Experimental evaluation of the avoidance reaction of Calanus finmarchicus

The avoidance reaction of Calanus finmarchicus copepodids wasobserved in the presence of quantifiable fluid mechanical signals.The experimental apparatus was a rotating cylinder of sea waterwith fixed obstacles (diameters 6 mm and 10 mm) which disturbedthe flow and stimulated avoidance. The experimental conditionsprevented unambiguous discrimination of signals; the amountand rate of fluid deformation appear to be the effective stimuli.Avoidance reactions began at a mean distance of 7 mm from thesmall obstacle and 6 mm from the large obstacle. Mean valuesfor fluid deformation for the small and large obstacles were0.12 and 0.24 mm/mm; fluid deformation rate values were 0.80and 1.0 sec^–1. There was less scatter for deformationvalues. The copepods preferentially avoided obstacles at anglesof about 60° with respect to the approach streamline. Avoidancespeeds averaged 16 cm.sec^–1; mean distance of avoidanceflights was 15 mm. ¹Contribution Number 4195 from the Woods Hole OceanographicInstitution. ²Present Address: Scripps Institution of Oceanography, La Jolla,California 92093. ³Present Address: Marathon Oil Company, Littleton, Colorado80160.     

Application of stereolithographic custom models for studying the impact of biofilms and mineral precipitation on fluid flow

Here we introduce the use of transparent experimental models fabricated by stereolithography for studying the impacts of biomass accumulation, minerals precipitation, and physical configuration of flow paths on liquid flow in fracture apertures. The internal configuration of the models ranged in complexity from simple geometric shapes to those that incorporate replicated surfaces of natural fractures and computationally derived fracture surfaces. High-resolution digital time-lapse imaging was employed to qualitatively observe the migration of colloidal and soluble dyes through the flow models. In this study, a Sphingomonas sp. and Sporosarcina (Bacillus) pasteurii influenced the fluid dynamics by physically altering flow paths. Microbial colonization and calcite deposition enhanced the stagnant regions adjacent to solid boundaries. Microbial growth and calcite precipitation occurred to a greater extent in areas behind the fabricated obstacles and less in high-velocity orifices.     

Application of Stereolithographic Custom Models for Studying the Impact of Biofilms and Mineral Precipitation on Fluid Flow

Here we introduce the use of transparent experimental models fabricated by stereolithography for studying the impacts of biomass accumulation, minerals precipitation, and physical configuration of flow paths on liquid flow in fracture apertures. The internal configuration of the models ranged in complexity from simple geometric shapes to those that incorporate replicated surfaces of natural fractures and computationally derived fracture surfaces. High-resolution digital time-lapse imaging was employed to qualitatively observe the migration of colloidal and soluble dyes through the flow models. In this study, a Sphingomonas sp. and Sporosarcina (Bacillus) pasteurii influenced the fluid dynamics by physically altering flow paths. Microbial colonization and calcite deposition enhanced the stagnant regions adjacent to solid boundaries. Microbial growth and calcite precipitation occurred to a greater extent in areas behind the fabricated obstacles and less in high-velocity orifices.     

Control of self-motion in dynamic fluids: fish do it differently from bees

To detect and avoid collisions, animals need to perceive and control the distance and the speed with which they are moving relative to obstacles. This is especially challenging for swimming and flying animals that must control movement in a dynamic fluid without reference from physical contact to the ground. Flying animals primarily rely on optic flow to control flight speed and distance to obstacles. Here, we investigate whether swimming animals use similar strategies for self-motion control to flying animals by directly comparing the trajectories of zebrafish (Danio rerio) and bumblebees (Bombus terrestris) moving through the same experimental tunnel. While moving through the tunnel, black and white patterns produced (i) strong horizontal optic flow cues on both walls, (ii) weak horizontal optic flow cues on both walls and (iii) strong optic flow cues on one wall and weak optic flow cues on the other. We find that the mean speed of zebrafish does not depend on the amount of optic flow perceived from the walls. We further show that zebrafish, unlike bumblebees, move closer to the wall that provides the strongest visual feedback. This unexpected preference for strong optic flow cues may reflect an adaptation for self-motion control in water or in environments where visibility is limited.     

Measurement and prediction of flow through a replica segment of a mildly atherosclerotic coronary artery of man

Pressure distributions were measured along a hollow vascular axisymmetric replica of a segment of the left circumflex coronary artery of man with mildly atherosclerotic diffuse disease. A large range of physiological Reynolds numbers from about 60 to 500, including hyperemic response, was spanned in the flow investigation using a fluid simulating blood kinematic viscosity. Predicted pressure distributions from the numerical solution of the Navier-Stokes equations were similar in trend and magnitude to the measurements. Large variations in the predicted velocity profiles occurred along the lumen. The influence of the smaller scale multiple flow obstacles along the wall (lesion variations) led to sharp spikes in the predicted wall shear stresses. Reynolds number similarity was discussed, and estimates of what time averaged in vivo pressure drop and shear stress might be were given for a vessel segment.     

