To preserve or to destroy, that is the question: the role of the cell wall integrity pathway in pollen tube growth

In plants, cell-shape is defined by the cell wall, a complex network of polymers located outside the plasma membrane. During cell growth, cell wall properties have to be adjusted, assuring cell expansion without compromising cell integrity. Plasma membrane-located receptors sense cell wall properties, transducing extracellular signals into intracellular cascades through the cell wall integrity (CWI) pathway that, in turn, leads to adjustments in the regulation and composition of the cell wall. Using pollen tube growth as a single celled model system, we describe the importance of RAPID ALKALINIZATION FACTOR (RALF) peptides as sensors of cell wall integrity. RALF peptides can mediate the communication between cell wall components and plasma membrane-localized receptor-like kinases (RLKs) of the CrRLK1L family. The subsequent activation of intracellular pathways regulates H+, Ca2+, and ROS levels in the cell and apoplast, thereby modulating cell wall integrity. Interestingly, the RALF-CrRLK1L module and some of the components working up- and downstream of the RLK is conserved in many other developmental and physiological signaling processes.