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Uterine Adenogenesis and Pregnancy: Multiple Roles for Foxa2 in Mice

Adenogenesis,the development of uterine glands, is a unique postnatal event required for successful reproduction in mammals. Uterine glands synthesize, secrete, and transport a complex array of molecules known collectively as histotroph into the uterine lumen to nourish the developing conceptus (embryo and associated placental membranes) [1, 2] (see Fig. 1). For example, the uterine gland knockout ewe model can neither cycle normally nor establish pregnancy; this indicates that functionality of adult uterine tissues is defined by events associated with uterine adenogenesis [3–5]. Genes required for uterine adenogenesis include Pax2, Lhx1 (also known as Lim1) and Emx2, genes in the abdominal-B Hoxa cluster, members of both Wnt and Hedgehog (Hh) gene families [5], prolactin, insulin-like growth factor 1 (IGF1), IGF2, fibroblast growth factor 7 (FGF7), FGF10, hepatocyte growth factor (HGF), cMet, and FGFR2IIIb [3]. Both estrogen receptor alpha (ESR1) and progesterone receptor (PGR) are expressed in nascent and proliferating uterine glandular epithelium (GE), but not in the uterine GE of lambs treated with a progestin that ablates uterine adenogenesis [6]. Secretions from uterine glands are required for conceptus survival and development in mammals [7–12]. In ungulates, secretions from uterine glands are absorbed by adjacent placental areolae for transcellular transport and release into the circulatory system of the conceptus to, for example, meet metabolic demands and stimulate hematopoiesis [12–17].