Cell-walls of growing plant cells

The plant primary cell wall is a three-dimensional interwoven network of cellulose microfibrils, cross-linked by xyloglucan and dispersed in a pectin matrix. It has been suggested that in the wall of growing plant cells, xyloglucan is bound to the rigid cellulose microfibrils by hydrogen bonds and holds the microfibrils together by forming molecular tethers, which is referred to as the ‘sticky network’ model. Plant growth occurs when these tethers are peeled from the microfibrils by expansins or broken by glycosidases or transglycosylases. A number of researchers have presented theoretical difficulties and observations inconsistent with this model and a new hypothesis has been proposed, claiming that the cellulose – xyloglucan cross-links may act as ‘scaffolds’ holding the microfibrils apart. Analogies with synthetic polymers suggests that the spacing between the cellulose microfibrils may be an important determinant of the mechanical properties of the cell wall and the results presented in this thesis support this hypothesis. Water contents of Acetobacter xylinus synthesized cellulose based cell wall analogues (as a mimic of primary cell wall) and sunflower hypocotyl cell walls were altered using high molecular weight polyethylene glycol (PEG) solution, and their extension under a constant load was measured using a creep extensiometer and showed that there were clear reduction (30-35%) in extensibility suggesting that water content of the wall and therefore the cell wall free volume directly influence wall extensibility. When hydration of A. xylinus cellulose composite pellicles was reduced using PEG 6000 solution and re-hydrated in buffer solution, followed by treatment with α-expansin or snail acetone powder extract, it was found that expansin and snail powder extracts caused a rapid rehydration of the composites and that the pellicles only returned to their original weights after these treatments, suggesting that expansin and snail powder can increase the free volume of the wall perhaps contributing to the increases in extensibility that they cause. Assays on cell wall fragments also indicated that expansin increased the cell wall free volume, demonstrated by changes of the turbidity of fragment suspensions. The role of pectic polysaccharide, RG-II, in cell wall biomechanics was also investigated using mechanical and biochemical testing of available Arabidopsis thaliana cell wall mutants and by incorporating RG-II (purified from red wine) with Acetobacter cellulose. It was demonstrated that RG-II significantly increased the hydration of cellulose composite; hydration rate was 15 -16% more than the composite without RG-II and thus increased the pellicle extensibility.

From the results, it is evidenced that cell wall extension is not only the consequences of breaking hydrogen bonds between cellulose microfibrils and xyloglucan by expansins or glycosidases and transglycosylases, but also a wider range of factors are involved including cell wall water content, cell wall free volume and the pectic polymers, especially RG-II.