
854:
816:"Retinoic acid controls the bilateral symmetry of somite formation in the mouse embryo"
791:
766:
739:
611:
449:
1120:
1089:
290:. The position of the organizing center is in turn determined by the regions in which
82:
1475:
1411:
1197:
1187:
1126:
1096:
1059:
1036:
1013:
986:
963:
910:
846:
796:
731:
618:
565:
483:
425:
384:
320:
767:"Mechanisms of retinoic acid signalling and its roles in organ and limb development"
743:
1350:
1313:
1294:
900:
892:
858:
838:
786:
778:
723:
413:
408:
307:
The mesoderm is one of the three germinal layers that appears in the third week of
1227:
478:
380:
176:
751:
1406:
1304:
673:
316:
130:
896:
727:
1469:
1355:
1299:
514:
509:
445:
376:
283:
842:
552:
Ruppert, E.E.; Fox, R.S.; Barnes, R.D. (2004). "Introduction to
Bilateria".
1370:
1325:
1320:
914:
850:
800:
735:
504:
404:
312:
296:
291:
221:
1365:
1290:
1232:
372:
352:
248:
244:
1166:
813:
1421:
1308:
1282:
1222:
1217:
1202:
976:
591:
589:
587:
585:
583:
581:
348:
340:
336:
256:
240:
217:
201:
1254:
432:
388:
356:
332:
782:
578:
24:
1380:
1342:
1259:
233:
229:
438:
295:
capability to induce the growth of other structures, such as the
278:
The mesoderm differentiates from the rest of the embryo through
1416:
1207:
1192:
392:
344:
268:
252:
878:
264:
368:
364:
1049:
142:
1008:
Bioregenerative engineering: principles and applications
977:
Kenderew, John
Cowdery; Lawrence, Eleanor, eds. (1994).
604:
482:
Cytology of normal mesothelium, with typical features.
674:"Paraxial Mesoderm: The Somites and Their Derivatives"
758:
520:
List of human cell types derived from the germ layers
315:. There are four important components, which are the
1086:
607:"Vertebrate Mesoderm Induction: From Frogs to Mice"
347:in occipital and caudal segments, and give rise to
255:. Mesoderm forms the muscles in a process known as
49:. Unsourced material may be challenged and removed.
1088:
1026:
1005:
662:(4th ed.). Lippincott Williams & Wilkins.
610:
553:
764:
598:
595:Langman's Medical Embryology, 11th edition. 2010.
547:
545:
543:
541:
539:
537:
535:
1467:
1087:Sherman, Lawrence S.; et al., eds. (2001).
657:
551:
263:(length-wise partitions); and forms part of the
395:of teeth, the kidneys, and the adrenal cortex.
642:
532:
439:Molecular regulation of somite differentiation
1152:
926:
924:
678:National Center for Biotechnology Information
1078:: CS1 maint: multiple names: authors list (
949:
713:
711:
709:
697:
638:
636:
634:
271:). Myogenesis is specifically a function of
999:
960:Organization of the early vertebrate embryo
930:
1159:
1145:
1095:(3rd ed.). Elsevier Health Sciences.
958:. In Zagris, Nikolas; et al. (eds.).
921:
141:
129:
16:Middle germ layer of embryonic development
952:"The formation of mesoderm and muscle in
904:
790:
706:
691:
651:
647:(ninth ed.). US: Sinauer Associates.
631:
473:
109:Learn how and when to remove this message
1027:McGeady, Thomas A.; et al. (2006).
666:
605:Kimelman, D. & Bjornson, C. (2004).
477:
464:
403:During the third week, a process called
311:. It is formed through a process called
228:of most animals. The outer layer is the
331:. The axial mesoderm gives rise to the
299:, the precursor to the nervous system.
1468:
1052:"Early Embryonic Mesoderm Development"
1029:"Establishment of the Basic Body Plan"
985:. John Wiley & Sons. p. 541.
765:Cunningham, T.J.; Duester, G. (2015).
560:(7th ed.). Brooks/Cole. pp.
470:of the urinary system and the gonads.
1140:
983:The encyclopedia of molecular biology
717:
419:
259:, septa (cross-wise partitions) and
47:adding citations to reliable sources
18:
1050:Pappaioannou, Virginia, E. (2004).
1002:"Early Embryonic Organ Development"
13:
943:
613:Gastrulation: from cells to embryo
335:. The paraxial mesoderm forms the
14:
1507:
1110:
216:is the middle layer of the three
1058:. Gulf Professional Publishing.
1056:Handbook of stem cells, Volume 1
868:from the original on 2022-10-09.
