Molecular organization and mechanical properties of the hyaluronan matrix surrounding the mammalian oocyte

Successful ovulation and oocyte fertilization are essential prerequisites for the beginning of life in sexually reproducing animals. In mammalian fertilization, the relevance of the protein coat surrounding the oocyte plasma membrane, known as zona pellucida, has been widely recognized, while, until not too long ago, the general belief was that the cumulus oophorus, consisting of follicle cells embedded in a hyaluronan rich extracellular matrix, was not essential. This opinion was based on in vitro fertilization procedures, in which a large number of sperms are normally utilized and the oocyte can be fertilized even if depleted of cumulus cells. Conversely, in vivo, only very few sperm cells reach the fertilization site, arguing against the possibility of a coincidental encounter with the oocyte. In the last two decades, proteins required for HA organization in the cumulus extracellular matrix have been identified and the study of fertility in mice deprived of the corresponding genes have provided compelling evidence that this jelly-like coat is critical for fertilization. This review focuses on the advances in understanding the molecular interactions making the cumulus environment suitable for oocyte and sperm encounter. Most of the studies on the molecular characterization of the cumulus extracellular matrix have been performed in the mouse and we will refer essentially to findings obtained in this animal model