On the bending of rectangular atomic monolayers along different directions: anab initiostudy.

We study the bending of rectangular atomic monolayers along different directions from first principles. Specifically, choosing the phosphorene, GeS, TiS3, and As2S3monolayers as representative examples, we perform Kohn-Sham density functional theory calculations to determine the variation in transverse flexoelectric coefficient and bending modulus with the direction of bending. We find that while the flexoelectric coefficient is nearly isotropic, there is significant and complex anisotropy in bending modulus that also differs between the monolayers, with extremal values not necessarily occurring along the principal directions. In particular, the commonly adopted orthotropic continuum plate model with uniform thickness fails to describe the observed variations in bending modulus for GeS, TiS3, and As2S3. We determine the direction-dependent effective thickness for use in such continuum models. We also show that the anisotropy in bending modulus is not associated with the rehybridization of atomic orbitals.

Mechanical properties and structure-function trade-offs in secondary xylem of young roots and stems.

Bending and torsional properties of young roots and stems were measured in nine woody angiosperms. The variation in mechanical parameters was correlated to wood anatomical traits and analysed with respect to the other two competing functions of xylem (namely storage and hydraulics). Compared with stems, roots exhibited five times greater flexibility in bending and two times greater flexibility in torsion. Lower values of structural bending and structural torsional moduli (Estr and Gstr, respectively) of roots compared with stems were associated with the presence of thicker bark and a greater size of xylem cells. Across species, Estr and Gstr were correlated with wood density, which was mainly driven by the wall thickness to lumen area ratio of fibres. Higher fractions of parenchyma did not translate directly into a lower wood density and reduced mechanical stiffness in spite of parenchyma cells having thinner, and in some cases less lignified, cell walls than fibres. The presence of wide, partially non-lignified rays contributed to low values of Estr and Gstr in Clematis vitalba. Overall, our results demonstrate that higher demands for mechanical stability in self-supporting stems put a major constraint on xylem structure, whereas root xylem can be designed with a greater emphasis on both storage and hydraulic functions.

HIV-1 matrix-31 membrane binding peptide interacts differently with membranes containing PS vs. PI(4,5)P2.

Efficient assembly of HIV-1 at the plasma membrane (PM) of the T-cell specifically requires PI(4,5)P2. It was previously shown that a highly basic region (HBR) of the matrix protein (MA) on the Gag precursor polyprotein Pr55Gag is required for membrane association. MA is N-terminally myristoylated, which enhances its affinity to membranes. In this work we used X-ray scattering and neutron reflectivity to determine how the physical properties and structure of lipid bilayers respond to the addition of binding domain peptides, either in the myristoylated form (MA31myr) or without the myristoyl group (MA31). Neutron reflectivity measurements showed the peptides predominantly located in the hydrocarbon interior. Diffuse X-ray scattering showed differences in membrane properties upon addition of peptides and the direction of the changes depended on lipid composition. The PI(4,5)P2-containing bilayers softened, thinned and became less ordered as peptide concentration increased. In contrast, POPS-containing bilayers with equivalent net charge first stiffened, thickened and became more ordered with increasing peptide concentration. As softening the host cell's PM upon contact with the protein lowers the free energy for membrane restructuring, thereby potentially facilitating budding of viral particles, our results suggest that the role of PI(4,5)P2 in viral assembly goes beyond specific stereochemical membrane binding. These studies reinforce the importance of lipids in virology.

Strong Resistance to Bending Observed for Nanoparticle Membranes.

We demonstrate how gold nanoparticle monolayers can be curled up into hollow scrolls that make it possible to extract both bending and stretching moduli from indentation by atomic force microscopy. We find a bending modulus that is 2 orders of magnitude larger than predicted by standard continuum elasticity, an enhancement we associate with nonlocal microstructural constraints. This finding opens up new opportunities for independent control of resistance to bending and stretching at the nanoscale.

Trehalose-Induced Variation in Mechanical Properties of Vesicles in Aqueous Solution.

The effect of the trehalose incorporation on the nanomechanical properties of dipalmitoylphosphatidylcholine vesicles was studied using atomic force microscope (AFM) on mica surface. The vesicles were prepared only with the variation in the trehalose concentration and adsorbed on the mica surface. After the morphology of the adsorbed vesicles was characterized, the behavior of an AFM tip into the vesicle was monitored using the plot of the tip displacement versus the tip deflection. It was observed that the breakthrough of the tip into the vesicles occurred two times. Each breakthrough represented each penetration of the tip into each layer. Force data prior to the first breakthrough fitted well with the Hertzian model to estimate Young's modulus and bending modulus of the vesicles. It was found that the Young's modulus and bending modulus decreased proportionally to the increase in the trehalose concentration up to 0.5 of trehalose to lipid. However, above 0.5, the moduli were a little varied with the increase. In the identical measurements at glucose, just a slight change in the moduli was observed with the increase in the glucose composition from 0 % glucose up to even 2:1 ratio of glucose:lipid. These results in the mechanical properties seem attributable to the osmotic and volumetric effects on the headgroup packing disruption.

