The multicellular female gametophyte, a unique feature of higher plants, provides us with an excellent experimental system to address fundamental questions in biology. During the past few years, we have gained significant insight into the mechanisms that control embryo sac polarity, gametophytic cell specification, and recognition between male and female gametophytic cells. An auxin gradient has been shown for the first time to function in the female gametophyte to regulate gametic cell fate, and key genes that control gametic cell fate have also been identified. This review provides an overview of these exciting discoveries with a focus on molecular and genetic data.