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Our study suggests that controlled alterations in CSC distribution and orchestrated movements contribute to the high density and bundling of cellulose microfibrils in SCWs.

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Plant cell walls are important in plant development and for textiles, wood products, and bioenergy.

Cellulose, the microfibrillar component of primary cell walls (PCWs) and secondary cell walls (SCWs), is formed by cellulose synthase complexes (CSCs) at the plasma membrane.

Here, we show that CSCs behave differently during PCW and SCW synthesis and form microfibrils with different organization.

During PCW synthesis, dispersed CSCs synthesize cellulose microfibrils with low aggregation, whereas during SCW synthesis, densely arranged groups of CSCs move coherently to synthesize highly aggregated microfibrils.

Both during PCW and SCW synthesis, PM-localized CSCs moved steadily along linear trajectories (Fig.

2 and Movie S1), which occurs during active cellulose synthesis (11, 12).

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Until recently, similar image analyses during SCW synthesis have been thwarted due to limitations in imaging SCW-synthesizing cells, which are positioned deep within plant tissues (21).

The inducible transdifferentiation of epidermal cells into SCW-synthesizing xylem-like cells now facilitates live-cell imaging of CSCs during SCW synthesis and allows for the comparison of CSC behavior during PCW and SCW synthesis (22).

Primary cell walls (PCWs) have a multilamellate arrangement of cellulose microfibrils to accommodate growth while maintaining structural integrity (1).

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