Rheology of saliva in health and disease.

BACKGROUND: Saliva is a complex fluid that lubricates the oropharynx and facilitates chewing, swallowing, and vocalization. Viscoelasticity is critical for the ability of saliva to fulfill these functions. Xerostomia, or a sensation of dry mouth, occurs in 17-26% of the population. Although many equate xerostomia with hyposalivation, high-risk patients frequently report oral dryness in the absence of decreased salivary flow. OBJECTIVE: This study aims to determine if xerostomia is associated with alterations in the rheological properties of saliva in addition to decreased salivary production. METHODS: The study population included patients with post-radiation xerostomia, patients with anticholinergic-induced xerostomia and healthy controls. Salivary volumetric flow rate was measured, shear viscosity was measured using oscillatory rheometry, and extensional viscosity was measured using capillary thinning methods. Groups were compared using descriptive statistics and univariate analysis. RESULTS: A total of 36 subjects were included: 15 with post-radiation xerostomia, 9 with anticholinergic-induced xerostomia and 12 controls. Salivary volumetric flow was significantly decreased in post-radiation and anticholinergic-induced patients compared to controls. On capillary thinning testing, saliva from xerostomia patients had significantly greater extensional viscosity compared to controls. However, saliva from the three groups showed no significant difference in the complex viscosity or the storage or loss modulus of saliva with oscillatory rheology. CONCLUSIONS: Xerostomia is associated with decreased salivary volumetric flow and quantitative changes in the rheologic properties of saliva.

Structure-property relationships from universal signatures of plasticity in disordered solids.

When deformed beyond their elastic limits, crystalline solids flow plastically via particle rearrangements localized around structural defects. Disordered solids also flow, but without obvious structural defects. We link structure to plasticity in disordered solids via a microscopic structural quantity, "softness," designed by machine learning to be maximally predictive of rearrangements. Experimental results and computations enabled us to measure the spatial correlations and strain response of softness, as well as two measures of plasticity: the size of rearrangements and the yield strain. All four quantities maintained remarkable commonality in their values for disordered packings of objects ranging from atoms to grains, spanning seven orders of magnitude in diameter and 13 orders of magnitude in elastic modulus. These commonalities link the spatial correlations and strain response of softness to rearrangement size and yield strain, respectively.

Running and tumbling with E. coli in polymeric solutions.

Run-and-tumble motility is widely used by swimming microorganisms including numerous prokaryotic and eukaryotic organisms. Here, we experimentally investigate the run-and-tumble dynamics of the bacterium E. coli in polymeric solutions. We find that even small amounts of polymer in solution can drastically change E. coli dynamics: cells tumble less and their velocity increases, leading to an enhancement in cell translational diffusion and a sharp decline in rotational diffusion. We show that suppression of tumbling is due to fluid viscosity while the enhancement in swimming speed is mainly due to fluid elasticity. Visualization of single fluorescently labeled DNA polymers reveals that the flow generated by individual E. coli is sufficiently strong to stretch polymer molecules and induce elastic stresses in the fluid, which in turn can act on the cell in such a way to enhance its transport. Our results show that the transport and spread of chemotactic cells can be independently modified and controlled by the fluid material properties.

Flagellar kinematics and swimming of algal cells in viscoelastic fluids.

The motility of microorganisms is influenced greatly by their hydrodynamic interactions with the fluidic environment they inhabit. We show by direct experimental observation of the bi-flagellated alga Chlamydomonas reinhardtii that fluid elasticity and viscosity strongly influence the beating pattern - the gait - and thereby control the propulsion speed. The beating frequency and the wave speed characterizing the cyclical bending are both enhanced by fluid elasticity. Despite these enhancements, the net swimming speed of the alga is hindered for fluids that are sufficiently elastic. The origin of this complex response lies in the interplay between the elasticity-induced changes in the spatial and temporal aspects of the flagellar cycle and the buildup and subsequent relaxation of elastic stresses during the power and recovery strokes.

Tunable capillary-induced attraction between vertical cylinders.

