Mechanisms of endosperm initiation.

KEY MESSAGE: Overview of developmental events and signalling during central cell maturation and early endosperm development with a focus on mechanisms of sexual and autonomous endosperm initiation. Endosperm is important for seed viability and global food supply. The mechanisms regulating the developmental transition between Female Gametophyte (FG) maturation and early endosperm development in angiosperms are difficult to study as they occur buried deep within the ovule. Knowledge of the molecular events underlying this developmental window of events has significantly increased with the combined use of mutants, cell specific markers, and plant hormone sensing reporters. Here, we review recent discoveries concerning the developmental events and signalling of FG maturation, fertilization, and endosperm development. We focus on the regulation of the initiation of endosperm development with and without fertilization in Arabidopsis and the apomict Hieracium, comparing this to what is known in monocots where distinct differences in developmental patterning may underlie alternative mechanisms of suppression and initiation. The Polycomb Repressive Complex 2 (PRC2), plant hormones, and transcription factors are iteratively involved in early fertilization-induced endosperm formation in Arabidopsis. Auxin increases and PRC2 complex inactivation can also induce fertilization-independent endosperm proliferation in Arabidopsis. Function of the PRC2 complex member FERTILIZATION-INDEPENDENT ENDOSPERM and two loci AutE and LOP are required for autonomous endosperm development in apomictic Hieracium. A comparative understanding of cues required for early endosperm development will facilitate genetic engineering approaches for the development of resilient seed crops, especially if an option for fertilization-independent endosperm formation was possible to combat stress-induced crop failure.