The Role of the Tight Junction in Paracellular Fluid Transport across Corneal Endothelium. Electro-osmosis as a Driving Force

The mechanism of epithelial fluid transport is controversial and remains unsolved. Experimental difficulties pose obstacles for work on a complex phenomenon in delicate tissues. However, the corneal endothelium is a relatively simple system to which powerful experimental tools can be applied. In recent years our laboratory has developed experimental evidence and theoretical insights that illuminate the mechanism of fluid transport across this leaky epithelium. Our evidence points to fluid being transported via the paracellular route by a mechanism requiring junctional integrity, which we attribute to electro-osmotic coupling at the junctions. Fluid movements can be produced by electrical currents. The direction of the movement can be reversed by current reversal or by changing junctional electrical charges by polylysine. Aquaporin 1 (AQP1) is the only AQP present in these cells, and its deletion in AQP1 null mice significantly affects cell osmotic permeability but not fluid transport, which militates against the presence of sizable water movements across the cell. By contrast, AQP1 null mice cells have reduced regulatory volume decrease (only 60% of control), which suggests a possible involvement of AQP1 in either the function or the expression of volume-sensitive membrane channels/transporters. A mathematical model of corneal endothelium predicts experimental results only when based on paracellular electro-osmosis, and not when transcellular local osmosis is assumed instead. Our experimental findings in corneal endothelium have allowed us to develop a novel paradigm for this preparation that includes: (1) paracellular fluid flow; (2) a crucial role for the junctions; (3) hypotonicity of the primary secretion; (4) an AQP role in regulation and not as a significant water pathway. These elements are remarkably similar to those proposed by the Hill laboratory for leaky epithelia.     

Numerical study of the impact of non-Newtonian blood behavior on flow over a two-dimensional backward facing step

Endothelial cell (EC) responsiveness to shear stress is essential for vasoregulation and plays a role in atherogenesis. Although blood is a non-Newtonian fluid, EC flow studies in vitro are typically performed using Newtonian fluids. The goal of the present study was to determine the impact of non-Newtonian behavior on the flow field within a model flow chamber capable of producing flow disturbance and whose dimensions permit Reynolds and Womersley numbers comparable to those present in vivo. We performed two-dimensional computational fluid dynamic simulations of steady and pulsatile laminar flow of Newtonian and non-Newtonian fluids over a backward facing step. In the non-Newtonian simulations, the fluid was modeled as a shear-thinning Carreau fluid. Steady flow results demonstrate that for Re in the range 50-400, the flow recirculation zone downstream of the step is 22-63% larger for the Newtonian fluid than for the non-Newtonian fluid, while spatial gradients of shear stress are larger for the non-Newtonian fluid. In pulsatile flow, the temporal gradients of shear stress within the flow recirculation zone are significantly larger for the Newtonian fluid than for the non-Newtonian fluid. These findings raise the possibility that in regions of flow disturbance, EC mechanotransduction pathways stimulated by Newtonian and non-Newtonian fluids may be different.     

The effects of grain geometry on waterflooding and viscous fingering in micro-fractures and porous media from a lattice Boltzmann method study

The study of fluid fronts formed in porous media is important for enhanced oil recovery. The purposes of this study are to simulate waterflooding and to investigate influential factors on the fluid front movement through a micro-fracture and through simple porous media with different grain geometries. This study used the Shan–Chen form of the Lattice Boltzmann Method (LBM). An increase in the velocity is found to result in viscous fingering, whereas an increase in the wettability of the displacing fluid and the dynamic viscosity ratio creates a piston form of the fluid front. In porous media with the same porosities, various geometries act differently as obstacles against fluid flow from the inlet to the outlet. By enlarging the cross-sectional area of grains in the fluid paths and making them more tortuous, narrower and more twisted films of viscous fingering are formed. The sweep efficiency was also determined under various conditions: with a fixed capillary number, neutral wettability and different viscosity ratios; and with a fixed capillary number, viscosity ratio of (1/3) and wet or non-wet conditions. In all cases, the best sweep efficiency was obtained with grains of diamond geometry. Generally, the least sweep efficiency occurs with grains of star geometry. Simulation results verified the strength and accuracy of LBM predictions.     