Bending properties of a macroalga: Adaptation of Peirce's cantilever test for in situ measurements of Laminaria digitata (Laminariaceae).

• Premise of the study: The mechanical properties of a plant are key variables governing the interaction between the plant and its environment. Thus, measuring variables such as the flexural rigidity (bending) of a plant element is necessary to understand and predict the plant-flow interaction. However, plant elements such as macrophyte blades can be relatively thin and flexible, thus difficult to characterize. Different adaptations of the classical 3-point bending tests can also affect the interpretation of the flexural rigidity of an element. A simple, robust, method is newly applied to a biomaterial and validated here as an alternative to measure flexural rigidity of thin, flexible plant elements.• Methods: Based on a bending test procedure developed for the textile industry, an apparatus for in-situ measurements was developed and compared with other normalized methods, then used in a field test on the blade of a marine macroalga (Laminaria digitata) to assess its suitability to measure the bending modulus of a biomaterial.• Key results: Results of the presented method on selected surrogate materials agree with a normalized cantilever method (ISO 9073-7:1998) and 3-point bending test (ISO 178:2010). Values determined for the bending moduli for blades of L. digitaria were in the typical range for algal material. The range of validity of the method is discussed.• Conclusion: By validating this method with existing norms, this study suggests a better approach to measure bending properties of different biomaterials in the field compared with more traditional bending tests and opens new possibilities.

Computing the influence of lipids and lipid complexes on membrane mechanics.

Methodology for extracting the spontaneous curvature, bending modulus, and neutral surface of a lipid bilayer is described. The "SPEX" method is a robust technique for computing the bilayer bending modulus while allowing for resolution of the spontaneous curvature of specific interacting lipids and complexes, and the dependence of spontaneous curvature on wavelength. The method is described referring to the publicly available MembraneAnalysis.jl software package.

Effect of drought stress on bending stiffness in petioles of Caladium bicolor (Araceae).

PREMISE OF THE STUDY: Cell turgor plays an important role in the mechanical stability of herbaceous plants. This study on petioles of Caladium bicolor 'Candyland' analyzes the correlation between flexural rigidity and cell turgor. The results offer new insights into the underlying form-structure-function relationship and the dependency of mechanical properties from water availability. METHODS: Bending modulus E of petioles is calculated from two-point bending tests, taking into account the tapering mode. The corresponding turgor of parenchyma cells during wilting is investigated by pressure probe tests. KEY RESULTS: Wilting petioles show highly significant lower values of E than petioles with sufficient water supply. These differences are also found when comparing well-watered petioles to drought-stressed petioles having parenchyma turgor values in the same range. These results indicate an additional mechanical system sensitive to drought stress. On the basis of analyses of the contribution of different petiolar tissues toward the axial second moment of area and by using experimentally determined and literature values of E for the different tissues, we were able to (1) recalculate E of the intact petiole and to compare it with experimental data and (2) quantitatively estimate the importance of the different tissues for flexural rigidity and E of the petiole. CONCLUSIONS: Our results show that the decrease in flexural rigidity of petioles of Caladium bicolor 'Candyland' during wilting results from (1) a water-loss-induced decrease in mechanical efficiency of collenchyma fibers and (2) turgor loss of parenchyma cells.

Flexural Properties in Edgewise Bending of LVL Reinforced with Woven Carbon Fibers.

