In plant tissues, cells are glued to each other by a pectic polysaccharide rich material known as middle lamella (ML). Along with many biological functions, the ML plays a crucial role in maintaining the structural integrity of plant tissues and organs, as it prevents the cells from separating or sliding against each other. The macromolecular organization and the material properties of the ML are different from those of the adjacent primary cell walls that envelop all plant cells and provide them with a stiff casing. Due to its nanoscale dimensions and the extreme challenge to access the structure for material characterization, the ML is poorly characterized in terms of its distinct material properties. This review explores the ML beyond its functionality as a gluing agent. The putative molecular interactions of constituent macromolecules within the ML and at the interface between ML and primary cell wall are discussed. The correlation between the spatiotemporal distribution of pectic polysaccharides in the different portions of the ML and the subcellular distribution of mechanical stresses within the plant tissue are analyzed.

Pectin is the most abundant component of primary cell walls in eudicot plants. The modification and degradation of pectin affects multiple processes during plant development, including cell expansion, organ initiation, and cell separation. However, the extent to which pectin degradation by polygalacturonases affects stem development and secondary wall formation remains unclear. Using an activation tag screen, we identified a transgenic Arabidopsis thaliana line with longer etiolated hypocotyls, which overexpresses a gene encoding a polygalacturonase. We designated this gene as POLYGALACTURONASE INVOLVED IN EXPANSION2 (PGX2), and the corresponding activation tagged line as PGX2 . PGX2 is widely expressed in young seedlings and in roots, stems, leaves, flowers, and siliques of adult plants. PGX2-GFP localizes to the cell wall, and PGX2 plants show higher total polygalacturonase activity and smaller pectin molecular masses than wild-type controls, supporting a function for this protein in apoplastic pectin degradation. A heterologously expressed, truncated version of PGX2 also displays polygalacturonase activity in vitro. Like previously identified PGX1 plants, PGX2 plants have longer hypocotyls and larger rosette leaves, but they also uniquely display early flowering, earlier stem lignification, and lodging stems with enhanced mechanical stiffness that is possibly due to decreased stem thickness. Together, these results indicate that PGX2 both functions in cell expansion and influences secondary wall formation, providing a possible link between these two developmental processes.

The fracture strength and fracture strain were significantly different along the major and minor growth directions, the wall fragment level modulus of elasticity anisotropy for a dehydrated cell wall was 1.23, suggesting a limited anisotropy of the cell wall itself compared with tissue-scale results.

The tensile properties of the ML might not have a major impact on the tissue-scale mechanical properties. This conclusion calls for further study of the ML, including characterization under shear loading conditions and elucidation of the contributions of other extracellular parameters, such as cell size and shape, to the overall tissue-level mechanical response.

No abstract