Review: Marsupials: placental mammals with a difference.

The placenta is the most varied organ within the Mammalia. There are many similarities, as well as some differences, between the marsupial embryo and those of eutherian mammals. The most striking difference is the lack of the inner cell mass in the blastocyst which consists solely of a single layer of trophoblast cells. The trophoblast expands and eventually becomes part of the definitive chorio-vitelline placenta. The degree of functional differentiation between the vascular and non-vascular parts of the yolk sac placenta differs between species in the relative surface area that is attached to the endometrium, in trophoblast thickness, in yolk sac fusion with the luminal epithelium and most markedly in the degree of invasiveness. In marsupials, placental physiology has been best studied in the tammar wallaby. Despite the lack of invasion in the tammar, there is nevertheless maternal recognition of pregnancy in response to trophoblast formation. Contrary to popular opinion, the tammar placenta also elaborates hormones: at term it secretes prostaglandin F2alpha and accumulates cortisol, and it expresses genes for hormones such as growth hormone, IGF2 and relaxin. As in eutherian mammals, genomic imprinting is important for placental function. Despite the relatively short period of placentation, it is clear that the trophoblast and the placenta it forms are as important for successful pregnancy in marsupial as in eutherian mammals. Marsupials are certainly placental mammals. However marsupials have an additional trick in their pouches, with the physiologically sophisticated and extended lactation that has allowed them to exchange the umbilical cord for the teat.