This paper presents the results of experimental testing of the bending strength and modulus of elasticity in edgewise bending of unreinforced and reinforced seven-layer LVL (laminated veneer lumber) poplar veneer panels. The aim of the research is to determine the influence of woven carbon fibers on the improvement of the bending properties and modulus of elasticity of LVL bending in the plane of the plate, as well as the influence of adhesives on the bending properties of the composite product, in order to test the potential of using this newly obtained material as a structural element. Bending was performed on small-scale samples. The main research task is the examination of three types of reinforcement, which differ from each other in position, orientation, and number of layers of reinforcement, using two different types of adhesives: epoxy adhesive and Melamine Urea Formaldehyde Resins (MUF). The composite material was produced in four different combinations in relation to the orientation and position of the reinforcement in the layup. The applied reinforcement is defined through three different configurations (EK1, EK2, and EK3) and a fourth control sample (EK4). Each configuration was produced by applying the two previously mentioned types of adhesives. The research findings showed that in the case of samples produced by applying CFRP (carbon fiber reinforced polymer) using epoxy adhesive, it significantly affected the increase in bending strength and flexural modulus of elasticity. The average improvement in bending strength is 32.9%, 33.2%, and 38.7%, i.e., the flexural modulus of elasticity is 54.1%, 50.7%, and 54.7%, respectively, for configurations EK1, EK2, and EK3, compared to control sample EK4. During the testing, the test samples from reinforced panels EK1 and EK2 showed partly plastic behavior up to the fracture point, while the diagram for the test samples from reinforced panels EK3 shows elastic behavior to a considerable extent, with a significantly smaller plastic behavior zone. This research proved the impossibility of using melamine-urea formaldehyde adhesive to form a composite product based on veneer and carbon fabric. The greatest contribution of this work is the experimentally verified and confirmed result of the possibility of applying poplar veneer to design structural elements in LVL using epoxy adhesive.

Bending modulus of the rippled graphene: the role of thickness.

Bending modulus is a key parameter to characterize the stiffness of materials. Commonly, it is believed that the bending modulus is closely related to the thickness as described by the thin plate theory. However, the thin plate theory fails in multilayer van der Waals materials like multilayer graphene, suggesting a more complex relationship between the bending modulus and thickness. Here, rippled graphene structures containing non-hexagonal carbon rings with different thicknesses are constructed to study the thickness-dependent bending modulus by the first-principles calculations. It is found that the bending modulus of rippled graphene depends on several factors, such as geometry, bending curvature, and thickness. Particularly, for the egg-tray graphene structures with similar structural pattern and bending curvature, i.e., eliminating the effects of structural pattern and bending curvature, the bending modulus could show a linear relationship to the thickness. Moreover, this linear relationship is very robust even in the case of changing the thickness through heteroatom doping.

Charting the twist-to-bend ratio of plant axes.

During the evolution of land plants many body plans have been developed. Differences in the cross-sectional geometry and tissue pattern of plant axes influence their flexural rigidity, torsional rigidity and the ratio of both of these rigidities, the so-called twist-to-bend ratio. For comparison, we have designed artificial cross-sections with various cross-sectional geometries and patterns of vascular bundles, collenchyma or sclerenchyma strands, but fixed percentages for these tissues. Our mathematical model allows the calculation of the twist-to-bend ratio by taking both cross-sectional geometry and tissue pattern into account. Each artificial cross-section was placed into a rigidity chart to provide information about its twist-to-bend ratio. In these charts, artificial cross-sections with the same geometry did not form clusters, whereas those with similar tissue patterns formed clusters characterized by vascular bundles, collenchyma or sclerenchyma arranged as one central strand, as a peripheral closed ring or as distributed individual strands. Generally, flexural rigidity increased the more the bundles or fibre strands were placed at the periphery. Torsional rigidity decreased the more the bundles or strands were separated and the less that they were arranged along a peripheral ring. The calculated twist-to-bend ratios ranged between 0.85 (ellipse with central vascular bundles) and 196 (triangle with individual peripheral sclerenchyma strands).

Synthesis of a Novel Hyperbranched Polyimide for Reinforcing Toughness and Insulating Properties of Bismaleimide Resin.

Bismaleimide (BMI) resin has great potential in aerospace, electronic, and machinery fields due to its extraordinary thermal stability. Owing to BMI's lower impact strength, various modified BMI resins have been prepared using CTBN, PEEK, fillers, and hyperbranched polymer to achieve higher impact strength. However, enhancement of toughness causes deterioration of other performance, such as Tg, thermal stability, and brittleness. In this work, BMI resin modified by hyperbranched polyimide (HBPI) was obtained. HBPI designed with flexible segments, unsaturated bonds, and a low degree of branching was synthesized. FT-IR and 13C-NMR were applied to confirm the successful fabrication of HBPI. The mechanical strength and dielectric properties of cured BMI resin containing various levels of HBPI were analyzed systematically. The impact and bending strength were improved significantly with increased HBPI content. When the content of HBPI is 40 wt.%, the impact strength and bending strength reach the maximum value of 32 kJ/mm and 88 MPa. In addition, the BMI cured with HBPI exhibits enhanced bending modulus to the value of 5.9 GPa. Furthermore, the dielectric strength of cured resin was improved to 28.3 kV/mm. The improved mechanical strength and enhanced dielectric properties are attributed to the increasing free volume induced by HBPI. These results indicate the promise of BMI resin modified by HBPI applied in insulating coatings and low dielectric laminates used in high frequency.

