190:
differences between CCs and MGCs. CCs primarily support growth and development of the oocyte whereas MGCs primarily serve an endocrine function and support the growth of the follicle. Cumulus cells aid in oocyte development and show higher expression of SLC38A3, a transporter for amino acids, and Aldoa, Eno1, Ldh1, Pfkp, Pkm2, and Tpi1, enzymes responsible for glycolysis. MGCs are more steroidogenically active and have higher levels of mRNA expression of steroidogenic enzymes such as cytochrome P450. MGCs produce an increasing amount of estrogen which leads to the
20:
150:
172:
The embryological origin of granulosa cells remains controversial. In the 1970s, evidence emerged that the first cells to make contact with the oogonia were of mesonephric origin. It was suggested that mesonephric cells already closely associated with the oogonia proliferated throughout development
189:
Cumulus cells (CC) surround the oocyte. They provide nutrients to the oocyte and influence the development of the oocyte in a paracrine fashion. Mural granulosa cells (MGC) line the follicular wall and surround the fluid-filled antrum. The oocyte secretes factors that determine the functional
181:(i.e., epithelial cells from the presumptive surface epithelium of the ovary). In 2013, it was proposed that both granulosa cells and the ovarian surface epithelial cells are instead derived from a precursor cell called gonadal-ridge epithelial-like cell.
194:. Following the LH surge, cumulus cells undergo cumulus expansion, in which they proliferate at a ten-fold higher rate than MGCs in response to FSH. During expansion CCs also produce a mucified matrix required for ovulation.
215:(number of cells per volume of culture medium) plays a critical role for the differentiation. A lower plating density makes granulosa cells exhibit estrogen production, while a higher plating density makes them appear as
691:
Chen L, Russell PT, Larsen WJ (January 1993). "Functional significance of cumulus expansion in the mouse: roles for the preovulatory synthesis of hyaluronic acid within the cumulus mass".
291:
Garzo, V. G.; Dorrington, J. H. (1984). "Aromatase activity in human granulosa cells during follicular development and the modulation by follicle-stimulating hormone and insulin".
586:
Li R, Norman RJ, Armstrong DT, Gilchrist RB (September 2000). "Oocyte-secreted factor(s) determine functional differences between bovine mural granulosa cells and cumulus cells".
551:
Eppig JJ, Pendola FL, Wigglesworth K, Pendola JK (August 2005). "Mouse oocytes regulate metabolic cooperativity between granulosa cells and oocytes: amino acid transport".
656:
Khamsi F, Roberge S (September 2001). "Granulosa cells of the cumulus oophorus are different from mural granulosa cells in their response to gonadotrophins and IGF-I".
267:
272:
865:
158:
141:. The progesterone may maintain a potential pregnancy and causes production of a thick cervical mucus that inhibits sperm entry into the uterus.
492:
Hummitzsch, K; Irving-Rodgers, HF; Hatzirodos, N; Bonner, W; Sabatier, L; Reinhardt, DP; Sado, Y; Ninomiya, Y; Wilhelm, D; Rodgers, RJ (2013).
1293:
1262:
1283:
367:
Upadhyay S, Zamboni L (1982). "Preliminary observations on the role of the mesonephros in the development of the adrenal cortex".
1288:
858:
1257:
728:"Cell plating density alters the ratio of estrogenic to progestagenic enzyme gene expression in cultured granulosa cells"
851:
621:
Diaz FJ, Wigglesworth K, Eppig JJ (April 2007). "Oocytes determine cumulus cell lineage in mouse ovarian follicles".
236:
772:"Increased DNA damage and repair deficiency in granulosa cells are associated with ovarian aging in rhesus monkey"
830: – Histology Learning System at Boston University — "Female Reproductive System: ovary, membrana granulosa"
824: – Histology Learning System at Boston University — "Female Reproductive System: ovary, cumulus oophorus"
1186:
106:
1047:
243:
declines with age. These changes at the DNA level in granulosa cells may contribute to ovarian aging.
