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Biomedical subjects

T J Pedley

Publications and source records attributed to T J Pedley.

At least 19 recordsLinked to original sources

Measurement of cell velocity distributions in populations of motile algae.

The self-propulsion of unicellular algae in still ambient fluid is studied using a previously reported laser-based tracking method, supplemented by new tracking software. A few hundred swimming cells are observed simultaneously and the average parameters of the cells' motility are calculated. The time-dependent, two-dimensional distribution of swimming velocities is measured and the three-dimensional distribution is recovered by assuming horizontal isotropy. The mean and variance of the cell turning angle are quantified, to estimate the reorientation time and rotational diffusivity of the bottom-heavy cell. The cells' phototactic and photokinetic responses to the laser light are evaluated. The results are generally consistent both with earlier assumptions about the nature of cell swimming and quantitative measurements, appropriately adjusted. The laser-based tracking method, which makes it possible to average over a large number of motile objects, is shown to be a powerful tool for the study of microorganism motility.

Animals↗

A model for time-dependent flow in (giraffe jugular) veins: uniform tube properties.

Computations are reported for a one-dimensional model of time-dependent flow in collapsible tubes representing long mammalian veins. The tubes are taken to have uniform intrinsic properties and we concentrate on the effect of longitudinal gravity. The main application is to the jugular vein of the upright giraffe, with given inflow rate from the head, a given pressure, slightly above the external, atmospheric pressure, at the downstream (vena caval) end, and a variety of initial conditions. We show that: (i) previously calculated steady flows are the long time limits of unsteady computations, although only after a considerable time in which slowly-decaying waves and elastic jumps propagate up and down, (ii) steady flows are indeed not found when the steady-flow analysis shown them not to exist, although the consequent unsteadiness is of such small amplitude as to be practically unimportant, (iii) the time taken for the flow to become steady when the neck is raised from the horizontal or the head-down position can be several seconds longer than the neck-raising time itself (3-7s). We also find that roll-waves do not develop despite having been previously predicted for long collapsible tubes. Further application is made to the effect of postural changes on human neck and leg veins.

Animals↗

The influence of turbulence on plankton predation strategies.

The importance of predation in regulating the size of competing plankton and larval fish populations has long been appreciated. However, it has only recently been recognized that turbulence must have a significant influence on predator-prey interactions because most rival species of microorganisms co-exist in oceanic or fast moving fresh water flows. Turbulence is likely to influence predation strategies in two ways. The extra energy imparted to a micro-organism from the flow field will enhance the number of encounters or "contacts" between predators and prey. At the same time, because the velocity of a predator relative to its potential prey will be increased, the time-scale over which a capture must be completed is reduced. Balancing the benefits of extra encounters with the drawbacks of more difficult captures, will dictate an optimal predation strategy, either foraging behaviour or ambush feeding, on the predator. This will depend on its own and the prey's swimming capabilities, as well as the characteristics of the turbulent environment. In this paper some previous work, examining the increased encounter rate in turbulence, will be extended to look at the capture problem. The main proposal is that the capture event should be encapsulated in a capture probability function, from which the optimal predation strategy can be derived. As an illustration, plausible capture probability functions will be postulated and the resulting predictions tested against numerical simulations carried out in a turbulent-like flow field. Good agreement between the predictions and the simulations is demonstrated.

Animal Nutritional Physiological Phenomena↗

Planktonic contact rates in homogeneous isotropic turbulence: theoretical predictions and kinematic simulations.

The key role played by turbulence in the environment of plankton and larval fish populations has become appreciated in recent years. In particular, the turbulent enhancement of encounter rates between different species of microorganisms, either swimming or passively advected by the flow, is well established. However, most of the current modelling approaches are rather ad hoc, giving rise to ambiguities in the specification of certain key parameters. In this paper, the encounter problem in a turbulent flow of large Reynolds number is re-examined from first principles and a number of new formulae will be established for different swimming strategies. The key innovation is the proposal of a model form for the conditional joint probability density function of predator and prey velocities when the organisms are separated by their given contact radius, R. Particular attention will be paid to the case when a microorganism follows a random trajectory, due to a combination of its own swimming and the action of the flow. The theoretical predictions are subsequently tested against corresponding quantities derived from a series of kinematic simulations of a turbulent-like flow field. Good agreement is demonstrated between the predictions and simulations.

