The Mammalian Egg’s Block to Polyspermy

  • Don P. Wolf


The eggs of most mammals at ovulation are metabolically relatively inert cells arrested in metaphase II of meiosis. In response to an activation stimulus normally provided by the fertilizing sperm but induced artificially by a number of parthenogenetic agents, the egg resumes meiosis, undergoes a cortical reaction, and becomes metabolically more active. Visible evidence for the resumption of meiosis involves the abstriction of a second polar body at approximately 30 min postactivation. In the cortical reaction, the egg undergoes the exocytotic release of its cortical granules, a process that results in formation of a new mosaic plasma membrane from the fusion of limiting cortical granule membranes with the egg plasma membrane. At the same time, cortical granule contents released into the perivitelline space come in contact with the egg plasma membrane and the zona pellucida. The cortical reaction and granule exocytosis are of primary interest to any discussion of polyspermy for cortical granule contents have been associated with polyspermy-preventing mechanisms in the eggs of animals of diverse species (for review, see Schuel, 1978, or Gulyas, 1980).


Zona Pellucida Cortical Granule Perivitelline Space Zona Hardening Cortical Reaction 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Austin, C. R., 1956, Cortical granules in hamster eggs, Exp. Cell Res. 10:533–540.CrossRefGoogle Scholar
  2. Austin, C. R., and Bishop, M. W. H., 1957, Fertilization in mammals, Biol. Rev. 32:296–349.CrossRefGoogle Scholar
  3. Baranska, W., Konwinski M., and Kujawa, M., 1975, Fine structure of the zona pellucida of unfertilized egg cells and embryos, J. Exp. Zool. 192:193–202.CrossRefGoogle Scholar
  4. Barros, C., and Yanagimachi, R., 1971, Induction of the zona reaction in golden hamster eggs by cortical granule material, Nature (London) 233:268–269.CrossRefGoogle Scholar
  5. Barros, C., and Yanagimachi, R., 1972, Polyspermy-preventing mechanisms in the golden hamster egg, J. Exp. Zool. 180:251–266.CrossRefGoogle Scholar
  6. Bedford, J. M., and Cross, N. L., 1978, Normal penetration of rabbit spermatozoa through a trypsin-and acrosin-resistant zona pellucida, J. Reprod. Fertil. 54:385–392.CrossRefGoogle Scholar
  7. Bleil, J. D., and Wassarman, P. M., 1980a, Structure and function of the zona pellucida: Identification and characterization of the proteins of the mouse oocyte’s zona pellucida, Dev. Biol. 76:185–202.CrossRefGoogle Scholar
  8. Bleil, J. D., and Wassarman, P. M., 1980b, Mammalian sperm-egg interaction: Identification of a glycoprotein in mouse egg zonae pellucidae possessing receptor activity for sperm, Cell 20:873–882.CrossRefGoogle Scholar
  9. Chang, M. C., and Hunt, D. M., 1956, Effects of proteolytic enzymes on the zona pellucida of fertilized and unfertilized mammalian eggs, Exp. Cell Res. 11:497–499.CrossRefGoogle Scholar
  10. Cooper, G. W., and Bedford, J. M., 1971, Charge density change in the vitelline surface following fertilization of the rabbit egg, J. Reprod. Fertil. 25:431–436.CrossRefGoogle Scholar
  11. Dunbar, B. S., and Raynor, B. D., 1980, Characterization of porcine zona pellucida antigens, Biol. Reprod. 22:941–954.CrossRefGoogle Scholar
  12. Dunbar, B. S., Wardrip, N. J., and Hedrick, J. L., 1980, Isolation, physicochemical properties, and macromolecular composition of zona pellucida from porcine oocytes, Biochemistry 19:356–365.CrossRefGoogle Scholar
  13. Eager, D. D., Johnson, M. H., and Thurley, K. W., 1976, Ultrastructural studies on the surface membrane of the mouse egg, J. Cell Sci. 22:345–353.Google Scholar
