Uncovering an ancestral green ménage à trois: Contributions of Chlamydomonas to the discovery of a broadly conserved triad of plant fertilization proteins

During sexual reproduction in the unicellular green alga Chlamydomonas, gametes undergo the conserved cellular events that define fertilization across the tree of life. After initial ciliary adhesion, plus and minus gametes attach to each other at plasma membrane sites specialized for fusion, their bilayers merge, and cell coalescence into a quadri-ciliated cell signals for nuclear fusion. Recent findings show that these conserved cellular events are driven by 3 conserved protein families, FUS1/GEX2, HAP2/GCS1, and KAR5/GEX1. New results also show that species-specific recognition in Chlamydomonas activates the ancestral, viral-like fusogen HAP2 to drive fusion; that the conserved nuclear envelope fusion protein KAR5/GEX1 is also essential for nuclear fusion in Arabidopsis; and that heterodimerization of BELL-KNOX proteins signals for nuclear fusion in Chlamydomonas through early diverging land plants. This review outlines how Chlamydomonas's Janus-like position in evolution along with the ease of working with its gametes have revealed broadly conserved mechanisms.