Animals↗

Modelling run-and-tumble chemotaxis in a shear flow.

The biased random walk undergone by chemotactic bacteria such as Escherichia coli will be influenced at the microscopic level by flow in the ambient medium. In this paper, we model swimming bacteria being advected and rotated by a simple shear flow. Under certain scaling assumptions, we obtain an advection-diffusion equation for cell density, when the chemotactic response is small, which shows a coupling between the rotation and chemotaxis. We also present an alternative method for calculating the chemotactic flux in an unbounded region which is valid for more general chemotactic responses.

Chemotaxis↗

Vascular endothelial cells minimize the total force on their nuclei.

The vascular endothelium is a cellular monolayer that lines the arterial walls. It plays a vital role in the initiation and development of atherosclerosis, an occlusive arterial disease responsible for 50% of deaths in the Western world. The focal nature of the disease suggests that hemodynamic forces are an important factor in its pathogenesis. This has led to the investigation of the effects of mechanical forces on the endothelial cells themselves. It has been found that endothelial cells do respond to stresses induced by the flowing blood; in particular, they elongate and align with an imposed flow direction. In this paper, we calculate the distribution of force exerted on a three-dimensional hump, representing the raised cell nucleus, by a uniform shear flow. It is found that, for a nonaxisymmetric ellipsoidal hump, the least total force is experienced when the hump is aligned with the flow. Furthermore, for a hump of fixed volume, there is a specific aspect ratio combination that results in the least total force upon the hump, (0.38:2.2:1.0; height:length:width). This is approximately the same as the average aspect ratio taken up by the cell nuclei in vivo (0.27:2.23:1.0). It is possible, therefore, that the cells respond to the flow in such a way as to minimize the total force on their nuclei.

Animals↗

Large-amplitude undulatory fish swimming: fluid mechanics coupled to internal mechanics.

The load against which the swimming muscles contract, during the undulatory swimming of a fish, is composed principally of hydrodynamic pressure forces and body inertia. In the past this has been analysed, through an equation for bending moments, for small-amplitude swimming, using Lighthill's elongated-body theory and a 'vortex-ring panel method', respectively, to compute the hydrodynamic forces. Those models are outlined in this review, and a summary is given of recent work on large-amplitude swimming that has (a) extended the bending moment equation to large amplitude, which involves the introduction of a new (though probably usually small) term, and (b) developed a large-amplitude vortex-ring panel method. The latter requires computation of the wake, which rolls up into concentrated vortex rings and filaments, and has a significant effect on the pressure on the body. Application is principally made to the saithe (Pollachius virens). The calculations confirm that the wave of muscle activation travels down the fish much more rapidly than the wave of bending.

Animals↗

Alteration of mean wall shear stress near an oscillating stagnation point.

The site opposite an end-to-side anastomosis, resulting from femoral bypass surgery, and the carotid sinus are two regions well known to be prone to fibrous intimal hyperplasia or atherogenesis, respectively. The blood flow at these two sites features a stagnation point, which oscillates in strength and position. Mathematical models are used to determine some of the features of such a flow; in particular, the mean wall shear stress is calculated. The positional oscillations cause a significant change in the distribution and magnitude of the mean wall shear stress from that of the well-studied case of a stagnation point that oscillates only in strength. It is therefore proposed that the recorded effect of time dependence in the flow upon atherogenesis could still be a result of the distribution of the mean and not the time-varying components of the wall shear stress.

Algorithms↗

An active membrane model for peristaltic pumping: Part I--Periodic activation waves in an infinite tube.

