title: The biology and dynamics of mammalian cortical granules.
abstract: Cortical granules are membrane bound organelles located in the cortex of unfertilized oocytes. Following fertilization, cortical granules undergo exocytosis to release their contents into the perivitelline space. This secretory process, which is calcium dependent and SNARE protein-mediated pathway, is known as the cortical reaction. After exocytosis, the released cortical granule proteins are responsible for blocking polyspermy by modifying the oocytes' extracellular matrices, such as the zona pellucida in mammals. Mammalian cortical granules range in size from 0.2 um to 0.6 um in diameter and different from most other regulatory secretory organelles in that they are not renewed once released. These granules are only synthesized in female germ cells and transform an egg upon sperm entry; therefore, this unique cellular structure has inherent interest for our understanding of the biology of fertilization. Cortical granules are long thought to be static and awaiting in the cortex of unfertilized oocytes to be stimulated undergoing exocytosis upon gamete fusion. Not till recently, the dynamic nature of cortical granules is appreciated and understood. The latest studies of mammalian cortical granules document that this organelle is not only biochemically heterogeneous, but also displays complex distribution during oocyte development. Interestingly, some cortical granules undergo exocytosis prior to fertilization; and a number of granule components function beyond the time of fertilization in regulating embryonic cleavage and preimplantation development, demonstrating their functional significance in fertilization as well as early embryonic development. The following review will present studies that investigate the biology of cortical granules and will also discuss new findings that uncover the dynamic aspect of this organelle in mammals.
authors.affiliation: Department of Life Science and Graduate Institute of Biotechnology, Private Chinese Culture University, Taipei, Republic of China. lm@faculty.pccu.edu.tw
authors.forename: Min
authors.initials: M
authors.lastname: Liu
mesh_terms: D000818:Animals; D002118:Calcium / Q000378:metabolism*; D003594:Cytoplasmic Granules / Q000378:metabolism*; D005089:Exocytosis; D005260:Female; D006801:Humans; D008954:Models, Biological; D009865:Oocytes / Q000378:metabolism*; D011247:Pregnancy; D050600:SNARE Proteins / Q000378:metabolism*
chemical_list: D050600:SNARE Proteins; D002118:Calcium
references.citation: Mol Reprod Dev. 2001 Nov;60(3):405-13
references.pmid: 11599052
references.citation: Cell Tissue Res. 1988 Mar;251(3):555-64
references.pmid: 3365751
references.citation: Hum Reprod. 1997 Dec;12(12):2773-80
references.pmid: 9455851
references.citation: Dev Biol. 1977 May;57(1):56-74
references.pmid: 863112
references.citation: Lab Invest. 1968 Jan;18(1):63-71
references.pmid: 4171243
references.citation: Exp Cell Res. 1956 Apr;10(2):533-40
references.pmid: 13317917
references.citation: Biol Reprod. 1974 Mar;10(2):125-49
references.pmid: 4462822
references.citation: Hum Reprod Update. 1996 Jan-Feb;2(1):29-42
references.pmid: 9079401
references.citation: Curr Top Pathol. 1976;62:9-27
references.pmid: 1009793
references.citation: J Electron Microsc Tech. 1991 Mar;17(3):319-35
references.pmid: 2045964
references.citation: Cell. 2003 Feb 21;112(4):519-33
references.pmid: 12600315
references.citation: Mol Reprod Dev. 1992 Feb;31(2):135-43
references.pmid: 1599682
references.citation: Curr Opin Cell Biol. 2002 Aug;14(4):488-95
references.pmid: 12383801
references.citation: J In Vitro Fert Embryo Transf. 1990 Jun;7(3):141-5
references.pmid: 2380620
references.citation: Reprod Biol Endocrinol. 2003 Nov 07;1:77
references.pmid: 14613547
references.citation: Biol Reprod. 1998 Jan;58(1):94-102
references.pmid: 9472928
