At the crossroads of epidemiology and biology: Bridging the gap between SARS-CoV-2 viral strain properties and epidemic wave characteristics.

The COVID-19 pandemic has given rise to numerous articles from different scientific fields (epidemiology, virology, immunology, airflow physics …) without any effort to link these different insights. In this review, we aim to establish relationships between epidemiological data and the characteristics of the virus strain responsible for the epidemic wave concerned. We have carried out this study on the Wuhan, Alpha, Delta and Omicron strains allowing us to illustrate the evolution of the relationships we have highlighted according to these different viral strains. We addressed the following questions. 1) How can the mean infectious dose (one quantum, by definition in epidemiology) be measured and expressed as an amount of viral RNA molecules (in genome units, GU) or as a number of replicative viral particles (in plaque-forming units, PFU)? 2) How many infectious quanta are exhaled by an infected person per unit of time? 3) How many infectious quanta are exhaled, on average, integrated over the whole contagious period? 4) How do these quantities relate to the epidemic reproduction rate R as measured in epidemiology, and to the viral load, as measured by molecular biological methods? 5) How has the infectious dose evolved with the different strains of SARS-CoV-2? We make use of state-of-the-art modelling, reviewed and explained in the appendix of the article (Supplemental Information, SI), to answer these questions using data from the literature in both epidemiology and virology. We have considered the modification of these relationships according to the vaccination status of the population.

Risk assessment for long- and short-range airborne transmission of SARS-CoV-2, indoors and outdoors.

Preventive measures to reduce infection are needed to combat the COVID-19 pandemic and prepare for a possible endemic phase. Current prophylactic vaccines are highly effective to prevent disease but lose their ability to reduce viral transmission as viral evolution leads to increasing immune escape. Long-term proactive public health policies must therefore complement vaccination with available nonpharmaceutical interventions aiming to reduce the viral transmission risk in public spaces. Here, we revisit the quantitative assessment of airborne transmission risk, considering asymptotic limits that considerably simplify its expression. We show that the aerosol transmission risk is the product of three factors: a biological factor that depends on the viral strain, a hydrodynamical factor defined as the ratio of concentration in viral particles between inhaled and exhaled air, and a face mask filtering factor. The short-range contribution to the risk, present both indoors and outdoors, is related to the turbulent dispersion of exhaled aerosols by air drafts and by convection (indoors), or by the wind (outdoors). We show experimentally that airborne droplets and CO2 molecules present the same dispersion. As a consequence, the dilution factor, and therefore the risk, can be measured quantitatively using the CO2 concentration, regardless of the room volume, the flow rate of fresh air, and the occupancy. We show that the dispersion cone leads to a concentration in viral particles, and therefore a short-range transmission risk, inversely proportional to the squared distance to an infected person and to the flow velocity. The aerosolization criterion derived as an intermediate result, which compares the Stokes relaxation time to the Lagrangian time-scale, may find application for a broad class of aerosol-borne pathogens and pollutants.

A lower-than-expected saltation threshold at Martian pressure and below.

Aeolian sediment transport is observed to occur on Mars as well as other extraterrestrial environments, generating ripples and dunes as on Earth. The search for terrestrial analogs of planetary bedforms, as well as environmental simulation experiments able to reproduce their formation in planetary conditions, are powerful ways to question our understanding of geomorphological processes toward unusual environmental conditions. Here, we perform sediment transport laboratory experiments in a closed-circuit wind tunnel placed in a vacuum chamber and operated at extremely low pressures to show that Martian conditions belong to a previously unexplored saltation regime. The threshold wind speed required to initiate saltation is only quantitatively predicted by state-of-the art models up to a density ratio between grain and air of [Formula: see text] but unexpectedly falls to much lower values for higher density ratios. In contrast, impact ripples, whose emergence is continuously observed on the granular bed over the whole pressure range investigated, display a characteristic wavelength and propagation velocity essentially independent of pressure. A comparison of these findings with existing models suggests that sediment transport at low Reynolds number but high grain-to-fluid density ratio may be dominated by collective effects associated with grain inertia in the granular collisional layer.