A model for the coupled problem of wall deformation and fluid flow, based on thin-shell and lubrication theories, and driven by a propagating wave of smooth muscle activation, is proposed for peristaltic pumping in the ureter. The model makes use of the available experimental data on the mechanical properties of smooth muscle and accounts for the soft material between the muscle layer and the vessel lumen. The main input is the activation wave of muscular contraction. Equations for the time-dependent problem in tubes of arbitrary length are derived and applied to the particular case of periodic activation waves in an infinite tube. Mathematical (small amplitude) and numerical analyses of this case are presented. Predictions on phase-lag in wall constriction with respect to peak activation wave, lumen occlusion due to thickening lumen material with contracting smooth muscle, and the general bolus shape are in qualitative agreement with observation. Some modifications to the mechanical, elastic, and hydrodynamic properties of the ureter that will make peristalsis less efficient, due for example to disease, are identified. In particular, the flow rate-pressure rise relationship in linear for weak to moderate activation waves, but as the lumen is squeezed shut, it is seen to be nonlinear in a way that increases pumping efficiency. In every case a ureter whose lumen can theoretically be squeezed shut is the one for which pumping is most efficient.

Animals↗

Blood pressure and flow rate in the giraffe jugular vein.

Experimental measurements in the jugular veins of upright giraffes have shown that the internal pressure is somewhat above atmospheric and increases with height above the heart. A simple model of steady viscous flow in an inverted U-tube shows that these observations are inconsistent with a model in which the blood vessels in the head and neck are effectively rigid and the system resembles a siphon. Instead, the observations indicate that the veins are collapsed and have a high resistance to flow. However, laboratory experiments with collapsible drain tubing in place of the down arm of the U-tube show internal pressure to be exactly atmospheric and uniform with height. A model of viscous flow in a collapsible tube with non-uniform properties is used to suggest that the observed pressure distribution may be a consequence of the intrinsic cross-sectional area and/or compliance of the veins increasing with distance towards the heart, or the external, tissue pressure falling. Finally, the effect of fluid inertia on steady flow in vertical collapsible tubes with uniform intrinsic properties is analysed, and it is shown that a phenomenon of flow limitation is theoretically possible, in which the supercritical flow in the collapsed vein cannot return to the presumably subcritical flow in the open vena cava, even with the help of an 'elastic jump', if the flow rate is too large. The computed critical flow-rate, of about 80 ml s-1, is about twice the flow-rate estimated to be present in the normal giraffe jugular vein. If there were circumstances in which flow limitation occurred in the jugular veins, it would mean that the cerebral blood flow would be limited by downstream conditions, not directly by local requirements.

Animals↗

The development of concentration gradients in a suspension of chemotactic bacteria.

When a suspension of bacterial cells of the species Bacillus subtilis is placed in a chamber with its upper surface open to the atmosphere complex bioconvection patterns are observed. These arise because the cells: (1) are denser than water; and (2) usually swim upwards, so that the density of an initially uniform suspension becomes greater at the top than the bottom. When the vertical density gradient becomes large enough, an overturning instability occurs which ultimately evolves into the observed patterns. The reason that the cells swim upwards is that they are aerotactic, i.e., they swim up gradients of oxygen, and they consume oxygen. These properties are incorporated in conservation equations for the cell (N) and oxygen (C) concentrations, and these are solved in the pre-instability phase of development when N and C depend only on the vertical coordinate and time. Numerical results are obtained for both shallow- and deep-layer chambers, which are intrinsically different and require different mathematical and numerical treatments. It is found that, for both shallow and deep chambers, a thin boundary layer, densely packed with cells, forms near the surface. Beneath this layer the suspension becomes severely depleted of cells. Furthermore, in the deep chamber cases, a discontinuity in the cell concentration arises between this cell-depleted region and a cell-rich region further below, where no significant oxygen concentration gradients develop before the oxygen is fully consumed. The results obtained from the model are in good qualitative agreement with the experimental observations.

Bacillus subtilis↗

High Reynolds number flow in tubes of complex geometry with application to wall shear stress in arteries.

The arterial systems of mammals contain many generations of branching, curved, elastic tubes, and the flow in them is unsteady. The flow details are important in determining the distribution of wall shear stress in arteries, a major factor in atherogenesis. The purpose of this paper is to point out that unsteady flow at realistic values of the Reynolds number (several hundred) and frequency parameter is much more complicated than a knowledge of steady flow in simpler geometries would suggest. Three-dimensional effects in curved, branched or indented tubes introduce secondary motions and horseshoe vortices, under which the wall shear can remain high for a considerable distance downstream. Unsteady flow in two-dimensional, non-uniform tubes can lead to regions of flow separation far from the non-uniformity, again associated with high wall shear, not low shear as for two-dimensional, steady, separated flow. Moreover, both two- and three-dimensional flows can be non-unique, so that the same driving pressures in the same geometry can lead to different flow patterns, depending on the initial conditions. There is great sensitivity to small geometric and temporal perturbations, making it very difficult to predict flow in one arterial bifurcation from a knowledge of another, or even at a particular site in one subject from a knowledge of another. Vessel elasticity is an additional complicating factor whose effects remain to be assessed in detail.

