Morphogenesis of complex plant cell shapes: the mechanical role of crystalline cellulose in growing pollen tubes.

Cellulose is the principal component of the load-bearing system in primary plant cell walls. The great resistance to tensile forces of this polysaccharide and its embedding in matrix components make the cell wall a material similar to a fiber composite. In the rapidly growing pollen tube, the amount of cellulose in the cell wall is untypically low. Therefore, we want to investigate whether the load-bearing function of cellulose is nevertheless important for the architecture of this cell. Enzymatic digestion with cellulase and inhibition of cellulose crystal formation with CGA (1-cyclohexyl-5-(2,3,4,5,6-pentafluorophenoxy)-1lambda4,2,4,6-thiatriazin-3-amine) resulted in the formation of tubes with increased diameter in Solanum chacoense and Lilium orientalis when present during germination. In pre-germinated tubes, application of both agents resulted in the transient arrest of growth accompanied by the formation of an apical swelling indicating a role in the mechanical stabilization of this cellular region. Once growth resumed in the presence of cellulase, however, the cell wall in the newly formed tube showed increased amounts of pectins, possibly to compensate for the reduced amount of cellulose. Scanning electron microscopy of pollen tubes subjected to digestion of matrix polysaccharides revealed the mechanical anisotropy of the cell wall. In both Lilium and Solanum, the angle of highest stability revealed by crack formation was significantly below 45 degrees , an indication that in the mature part of the cell cellulose may not the main stress-bearing component against turgor pressure induced tensile stress in circumferential direction.