The "closed loop model" in controlling mRNA translation during development.

Translational control is particularly important in situations where the correlation of a distinct mRNA and the abundance of the corresponding protein might be low. This is the case for instance during oocyte maturation, shortly before the GVBD when the chromatin is condensed, until the embryonic genome is activated. In these situations, gene expression relies on the activation of maternal mRNAs which were stored stably in a dormant form. The most sophisticated model for translational initiation at present is the so-called "closed loop" model, where a circularization of the mRNA is mediated by associated 5'-cap- and 3'-poly(A) binding proteins. Depending on differential interactions, this event can result in translational stimulation or repression. Several studies describe correlated regulation mechanisms in model organisms like mouse or Xenopus, but data addressing translational regulation in farm animals are rare. Cytoplasmic mRNA activating or repressing factors, however, might contribute to achieve developmental competence in bovine or porcine oocytes. Recently we showed that, in the pig, embryonic signals can modify essential components of the mRNA-5'-translation initiation complex in the uterine luminal epithelium at the time of implantation. In accordance with the closed loop model of translational initiation, this review focuses on the regulatory impact of 5'-mRNA end associated proteins (components of the mRNA-cap binding complex) and 3'-end associated proteins (components of the poly(A) binding complex) during in vitro maturation of cattle and pig oocytes, early embryonic development and in the pig uterine epithelia.