910:
837:
173:
to form the granulosa cell layer. Recently, this hypothesis has been challenged with some thorough
1019:
930:
177:. Sawyer et al. hypothesized that in sheep most of the granulosa cells develop from cells of the
1042:
153:
Section of vesicular ovarian follicle of a cat. X 50. Membrana granulosa labeled at upper left.
72:
252:
134:
827:
821:
1245:
1235:
833:
505:
8:
1024:
977:
969:
509:
942:
892:
796:
771:
528:
493:
474:
392:
344:
327:
262:
1168:
1034:
947:
920:
801:
749:
744:
727:
708:
673:
638:
603:
568:
533:
466:
384:
349:
308:
304:
220:
396:
1219:
1084:
1079:
1074:
959:
952:
925:
875:
791:
783:
770:
Zhang D, Zhang X, Zeng M, Yuan J, Liu M, Yin Y, Wu X, Keefe DL, Liu L (July 2015).
739:
700:
665:
630:
595:
560:
523:
513:
478:
456:
423:
376:
339:
300:
126:
83:
79:
75:
843:
564:
1102:
1094:
987:
982:
900:
883:
518:
212:
130:
91:
461:
444:
23:
Pig oocyte surrounded by granulosa cells. Fluorescence microscopy, colored with
1066:
997:
915:
905:
599:
232:
787:
1277:
1156:
1146:
1057:
937:
166:
1240:
1206:
1196:
1134:
805:
753:
704:
677:
642:
607:
572:
537:
470:
428:
411:
216:
202:
138:
40:
712:
669:
388:
380:
353:
312:
109:(FSH) from the anterior pituitary, stimulating granulosa cells to convert
1214:
1129:
412:"The role of the mesonephros in the development of the sheep fetal ovary"
178:
114:
98:
491:
1191:
1176:
634:
240:
443:
Sawyer H, Smith P, Heath D, Juengel J, Wakefield S, McNatty K (2002).
1181:
992:
257:
174:
122:
118:
110:
184:
19:
1252:
1124:
207:
191:
44:
445:"Formation of ovarian follicles during fetal development in sheep"
1011:
494:"A new model of development of the mammalian ovary and follicles"
162:
97:
The major functions of granulosa cells include the production of
101:, as well as myriad growth factors thought to interact with the
550:
105:
during its development. The sex steroid production begins with
102:
87:
60:
52:
48:
328:"Histogenesis and organogenesis of the gonad in human embryos"
1141:
1116:
56:
149:
24:
585:
239:
increase in granulosa cells with age, and the ability to
133:. However, after ovulation the granulosa cells turn into
144:
442:
416:
Annales de
Biologie Animale, Biochimie, et Biophysique
47:
that is closely associated with the developing female
725:
620:
268:
List of human cell types derived from the germ layers
409:
873:
273:
List of distinct cell types in the adult human body
769:
159:development of the urinary and reproductive organs
82:), granulosa cells advance to form a multilayered
690:
185:Cumulus cells (CC) vs mural granulosa cells (MGC)
1275:
726:Portela VM, Zamberlam G, Price CA (April 2010).
366:
290:
859:
655:
410:Zamboni, L.; Bezard, J.; Mauleon, P. (1979).