  14. Flechon, J.-E., Huneau, D., Solari, A., and Thibault, C., 1975, Réaction corticale et blocage de la polyspermie dans l’oeuf de lapine, Ann. Biol. Anim. Biochim. Biophys. 15:9–18.CrossRefGoogle Scholar
  15. Foerder, C. A., and Shapiro, B. M., 1977, Release of ovoperoxidase from sea urchin eggs hardens the fertilization membrane with tyrosine crosslinks, Proc. Natl. Acad. Sci. USA 79: 4214–4218.CrossRefGoogle Scholar
  16. Fraser, L. R., Dandekar, P. V., and Gordon, M. R. 1972, Loss of cortical granules in rabbit eggs exposed to spermatozoa in vitro, J. Reprod. Fertil. 29:295–297.CrossRefGoogle Scholar
  17. Fukuda, Y., and Chang, M. C., 1978, The time of cortical granule breakdown and sperm penetration in mouse and hamster eggs inseminated in vitro, Biol. Reprod. 19:261–266.CrossRefGoogle Scholar
  18. Gabel, C. A., Eddy, E. M., and Shapiro, B. M., 1979, After fertilization, sperm surface components remain as a patch in sea urchin and mouse embryos, Cell 18:207–215.CrossRefGoogle Scholar
  19. Gordon, M., Fraser, L. R., and Dandekar, P. V., 1975, The effect of ruthenium red and concanavalin A on the vitelline surface of fertilized and unfertilized rabbit ova, Anat. Rec. 181:95–112.CrossRefGoogle Scholar
  20. Gould, K., Zaneveld, L. J. D., Srivastava, P. N., and Williams, W. L., 1971, Biochemical changes in the zona pellucida of rabbit ova induced by fertilization and sperm enzymes, Proc. Soc. Exp. Biol. Med 136:6–10.Google Scholar
  21. Grey, R. D., Wolf, D. P., and Hedrick, J. L., 1974, Formation and structure of the fertilization envelope in Xenopus laevis, Dev. Biol. 36:44–61.CrossRefGoogle Scholar
  22. Gulyas, B. J., 1980, Cortical granules of mammalian eggs, Int. Rev. Cytol. 63:357–392.CrossRefGoogle Scholar
  23. Gulyas, B. J., and Schmell, E. D., 1980, Ovoperoxidase activity in ionophore treated mouse eggs. I. Electron microscopic localization, Gamete Res. 3:267.CrossRefGoogle Scholar
  24. Gwatkin, R. B. L., Fertilization Mechanisms in Man and Mammals, Plenum Press, New York.Google Scholar
  25. Gwatkin, R. B. L., Williams, D. T., Hartmann, J. F., and Kniazuk, M., 1973, The zona reaction of hamster and mouse eggs: Production in vitro by a trypsin-like protease from cortical granules, J. Reprod. Fertil. 32:259–265.CrossRefGoogle Scholar
  26. Gwatkin, R. B. L., Rasmusson, G. H., and Williams, D. T., 1976, Induction of the cortical reaction in hamster eggs by membrane-active agents, J. Reprod. Fertil. 47:299–303.CrossRefGoogle Scholar
  27. Gwatkin, R. B. L., Williams, D. T., and Meyenhofer, M., 1979, Isolation of bovine zonae pellucidae from ovaries with collagenase: Antigenic and sperm receptor properties, Gamete Res. 2:187–192.CrossRefGoogle Scholar
  28. Hagiwara, S., and Jaffe, L. A., 1979, Electrical properties of egg cell membranes, Annu. Rev. Biophys. Bioeng. 8:385–416.CrossRefGoogle Scholar
  29. Hall, H. G., 1978, Hardening of the sea urchin fertilization envelope by peroxidase-catalyzed phenolic coupling of tyrosines, Cell 15:343–355.CrossRefGoogle Scholar
  30. Hanada, A., and Chang, M. C., 1972. Penetration of zona-free rat eggs by spermatozoa of different species, Biol. Reprod. 6:300–309.Google Scholar
  31. Hanada, A., and Chang, M. C., 1976, Penetration of hamster and rabbit zona-free eggs by rat and mouse spermatozoa with special reference to sperm capacitation, J. Reprod. Fertil. 46:239–241.CrossRefGoogle Scholar
  32. Hartmann, J. F., and Gwatkin, R. B. L., 1971, Alteration of sites on the mammalian sperm surface following capacitation, Nature (London) 234:479–481.CrossRefGoogle Scholar
