Recent insights into the physical biology of plant cell walls are reviewed, summarizing the essential differences between primary and secondary cell walls and identifying crucial gaps in our knowledge of their structure and biomechanics. Unexpected parallels are identified between the mechanism of expansion of primary cell walls during growth and the mechanisms by which hydrated wood deforms under external tension. There is a particular need to revise current “cartoons” of plant cell walls to be more consistent with data from diverse approaches and to go beyond summarizing limited aspects of cell walls, serving instead as guides for future experiments and for the application of new techniques.

Cellulose is the most familiar and most abundant strong biopolymer, but the reasons for its outstanding mechanical performance are not well understood. Each glucose unit in a cellulose chain is joined to the next by a covalent C–O–C linkage flanked by two hydrogen bonds. This geometry suggests some form of cooperativity between covalent and hydrogen bonding. Using infrared spectroscopy and X-ray diffraction, we show that mechanical tension straightens out the zigzag conformation of the cellulose chain, with each glucose unit pivoting around a fulcrum at either end. Straightening the chain leads to a small increase in its length and is resisted by one of the flanking hydrogen bonds. This constitutes a simple form of molecular leverage with the covalent structure providing the fulcrum and gives the hydrogen bond an unexpectedly amplified effect on the tensile stiffness of the chain. The principle of molecular leverage can be directly applied to certain other carbohydrate polymers, including the animal polysaccharide chitin. Related but more complex effects are possible in some proteins and nucleic acids. The stiffening of cellulose by this mechanism is, however, in complete contrast to the way in which hydrogen bonding provides toughness combined with extensibility in protein materials like spider silk.

Evidence has emerged that SARS-CoV-2, the coronavirus that causes COVID-19, can be transmitted airborne in aerosol particles as well as in larger droplets or by surface deposits. This minireview outlines the underlying aerosol science, making links to aerosol research in other disciplines. SARS-CoV-2 is emitted in aerosol form during normal breathing by both asymptomatic and symptomatic people, remaining viable with a half-life of up to about an hour during which air movement can carry it considerable distances, although it simultaneously disperses. The proportion of the droplet size distribution within the aerosol range depends on the sites of origin within the respiratory tract and on whether the distribution is presented on a number or volume basis. Evaporation and fragmentation reduce the size of the droplets, whereas coalescence increases the mean droplet size. Aerosol particles containing SARS-CoV-2 can also coalesce with pollution particulates, and infection rates correlate with pollution. The operation of ventilation systems in public buildings and transportation can create infection hazards via aerosols, but provides opportunities for reducing the risk of transmission in ways as simple as switching from recirculated to outside air. There are also opportunities to inactivate SARS-CoV-2 in aerosol form with sunlight or UV lamps. The efficiency of masks for blocking aerosol transmission depends strongly on how well they fit. Research areas that urgently need further experimentation include the basis for variation in droplet size distribution and viral load, including droplets emitted by “superspreader” individuals; the evolution of droplet sizes after emission, their interaction with pollutant aerosols and their dispersal by turbulence, which gives a different basis for social distancing.

It is now established that crystalline cellulose is held together not just by hydrogen bonding, but also by dispersion forces and by electrostatic attraction modulated by stereoelectronic factors such as the exo-anomeric effect. The surface chains of native cellulose microfibrils differ in C6 conformation from crystalline cellulose and therefore form different hydrogen bonds, both outward and inward. Dispersion and electrostatic forces, influenced by cellulose conformation, also operate at the microfibril surface. The surface conformation depends on whether cellulose interacts with water, with the surfaces of other microfibrils or with non-cellulosic polymers. Cellulose-water binding competes with other binding interactions, so that diverse surface interactions are finely balanced in free energy, difficult to simulate, and dependent on local details of water structuring about which little is known, especially in the presence of dispersed chains of hemicellulosic or pectic polymers. An example is the influence of hydration on the aggregation of microfibrils as measured by neutron scattering, which is large for primary-wall cellulose and small for hardwood microfibrils. There are many consequent uncertainties about the surface interactions of hydrated cellulose microfibrils, for example how pectins associate with cellulose or why cellulose-xylan interfaces resist hydration. Evidence from a range of experimental technologies, alongside simulations, will be needed to resolve these uncertainties. The practical implications are wide-ranging, from the mechanism of plant growth and the mechanical resilience of wood to the development of novel, wood-based building materials.

Cellulose from grasses and cereals makes up much of the potential raw material for biofuel production. It is not clear if cellulose microfibrils from grasses and cereals differ in structure from those of other plants. The structures of the highly oriented cellulose microfibrils in the cell walls of the internodes of the bamboo Pseudosasa amabilis are reported. Strong orientation facilitated the use of a range of scattering techniques. Small-angle neutron scattering provided evidence of extensive aggregation by hydrogen bonding through the hydrophilic edges of the sheets of chains. The microfibrils had a mean centre-to-centre distance of 3.0 nm in the dry state, expanding on hydration. The expansion on hydration suggests that this distance between centres was through the hydrophilic faces of adjacent microfibrils. However in the other direction, perpendicular to the sheets of chains, the mean, disorder-corrected Scherrer dimension from wide-angle X-ray scattering was 3.8 nm. It is possible that this dimension is increased by twinning (crystallographic coalescence) of thinner microfibrils over part of their length, through the hydrophobic faces. The wide-angle scattering data also showed that the microfibrils had a relatively large intersheet d-spacing and small monoclinic angle, features normally considered characteristic of primary-wall cellulose. Bamboo microfibrils have features found in both primary-wall and secondary-wall cellulose, but are crystallographically coalescent to a greater extent than is common in celluloses from other plants. The extensive aggregation and local coalescence of the microfibrils are likely to have parallels in other grass and cereal species and to influence the accessibility of cellulose to degradative enzymes during conversion to liquid biofuels

Cellulose microfibrils in conifers, as in other woody materials, are aggregated into loose bundles called macrofibrils. The centre-to-centre spacing of the microfibrils within these macrofibrils can be estimated from the position of a broad diffraction peak in small-angle neutron scattering (SANS) after deuteration. A known spacing of 3.0 nm, increasing with moisture content, is consistent with direct microfibril to microfibril contact. However recent evidence indicates that conifer microfibrils are partially coated with bound xylan chains, and possibly with lignin and galactoglucomannan, implying a wider centre-to-centre spacing as found in angiosperm wood. Delignification of spruce wood allowed a weak SANS peak to be observed without measurable change in spacing. By deuterating spruce wood in mildly alkaline D₂O and then re-equilibrating with ambient H₂O, deuterium atoms were trapped in a position that gave a 3.8 nm microfibril spacing under dry conditions as in angiosperm wood, instead of the 3.0 nm spacing normally observed in conifers. After conventional vapour deuteration of spruce wood a minor peak at 3.8 nm could be fitted in addition to the 3.0 nm peak. These observations are consistent with some microfibril segments being separated by bound xylan chains as in angiosperms, in addition to the microfibril segments that are in direct contact.