23:
1054:. In Lanza, Robert Paul (ed.).
872:
807:
371:and cartilage, dura mater, the
343:of the head, and organize into
34:needs additional citations for
609:. In Stern, Claudio D. (ed.).
398:
282:, after which the mesoderm is
147:Section through a human embryo
135:Tissues derived from mesoderm.
1:
525:
302:
232:, and the inner layer is the
933:Textbook of human embryology
7:
1444:Splanchnopleuric mesenchyme
1389:Splanchnopleuric mesenchyme
1168:Human embryonic development
617:. CSHL Press. p. 363.
498:
10:
1512:
885:Tissue Engineering. Part A
423:
1397:
1379:
1341:
1281:
1268:
1241:
1175:
1012:. John Wiley & Sons.
897:10.1089/ten.tea.2008.0351
728:10.1007/s00429-006-0119-8
700:Color atlas of embryology
658:Dudek, Ronald W. (2009).
226:development of the embryo
194:
182:
170:
165:
157:
152:
140:
128:
123:
1449:Somatopleuric mesenchyme
1361:Somatopleuric mesenchyme
1170:in the first three weeks
771:Nat. Rev. Mol. Cell Biol
843:10.1126/science.1108363
643:Scott, Gilbert (2010).
329:lateral plate mesoderms
280:intercellular signaling
1333:Regional specification
722:. 211 Suppl 1: 21–30.
720:Anatomy and Embryology
660:High-yield. Embryology
492:lateral plate mesoderm
487:
474:Lateral plate mesoderm
196:Anatomical terminology
1486:Developmental biology
1439:Intraembryonic coelom
1033:Veterinary embryology
950:Gurdon, J.B. (1995).
935:. I.K. International.
698:Drew, Ulrich (1993).
645:Developmental biology
481:
465:Intermediate mesoderm
339:, which give rise to
309:embryonic development
220:that develops during
1127:Embryology at Temple
1000:Liu, Shu Q. (2007).
979:"Mesoderm Induction"
931:Kumar, Rani (2008).
556:Invertebrate Zoology
267:(the rest being the
251:. Mesothelium lines
43:improve this article
1035:. Wiley-Blackwell.
835:2005Sci...308..563V
239:The mesoderm forms
1117:Embryology at UNSW
488:
456:Many studies with
450:bilateral symmetry
224:in the very early
1463:
1462:
1459:
1458:
1188:Oocyte activation
1102:978-0-443-06583-5
1065:978-0-12-436642-8
1042:978-1-4051-1147-8
1019:978-0-471-70907-7
992:978-0-632-02182-6
969:978-0-306-45132-4
829:(5721): 563–566.
702:. German: Thieme.
624:978-0-87969-707-5
571:978-0-03-025982-1
426:Paraxial mesoderm
420:Paraxial mesoderm
385:white blood cells
288:organizing center
210:
209:
205:
119:
118:
111:
93:
1503:
1351:Surface ectoderm
1314:Primitive groove
1295:Primitive streak
1279:
1278:
1161:
1154:
1147:
1138:
1137:
1106:
1094:
1091:Human embryology
1083:
1077:
1069:
1046:
1023:
1011:
996:
973:
937:
936:
928:
919:
918:
908:
891:(8): 1897–1907.
876:
870:
869:
867:
820:
811:
805:
804:
794:
762:
756:
755:
715:
704:
703:
695:
689:
688:
686:
684:
670:
664:
663:
655:
649:
648:
640:
629:
628:
616:
602:
596:
593:
576:
575:
559:
549:
414:prechordal plate
409:primitive streak
202:edit on Wikidata
199:
145:
133:
121:
120:
114:
107:
103:
100:
94:
92:
51:
27:
19:
1511:
1510:
1506:
1505:
1504:
1502:
1501:
1500:
1466:
1465:
1464:
1455:
1393:
1375:
1337:
1270:
1264:
1243:
1237:
1228:Inner cell mass
1171:
1165:
1131:EMBIII97/sld039
1113:
1103:
1071:
1070:
1066:
1043:
1020:
993:
970:
946:
944:Further reading
941:
940:
929:
922:
877:
873:
865:
818:
812:
808:
783:10.1038/nrm3932
763:
759:
716:
707:
696:
692:
682:
680:
672:
671:
667:
656:
652:
641:
632:
625:
603:
599:
594:
579:
572:
550:
533:
528:
501:
476:
467:
441:
428:
422:
401:
381:red blood cells
305:
206:
148:
136:
115:
104:
98:
95:
52:
50:
40:
28:
17:
12:
11:
5:
1509:
1499:
1498:
1493:
1488:
1483:
1478:
1461:
1460:
1457:
1456:
1454:
1453:
1452:
1451:
1446:
1441:
1431:
1426:
1425:
1424:
1419:
1409:
1407:Axial mesoderm
1403:
1401:
1395:
1394:
1392:
1391:
1385:
1383:
1377:
1376:
1374:
1373:
1368:
1363:
1358:
1353:
1347:
1345:
1339:
1338:
1336:
1335:
1330:
1329:
1328:
1318:
1317:
1316:
1311:
1305:Primitive node
1302:
1287:
1285:
1276:
1266:
1265:
1263:
1262:
1257:
1251:
1249:
1239:
1238:
1236:
1235:
1230:
1225:
1220:
1215:
1210:
1205:
1200:
1195:
1190:
1185:
1179:
1177:
1173:
1172:
1164:
1163:
1156:
1149:
1141:
1135:
1134:
1124:
1112:
1111:External links
1109:
1108:
1107:
1101:
1084:
1064:
1047:
1041:
1024:
1018:
997:
991:
974:
968:
945:
942:
939:
938:
920:
871:
806:
777:(2): 110–123.
