As one of the most abundant polysaccharides on Earth, xylan will provide more than a third of the sugars for lignocellulosic biofuel production when using grass or hardwood feedstocks. Xylan is characterized by a linear β(1,4)-linked backbone of xylosyl residues substituted by glucuronic acid, 4-O-methylglucuronic acid or arabinose, depending on plant species and cell types. The biological role of these decorations is unclear, but they have a major influence on the properties of the polysaccharide. Despite the recent isolation of several mutants with reduced backbone, the mechanisms of xylan synthesis and substitution are unclear. We identified two Golgi-localized putative glycosyltransferases, GlucUronic acid substitution of Xylan (GUX)-1 and GUX2 that are required for the addition of both glucuronic acid and 4-O-methylglucuronic acid branches to xylan in Arabidopsis stem cell walls. The gux1 gux2 double mutants show loss of xylan glucuronyltransferase activity and lack almost all detectable xylan substitution. Unexpectedly, they show no change in xylan backbone quantity, indicating that backbone synthesis and substitution can be uncoupled. Although the stems are weakened, the xylem vessels are not collapsed, and the plants grow to normal size. The xylan in these plants shows improved extractability from the cell wall, is composed of a single monosaccharide, and requires fewer enzymes for complete hydrolysis. These findings have implications for our understanding of the synthesis and function of xylan in plants. The results also demonstrate the potential for manipulating and simplifying the structure of xylan to improve the properties of lignocellulose for bioenergy and other uses.bioenergy | glucuronoxylan | glycosyltransferase | plant cell wall | polysaccharide
SUMMARYMannans are hemicellulosic polysaccharides that have previously been implicated as structural constituents of cell walls and as storage reserves but which may serve other functions during plant growth and development. Several members of the Arabidopsis cellulose synthase-like A (CSLA) family have previously been shown to synthesise mannan polysaccharides in vitro when heterologously expressed. It has also been found that CSLA7 is essential for embryogenesis, suggesting a role for the CSLA7 product in development. To determine whether the CSLA proteins are responsible for glucomannan synthesis in vivo, we characterised insertion mutants in each of the nine Arabidopsis CSLA genes and several double and triple mutant combinations. csla9 mutants showed substantially reduced glucomannan, and triple csla2csla3csla9 mutants lacked detectable glucomannan in stems. Nevertheless, these mutants showed no alteration in stem development or strength. Overexpression of CSLA2, CSLA7 and CSLA9 increased the glucomannan content in stems. Increased glucomannan synthesis also caused defective embryogenesis, leading to delayed development and occasional embryo death. The embryo lethality of csla7 was complemented by overexpression of CSLA9, suggesting that the glucomannan products are similar. We conclude that CSLA2, CSLA3 and CSLA9 are responsible for the synthesis of all detectable glucomannan in Arabidopsis stems, and that CSLA7 synthesises glucomannan in embryos. These results are inconsistent with a substantial role for glucomannan in wall strength in Arabidopsis stems, but indicate that glucomannan levels affect embryogenesis. Together with earlier heterologous expression studies, the glucomannan deficiency observed in csla mutant plants demonstrates that the CSLA family encodes glucomannan synthases.
Tape springs are straight, thin-walled strips with a curved cross-section. Following recent proposals for large deployable structures exploiting the structural simplicity and robustness of such springs as deployment actuators, the paper investigates the dynamic deployment of a tape spring that is either coiled around a circular hub, or folded into a zigzag pattern. It is shown that in both cases the spring deforms by forming an elastically deformed region with zero transverse curvature and uniform longitudinal curvature. The process of formation and growth of a fold belongs to a wide class of propagating instabilities. It is characterized by a high peak moment and a lower propagation moment. A compact characterization of the moment-rotation relationship for an elastic fold is presented. A key feature is that the bending moment on either side of a fold moving along a uniform tape spring, away from any end supports, is constant, whereas this moment increases near a support. Compact and accurate two-dimensional theories are developed to simulate the self-actuated deployment of tape springs. It is shown that conservative energy formulations are appropriate for coiled springs, where the velocity field is smooth, but not for springs with localized folds. To simulate the motion of such localized folds a non-conservative impulsemomentum formulation is proposed, and it is found that this model can accurately predict both the steady motion of the folds along the tape spring and their rebound against the end supports.
This study is concerned with the equilibrium shapes of orthotropic, elliptical plates and shells deforming elastically without initial stresses. The aim is to explore potential bistable configurations and their dependencies on material parameters and initial shape for elucidating novel morphing structures. A strain energy formulation gives way to a compact set of governing equations of deformation, which can be solved in closed form for some isotropic and orthotropic conditions. It is shown that bistability depends on the change in Gaussian curvature of the shell, in particular, for initially untwisted shells, isotropy precludes bistability, where there is negative initial Gaussian curvature, but orthotropic materials yield bistability irrespective of the sign of the initial Gaussian curvature. This improved range of performance stems from increasing the independent shear modulus, which imparts sufficient torsional rigidity to stabilize against perturbations in the deformed state. It is also shown that the range of bistable configurations for initially twisted shells generally diminishes as the degree of twist increases.
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