Mechanics and Energetics of Electromembranes.

The recent discovery of electroactive polymers has shown great promises in the field of soft robotics and was logically followed by experimental, numerical, and theoretical developments. Most of these studies were concerned with systems entirely covered by electrodes. However, there is a growing interest for partially active polymers, in which the electrode covers only one part of the membrane. Indeed, such actuation can trigger buckling instabilities and so represents a route toward the control of three-dimensional shapes. Here, we study theoretically the behavior of such partially active electroactive polymer. We address two problems: (1) the electrostatic elastica including geometric nonlinearities and partially electroactive strip using a variational approach. We propose a new interpretation of the equations of deformation, by drawing analogies with biological growth, in which the effect of the electric voltage is seen as a change in the reference stress-free state. (2) We explain the nature of the distribution of electrostatic forces on this simple system, which is not trivial. In particular, we find that edge effects are playing a major role in this problem.

A flexible rheometer design to measure the visco-elastic response of soft solids over a wide range of frequency.

We present a flexible setup for determining the rheology of visco-elastic materials which is based on the mechanical response of a magnet deposited at the surface of a slab of material and excited electromagnetically. An interferometric measurement of the magnet displacement allows one to reach an excellent accuracy over a wide range of frequency. Except for the magnet, there is no contact between the material under investigation and the apparatus. At low frequency, inertial effects are negligible so that the mechanical response, obtained through a lock-in amplifier, directly gives the material complex modulus. At high frequency, damped waves are emitted and the rheology must be extracted numerically from a theoretical model. To validate the design, the instrument was used to measure the rheology of a test polydimethylsiloxane gel which presents an almost perfect scale free response at high frequency.

Peeling an elastic film from a soft viscoelastic adhesive: experiments and scaling laws.

The functionality of adhesives relies on their response under the application of a load. Yet, it has remained a challenge to quantitatively relate the macroscopic dynamics of peeling to the dissipative processes inside the adhesive layer. Here we investigate the peeling of a reversible adhesive made of a polymer gel, measuring the relationship between the peeling force, the peeling velocity, and the geometry of the interface at small-scale. Experiments are compared to a theory based on the linear viscoelastic response of the adhesive, augmented with an elastocapillary regularization approach. This theory, fully quantitative in the limit of small surface deformations, demonstrates the emergence of a "wetting" angle at the contact line and exhibits scaling laws for peeling which are in good agreement with the experimental results. Our findings provide a new strategy for design of reversible adhesives, by quantitatively combining wetting, geometry and dissipation.

Paradox of Contact Angle Selection on Stretched Soft Solids.

The interfacial mechanics of soft elastic networks plays a central role in biological and technological contexts. Yet, effects of solid capillarity have remained controversial, primarily due to the strain-dependent surface energy. Here we derive the equations that govern the selection of contact angles of liquid drops on elastic surfaces from variational principles. It is found that the substrate's elasticity imposes a nontrivial condition that relates pinning, hysteresis, and contact line mobility to the so-called Shuttleworth effect. We experimentally validate our theory for droplets on a silicone gel, revealing an enhanced contact line mobility when stretching the substrate.

Thermally activated motion of a contact line over defects.

At the nanometer scale, the motion of a contact line separating a dry from a wet region is limited by the presence of surface heterogeneities that pin it. Here we revisit the seminal model proposed by Joanny and de Gennes to include the influence of thermal noise and viscosity using a Langevin model with two degrees of freedom: the average position of the contact line and its distortion. We identify the conditions under which the dynamics in a velocity-driven experiment can in fact be described by a constant forcing at small scale. We then relate the asymptotic properties of the relation between force and contact line velocity to the properties of the defects. In particular, we show that Kramers' approximation misses the strong asymmetry between advancing and receding directions. Finally, we show how to use the model to fit experimental data and extract the salient features of the surface energy landscape.

Dynamical theory of the inverted cheerios effect.