Deformation of a fluid interface caused by the presence of objects at the interface can lead to large lateral forces between objects. We explore these fluid-mediated attractive force between partially submerged vertical cylinders. Forces are experimentally measured by slowly separating cylinder pairs and cylinder triplets after capillary rise is initially established for cylinders in contact. For cylinder pairs, numerical computations and a theoretical model are found to be in good agreement with measurements. The model provides insight into the relative importance of the contributions to the total force. For small separations, the lateral force is dominated by the fluid pressure acting over the wetted cylinder surfaces. At large separations, the surface tension acting along the contact line dominates the lateral force. A crossover between the two regimes occurs at a separation of around half of a capillary length. The experimentally measured forces between cylinder triplets are also in good agreement with numerical computations, and we show that pairwise contributions account for nearly all of the attractive force between triplets. For cylinders with an equilibrium capillary rise height greater than the height of the cylinder, we find that the attractive force depends on the height of the cylinders above the submersion level, which provides a means to create precisely controlled tunable cohesive forces between objects deforming a fluid interface.

Rheology of soft colloids across the onset of rigidity: scaling behavior, thermal, and non-thermal responses.

We study the rheological behavior of colloidal suspensions composed of soft sub-micron-size hydrogel particles across the liquid-solid transition. The measured stress and strain-rate data, when normalized by thermal stress and time scales, suggest our systems reside in a regime wherein thermal effects are important. In a different vein, critical point scaling predictions for the jamming transition, typical in athermal systems, are tested. Near dynamic arrest, the suspensions exhibit scaling exponents similar to those reported in Nordstrom et al., Phys. Rev. Lett., 2010, 105, 175701. The observation suggests that our system exhibits a glass transition near the onset of rigidity, but it also exhibits a jamming-like scaling further from the transition point. These observations are thought-provoking in light of recent theoretical and simulation findings, which show that suspension rheology across the full range of microgel particle experiments can exhibit both thermal and athermal mechanisms.

Nonlinear elastic instability in channel flows at low Reynolds numbers.

It is presently believed that flows of viscoelastic polymer solutions in geometries such as a straight pipe or channel are linearly stable. Here we present experimental evidence that such flows can be nonlinearly unstable and can exhibit a subcritical bifurcation. Velocimetry measurements are performed in a long, straight microchannel; flow disturbances are introduced at the entrance of the channel system by placing a variable number of obstacles. Above a critical flow rate and a critical size of the perturbation, a sudden onset of large velocity fluctuations indicates the presence of a nonlinear subcritical instability. Together with the previous observations of hydrodynamic instabilities in curved geometries, our results suggest that any flow of polymer solutions becomes unstable at sufficiently high flow rates.

Rheology of human blood plasma: viscoelastic versus Newtonian behavior.

We investigate the rheological characteristics of human blood plasma in shear and elongational flows. While we can confirm a Newtonian behavior in shear flow within experimental resolution, we find a viscoelastic behavior of blood plasma in the pure extensional flow of a capillary breakup rheometer. The influence of the viscoelasticity of blood plasma on capillary blood flow is tested in a microfluidic device with a contraction-expansion geometry. Differential pressure measurements revealed that the plasma has a pronounced flow resistance compared to that of pure water. Supplementary measurements indicate that the viscoelasticity of the plasma might even lead to viscoelastic instabilities under certain conditions. Our findings show that the viscoelastic properties of plasma should not be ignored in future studies on blood flow.

Undulatory locomotion of Caenorhabditis elegans on wet surfaces.

The physical and biomechanical principles that govern undulatory movement on wet surfaces have important applications in physiology, physics, and engineering. The nematode Caenorhabditis elegans, with its highly stereotypical and functionally distinct sinusoidal locomotory gaits, is an excellent system in which to dissect these properties. Measurements of the main forces governing the C. elegans crawling gait on lubricated surfaces have been scarce, primarily due to difficulties in estimating the physical features at the nematode-gel interface. Using kinematic data and a hydrodynamic model based on lubrication theory, we calculate both the surface drag forces and the nematode's bending force while crawling on the surface of agar gels within a preexisting groove. We find that the normal and tangential surface drag coefficients during crawling are ∼222 and 22, respectively, and the drag coefficient ratio is ∼10. During crawling, the calculated internal bending force is time-periodic and spatially complex, exhibiting a phase lag with respect to the nematode's body bending curvature. This phase lag is largely due to viscous drag forces, which are higher during crawling as compared to swimming in an aqueous buffer solution. The spatial patterns of bending force generated during either swimming or crawling correlate well with previously described gait-specific features of calcium signals in muscle. Further, our analysis indicates that one may be able to control the motility gait of C. elegans by judiciously adjusting the magnitude of the surface drag coefficients.

Undulatory swimming in viscoelastic fluids.