references.citation: Am J Physiol. 1998 Jun;274(6):C1496-500
references.pmid: 9696691
references.citation: Dev Biol. 1989 Jun;133(2):385-92
references.pmid: 2731635
references.citation: Anat Rec. 1967 Dec;159(4):431-46
references.pmid: 5626358
references.citation: J Cell Biol. 1988 Feb;106(2):345-53
references.pmid: 3123497
references.citation: Dev Biol. 1974 Oct;40(2):292-7
references.pmid: 4473390
references.citation: J Biol Chem. 2002 May 17;277(20):17804-10
references.pmid: 11882657
references.citation: Mol Reprod Dev. 1996 Dec;45(4):521-34
references.pmid: 8956291
references.citation: Dev Biol. 1989 May;133(1):295-304
references.pmid: 2785065
references.citation: Biochem Cell Biol. 1998;76(5):779-85
references.pmid: 10353711
references.citation: Exp Cell Res. 1971 Dec;69(2):452-6
references.pmid: 5163638
references.citation: Cell Tissue Res. 1980;205(1):67-76
references.pmid: 7189137
references.citation: Science. 2010 Jul 9;329(5988):216-9
references.pmid: 20616279
references.citation: Biol Reprod. 2011 Feb;84(2):327-35
references.pmid: 20944080
references.citation: Annu Rev Biochem. 1999;68:863-911
references.pmid: 10872468
references.citation: Biol Reprod. 1993 Jun;48(6):1251-7
references.pmid: 8318579
references.citation: Endocrinology. 1989 Jan;124(1):187-94
references.pmid: 2462486
references.citation: Int Rev Cytol. 1980;63:357-92
references.pmid: 395132
references.citation: Front Biosci. 2001 Jul 01;6:D792-806
references.pmid: 11438440
references.citation: J Reprod Fertil. 1990 May;89(1):253-9
references.pmid: 2374118
references.citation: Aust J Biol Sci. 1953 Nov;6(4):693-705
references.pmid: 13126047
references.citation: Am J Anat. 1974 Aug;140(4):577-82
references.pmid: 4844442
references.citation: Anat Rec. 1971 Nov;171(3):347-68
references.pmid: 4108333
references.citation: Am J Anat. 1974 Apr;139(4):535-65
references.pmid: 4274178
references.citation: Biol Reprod. 1977 Feb;16(1):104-11
references.pmid: 188498
references.citation: Dev Biol. 1988 Nov;130(1):184-97
references.pmid: 3141231
references.citation: Dev Biol. 1988 Jan;125(1):1-7
references.pmid: 3334712
references.citation: J Exp Zool. 1988 Apr;246(1):81-93
references.pmid: 3133448
references.citation: Cell. 1978 Oct;15(2):343-55
references.pmid: 569021
references.citation: Adv Morphog. 1966;5:131-79
references.pmid: 4890505
references.citation: Theriogenology. 2011 Jan 1;75(1):80-9
references.pmid: 20843544
references.citation: Am J Anat. 1976 Oct;147(2):203-18
references.pmid: 987704
references.citation: Biochemistry. 1998 Jul 21;37(29):10354-62
references.pmid: 9671503
references.citation: J Submicrosc Cytol. 1986 Apr;18(2):233-47
references.pmid: 3712508
references.citation: Cell. 1985 Dec;43(2 Pt 1):551-8
references.pmid: 3935323
references.citation: Nature. 1992 Jun 18;357(6379):589-93
references.pmid: 1608469
references.citation: J Exp Zool. 1981 Apr;216(1):63-79
references.pmid: 7288389
references.citation: J Cell Sci. 1988 Sep;91 ( Pt 1):139-44
references.pmid: 3267017
references.citation: J Cell Biol. 1994 Jun;125(5):1157-63
references.pmid: 8195297
references.citation: J Reprod Fertil. 1973 Feb;32(2):259-65
references.pmid: 4734596
references.citation: Nature. 1998 May 7;393(6680):14-5
references.pmid: 9590682
references.citation: Mol Reprod Dev. 1999 Feb;52(2):183-8
references.pmid: 9890749
references.citation: Mech Dev. 1998 Jan;70(1-2):77-89
references.pmid: 9510026
references.citation: Biol Reprod Suppl. 1970;2:44-63
references.pmid: 12254593
references.citation: Hum Reprod. 2004 Aug;19(8):1842-55
references.pmid: 15218004
references.citation: J Cell Biol. 1980 Mar;84(3):618-32
references.pmid: 7188942
references.citation: Fertil Steril. 1983 Dec;40(6):748-54
references.pmid: 6418574
references.citation: Hum Reprod. 1992 Aug;7(7):1004-9
references.pmid: 1430117
references.citation: Mol Reprod Dev. 1998 Nov;51(3):295-303
references.pmid: 9771650