757:
705:
690:
665:
650:
630:
623:
597:
577:
570:
530:
529:
527:
524:
523:
522:
517:
512:
507:
500:
497:
484:Wright's stain
475:
472:
466:
463:
440:
437:
424:Main article:
421:
418:
400:
397:
304:
301:
208:
207:
198:
192:
191:
186:
180:
179:
174:
168:
167:
163:
162:
159:
155:
154:
150:
149:
146:
138:
137:
134:
126:
125:
117:
116:
31:
29:
22:
15:
9:
6:
4:
3:
2:
1508:
1497:
1494:
1492:
1489:
1487:
1484:
1482:
1479:
1477:
1474:
1473:
1471:
1450:
1447:
1445:
1442:
1440:
1437:
1436:
1435:
1434:Lateral plate
1432:
1430:
1427:
1423:
1420:
1418:
1415:
1414:
1413:
1410:
1408:
1405:
1404:
1402:
1400:
1396:
1390:
1387:
1386:
1384:
1382:
1378:
1372:
1369:
1367:
1364:
1362:
1359:
1357:
1356:Neuroectoderm
1354:
1352:
1349:
1348:
1346:
1344:
1340:
1334:
1331:
1327:
1324:
1323:
1322:
1319:
1315:
1312:
1310:
1306:
1303:
1301:
1300:Primitive pit
1298:
1297:
1296:
1292:
1289:
1288:
1286:
1284:
1280:
1277:
1274:
1267:
1261:
1258:
1256:
1253:
1252:
1250:
1247:
1240:
1234:
1231:
1229:
1226:
1224:
1221:
1219:
1216:
1214:
1211:
1209:
1206:
1204:
1201:
1199:
1196:
1194:
1191:
1189:
1186:
1184:
1183:Fertilization
1181:
1180:
1178:
1174:
1169:
1162:
1157:
1155:
1150:
1148:
1143:
1142:
1139:
1133:
1132:
1128:
1125:
1123:
1122:
1118:
1115:
1114:
1104:
1098:
1093:
1092:
1085:
1081:
1075:
1067:
1061:
1057:
1053:
1048:
1044:
1038:
1034:
1030:
1025:
1021:
1015:
1010:
1009:
1003:
998:
994:
988:
984:
980:
975:
971:
965:
961:
957:
955:
948:
947:
934:
927:
925:
916:
912:
907:
902:
898:
894:
890:
886:
882:
875:
864:
860:
856:
852:
848:
844:
840:
836:
832:
828:
824:
817:
810:
802:
798:
793:
788:
784:
780:
776:
772:
768:
761:
753:
749:
745:
741:
737:
733:
729:
725:
721:
714:
712:
710:
701:
694:
679:
675:
669:
661:
654:
646:
639:
637:
635:
626:
620:
615:
614:
608:
601:
592:
590:
588:
586:
584:
582:
573:
567:
563:
558:
557:
548:
546:
544:
542:
540:
538:
536:
531:
521:
518:
516:
515:Triploblastic
513:
511:
510:Organogenesis
508:
506:
503:
502:
496:
493:
485:
480:
471:
462:
459:
454:
451:
447:
446:retinoic acid
436:
434:
427:
417:
415:
410:
406:
396:
394:
390:
386:
382:
378:
377:blood vessels
374:
370:
366:
360:
358:
354:
350:
346:
342:
338:
334:
330:
326:
322:
318:
314:
310:
300:
298:
293:
289:
285:
281:
276:
274:
270:
266:
262:
258:
254:
250:
246:
242:
237:
235:
231:
227:
223:
219:
215:
203:
197:
193:
190:
187:
185:
181:
178:
175:
173:
169:
164:
160:
156:
151:
144:
139:
132:
127:
122:
113:
110:
102:
91:
88:
84:
81:
77:
74:
70:
67:
63:
60: –
59:
55:
54:Find sources:
48:
44:
38:
37:
32:This article
30:
26:
21:
20:
1496:Gastrulation
1429:Intermediate
1398:
1371:Neural crest
1326:Gastrulation
1129:
1121:Notes/skmus6
1119:
1090:
1055:
1032:
1007:
982:
962:. Springer.
