The evolution of pregnancy

Although viviparity has evolved many times in the animal kingdom, it remains relatively uncommon-scorpions and therian mammals being rare examples of entirely viviparous major taxa. Viviparity is a specialised form of intra-species parasitism which biases parental investment towards fertilised eggs, temporally spreads that investment, and also temporarily protects offspring from many selection pressures. Importantly, the mammalian viviparity appeared at a relatively late stage in the process of vertebrate evolution. Because of this, viviparity was 'superimposed' on complex pre-existing cardiovascular, respiratory, metabolic and immune systems, and has altered them dramatically. Also, pregnancy has exerted pervasive effects on gene expression in mammals, including genetic imprinting, X inactivation, sex determination, and the ectopic expression in the extra-embryonic membranes of many genes previously expressed in the gonads, brain, pituitary and immune system. Finally, although lactation probably pre-dated viviparity in mammalian evolution, the two have co-evolved as alternative strategies of offspring nutrition ever since.