Recent experiments have shown that liquid drops on highly deformable substrates exhibit mutual interactions. This is similar to the Cheerios effect, the capillary interaction of solid particles at a liquid interface, but now the roles of solid and liquid are reversed. Here we present a dynamical theory for this inverted Cheerios effect, taking into account elasticity, capillarity and the viscoelastic rheology of the substrate. We compute the velocity at which droplets attract, or repel, as a function of their separation. The theory is compared to a simplified model in which the viscoelastic dissipation is treated as a localized force at the contact line. It is found that the two models differ only at small separation between the droplets, and both of them accurately describe experimental observations.

Giant ripples on comet 67P/Churyumov-Gerasimenko sculpted by sunset thermal wind.

Explaining the unexpected presence of dune-like patterns at the surface of the comet 67P/Churyumov-Gerasimenko requires conceptual and quantitative advances in the understanding of surface and outgassing processes. We show here that vapor flow emitted by the comet around its perihelion spreads laterally in a surface layer, due to the strong pressure difference between zones illuminated by sunlight and those in shadow. For such thermal winds to be dense enough to transport grains-10 times greater than previous estimates-outgassing must take place through a surface porous granular layer, and that layer must be composed of grains whose roughness lowers cohesion consistently with contact mechanics. The linear stability analysis of the problem, entirely tested against laboratory experiments, quantitatively predicts the emergence of bedforms in the observed wavelength range and their propagation at the scale of a comet revolution. Although generated by a rarefied atmosphere, they are paradoxically analogous to ripples emerging on granular beds submitted to viscous shear flows. This quantitative agreement shows that our understanding of the coupling between hydrodynamics and sediment transport is able to account for bedform emergence in extreme conditions and provides a reliable tool to predict the erosion and accretion processes controlling the evolution of small solar system bodies.

A moving contact line as a rheometer for nanometric interfacial layers.

How a liquid drop sits or moves depends on the physical and mechanical properties of the underlying substrate. This can be seen in the hysteresis of the contact angle made by a drop on a solid, which is known to originate from surface heterogeneities, and in the slowing of droplet motion on deformable solids. Here, we show how a moving contact line can be used to characterize a molecularly thin polymer layer on a solid. We find that the hysteresis depends on the polymerization index and can be optimized to be vanishingly small (<0.07°). The mechanical properties are quantitatively deduced from the microscopic contact angle, which is proportional to the speed of the contact line and the Rouse relaxation time divided by the layer thickness, in agreement with theory. Our work opens the prospect of measuring the properties of functionalized interfaces in microfluidic and biomedical applications that are otherwise inaccessible.

Liquid drops attract or repel by the inverted Cheerios effect.

Solid particles floating at a liquid interface exhibit a long-ranged attraction mediated by surface tension. In the absence of bulk elasticity, this is the dominant lateral interaction of mechanical origin. Here, we show that an analogous long-range interaction occurs between adjacent droplets on solid substrates, which crucially relies on a combination of capillarity and bulk elasticity. We experimentally observe the interaction between droplets on soft gels and provide a theoretical framework that quantitatively predicts the interaction force between the droplets. Remarkably, we find that, although on thick substrates the interaction is purely attractive and leads to drop-drop coalescence, for relatively thin substrates a short-range repulsion occurs, which prevents the two drops from coming into direct contact. This versatile interaction is the liquid-on-solid analog of the "Cheerios effect." The effect will strongly influence the condensation and coarsening of drops on soft polymer films, and has potential implications for colloidal assembly and mechanobiology.

Defects at the Nanoscale Impact Contact Line Motion at all Scales.

The contact angle of a liquid drop moving on a real solid surface depends on the speed and direction of motion of the three-phase contact line. Many experiments have demonstrated that pinning on surface defects, thermal activation and viscous dissipation impact contact line dynamics, but so far, efforts have failed to disentangle the role of each of these dissipation channels. Here, we propose a unifying multiscale approach that provides a single quantitative framework. We use this approach to successfully account for the dynamics measured in a classic dip-coating experiment performed over an unprecedentedly wide range of velocity. We show that the full contact line dynamics up to the liquid film entrainment threshold can be parametrized by the size, amplitude and density of nanometer-scale defects. This leads us to reinterpret the contact angle hysteresis as a dynamical crossover rather than a depinning transition.

