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EA1-like peptides do not exist in eudicotyledonous plant species. In the eudicot model plant Torenia fournieri cysteine-rich defensin-like proteins were identified as LUREs to attract own pollen tubes in a species-preferential manner.6 LUREs have been identified recently also in Torenia concolor (TcCRP1) and in Arabidopsis species (AtLURE1 and AlLURE1).7,8 In a recent publication we could show that EA1 acts as a species-preferential direct attractant of maize pollen tubes in vitro. Arabidopsis ovules secreting the peptide are capable to guide maize pollen tubes in a species-preferential manner toward the micropylar opening of the ovule.9
Until now only few ligand-receptor pairs for cell-to-cell communication are described in plants and they mainly involve receptor-like kinases (RLKs). One of the best characterized RLKs is CLAVATA1 (CLV1) which is bound by the signaling peptide CLAVATA3 (CLV3) to control stem cell fate in the shoot apical meristem.10 Other signaling peptides include EPIDERMAL PATTERNING FACTOR 1/2 (EPF1/2) which is regulating stomatal patterning via interaction with the ERECTA-family RLKs11 and INFLORESCENCE DEFICIENT IN ABSCISSION (IDA) which is signaling through the RLKs HAESA (HAE) and HAESA-LIKE2 (HSL2) during floral abscission and lateral root emergence.12,13 In the Brassicacea binding of the pollen-coat protein S-Locus Cysteine-Rich/S-Protein-11 (SCR/SP11) to the stigma-specific S-locus Receptor Kinase (SRK) regulates sporophytic self-incompatibility, which is a reproductive strategy for preventing self-fertilization and allowing genetic diversity to be maintained.14 Although it still has to be shown whether the EA1 peptide also acts through a proteinaceous membrane receptor, it is conceivable that it may be recognized either by a tip-localized transmembrane RLK protein or an ion channel. The RLK superfamily forms a large receptor group in plants with more than 100 RLKs expressed in pollen of Arabidopsis.15,16 A subsequent signal transduction pathway could lead to reorientation of the polar tip growth toward the source of the attractant, since the direction of pollen tube growth is controlled at its tip involving Rho GTPase signaling.16 The receptor might also represent an ion channel, which have been shown to be involved in pollen tube growth and guidance.17 To avoid circular tube growth, one would expect such a reorientation signal to be transient and short for fine-tuning of the growth direction. A ligand-receptor complex thus must be inactivated, dissociated, degraded or internalized very shortly after signal transduction pathway(s) are initiated. Usually disassociation and inactivation of peptide ligands occurs in endosomes.18-20 As the fluorophore Dylight 488 is stable from pH 4 to pH 9, its insensitivity against the acid environment inside endosomal vesicles is consistent with the remaining strong signal inside the pollen tube apex after removing excess of labeled peptide from the pollen tubes (Fig. 2). 041b061a72