Animals↗

Gas flow and mixing in the airways.

OBJECTIVE: To survey the current state of scientific knowledge of gas flow and mixing in pulmonary airways, especially at high frequencies. DATA SOURCES: Results from the authors' own laboratory studies and Western bioengineering literature on respiratory fluid mechanics. STUDY SELECTION: This survey concentrates on understanding the principal physical mechanisms that underlie the enhancement of airway gas transport in high-frequency oscillation. The results of experimental, computational, and mathematical studies are described. DATA SYNTHESIS: The topic was covered by seven presenters at the Münster Meeting on High Frequency Ventilation, January 31 to February 2, 1993. Six of these presentations are summarized in the six sections of this paper under the following headings: Introductory Survey; Three-Dimensional Numerical Simulation of Inspiratory and Expiratory Flows in Small Airways; Computational and Experimental Models of High-Frequency Oscillation; Unsteady Gas Mixing in Airways; Gas Dispersion From the Lagrangian Viewpoint; and Soluble Gas Mass Transfer in Tubes and Airways. CONCLUSIONS: The dominant mechanism for the enhancement of gas transport along airways at high frequency is likely to lie in the coupling of the secondary motions caused by airway curvature with the oscillatory longitudinal flow. However, laboratory experiments and theoretical analyses have led only to an accurate simulation of the phenomena in idealized geometries, not in real lungs.

Animals↗

The heart works against gravity.

The circulatory systems of vertebrate animals are closed, and blood leaves and returns to the heart at the same level. It is often concluded, therefore, that the heart works only against the viscous resistance of the system, not against gravity, even in vascular loops above the heart in which the siphon principle operates. However, we argue that the siphon principle does not assist blood flow in superior vascular loops if any of the descending vasculature is collapsible. If central arterial blood pressure is insufficient to support a blood column between the heart and the head, blood flow ceases because of vascular collapse. Furthermore, the siphon principle does not assist the heart even when a continuous stream of blood is flowing in a superior loop. The potential energy gained by blood as it is pumped to the head is lost to friction in partially collapsed descending vessels and thus is not regained. Application of the Poiseuille equation to flow in collapsible vessels is limited; resistance depends on flow rate in partially collapsed vessels with no transmural pressure difference, but flow rate is independent of resistance. Thus the pressure developed by the heart to establish a given flow rate is independent of the resistance occurring in the partially collapsed vessels. The pressure depends only on the height of the blood column and the resistance in the noncollapsed parts of the system. Simple laboratory models, involving water flow in collapsible tubing, dispel the idea that the siphon principle facilitates blood flow and suggest that previously published results may have been affected by experimental artifact.

Animals↗

Role of pulmonary surfactant in airway closure: a computational study.

A numerical model that simulates airway closure by liquid bridging during expiration has been developed. The effects of both surfactant and time-varying geometry have been included; the model determines the liquid layer flow resulting from a surface tension (Rayleigh) instability, and the computation traces the film's development to closure, yielding pressure, velocity, surface shape, and surfactant concentration distributions. It is found that surfactant is effective in retarding or eliminating liquid bridging through the reduction of the mean surface tension and the action of surface tension gradients. The former effect is also critical in minimizing the magnitude of the negative pressure in the liquid layer and thus presumably in reducing the tendency for airway compliant collapse.

Animals↗

Longitudinal tension variation in collapsible channels: a new mechanism for the breakdown of steady flow.

