Male gametogenesis and germline specification in flowering plants

During angiosperm male gametophyte development, the male germline is segregated by an asymmetric cell division of the haploid microspore. This review encompasses recent advances in understanding the genetic and molecular mechanisms involved in generating the male germline from this pluripotent germline initial and in specifying the production of the twin sperm cells required for double fertilization. Genetic studies and access to the transcriptome of isolated gametes have enabled remarkable progress in understanding some of the key regulators that control and integrate germ cell cycle progression with germline specification, and an emerging regulatory model is presented. Rapid advances have also been made in understanding epigenetic regulation and small RNA pathways in the male gametophyte and germline that impact on genome integrity and gamete development, traits that are shared with animal germlines. The review concludes with a perspective of the outstanding issues and directions of future research that will further our understanding of germline specification and the gametophytic control of pollen development.