Rapid alkalinization factor 22 has a structural and signalling role in root hair cell wall assembly.

Pressurized cells with strong walls make up the hydrostatic skeleton of plants. Assembly and expansion of such stressed walls depend on a family of secreted RAPID ALKALINIZATION FACTOR (RALF) peptides, which bind both a membrane receptor complex and wall-localized LEUCINE-RICH REPEAT EXTENSIN (LRXs) in a mutually exclusive way. Here we show that, in root hairs, the RALF22 peptide has a dual structural and signalling role in cell expansion. Together with LRX1, it directs the compaction of charged pectin polymers at the root hair tip into periodic circumferential rings. Free RALF22 induces the formation of a complex with LORELEI-LIKE-GPI-ANCHORED PROTEIN 1 and FERONIA, triggering adaptive cellular responses. These findings show how a peptide simultaneously functions as a structural component organizing cell wall architecture and as a feedback signalling molecule that regulates this process depending on its interaction partners. This mechanism may also underlie wall assembly and expansion in other plant cell types.

Plant cell wall patterning and expansion mediated by protein-peptide-polysaccharide interaction.

Assembly of cell wall polysaccharides into specific patterns is required for plant growth. A complex of RAPID ALKALINIZATION FACTOR 4 (RALF4) and its cell wall-anchored LEUCINE-RICH REPEAT EXTENSIN 8 (LRX8)-interacting protein is crucial for cell wall integrity during pollen tube growth, but its molecular connection with the cell wall is unknown. Here, we show that LRX8-RALF4 complexes adopt a heterotetrametric configuration in vivo, displaying a dendritic distribution. The LRX8-RALF4 complex specifically interacts with demethylesterified pectins in a charge-dependent manner through RALF4's polycationic surface. The LRX8-RALF4-pectin interaction exerts a condensing effect, patterning the cell wall's polymers into a reticulated network essential for wall integrity and expansion. Our work uncovers a dual structural and signaling role for RALF4 in pollen tube growth and in the assembly of complex extracellular polymers.

The differential tolerance of C3 and C4 cereals to aluminum toxicity is faded under future CO2 climate.

Industrial activities have led to a gradual and global increase in soil aluminum (Al) and atmospheric CO2 concentrations. Al bioavailability strongly depends on the soil pH, which in turn is affected by atmospheric CO2 levels. In spite of the concurrent impact which Al and elevated CO2 (eCO2) could have on plants, their interaction and how it might affect the growth of economically important crop species has not been investigated. Here, we have investigated the combined impact of soil Al and eCO2 exposure on key C3 (wheat, oat) and C4 (maize, sorghum) crops, at the physiological and biochemical level. Compared to C3 plants, C4 plants accumulated less Al by stimulating soil Al retention through exudation of root organic acids. Consequently, Al-exposed C4 plants maintained photosynthetic performance and anti-oxidative capacity. Exposure to eCO2 reduced the stress responses of C3 and C4 crops to Al exposure. Elevated CO2 decreased Al accumulation and oxidative damage in all cereals, and ameliorated C3 plant growth. This was reflected on the biochemical level, where eCO2 inhibited ROS production and restored RuBisCo activity in C3 crops only. Overall, our data suggest that, compared to C3 crops, C4 cereals are more tolerant to soil Al exposure under current ambient CO2 (aCO2) levels whereas future eCO2 levels might stimulate Al tolerance in C3 crops.