The GhCEWT1-GhCEWT2-GhCes4D/GhCOBL4D module orchestrates plant cell elongation and cell wall thickness

Cell elongation defines cell size and shape, whereas the cell wall supports and protects it. However, the mechanism regulating cell elongation and cell wall thickness remains unknown. Here, taking advantage of a model for both cell elongation and cell wall biogenesis, cotton fiber, we identified a basic-helix-loop-helix (bHLH) factor, GhCEWT1, that contributes to both fiber cell elongation and cell wall thickness. Loss of function of GhCEWT1 reduced the fiber length and cell wall thickness. GhCEWT1 induced transcription of GhCEWT2. We also identified two target genes of GhCEWT2, cellulose synthase 4D (GhCes4D) and COBRA-LIKE 4D (GhCOBL4D). GhCEWT2 enhanced the transcription of GhCes4D and GhCOBL4D. GhCOBL4D overexpression significantly enhanced cotton fiber cell length and cell wall thickness. Our results revealed a GhCEWT1-GhCEWT2-GhCes4D/GhCOBL4D cascade functioning in both fiber cell elongation and cell wall thickness. These findings provide a comprehensive understanding of plant cell elongation and cell wall formation, as well as a theoretical basis for boosting the biomass on Earth.