293:American Journal of Obstetrics and Gynecology
765:
763:
1263:List of human endocrine organs and actions
866:
852:
836:at the U.S. National Library of Medicine
795:
760:
743:
527:
517:
460:
427:
343:
66:
148:
18:
325:
1276:
847:
145:Embryology of ovarian granulosa cells
78:, and later in follicle development (
205:of granulosa cells can be performed
13:
14:
1305:
1294:Mammal female reproductive system
815:
745:10.1016/j.fertnstert.2009.01.151
226:
1284:Steroid hormone secreting cells
719:
684:
649:
197:
16:Mammal reproductive system cell
614:
579:
544:
485:
436:
403:
360:
319:
284:
1:
1289:Human female endocrine system
565:10.1095/biolreprod.105.041798
278:
519:10.1371/journal.pone.0055578
305:10.1016/0002-9378(84)90769-5
107:follicle-stimulating hormone
7:
462:10.1095/biolreprod66.4.1134
246:
10:
1310:
600:10.1095/biolreprod63.3.839
165:become invaginated in the
1228:
1205:
1167:
1115:
1093:
1065:
1056:
1033:
1010:
968:
891:
882:
828:Histology image: 14808loa
822:Histology image: 18404loa
788:10.1007/s10815-015-0483-5
838:Medical Subject Headings
237:DNA double-strand breaks
94:(or Graafian) follicle.
931:Somatomammotrophic cell
90:in the preovulatory or
705:10.1002/mrd.1080340114
241:repair such DNA breaks
154:
135:granulosa lutein cells
67:Structure and function
28:
776:J Assist Reprod Genet
670:10.1677/joe.0.1700565
381:10.1002/ar.1092020112
253:Granulosa cell tumour
152:
22:
1246:Organ of Zuckerkandl
1236:Enteroendocrine cell
429:10.1051/rnd:19790801
1025:Parafollicular cell
510:2013PLoSO...855578H
1169:Islets of pancreas
943:Corticotropic cell
635:10.1242/jcs.000968
263:Membrana granulosa
221:theca lutein cells
155:
29:
1271:
1270:
1111:
1110:
1035:Parathyroid gland
1006:
1005:
948:Gonadotropic cell
921:Somatotropic cell
629:(Pt 8): 1330–40.
113:(coming from the
1301:
1220:Corpora arenacea
1085:Zona reticularis
1080:Zona fasciculata
1075:Zona glomerulosa
1063:
1062:
960:Chromophobe cell
953:Thyrotropic cell
889:
888:
876:endocrine system
868:
861:
854:
845:
844:
810:
809:
799:
767:
758:
757:
747:
723:
717:
716:
688:
682:
681:
653:
647:
646:
618:
612:
611:
583:
577:
576:
548:
542:
541:
531:
521:
489:
483:
482:
464:
440:
434:
433:
431:
407:
401:
400:
364:
358:
357:
347:
326:Satoh M (1991).
323:
317:
316:
288:
127:follicular phase
86:surrounding the
84:cumulus oophorus
80:folliculogenesis
76:ovarian follicle
1309:
1308:
1304:
1303:
1302:
1300:
1299:
1298:
1274:
1273:
1272:
1267:
1224:
1201:
1163:
1107:
1103:Chromaffin cell
1089:
1052:
1029:
1020:Follicular cell
1002:
983:Median eminence
964:
901:Pars intermedia
884:Pituitary gland
878:
874:Anatomy of the
872:
834:Granulosa+Cells
818:
813:
768:
761:
724:
720:
689:
685:
654:
650:
619:
615:
584:
580:
549:
545:
490:
486:
441:
437:
422:(4B): 1153–78.
408:
404:
365:
361:
324:
320:
289:
285:
281:
249:
229:
213:Plating density
200:
187:
147:
131:menstrual cycle
69:
55:or egg) in the
37:follicular cell
17:
12:
11:
5:
1307:
1297:
1296:
1291:
1286:
1269:
1268:
1266:
1265:
1260:
1255:
1250:
1249:
1248:
1238:
1232:
1230:
1226:
1225:
1223:
1222:
1217:
1211:
1209:
1203:
1202:
1200:
1199:
1194:
1189:
1184:
1179:
1173:
1171:
1165:
1164:
1162:
1161:
1160:
1159:
1154:
1152:Granulosa cell
1149:
1139:
1138:
1137:
1132:
1121:
1119:
1113:
1112:
1109:
1108:
1106:
1105:
1099:
1097:
1091:
1090:
1088:
1087:
1082:
1077:
1071:
1069:
1060:
1054:
1053:
1051:
1050:
1045:
1039:
1037:
1031:
1030:
1028:
1027:
1022:
1016:
1014:
1008:
1007:
1004:
1003:
1001:
1000:
998:Herring bodies
995:
990:
985:
980:
974:
972:
966:
965:
963:
962:
957:
956:
955:
950:
945:
935:
934:
933:
928:
926:Prolactin cell
923:
916:Acidophil cell
913:
908:
906:Pars tuberalis
903:
897:
895:
886:
880:
879:
871:
870:
863:
856:
848:
842:
841:
831:
825:
817:
816:External links
814:
812:
811:
782:(7): 1069–78.
