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. 2022 May;12(5):210208.
doi: 10.1098/rsob.210208. Epub 2022 May 4.

Secondary cell wall patterning-connecting the dots, pits and helices

Affiliations

Secondary cell wall patterning-connecting the dots, pits and helices

Huizhen Xu et al. Open Biol. 2022 May.

Abstract

All plant cells are encased in primary cell walls that determine plant morphology, but also protect the cells against the environment. Certain cells also produce a secondary wall that supports mechanically demanding processes, such as maintaining plant body stature and water transport inside plants. Both these walls are primarily composed of polysaccharides that are arranged in certain patterns to support cell functions. A key requisite for patterned cell walls is the arrangement of cortical microtubules that may direct the delivery of wall polymers and/or cell wall producing enzymes to certain plasma membrane locations. Microtubules also steer the synthesis of cellulose-the load-bearing structure in cell walls-at the plasma membrane. The organization and behaviour of the microtubule array are thus of fundamental importance to cell wall patterns. These aspects are controlled by the coordinated effort of small GTPases that probably coordinate a Turing's reaction-diffusion mechanism to drive microtubule patterns. Here, we give an overview on how wall patterns form in the water-transporting xylem vessels of plants. We discuss systems that have been used to dissect mechanisms that underpin the xylem wall patterns, emphasizing the VND6 and VND7 inducible systems, and outline challenges that lay ahead in this field.

Keywords: cell wall patterning; cellulose; microtubules; plant cell wall; xylem.

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Conflict of interest statement

We declare we have no competing interests.

Figures

Figure 1.
Figure 1.
Examples of secondary cell wall pattern types. From left to right: annular (cyan), helical (yellow), reticulate (orange), scalariform (red), opposite-pitted (purple) and alternate-pitted (blue).
Figure 2.
Figure 2.
Systems to study cell wall patterns. (a) Hormones and hormone-related molecules can induce xylem transdifferentiation. (b) Adding dexamethasone to the VND6/7-glucocorticoid receptor (GR) systems induces metaxylem (VND6) and protoxylem (VND7; image from A. thaliana hypocotyl cell) formation.
Figure 3.
Figure 3.
Drivers of xylem cell wall patterns. (a) ROP domains formation in the presence of a primary wall-like microtubule cytoskeleton. Microtubule ends are targeted in the active ROP domains, while microtubules remain unperturbed outside the ROP domains. (b) Active ROP domains are shaped in a microtubule-dependent manner. (c) SCW synthesis machinery (CESA complexes) is recruited. (d) Patterned deposition of SCW material.
Figure 4.
Figure 4.
A schematic model of regulation of secondary cell wall development in metaxylem vessels. Light green domains (ovals) indicate plasma membrane domains marked with activated ROP11. Double arrowheads indicate interactions. Red arrows indicate promotion of bundling and/or polymerization. Light blue bars indicate elimination of microtubules or activated ROPs.

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