The influence of viscous hydrodynamics on the fish lateral-line system

Fish exhibit many behaviors that involve sensing water flows with their lateral-line system. In many situations, viscosity affects how the flow interacts with the body of the fish and the neuromasts of the lateral line. Here we discuss how viscosity influences the stimulus to the fish lateral-line system. The movement of a fish's body creates flows that can interfere with the detection of external signals, but these flows can also serve as a source of information about nearby obstacles and the fish's own hydrodynamic performance. The viscous boundary layer on the surface of the skin alters external signals by attenuating the low-frequency components of stimuli. The stimulus to each neuromast depends on the interaction of the fluid surrounding the neuromast and the structural properties of that neuromast, including the number of mechanosensory hair cells it contains. A consideration of the influences of viscosity on flow, at both the whole-body and receptor levels, offers the promise of a more comprehensive understanding of the signals involved in behaviors mediated by the lateral-line system.     

Lateral diffusion and percolation in two-phase, two-component lipid bilayers. Topology of the solid-phase domains in-plane and across the lipid bilayer.

Fluorescence recovery after photobleaching (FRAP) has recently been used to examine the percolation properties of coexisting phases in two-component, two-phase phosphatidylcholine bilayers [Vaz, W. L. C., Melo, E. C. C., & Thompson, T. E. (1989) Biophys. J. 56, 869-876]. We now report the use of FRAP to study two additional problems in similar systems. The first is the effect of solid-phase obstacles on the lateral diffusion in the fluid phase. The second is the question of whether or not, in a single bilayer, solid-phase domains in one monolayer are exactly superimposed on solid domains in the apposing monolayer. To address the first problem, the lateral diffusion of N-(7-nitrobenzoxa-2,3-diazol-4-yl)-1-palmitoyl-2-oleoylphosp hatidylethanolamine (NBD-POPE), a probe soluble only in the fluid phase when solid and fluid phases coexist, has been studied in the mixture N-lignoceroyldihydrogalactosylceramide (LigGalCer)/dipalmitoylphosphatidylcholine (DPPC). Percolation of the fluid phase occurs at a high mass fraction of solid phase. This indicates that the solid domains have a centrosymmetric shape, a characteristic which makes this a good experimental system to test theoretical simulations of diffusion in an archipelago. It is shown that agreement between theory and experiment is poor, a result that had already been observed when the obstacles were integral membrane proteins. We develop an effective-medium model for diffusion in two-phase systems which explains both our results and those obtained with integral proteins. The distinctive feature of the model is the consideration of an annular region around the obstacles where the lipids are more ordered than in the bulk fluid phase. The diffusion coefficient is then calculated by extending the free area model to two-phase systems, taking these annuli into account. The second question, the organization of the solid-phase domains across the lipid bilayer, is examined in the systems LigGalCer/DPPC and dimyristoylphosphatidylcholine (DMPC)/distearoylphosphatidylcholine (DSPC) by comparing the diffusion of a fluid-phase-soluble, gel-phase-insoluble lipid derivative which spans the two monolayers of a bilayer (NBD-membrane-spanning-phosphatidylethanolamine, NBD-msPE) with that of a probe which is restricted to a single monolayer. In LigGalCer/DPPC, 20:80, the distribution of solid domains in one of the monolayers is independent of the distribution in the apposing monolayer. In contrast, in DMPC/DSPC, 50:50, the solid domains in one monolayer are exactly superimposed upon the solid domains existing in the apposing monolayer.     

The influence of the non-Newtonian properties of blood on the flow in large arteries: unsteady flow in a 90 degrees curved tube.

A numerical and experimental investigation of unsteady entry flow in a 90 degrees curved tube is presented to study the impact of the non-Newtonian properties of blood on the velocity distribution. The time-dependent flow rate for the Newtonian and the non-Newtonian blood analog fluid were identical. For the numerical computation, a Carreau-Yasuda model was employed to accommodate the shear thinning behavior of the Xanthan gum solution. The viscoelastic properties were not taken into account. The experimental results indicate that significant differences between the Newtonian and non-Newtonian fluid are present. The numerical results for both the Newtonian and the non-Newtonian fluid agree well with the experimental results. Since viscoelasticity was not included in the numerical code, shear thinning behavior of the blood analog fluid seems to be the dominant non-Newtonian property, even under unsteady flow conditions. Finally, a comparison between the non-Newtonian fluid model and a Newtonian fluid at a rescaled Reynolds number is presented. The rescaled Reynolds number, based on a characteristic rather than the high-shear rate viscosity of the Xanthan gum solution, was about three times as low as the original Reynolds number. Comparison reveals that the character of flow of the non-Newtonian fluid is simulated quite well by using the appropriate Reynolds number.     

