Fertilization comprises a series of precisely orchestrated steps that culminate in the fusion of male and female gametes. The most intimate steps during fertilization encompass gamete recognition, adhesion and fusion. In animals, some binding-effector proteins and enzymes have been identified that act on the cell surfaces of the gametes to regulate gamete compatibility and fertilization success. In contrast, exploring plant gamete interaction during double fertilization, a characteristic trait of flowering plants, has been hampered for a long time because of the protected location of the female gametes and technical limitations. Over the last couple of years, however, the use of advanced methodologies, new imaging tools and new mutants has provided deeper insights into double fertilization, at both the cellular and the molecular level, especially for the model plant Arabidopsis thaliana. Most likely, one consequence of inventing double fertilization may be the co-evolution of special molecular mechanisms to govern each successful sperm delivery and efficient gamete recognition and fusion. In vivo imaging of double fertilization and the recent discovery of numerous female-gametophyte-specific expressed genes encoding small secreted proteins, some of whom were found to be essential for the fertilization process, support this hypothesis. Nevertheless, recent findings indicate that at least the membrane-merger step in plant gamete interaction may rely on an ancient and widely used gamete fusion system.

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