  33. Hirao, Y., and Yanagimachi, R., 1978, Effects of various enzymes on the ability of hamster egg plasma membranes to fuse with spermatozoa, Gamete Res. 1:3–12.CrossRefGoogle Scholar
  34. Inoue, M., and Wolf, D. P., 1974, Comparative solubility properties of the zonae pellucidae of unfertilized and fertilized mouse ova, Biol. Reprod. 11:558–565.CrossRefGoogle Scholar
  35. Inoue, M., and Wolf, D. P., 1975a, Fertilization-associated changes in the murine zona pellucida: A time sequence study, Biol. Reprod. 13:546–551.CrossRefGoogle Scholar
  36. Inoue, M., and Wolf, D. P., 1975b, Comparative solubility properties of rat and hamster zonae pellucidae, Biol Reprod. 12:535–540.CrossRefGoogle Scholar
  37. Jackowski, S., and Dumont, J. N., 1979, Surface alterations of the mouse zona pellucida and ovum following in vivo fertilization: Correlation with the cell cycle, Biol. Reprod. 20:150–161.CrossRefGoogle Scholar
  38. Jackowski, S., Leibo, S. P., and Mazur, P., 1980, Glycerol permeabilities of fertilized and unfertilized mouse ova, J. Exp. Zool. 212:329–341.CrossRefGoogle Scholar
  39. Johnson, M., and Edidin, M., 1978, Lateral diffusion in plasma membrane of mouse egg is restricted after fertilization, Nature (London) 272:448–450.CrossRefGoogle Scholar
  40. Kaleta, E., 1979, Sperm penetration in vitro into ovarian and tubal oocytes from mice of the inbred KE and C57 strains, Gamete Res. 2:99–104.CrossRefGoogle Scholar
  41. Longo, F. J., 1974, Ultrastructural changes in rabbit eggs aged in vivo, Biol. Reprod. 11:22–39.CrossRefGoogle Scholar
  42. Nicolson, G. L., Yanagimachi, R., and Yanagimachi, R., 1975, Ultrastructural localization of lectin-binding sites on the zonae pellucidae and plasma membranes of mammalian eggs, J. Cell Biol. 66:263–274.CrossRefGoogle Scholar
  43. Nicosia, S. V., Wolf, D. P., and Inoue, M., 1977, Cortical granule distribution and cell surface characteristics in mouse ova, Dev. Biol. 57:56–74.CrossRefGoogle Scholar
  44. Nicosia, S. V., Wolf, D. P., and Mastroianni, L., Jr. 1978, Surface topography of mouse eggs before and after insemination, Gamete Res. 1:145–155.CrossRefGoogle Scholar
  45. Nitsch, B., Brück, H.-J., and Palm, S., 1977, Simple methods for observing cortical granules in living mammalian eggs, Experientia 33:1252–1253.CrossRefGoogle Scholar
  46. Kiwa, K., and Chang, M. C., 1975, Requirement of capacitation for sperm penetration of zona-free rat eggs, J. Reprod. Fertil. 44:305–308.CrossRefGoogle Scholar
  47. Oh, Y. K., and Brackett, B. G., 1975, Ultrastructure of rabbit ova recovered from ovarian follicles and inseminated in vitro, Fertil. Steril. 26:665–685.Google Scholar
  48. Okamoto, H., Takahashi, K., and Yamashita, N., 1977, Ionic currents through the membrane of the mammalian oocyte and their comparison with those in the tunicate and sea urchin, J. Physiol. 267:465–495.Google Scholar
  49. Overstreet, J. W., and Bedford, J. M., 1974, Comparison of the penetrability of the egg vestments in follicular oocytes, unfertilized and fertilized ova of the rabbit, Dev. Biol. 41:185–192.CrossRefGoogle Scholar
  50. Pavlok, A., and McLaren, A., 1972, The role of cumulus cells and the zona pellucida in fertilization of mouse eggs in vitro, J. Reprod. Fertil. 29:91–97.CrossRefGoogle Scholar
  51. Pienkowski, M., 1974, Study of the growth regulation of preimplantation mouse embryos using concanavalin A, Proc. Soc. Exp. Biol Med. 145:464–469.Google Scholar
  52. Powers, R. D., and Tupper, J. T., 1974, Some electrophysiological and permeability properties of the mouse egg, Dev. Biol. 38:320–331.CrossRefGoogle Scholar
  53. Powers, R. D., and Tupper, J. T., 1977, Developmental changes in membrane transport and permeability in the early mouse embryo, Dev. Biol. 56:305–315.CrossRefGoogle Scholar