Prolonged effect of fluid flow stress on the proliferative activity of mesothelial cells after abrupt discontinuation of fluid streaming

Encapsulating peritoneal sclerosis (EPS) often develops after transfer to hemodialysis and transplantation. Both termination of peritoneal dialysis (PD) and transplantation-related factors are risks implicated in post-PD development of EPS, but the precise mechanism of this late-onset peritoneal fibrosis remains to be elucidated. We previously demonstrated that fluid flow stress induced mesothelial proliferation and epithelial-mesenchymal transition via mitogen-activated protein kinase (MAPK) signaling. Therefore, we speculated that the prolonged bioactive effect of fluid flow stress may affect mesothelial cell kinetics after cessation of fluid streaming. To investigate how long mesothelial cells stay under the bioactive effect brought on by fluid flow stress after removal of the stress, we initially cultured mesothelial cells under fluid flow stress and then cultured the cells under static conditions. Mesothelial cells exposed to fluid flow stress for a certain time showed significantly high proliferative activity compared with static conditions after stoppage of fluid streaming. The expression levels of protein phosphatase 2A, which dephosphorylates MAPK, in mesothelial cells changed with time and showed a biphasic pattern that was dependent on the duration of exposure to fluid flow stress. There were no differences in the fluid flow stress-related bioactive effects on mesothelial cells once a certain time had passed. The present findings show that fluid flow stress exerts a prolonged bioactive effect on mesothelial cells after termination of fluid streaming. These findings support the hypothesis that a history of PD for a certain period could serve as a trigger of EPS after stoppage of PD.     

On the physics of CNS memory

The effects of geometry, type of fluid and properties of the desmotubule membrane on the fluid transport in plasmodesma are discussed from a hydrodynamics viewpoint. It is shown that the “necking” of the ends of plasmodesma has a profound effect on the volume flow rates reducing them by several orders of magnitude. Most of the pressure drop occurs across the “neck” regions. There appears to be little significant difference in the volume flow rates if we consider a Newtonian or powerlaw fluid or if we allow the desmotubule membrane to be permeable or slightly flexible, at least in comparison to the dominating feature of plasmodesma geometry.     

The biased lamellipodium development and microtubule organizing center position in vascular endothelial cells migrating under the influence of fluid flow*

Summary— To analytically study the morphological responses of vascular endothelial cells (ECs) to fluid flow, we designed a parallel plate flow culture chamber in which cells were cultured under fluid shear stress ranging from 0.01 to 2.0 Pa for several days. Via a viewing window of the chamber, EC responses to known levels of fluid shear stress were monitored either by direct observations or by a video-enhanced time-lapse microscopy. Among the responses of cultured ECs to flow, morphological responses take from hours to days to be fully expressed, except for the fluid shear stress-dependent motility pattern change we reported earlier which could be detected within 30 min of flow changes. We report here that ECs exposed to more than 1.0 Pa of fluid shear shear stress have developed lamellipodia in the direction of flow in 10 min. This is the fastest structurally identifiable EC response to fluid shear stress. This was a reversible response. When the flow was stopped or reduced to the level which exerted less than 0.1 Pa of fluid shear stress, the biased lamellipodium development was lost within several minutes. The microtubule organizing center was located posterior to the nucleus in ECs under the influence of flow. However, this position was established only in ECs responding to fluid shear stress for longer than 1 h, indicating that positioning of the microtubule organizing center was not the reason for, but rather the result of, the biased lamellipodium response. Colcemid-treated ECs responded normally to flow, indicating that microtubules were not involved in both flow sensing and the flow-induced, biased lamellipodium development.     

The effect of turbulent motion on the diffusion of microorganisms

The effect of turbulent fluid motion on the diffusion of simple organisms is discussed. The net reproduction rate and the turbulent flow are assumed to be Gaussian-correlated random variables. For homogeneous istropic turbulence, simple equations for the average concentration of the organisms are derived in terms of the energy density of the fluid. It is shown that the effective diffusivity generated by the motion is positive-definite, and is independent of the helicity of the flow.     

The influence of the non-Newtonian properties of blood on the flow in large arteries: steady flow in a carotid bifurcation model.