Solid capillarity: when and how does surface tension deform soft solids?

Soft solids differ from stiff solids in an important way: their surface stresses can drive large deformations. Based on a topical workshop held in the Lorentz Center in Leiden, this Opinion highlights some recent advances in the growing field of solid capillarity and poses key questions for its advancement.

Non-local rheology in dense granular flows: Revisiting the concept of fluidity.

The aim of this article is to discuss the concepts of non-local rheology and fluidity, recently introduced to describe dense granular flows. We review and compare various approaches based on different constitutive relations and choices for the fluidity parameter, focusing on the kinetic elasto-plastic model introduced by Bocquet et al. (Phys. Rev. Lett 103, 036001 (2009)) for soft matter, and adapted for granular matter by Kamrin et al. (Phys. Rev. Lett. 108, 178301 (2012)), and the gradient expansion of the local rheology µ(I) that we have proposed (Phys. Rev. Lett. 111, 238301 (2013)). We emphasise that, to discriminate between these approaches, one has to go beyond the predictions derived from linearisation around a uniform stress profile, such as that obtained in a simple shear cell. We argue that future tests can be based on the nature of the chosen fluidity parameter, and the related boundary conditions, as well as the hypothesis made to derive the models and the dynamical mechanisms underlying their dynamics.

Direct numerical simulations of aeolian sand ripples.

Aeolian sand beds exhibit regular patterns of ripples resulting from the interaction between topography and sediment transport. Their characteristics have been so far related to reptation transport caused by the impacts on the ground of grains entrained by the wind into saltation. By means of direct numerical simulations of grains interacting with a wind flow, we show that the instability turns out to be driven by resonant grain trajectories, whose length is close to a ripple wavelength and whose splash leads to a mass displacement toward the ripple crests. The pattern selection results from a compromise between this destabilizing mechanism and a diffusive downslope transport which stabilizes small wavelengths. The initial wavelength is set by the ratio of the sediment flux and the erosion/deposition rate, a ratio which increases linearly with the wind velocity. We show that this scaling law, in agreement with experiments, originates from an interfacial layer separating the saltation zone from the static sand bed, where momentum transfers are dominated by midair collisions. Finally, we provide quantitative support for the use of the propagation of these ripples as a proxy for remote measurements of sediment transport.

Capillarity of soft amorphous solids: a microscopic model for surface stress.

The elastic deformation of a soft solid induced by capillary forces crucially relies on the excess stress inside the solid-liquid interface. While for a liquid-liquid interface this "surface stress" is strictly identical to the "surface free energy," the thermodynamic Shuttleworth equation implies that this is no longer the case when one of the phases is elastic. Here we develop a microscopic model that incorporates enthalpic interactions and entropic elasticity, based on which we explicitly compute as the surface stress and surface free energy. It is found that the compressibility of the interfacial region, through the Poisson ratio near the interface, determines the difference between surface stress and surface energy. We highlight the consequence of this finding by comparing with recent experiments and simulations on partially wetted soft substrates.

Aeolian and subaqueous bedforms in shear flows.

A sediment bed sheared by an unbounded flow is unconditionally unstable towards the growth of bedforms called ripples under water and dunes in the aeolian case. We review here the dynamical mechanisms controlling this linear instability, putting the emphasis on testing models against field and laboratory measurements. We then discuss the role of nonlinearities and the influence of finite size effects, namely the depth of the atmospheric boundary layer in the aeolian case and the water depth in the case of rivers.

Nonlocal rheology of granular flows across yield conditions.

The rheology of dense granular flows is studied numerically in a shear cell controlled at constant pressure and shear stress, confined between two granular shear flows. We show that a liquid state can be achieved even far below the yield stress, whose flow can be described with the same rheology as above the yield stress. A nonlocal constitutive relation is derived from dimensional analysis through a gradient expansion and calibrated using the spatial relaxation of velocity profiles observed under homogeneous stresses. Both for frictional and frictionless grains, the relaxation length is found to diverge as the inverse square root of the distance to the yield point, on both sides of that point.