LACHESIS Restricts Gametic Cell Fate in the Female Gametophyte of Arabidopsis

Abstract

In flowering plants, the egg and sperm cells form within haploid gametophytes. The female gametophyte of Arabidopsis consists of two gametic cells, the egg cell and the central cell, which are flanked by five accessory cells. Both gametic and accessory cells are vital for fertilization; however, the mechanisms that underlie the formation of accessory versus gametic cell fate are unknown. In a screen for regulators of egg cell fate, we isolated the lachesis (lis) mutant which forms supernumerary egg cells. In lis mutants, accessory cells differentiate gametic cell fate, indicating that LIS is involved in a mechanism that prevents accessory cells from adopting gametic cell fate. The temporal and spatial pattern of LIS expression suggests that this mechanism is generated in gametic cells. LIS is homologous to the yeast splicing factor PRP4, indicating that components of the splice apparatus participate in cell fate decisions. The selection and specification of the egg cell determine the number of eggs produced by an animal or plant, which in turn dictates how many offspring that organism can produce. In most higher plants, the egg cell forms in a specialized structure consisting of four different cell types. Two cells, the egg cell and the central cell, are fertilized by sperm cells and develop into the embryo proper and the nutritive tissue (endosperm), respectively. These two gametic cells are flanked by accessory cells; but why do some cells become gametic while others differentiate into accessory cells? To answer this question, we looked for mutants in which this process is disturbed. In the lachesis mutant, accessory cells become extra egg cells. Interestingly, it seems that the misspecification of these accessory cells results from defects in the gametic cells. This suggests that accessory cells monitor the state of the gametic cells to act as a backup if required, ensuring the formation of the key reproductive cells.