Mechanical Characterization of Two-Segment Free-Standing ZnO Nanowires Using Lateral Force Microscopy.

Mechanical characterization of quasi one-dimensional nanostructures is essential for the design of novel nanoelectromechanical systems. However, the results obtained on basic mechanical quantities, such as Young's modulus and fracture strength, show significant standard deviation in the literature. This is partly because of diversity in the quality of the nanowire, and partly because of inappropriately performed mechanical tests and simplified mechanical models. Here we present orientation-controlled bending and fracture studies on wet chemically grown vertical ZnO nanowires, using lateral force microscopy. The lateral force signal of the atomic force microscope was calibrated by a diamagnetic levitation spring system. By acquiring the bending curves of 14 nanowires, and applying a two-segment mechanical model, an average bending modulus of 108 ± 17 GPa was obtained, which was 23% lower than the Young's modulus of bulk ZnO in the [0001] direction. It was also found that the average fracture strain and stress inside the nanowire was above 3.1 ± 0.3 % and 3.3 ± 0.3 GPa, respectively. However, the fracture of the nanowires was governed by the quality of the nanowire/substrate interface. The demonstrated technique is a relatively simple and productive way for the accurate mechanical characterization of vertical nanowire arrays.

Effect of the Membrane Composition of Giant Unilamellar Vesicles on Their Budding Probability: A Trade-Off between Elasticity and Preferred Area Difference.

The budding and division of artificial cells engineered from vesicles and droplets have gained much attention in the past few decades due to an increased interest in designing stimuli-responsive synthetic systems. Proper control of the division process is one of the main challenges in the field of synthetic biology and, especially in the context of the origin of life studies, it would be helpful to look for the simplest chemical and physical processes likely at play in prebiotic conditions. Here we show that pH-sensitive giant unilamellar vesicles composed of mixed phospholipid/fatty acid membranes undergo a budding process, internally fuelled by the urea-urease enzymatic reaction, only for a given range of the membrane composition. A gentle interplay between the effects of the membrane composition on the elasticity and the preferred area difference of the bilayer is responsible for the existence of a narrow range of membrane composition yielding a high probability for budding of the vesicles.

Value of models for membrane budding.

The budding of membranes and curvature generation is common to many forms of trafficking in cells. Clathrin-mediated endocytosis, as a prototypical example of trafficking, has been studied in great detail using a variety of experimental systems and methods. Recently, advances in experimental methods have led to great strides in insights on the molecular mechanisms and the spatiotemporal dynamics of the protein machinery associated with membrane curvature generation. These advances have been ably supported by computational models, which have given us insights into the underlying mechanical principles of clathrin-mediated endocytosis. On the other hand, targeted experimental perturbation of membranes has lagged behind that of proteins in cells. In this area, modeling is especially critical to interpret experimental measurements in a mechanistic context. Here, we discuss the contributions made by these models to our understanding of endocytosis and identify opportunities to strengthen the connections between models and experiments.

Elastic moduli of lipid membranes: Reproducibility of AFM measures.

Membrane elastic properties play a major role in membrane remodeling events, such as vesicle fusion and fission. They are also crucial in drug delivery by liposomes. Different experimental techniques are available to measure elastic properties. Among them, atomic force microscopy (AFM) presents the unique advantage of being directly applicable to nano-sized liposomes. Unfortunately, different AFM measures reported in the literature show little agreement among each other and are difficult to compare with measures of bending modulus obtained by other experimental techniques or by molecular simulations. In this work we determine the bending rigidity of Egg PC liposomes in terms of Young modulus via AFM measurements, using two different tip shapes and different cantilever force constants. We interpret the measures using the Hertz and Shell models, and observe a clear dependency of the Young modulus values on the tip properties and on the interpretative theory. The effect of the AFM tip shape is less important than the effect of the cantilever force constant, and the mathematical model has a major effect on the interpretation of the data. The Shell theory provides the closest agreement between AFM data and other experimental data for the membrane bending modulus. Finally, we compare the results to calculations of bending modulus from molecular dynamics simulations of membrane buckles. Simulations provide values of bending modulus consistent with literature data, but the agreement with AFM experiments is reasonable only for some specific experimental conditions.

An Analytical Model for Estimating Alveolar Wall Elastic Moduli From Lung Tissue Uniaxial Stress-Strain Curves.

