Peptide signalling during the pollen tube journey and double fertilization.

Flowering seed plants (angiosperms) have evolved unique ways to protect their gametes from pathogen attack and from drying out. The female gametes (egg and central cell) are deeply embedded in the maternal tissues of the ovule inside the ovary, while the male gametes (sperm cells) are enclosed in the vegetative pollen tube cell. After germination of the pollen tube at the surface of papilla cells of the stigma the two immobile sperm cells are transported deep inside the sporophytic maternal tissues to be released inside the ovule for double fertilization. Angiosperms have evolved a number of hurdles along the pollen tube journey to prevent inbreeding and fertilization by alien sperm cells, and to maximize reproductive success. These pre-zygotic hybridization barriers require intensive communication between the male and female reproductive cells and the necessity to distinguish self from non-self interaction partners. General molecules such as nitric oxide (NO) or gamma-aminobutyric acid (GABA) therefore appear to play only a minor role in these species-specific communication events. The past 20 years have shown that highly polymorphic peptides play a leading role in all communication steps along the pollen tube pathway and fertilization. Here we review our current understanding of the role of peptides during reproduction with a focus on peptide signalling during self-incompatibility, pollen tube growth and guidance as well as sperm reception and gamete activation.

The role of diverse LURE-type cysteine-rich peptides as signaling molecules in plant reproduction.

In angiosperm sexual reproduction, the male pollen tube undergoes a series of interactions with female tissues. For efficient growth and precise guidance, the pollen tube perceives extracellular ligands. In recent decades, various types of secreted cysteine-rich peptides (CRPs) have been identified as peptide ligands that regulate diverse angiosperm reproduction processes, including pollen tube germination, growth, guidance, and rupture. Notably, in two distant core eudicot plants, multiple LURE-type CRPs were found to be secreted from egg-accompanying synergid cells, and these CRPs act as a cocktail of pollen tube attractants for the final step of pollen tube guidance. LURE-type CRPs have species-preferential activity, even among close relatives, and exhibit remarkably divergent molecular evolution with conserved cysteine frameworks, demonstrating that they play a key role in species recognition in pollen tube guidance. In this review, I focus on "reproductive CRPs," particularly LURE-type CRPs, which underlie common but species-specific mechanisms in angiosperm sexual reproduction, and discuss their action, functional regulation, receptors, and evolution.