Laser Doppler anemometry experiments and finite element simulations of steady flow in a three dimensional model of the carotid bifurcation were performed to investigate the influence of non-Newtonian properties of blood on the velocity distribution. The axial velocity distribution was measured for two fluids: a non-Newtonian blood analog fluid and a Newtonian reference fluid. Striking differences between the measured flow fields were found. The axial velocity field of the non-Newtonian fluid was flattened, had lower velocity gradients at the divider wall, and higher velocity gradients at the non-divider wall. The flow separation, as found with the Newtonian fluid, was absent. In the computations, the shear thinning behavior of the analog blood fluid was incorporated through the Carreau-Yasuda model. The viscoelastic properties of the fluid were not included. A comparison between the experimental and numerical results showed good agreement, both for the Newtonian and the non-Newtonian fluid. Since only shear thinning was included, this seems to be the dominant non-Newtonian property of the blood analog fluid under steady flow conditions.     

Lens intracellular hydrostatic pressure is generated by the circulation of sodium and modulated by gap junction coupling

We recently modeled fluid flow through gap junction channels coupling the pigmented and nonpigmented layers of the ciliary body. The model suggested the channels could transport the secretion of aqueous humor, but flow would be driven by hydrostatic pressure rather than osmosis. The pressure required to drive fluid through a single layer of gap junctions might be just a few mmHg and difficult to measure. In the lens, however, there is a circulation of Na(+) that may be coupled to intracellular fluid flow. Based on this hypothesis, the fluid would cross hundreds of layers of gap junctions, and this might require a large hydrostatic gradient. Therefore, we measured hydrostatic pressure as a function of distance from the center of the lens using an intracellular microelectrode-based pressure-sensing system. In wild-type mouse lenses, intracellular pressure varied from ～330 mmHg at the center to zero at the surface. We have several knockout/knock-in mouse models with differing levels of expression of gap junction channels coupling lens fiber cells. Intracellular hydrostatic pressure in lenses from these mouse models varied inversely with the number of channels. When the lens' circulation of Na(+) was either blocked or reduced, intracellular hydrostatic pressure in central fiber cells was either eliminated or reduced proportionally. These data are consistent with our hypotheses: fluid circulates through the lens; the intracellular leg of fluid circulation is through gap junction channels and is driven by hydrostatic pressure; and the fluid flow is generated by membrane transport of sodium.     

Effect of the glycocalyx layer on transmission of interstitial flow shear stress to embedded cells

In this paper, a simple theoretical model is developed to describe the transmission of force from interstitial fluid flow to the surface of a cell covered by a proteoglycan / glycoprotein layer (glycocalyx) and embedded in an extracellular matrix. Brinkman equations are used to describe flow through the extracellular matrix and glycocalyx layers and the solid mechanical stress developed in the glycocalyx by the fluid flow loading is determined. Using reasonable values for the Darcy permeability of extracellular matrix and glycocalyx layers and interstitial flow velocity, we are able to estimate the fluid and solid shear stresses imposed on the surface of embedded vascular, cartilage and tumor cells in vivo and in vitro. The principal finding is that the surface solid stress is typically one to two orders of magnitude larger than the surface fluid stress. This indicates that interstitial flow shear stress can be sensed by the cell surface glycocalyx, supporting numerous recent observations that interstitial flow can induce mechanotransduction in embedded cells. This study may contribute to understanding of interstitial flow-related mechanobiology in embryogenesis, tumorigenesis, tissue physiology and diseases and has implications in tissue engineering.     

Analysis of the flow field induced by the sessile peritrichous ciliate Opercularia asymmetrica

The feeding mechanism of the sessile protozoon Opercularia asymmetrica (Oligohymenophorea, Peritrichia) relies on the cilia beat generating a flow field that convectively transports suspended particles and dissolved substances to the oral cavity of the organism. By use of optical micro-flow measurement and theoretical methods the flow environment of two neighbouring peritrichous ciliate cells is studied. Both, yeast cells (Saccharomyces cerevisiae) and artificial flow tracers are used for the visualisation of the flow field. Artificial tracers are rejected by the protozoa and deviate from the fluid path lines, while yeast cells follow the flow almost perfectly. This is shown through a dimensional analysis of the involved hydrodynamic forces on the tracers. The measured flow field exhibits maximum velocities of 25 microm/s at around 20 microm distance ahead of an individual ciliate. The flow field extends 200 microm from the location of the ciliate. A nicking motion of the protozoon is observed and found not to obey any periodic law. Multiples of protozoa exhibit most commonly an alternating cilia beat regime generating a non-stationary flow field. It can be shown through theoretical methods that fluid exchange is enhanced in this alternating regime compared to a flow field generated by a single ciliate. Fluid exchange depends on the distance of the ciliates from each other and on the alteration frequency of the cilia beat. The comparison of an analytical Stokes' flow solution with the observed fluid flow serves to determine the force required to maintain the flow field against viscous dissipation. The force magnitude is in the order of magnitude of 10-100 pN.