759:
732:Fertil. Steril
718:
693:Mol Reprod Dev
683:
648:
613:
578:
543:
484:
455:(4): 1134–50.
435:
402:
375:(1): 105–111.
359:
318:
299:(5): 657–662.
282:
280:
277:
276:
275:
270:
265:
260:
255:
248:
245:
231:In the female
228:
225:
199:
196:
186:
183:
146:
143:
68:
65:
33:granulosa cell
15:
9:
6:
4:
3:
2:
1306:
1295:
1292:
1290:
1287:
1285:
1282:
1281:
1279:
1264:
1261:
1259:
1256:
1254:
1251:
1247:
1244:
1243:
1242:
1239:
1237:
1234:
1233:
1231:
1227:
1221:
1218:
1216:
1213:
1212:
1210:
1208:
1204:
1198:
1195:
1193:
1190:
1188:
1185:
1183:
1180:
1178:
1175:
1174:
1172:
1170:
1166:
1158:
1157:Corpus luteum
1155:
1153:
1150:
1148:
1147:Theca interna
1145:
1144:
1143:
1140:
1136:
1133:
1131:
1128:
1127:
1126:
1123:
1122:
1120:
1118:
1114:
1104:
1101:
1100:
1098:
1096:
1092:
1086:
1083:
1081:
1078:
1076:
1073:
1072:
1070:
1068:
1064:
1061:
1059:
1058:Adrenal gland
1055:
1049:
1046:
1044:
1041:
1040:
1038:
1036:
1032:
1026:
1023:
1021:
1018:
1017:
1015:
1013:
1009:
999:
996:
994:
991:
989:
986:
984:
981:
979:
976:
975:
973:
971:
967:
961:
958:
954:
951:
949:
946:
944:
941:
940:
939:
938:Basophil cell
936:
932:
929:
927:
924:
922:
919:
918:
917:
914:
912:
911:Pars distalis
909:
907:
904:
902:
899:
898:
896:
894:
890:
887:
885:
881:
877:
869:
864:
862:
857:
855:
850:
849:
846:
839:
835:
832:
829:
826:
823:
820:
819:
807:
803:
798:
793:
789:
785:
781:
777:
773:
766:
764:
755:
751:
746:
741:
738:(6): 2050–5.
737:
733:
729:
722:
714:
710:
706:
702:
698:
694:
687:
679:
675:
671:
667:
664:(3): 565–73.
663:
659:
652:
644:
640:
636:
632:
628:
624:
617:
609:
605:
601:
597:
594:(3): 839–45.
593:
589:
582:
574:
570:
566:
562:
558:
554:
547:
539:
535:
530:
525:
520:
515:
511:
507:
504:(2): e55578.
