Cellular Lignin Distribution Patterns and their Physiological Relevance
Sammanfattning: The diverse morphological shapes of plants are made possible by the structural rigidity provided by cell walls. In order to support vertical growth and long distance water transport, cell walls need to resist a variety of biological and physical stresses. Lignin, a cell wall polyphenolic unique to vascular plants, has long been considered to structurally support the cell walls of xylem vessels and other specialised cell types against these forces. Lignin is a complex polymer whose monomeric composition and biochemical properties vary widely between different species, tissues and cell types. However, the precise characterisation of this micro-scale variation poses considerable methodological hurdles. As a result, it has yet to be understood how differences in lignin composition contribute to the cell-type specific functions of the cell wall. In the works presented herein, we optimise and validate the Wiesner test and Raman microspectroscopy for the quantitative characterisation of lignin in situ and use these techniques to show how cell-type specific genetic regulation of lignification is crucial for cell wall function. Using synthetic lignin monomers and polymers, as well as genetically altered Arabidopsis and Populus plants in conjunction with biochemical lignin composition analyses, we establish the Wiesner test as a specific high-resolution method to quantify coniferaldehyde (I), and show that Raman microspectroscopy allows the relative quantification of total lignin, guaiacyl lignin subunits (G-units), coniferyl alcohol and syringyl lignin subunits (S-units) (II). We then use these methods to characterise cell-autonomous and cell-cell cooperative lignification patterns and show that cell walls of different vessel types depend on distinct amounts of lignin and specific G-units for structural reinforcement (III). S-unit incorporation into vessel lignin and increased adjacency to neighbouring vessels on the other hand compromise their resistance to collapse (III). Altogether, we provide evidence for a lignification process consisting of a fine scale, cell-type specific regulatory network of lignin biosynthesis, cell-to-cell cooperative monomer supply, and cell wall layer specific monomer incorporation. Crucially, it is this dynamic small-scale regulation that allows lignified plant cell walls to fulfil their cell-type specific functions.
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