959:
953:
932:
888:
884:
874:
826:
822:
809:
774:
770:
760:
719:
699:
693:
681:. Retrieved
668:
659:
653:
644:
612:
600:
555:
505:Histogenesis
489:
468:
457:
455:
442:
429:
405:gastrulation
402:
361:
325:intermediate
313:gastrulation
306:
297:neural plate
292:beta-catenin
277:
249:coelomocytes
238:
222:gastrulation
213:
211:
105:
96:
86:
79:
72:
65:
53:
41:Please help
36:verification
33:
1481:Germ layers
1366:Neurulation
1291:Archenteron
1283:Germ layers
1233:Trophoblast
399:Development
373:endothelium
353:neural tube
349:sclerotomes
337:somitomeres
261:mesenteries
245:mesothelium
218:germ layers
166:Identifiers
1491:Embryology
1470:Categories
1422:Somitomere
1309:Blastopore
1273:Trilaminar
1223:Blastocyst
1218:Blastocoel
1213:Cavitation
1203:Blastomere
526:References
433:neuromeres
341:mesenchyme
303:Definition
273:mesenchyme
257:myogenesis
241:mesenchyme
69:newspapers
58:"Mesoderm"
1255:Hypoblast
1246:Bilaminar
1074:cite book
752:212010706
683:April 15,
389:microglia
357:epidermis
333:notochord
284:polarized
1476:Mesoderm
1412:Paraxial
1399:Mesoderm
1381:Endoderm
1343:Ectoderm
1321:Gastrula
1260:Epiblast
1198:Cleavage
915:19196144
863:Archived
851:15731404
801:25560970
748:ProQuest
744:24633902
736:17024302
499:See also
321:paraxial
234:endoderm
230:ectoderm
214:mesoderm
124:Mesoderm
99:May 2022
1269:Week 3
1242:Week 2
954:Xenopus
906:2792108
859:5713738
831:Bibcode
823:Science
792:4636111
562:217–218
458:Xenopus
345:somites
269:gametes
253:coeloms
177:D008648
153:Details
83:scholar
1417:Somite
1208:Morula
1193:Zygote
1176:Week 1
1099:
1062:
1039:
1016:
989:
966:
913:
903:
857:
849:
799:
789:
750:
742:
734:
621:
568:
393:dentin
391:, the
355:, and
327:, and
286:by an
265:gonads
85:
78:
71:
64:
56:
866:(PDF)
855:S2CID
819:(PDF)
740:S2CID
317:axial
200:[
189:69072
90:JSTOR
76:books
1097:ISBN
1080:link
1060:ISBN
1037:ISBN
1014:ISBN
987:ISBN
964:ISBN
911:PMID
847:PMID
797:PMID
732:PMID
685:2013
619:ISBN
566:ISBN
490:The
369:bone
365:skin
247:and
212:The
172:MeSH
158:Days
62:news
901:PMC
893:doi
839:doi
827:308
787:PMC
779:doi
724:doi
375:of
319:,
184:FMA
45:by
1472::
1076:}}
1072:{{
1031:.
1004:.
981:.
923:^
909:.
899:.
889:15
887:.
883:.
861:.
853:.
845:.
837:.
825:.
821:.
795:.
785:.
775:16
773:.
769:.
746:.
738:.
730:.
708:^
676:.
633:^
580:^
564:.
534:^
387:,
383:,
379:,
367:,
323:,
275:.
243:,
236:.
161:16
1307:/
1293:/
1275:)
1271:(
1248:)
1244:(
1160:e
1153:t
1146:v
1105:.
1082:)
1068:.
1045:.
1022:.
995:.
972:.
956:"
917:.
895::
841::
833::
803:.
781::
754:.
726::
687:.
627:.
574:.
486:.
204:]
112:)
106:(
101:)
97:(
87:·
80:·
73:·
66:·
39:.
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.