references.citation: J Exp Zool. 1998 Jan 1;280(1):91-6
references.pmid: 9437855
references.citation: Am J Anat. 1969 Jun;125(2):177-215
references.pmid: 5788458
references.citation: Mol Reprod Dev. 1993 Jan;34(1):65-72
references.pmid: 8418819
references.citation: Biol Reprod. 1992 Nov;47(5):709-15
references.pmid: 1477197
references.citation: Mol Reprod Dev. 1993 Dec;36(4):462-8
references.pmid: 8305208
references.citation: Dev Biol. 1992 Jul;152(1):145-51
references.pmid: 1628752
references.citation: Anat Rec. 1975 Jan;181(1):95-111
references.pmid: 45880
references.citation: Cell. 1995 Jun 30;81(7):1095-104
references.pmid: 7600577
references.citation: Biol Reprod. 1995 Apr;52(4):753-8
references.pmid: 7779997
references.citation: Dev Biol. 1976 Nov;54(1):52-60
references.pmid: 1036730
references.citation: Chem Biol. 1999 Jan;6(1):1-10
references.pmid: 9889149
references.citation: Gamete Res. 1988 Jan;19(1):1-18
references.pmid: 3198040
references.citation: Acta Pharmacol Sin. 2006 Oct;27(10):1353-8
references.pmid: 17007743
references.citation: Int J Dev Biol. 2008;52(5-6):585-94
references.pmid: 18649272
references.citation: J Biol Chem. 1996 May 10;271(19):11588-94
references.pmid: 8626722
references.citation: J Reprod Fertil. 1971 Jun;25(3):431-6
references.pmid: 5579695
references.citation: Dev Biol. 1997 Jun 1;186(1):16-26
references.pmid: 9188749
references.citation: Biol Reprod. 1999 Nov;61(5):1340-6
references.pmid: 10529283
references.citation: Biol Reprod. 2002 Apr;66(4):1193-202
references.pmid: 11906941
references.citation: Ann N Y Acad Sci. 1985;442:88-95
references.pmid: 3860066
references.citation: Am J Clin Pathol. 1952 Mar;22(3):277-81
references.pmid: 14902736
references.citation: Proc Natl Acad Sci U S A. 1999 Oct 12;96(21):11830-5
references.pmid: 10518536
references.citation: J Morphol. 1975 Nov;147(3):251-74
references.pmid: 1185788
references.citation: J Cell Biol. 1996 Dec;135(6 Pt 2):1741-7
references.pmid: 8991087
references.citation: Microsc Res Tech. 2003 Jul 1;61(4):342-8
references.pmid: 12811739
references.citation: Mol Reprod Dev. 1994 Dec;39(4):439-48
references.pmid: 7893493
references.citation: Fertil Steril. 1995 Dec;64(6):1154-61
references.pmid: 7589669
references.citation: Mol Reprod Dev. 2001 Feb;58(2):223-35
references.pmid: 11139235
references.citation: J Ultrastruct Res. 1965 Jun;12(5):592-610
references.pmid: 4953463
references.citation: Dev Biol. 1993 Jul;158(1):62-78
references.pmid: 8392472
references.citation: Development. 1994 Jul;120(7):1851-9
references.pmid: 7924992
references.citation: J Biol Chem. 2003 Sep 5;278(36):34189-202
references.pmid: 12799386
references.citation: Dev Biol. 1992 Jan;149(1):80-9
references.pmid: 1728596
references.citation: Biol Reprod. 1998 Sep;59(3):655-60
references.pmid: 9716566
references.citation: Cells Tissues Organs. 2001;168(1-2):36-45
references.pmid: 11114585
references.citation: Dev Biol. 1974 Nov;41(1):185-92
references.pmid: 4474098
references.citation: Nature. 1975 Sep 25;257(5524):321-2
references.pmid: 1172195
references.citation: Dev Biol. 1980 Apr;76(1):185-202
references.pmid: 7380091
references.citation: Biol Reprod. 1984 Nov;31(4):837-48
references.pmid: 6509147
references.citation: Biol Reprod. 1990 Nov;43(5):870-6
references.pmid: 2291919
references.citation: Mol Reprod Dev. 1995 Jul;41(3):368-73
references.pmid: 8588937
references.citation: Nature. 1976 May 6;261(5555):68-71
references.pmid: 944858
references.citation: Development. 1988 Mar;102(3):595-604
references.pmid: 3141127
references.citation: Am J Obstet Gynecol. 2010 Aug;203(2):151.e1-7
references.pmid: 20579967
references.citation: Dev Biol. 1989 Mar;132(1):103-12
references.pmid: 2492959
references.citation: J Exp Zool. 1961 Dec;148:185-203
references.pmid: 14008938
references.citation: J Cell Biol. 1985 Mar;100(3):938-46
references.pmid: 3972903
references.citation: Biol Reprod. 2001 Mar;64(3):879-89
