Alteration in Secondary Wall Deposition by Overexpression of the Fragile Fiber1 Kinesin-Like Protein in Arabidopsis

Secondary walls in fibers and vessels are typically deposited in three distinct layers, which are formed by the successive re-orientation of cellulose microfibrils. Although cortical microtubules have been implicated in this process, the underlying mechanisms for the formation of three distinct wall layers are not known. The Fragile Fiber1 (FRA1) kinesin-like protein has been previously shown to be involved in the oriented deposition of cellulose microfibrils and important for cell wall strength in Arabidopsis thallana. In the present report, we investigated the expression pattern of the FRA1 gene and studied the effects of FRA1 overexpression on secondary wall deposition. The FRA1 gene was found to be expressed not only in cells undergoing secondary wall deposition including developing interfascicular fibers and xylem cells, but also in dividing cells and expanding/elongating parenchyma cells. Overexpression of FRA1 caused a severe reduction in the thickness of secondary walls in interfascicular fibers and deformation of vessels, which are accompanied with a marked decrease in stem strength. Close examination of secondary walls revealed that unlike the wild-type walls having three typical layers with the middle layer being the thickest, the secondary walls in FRA1 overexpressors exhibited an increased number of layers, all of which had a similar width. Together, these results provide further evidence implicating an important role of the FRA1 kinesin-like protein in the ordered deposition of secondary walls, which determines the strength of fibers and vessels.