An importin-beta-like protein mediates lignin-modification-induced dwarfism in Arabidopsis

Perturbation of lignin biosynthesis often results in severe growth and developmental defects in plants, which imposes practical limitations to genetic enhancement of lignocellulosic biomass for biofuel production. Currently, little information is known about the cellular and genetic mechanisms of this important phenomenon. Here we show that defects in both cell division and cell expansion underlie the dwarfism of an Arabidopsis lignin mutant ref8, and report the identification of a GROWTH INHIBITION RELIEVED 1 (GIR1) gene from a suppressor screen. GIR1 encodes an importin-beta-like protein required for the nuclear import of MYB4, a transcriptional repressor of phenylpropanoid metabolism. Disruption of GIR1 and MYB4 similarly alleviates the cellular defects and growth inhibition in ref8, suggesting that the growth rescue effect of gir1 is likely due to compromised MYB4 transport and function. Importantly, the phenylpropanoid perturbation is not alleviated in gir1 ref8 and myb4 ref8, suggesting that the function of MYB4 in growth inhibition of lignin-modified plants is likely to be distinct from its known role in transcriptional regulation of phenylpropanoid biosynthetic genes. This study also provides evidence that lignin-modification-induced dwarfism is not merely due to compromised water transport brought about by lignin deficiency, as gir1 has no effect on the growth inhibition of other lignin mutants that show the collapsed xylem phenotype.