The effects of fluid elasticity on the swimming behavior of the nematode Caenorhabditis elegans are experimentally investigated by tracking the nematode's motion and measuring the corresponding velocity fields. We find that fluid elasticity hinders self-propulsion. Compared to Newtonian solutions, fluid elasticity leads to up to 35% slower propulsion. Furthermore, self-propulsion decreases as elastic stresses grow in magnitude in the fluid. This decrease in self-propulsion in viscoelastic fluids is related to the stretching of flexible molecules near hyperbolic points in the flow.

Material properties of Caenorhabditis elegans swimming at low Reynolds number.

Undulatory locomotion, as seen in the nematode Caenorhabditis elegans, is a common swimming gait of organisms in the low Reynolds number regime, where viscous forces are dominant. Although the nematode's motility is expected to be a strong function of its material properties, measurements remain scarce. Here, the swimming behavior of C. elegans is investigated in experiments and in a simple model. Experiments reveal that nematodes swim in a periodic fashion and generate traveling waves that decay from head to tail. The model is able to capture the experiments' main features and is used to estimate the nematode's Young's modulus E and tissue viscosity eta. For wild-type C. elegans, we find E approximately 3.77 kPa and eta approximately -860 Pa.s; values of eta for live C. elegans are negative because the tissue is generating rather than dissipating energy. Results show that material properties are sensitive to changes in muscle functional properties, and are useful quantitative tools with which to more accurately describe new and existing muscle mutants.

Microfluidic rheology of soft colloids above and below jamming.

The rheology near jamming of a suspension of soft colloidal spheres is studied using a custom microfluidic rheometer that provides the stress versus strain rate over many decades. We find non-Newtonian behavior below the jamming concentration and yield-stress behavior above it. The data may be collapsed onto two branches with critical scaling exponents that agree with expectations based on Hertzian contacts and viscous drag. These results support the conclusion that jamming is similar to a critical phase transition, but with interaction-dependent exponents.

Stretching fields and mixing near the transition to nonperiodic two-dimensional flow.

Although time-periodic fluid flows sometimes produce mixing via Lagrangian chaos, the additional contribution to mixing caused by nonperiodicity has not been quantified experimentally. Here, we do so for a quasi-two-dimensional flow generated by electromagnetic forcing. Several distinct measures of mixing are found to vary continuously with the Reynolds number, with no evident change in magnitude or slope at the onset of nonperiodicity. Furthermore, the scaled probability distributions of the mean Lyapunov exponent have the same form in the periodic and nonperiodic flow states.

Polymeric filament thinning and breakup in microchannels.

The effects of elasticity on filament thinning and breakup are investigated in microchannel cross flow. When a viscous solution is stretched by an external immiscible fluid, a low 100 ppm polymer concentration strongly affects the breakup process, compared to the Newtonian case. Qualitatively, polymeric filaments show much slower evolution, and their morphology features multiple connected drops. Measurements of filament thickness show two main temporal regimes: flow- and capillary-driven. At early times both polymeric and Newtonian fluids are flow-driven, and filament thinning is exponential. At later times, Newtonian filament thinning crosses over to a capillary-driven regime, in which the decay is algebraic. By contrast, the polymeric fluid first crosses over to a second type of flow-driven behavior, in which viscoelastic stresses inside the filament become important and the decay is again exponential. Finally, the polymeric filament becomes capillary-driven at late times with algebraic decay. We show that the exponential flow thinning behavior allows a measurement of the extensional viscosities of both Newtonian and polymeric fluids.

Elastic instabilities of polymer solutions in cross-channel flow.

When polymer molecules pass near the hyperbolic point of a microchannel cross flow, they are strongly stretched. As the strain rate is varied at low Reynolds number (< 10(-2)), tracer and particle-tracking experiments show that molecular stretching produces two flow instabilities: one in which the velocity field becomes strongly asymmetric, and a second in which it fluctuates nonperiodically in time. The flow is strongly perturbed even far from the region of instability, and this phenomenon can be used to produce mixing.

Predicting the progress of diffusively limited chemical reactions in the presence of chaotic advection.

The effects of chaotic advection and diffusion on fast chemical reactions in two-dimensional fluid flows are investigated using experimentally measured stretching fields and fluorescent monitoring of the local concentration. Flow symmetry, Reynolds number, and mean path length affect the spatial distribution and time dependence of the reaction product. A single parameter lambdaN , where lambda is the mean Lyapunov exponent N and is the number of mixing cycles, can be used to predict the time-dependent total product for flows having different dynamical features.