The non-linear stress-strain behavior of uniaxially-stretched lung parenchyma is thought to be an emergent phenomenon arising from the ensemble behavior of its microscopic constituents. Such behavior includes the alignment and elongation of randomly oriented alveolar walls with initially flaccid fibers in the direction of strain. To account for the link between microscopic wall behavior and the macroscopic stress-strain curve, we developed an analytical model that represents both alignment and elongation of alveolar walls during uniaxial stretching. The model includes the kinetics and mechanical behavior of randomly oriented elastic alveolar walls that have a bending stiffness at their intersections. The alignment and stretch of the walls following incremental stretch of the tissue were determined based on energy minimization, and the total stress was obtained by differentiating the total energy density with respect to strain. The stress-strain curves predicted by the model were comparable to curves generated by a previously published numerical alveolar network model. The model was also fit to experimentally measured stress-strain curves in parenchymal strips obtained from mice with decreased lung collagen content, and from young and aged mice. This yielded estimates for the elastic modulus of an alveolar wall, which increased with age from 4.4 to 5.9 kPa (p = 0.043), and for the elastic modulus of fibers within the wall, which increased with age from 311 to 620 kPa (p = 0.001). This demonstrates the possibility of estimating alveolar wall mechanical properties in biological soft tissue from its macroscopic behavior given appropriate assumptions about tissue structure.

Mechanical Characterization of Liposomes and Extracellular Vesicles, a Protocol.

Both natural as well as artificial vesicles are of tremendous interest in biology and nanomedicine. Small vesicles (<200 nm) perform essential functions in cell biology and artificial vesicles (liposomes) are used as drug delivery vehicles. Atomic Force Microscopy (AFM) is a powerful technique to study the structural properties of these vesicles. AFM is a well-established technique for imaging at nanometer resolution and for mechanical measurements under physiological conditions. Here, we describe the procedure of AFM imaging and force spectroscopy on small vesicles. We discuss how to image vesicles with minimal structural disturbance, and how to analyze the data for accurate size and shape measurements. In addition, we describe the procedure for performing nanoindentations on vesicles and the subsequent data analysis including mechanical models used for data interpretation.

Material properties of interfacial films of mucoid and nonmucoid Pseudomonas aeruginosa isolates.

Chronic lung infection with bacterial biofilms is a leading cause of death in cystic fibrosis (CF) patients. Pseudomonas aeruginosa, one of the many species colonizing the lung airways, can undergo pathoadaptation, leading to a mucoid phenotype with interesting material properties. We hypothesize that the surface properties and extracellular materials of mucoid P. aeruginosa cells greatly influence the mechanical behavior of their films at fluid interfaces. In this study, we investigate the interfacial properties of films formed by nonmucoid (PANT) and mucoid (PASL) strains of P. aeruginosa isolated from CF patients. We use pendant drop elastometry to analyze the interfacial response of the films formed by PANT and PASL at the hexadecane-water interface. The dynamic rheological analyses of the films highlight the distinctive signature of the mucoid strains at fluid interfaces. The mucoid films exhibit greater relaxation following a compressive strain than a tensile one, while a full hysteresis response is achieved by the nonmucoid films; this indicates that the material properties of the PANT films are conserved under both compression and tension. The wrinkling and shape analyses of the interfacial bacterial films elucidate that the mucoid strain exhibits remarkable viscoelastic properties, enabling the remodeling of the living films and dissipation of the compressive stress. The comparative analysis of the material properties of mucoid and nonmucoid P. aeruginosa cells indicates that mucoid switch can play an important role in protecting the bacteria from interfacial stresses. Further characterization of interfacial bacterial films will provide new insights into the development of methods for controlling interfacial films of bacteria.

Highlighting the effect of amyloid beta assemblies on the mechanical properties and conformational stability of cell membrane.

Alzheimer disease (AD) is the most common cause of dementia, characterized by a progressive decline in cognitive function due to the abnormal aggregation and deposition of Amyloid beta (Aß) fibrils in the brain of patients. In this context, the molecular mechanisms of protein misfolding and aggregation that are known to induce significant biophysical alterations in cells, including destabilization of plasma membranes, remain partially unclear. Physical interaction between the Aß assemblies and the membrane leads to the disruption of the cell membrane in multiple ways including, surface carpeting, generation of transmembrane channels and detergent-like membrane dissolution. Understanding the impact of amyloidogenic protein in different stages of aggregation with the plasma membrane, plays a crucial role to fully elucidate the pathological mechanisms of AD. Within this framework, computer simulations represent a powerful tool able to shed lights on the interactions governing the structural influence of Aß proteins on biological membrane. In this study, molecular dynamics (MD) simulations have been performed in order to characterize how POPC bilayer conformational and mechanical properties are affected by the interaction with Aß11-42 peptide, oligomer and fibril.