references.pmid: 11207204
references.citation: Front Biosci. 1999 Jan 15;4:D114-31
references.pmid: 9889181
references.citation: Dev Growth Differ. 1997 Aug;39(4):523-9
references.pmid: 9352207
references.citation: Z Zellforsch Mikrosk Anat. 1968;92(2):275-95
references.pmid: 4907212
references.citation: Reprod Biol Endocrinol. 2005 Sep 01;3:42
references.pmid: 16137333
references.citation: J Exp Zool. 1980 Jul;213(1):137-59
references.pmid: 7452194
references.citation: Dev Biol. 2004 Jun 1;270(1):246-60
references.pmid: 15136153
references.citation: Exp Cell Res. 1967 Jun;46(3):553-70
references.pmid: 4951764
references.citation: Dev Biol. 1980 May;76(2):418-27
references.pmid: 7190108
references.citation: Hum Reprod. 1990 Jan;5(1):66-70
references.pmid: 2324245
references.citation: J Ultrastruct Res. 1967 Aug;19(3):354-81
references.pmid: 6053018
references.citation: Dev Biol. 1974 Jan;36(1):44-61
references.pmid: 4822839
references.citation: Dev Biol. 1983 Jan;95(1):244-8
references.pmid: 6825929
references.citation: Dev Biol. 2003 May 1;257(1):166-76
references.pmid: 12710965
references.citation: Proc Natl Acad Sci U S A. 1977 Oct;74(10):4214-8
references.pmid: 270665
references.citation: Mol Reprod Dev. 1992 May;32(1):28-32
references.pmid: 1515147
references.citation: Zygote. 1996 May;4(2):145-9
references.pmid: 8913028
references.citation: Hum Reprod. 1992 Aug;7(7):999-1003
references.pmid: 1430145
references.citation: Cell. 1998 Mar 20;92(6):759-72
references.pmid: 9529252
references.citation: Int Rev Cytol. 1982;78:257-360
references.pmid: 6216222
references.citation: J Biol Chem. 1995 Jul 7;270(27):15926-9
references.pmid: 7608143
references.citation: Biol Reprod. 2000 Aug;63(2):575-81
references.pmid: 10906067
references.citation: Adv Morphog. 1966;5:221-325
references.pmid: 4891032
references.citation: Mol Reprod Dev. 2008 May;75(5):915-24
references.pmid: 17948251
references.citation: Biol Reprod. 1997 Jun;56(6):1376-82
references.pmid: 9166688
references.citation: J Cell Biol. 1993 Dec;123(6 Pt 1):1431-40
references.pmid: 8253842
references.citation: Mol Reprod Dev. 2001 May;59(1):78-89
references.pmid: 11335949
references.citation: Development. 2002 Sep;129(18):4315-25
references.pmid: 12183383
references.citation: J Reprod Fertil. 1994 Nov;102(2):361-9
references.pmid: 7861389
references.citation: J Electron Microsc Tech. 1991 Mar;17(3):294-318
references.pmid: 2045963
references.citation: Rev Reprod. 1998 Jan;3(1):7-12
references.pmid: 9509984
references.citation: Biol Reprod. 1994 Mar;50(3):467-73
references.pmid: 8167217
references.citation: Biol Reprod. 1977 Oct;17(3):350-4
references.pmid: 901888
references.citation: Mol Hum Reprod. 1998 Mar;4(3):289-94
references.pmid: 9570275
references.citation: Biol Reprod. 1989 Feb;40(2):379-93
references.pmid: 2720033
references.citation: Dev Biol. 1998 Aug 1;200(1):103-15
references.pmid: 9698460
references.citation: J Reprod Fertil Suppl. 1989;38:49-62
references.pmid: 2677349
references.citation: Anat Rec. 1993 Dec;237(4):518-26
references.pmid: 8311265
references.citation: Mol Reprod Dev. 2001 Dec;60(4):517-34
references.pmid: 11746963
references.citation: Dev Biol. 1990 Jan;137(1):46-55
references.pmid: 2104813
references.citation: Dev Biol. 1990 Oct;141(2):381-92
references.pmid: 1698670
references.citation: J Reprod Fertil Suppl. 1990;42:177-88
references.pmid: 2077122
references.citation: Mol Reprod Dev. 2002 Dec;63(4):451-8
references.pmid: 12412047
references.citation: Nature. 1974 Nov 1;252(5478):41-3
references.pmid: 4473722
references.citation: Biol Reprod. 2000 Apr;62(4):979-87
references.pmid: 10727267
references.citation: Cell Tissue Res. 2001 Feb;303(2):271-5
references.pmid: 11291773
references.citation: Cell. 1985 May;41(1):313-24
references.pmid: 2986849
references.citation: Arch Med Res. 2001 Nov-Dec;32(6):614-8
references.pmid: 11750738