  54. Repin, V. S., and Akimova, I. M., 1976, Microelectrophoretic analysis of protein composition of zonae pellucidae of mammalian oocytes and zygotes, Biochemistry (USSR) 41:39–45.Google Scholar
  55. Saling, P. M., Storey, B. T., and Wolf, D. P., 1978, Calcium-dependent binding of mouse epididymal spermatozoa to the zona pellucida, Dev. Biol. 65:515–525.CrossRefGoogle Scholar
  56. Sato, K., 1979, Polyspermy-preventing mechanisms in mouse eggs fertilized in vitro, J. Exp. Zool. 210:353–359.CrossRefGoogle Scholar
  57. Schmell, E. D., and Gulyas, B. J., Ovoperoxidase activity in ionophore treated mouse eggs. II. Evidence for the enzyme’s role in hardening the zona pellucida, Gamete Res. 3:279.Google Scholar
  58. Schuel, H., 1978, The role of cortical granules in fertilization, Gamete Res. 1:299–382.CrossRefGoogle Scholar
  59. Solter, D., 1977, Organization and the antigenic properties of the egg membrane, in: Immunobiology of Gametes (M. Edidin and M. H. Johnson, eds.), Cambridge University Press, London, pp. 207–234.Google Scholar
  60. Szollosi, D., 1967, Development of cortical granules and the cortical reaction in rat and hamster eggs, Anat. Rec. 159:431–446.CrossRefGoogle Scholar
  61. Toyoda, Y., and Chang, M. C., 1968, Sperm penetration of rat eggs in vitro after dissolution of zona pellucida by chymotrypsin, Nature (London) 220:589–591.CrossRefGoogle Scholar
  62. Usui, N., and Yanagimachi, R., 1976, Behavior of hamster sperm nuclei incorporated into eggs at various stages of maturation, fertilization and early development, J. Ultrastruct. Res. 57:276–288.CrossRefGoogle Scholar
  63. Wabik-Sliz, B., 1979, Number of cortical granules in mouse oocytes from inbred strains differing in efficiency of fertilization, Biol. Reprod. 21:89–97.CrossRefGoogle Scholar
  64. Wartenberg, H., and Stegner, H. E., 1960, Über die elektronenmikroskopische Feinstruktur des Menschlichen Ovarialeis, Z. Zellforsch. Mikrosk. Anat. 52:450–474.CrossRefGoogle Scholar
  65. Wolf, D. P., 1978, The block to sperm penetration in zona-free mouse eggs, Dev. Biol. 64:1–10.CrossRefGoogle Scholar
  66. Wolf, D. P., and Hamada, M., 1977, Induction of zonal and oolemmal blocks to sperm penetration in mouse eggs with cortical granule exudate, Biol. Reprod. 17:350–354.CrossRefGoogle Scholar
  67. Wolf, D. P., and Hamada, M., 1979, Sperm binding to the mouse egg plasmalemma, Biol. Reprod. 21:205–211.CrossRefGoogle Scholar
  68. Wolf, D. P., Inoue, M., and Stark, R. A., 1976, Penetration of zona-free mouse eggs, Biol. Reprod. 15:213–221.CrossRefGoogle Scholar
  69. Wolf, D. P., Nicosia, S. V., and Hamada, M., 1979, Premature cortical granule loss does not prevent sperm penetration of mouse eggs, Dev. Biol. 71:22–32.CrossRefGoogle Scholar
  70. Yanagimachi, R., 1977, Specificity of sperm-egg interactions, in: Immunobiology of Gametes (M. Edidin and M. H. Johnson, eds.), Cambridge University Press, London, pp. 255–295.Google Scholar
  71. Yanagimachi, R., Yanagimachi, H., and Rogers, B. J., 1976, The use of zona-free animal ova as a test-system for the assessment of the fertilizing capacity of human spermatozoa, Biol. Reprod. 15:471–476.CrossRefGoogle Scholar
  72. Yu. S.-F., Wolf, D. P., 1981, Polyspermic mouse eggs can dispose of supernumerary sperm, Dev. Biol. 82:203–210.CrossRefGoogle Scholar

Copyright information

© Plenum Press, New York 1981

Authors and Affiliations

  • Don P. Wolf
    • 1
  1. 1.Division of Reproductive Biology, Departments of Obstetrics and Gynecology, and Biochemistry and BiophysicsUniversity of Pennsylvania School of MedicinePhiladelphiaUSA

Personalised recommendations