503:
499:
495:
488:
480:
476:
472:
468:
463:
458:
454:
450:
446:
439:
430:
425:
421:
417:
413:
406:
398:
394:
390:
386:
382:
378:
374:
370:
363:
355:
351:
346:
341:
337:
333:
329:
322:
314:
310:
306:
302:
298:
294:
287:
283:
274:
271:
269:
266:
264:
261:
259:
256:
254:
251:
250:
244:
242:
238:
234:
233:rhesus monkey
227:Ovarian aging
224:
222:
218:
214:
210:
209:
204:
195:
193:
182:
180:
176:
170:
168:
167:gonadal ridge
164:
160:
151:
142:
140:
137:that produce
136:
132:
128:
124:
120:
116:
112:
108:
104:
100:
95:
93:
89:
85:
81:
77:
74:
64:
62:
58:
54:
50:
46:
42:
38:
34:
26:
21:
1207:Pineal gland
1197:Epsilon cell
1151:
1135:Sertoli cell
1048:Oxyphil cell
978:Pars nervosa
779:
775:
735:
731:
721:
699:(1): 87–93.
696:
692:
686:
661:
658:J Endocrinol
657:
651:
626:
622:
616:
591:
587:
581:
559:(2): 351–7.
556:
552:
546:
501:
497:
487:
452:
448:
438:
419:
415:
405:
372:
368:
362:
335:
331:
321:
296:
292:
286:
230:
217:progesterone
206:
203:Cell culture
201:
198:Cell culture
188:
171:
156:
139:progesterone
115:thecal cells
99:sex steroids
96:
70:
41:somatic cell
36:
32:
30:
1258:Development
1241:Paraganglia
1215:Pinealocyte
1130:Leydig cell
588:Biol Reprod
553:Biol Reprod
449:Biol Reprod
179:mesothelium
125:during the
51:(called an
1278:Categories
1192:Delta cell
1177:Alpha cell
1043:Chief cell
623:J Cell Sci
338:: 85–107.
279:References
219:producing
73:primordial
1182:Beta cell
993:Pituicyte
970:Posterior
258:Ovulation
175:histology
123:aromatase
119:estradiol
111:androgens
1253:Placenta
1125:Testicle
893:Anterior
806:25957622
754:19324349
678:11524236
643:17389684
608:10952929
573:15843493
538:23409002
498:PLOS ONE
471:11906935
397:40498196
369:Anat Rec
247:See also
208:in vitro
192:LH surge
45:sex cord
1187:PP cell
1095:Medulla
1012:Thyroid
797:4531862
713:8418823
529:3567121
506:Bibcode
479:2271384
389:7059014
354:1769902
345:1260417
313:6422764
163:oogonia
157:In the
129:of the
71:In the
61:mammals
43:of the
1117:Gonads
1067:Cortex
840:(MeSH)
804:
794:
752:
711:
676:
641:
606:
571:
536:
526:
477:
469:
395:
387:
352:
342:
332:J Anat
311:
161:, the
103:oocyte
92:antral
88:oocyte
53:oocyte
49:gamete
1229:Other
1142:Ovary
988:Stalk
475:S2CID
393:S2CID
117:) to
57:ovary
39:is a
802:PMID
750:PMID
709:PMID
674:PMID
639:PMID
604:PMID
569:PMID
534:PMID
467:PMID
385:PMID
350:PMID
309:PMID
25:DAPI
792:PMC
784:doi
740:doi
701:doi
666:doi
662:170
631:doi
627:120
596:doi
561:doi
524:PMC
514:doi
457:doi
424:doi
377:doi
373:202
340:PMC
336:177
301:doi
297:148
121:by
59:of
35:or
1280::
800:.
790:.
780:32
778:.
774:.
762:^
748:.
736:93
734:.
730:.
707:.
697:34
695:.
672:.
660:.
637:.
625:.
602:.
592:63
590:.
567:.
557:73
555:.
532:.
522:.
512:.
500:.
496:.
473:.
465:.
453:66
451:.
447:.
420:19
418:.
414:.
391:.
383:.
371:.
348:.
334:.
330:.
307:.
295:.
235:,
223:.
211:.
169:.
63:.
31:A
867:e
860:t
853:v
808:.
786::
756:.
742::
715:.
703::
680:.
668::
645:.
633::
610:.
598::
575:.
563::
540:.
516::
508::
502:8
481:.
459::
432:.
426::
399:.
379::
356:.
315:.
303::
27:.
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