There are several mechanisms potentially involved in the breakdown of steady fluid flow in a collapsible tube under external pressure. Here we investigate one that has received little attention in the past: the fact that the longitudinal tension in the tube wall, T, decreases with distance downstream as a consequence of the viscous shear stress exerted by the fluid. If the tube is long enough, or the initial tension small enough, T may fall to zero before the end of the collapsible tube, and unsteady motion would presumably then ensue; this is what we mean by "breakdown." We study the phenomenon theoretically, when the flow Reynolds number is of order one, using lubrication theory in a symmetric two-dimensional channel in which the collapsible tube is replaced by membranes occupying a segment of each wall. The resulting nonlinear ordinary differential equations are solved numerically for values of the dimensionless parameters that cover all the qualitatively different types of solution (e.g., in which the channel is distended over all its length, collapsed over all its length, or distended in the upstream part and collapsed downstream). Reducing the longitudinal tension has a marked effect on the shape of the collapsible segment, causing it to become much more deformed for the same flow rate and external pressure. Indeed, the wall slope is predicted to become very large when the downstream tension is very small, so the model is not self-consistent then. Nevertheless, the parameter values for which T becomes zero are mapped out and are expected to be qualitatively useful.(ABSTRACT TRUNCATED AT 250 WORDS)

Blood Flow Velocity↗

Adaptive modeling of the human rib cage in median sternotomy.

This paper describes a limited computer-analyzed kinematic model of the rib cage that can be adapted to individual subjects. Also described is its validation and use in assessing the changes in chest wall shape after coronary artery bypass graft (CABG) surgery in 12 patients. The positions of a small number of anatomic locations on the thoracic spine, ribs, manubrium, and sternum are measured from lateral and posterior-anterior chest radiographs. The computer program puts these two views together removing the magnification and reconstructs any missing points to give a three-dimensional picture of the rib cage to which mathematical models of the bones are scaled. The patients had chest radiographs taken at total lung capacity (TLC) and residual volume (RV) to investigate the source of the restrictive ventilatory defect that follows CABG. The predictions from the model were tested by comparing full-sized computer plots with the actual chest radiographs. The estimates of the bony structures were accurate to +/- 3 degrees for orientations and +/- 6 mm for positions. We found reduced rib motion both "pump-handle" (theta) and "bucket handle" (psi) going from theta, psi left, psi right = 9 degrees, 10 degrees, 14 degrees to 4 degrees, 10 degrees, 9 degrees, respectively, after surgery with P less than 0.025, 0.42, 0.07. The angles were measured from the horizontal and increased caudally. There was also reduction in the range of angles subtended by the arc of the thoracic vertebrae between TLC and RV, which went from 12 degrees to -1 degrees (P less than 0.015). These data explain the fall in lung volumes that follow CABG and provide insight into the contribution made by the ribs and spine in full inspiration and full expiration.

Coronary Artery Bypass↗

A new continuum model for suspensions of gyrotactic micro-organisms.

A new continuum model is formulated for dilute suspensions of swimming micro-organisms with asymmetric mass distributions. Account is taken of randomness in a cell's swimming direction, p, by postulating that the probability density function for p satisfies a Fokker-Planck equation analogous to that obtained for colloid suspensions in the presence of rotational Brownian motion. The deterministic torques on a cell, viscous and gravitational, are balanced by diffusion, represented by an isotropic rotary diffusivity Dr, which is unknown a priori, but presumably reflects stochastic influences on the cell's internal workings. When the Fokker-Planck equation is solved, macroscopic quantities such as the average cell velocity Vc, the particle diffusivity tensor D and the effective stress tensor sigma can be computed; Vc and D are required in the cell conservation equation, and sigma in the momentum equation. The Fokker-Planck equation contains two dimensionless parameters, lambda and epsilon; lambda is the ratio of the rotary diffusion time Dr-1 to the torque relaxation time B (balancing gravitational and viscous torques), while epsilon is a scale for the local vorticity or strain rate made dimensionless with B. In this paper we solve the Fokker-Planck equation exactly for epsilon = 0 (lambda arbitrary) and also obtain the first-order solution for small epsilon. Using experimental data on Vc and D obtained with the swimming alga, Chlamydomonas nivalis, in the absence of bulk flow, the epsilon = 0 results can be used to estimate the value of lambda for that species (lambda approximately 2.2; Dr approximately 0.13 s-1). The continuum model for small epsilon is then used to reanalyse the instability of a uniform suspension, previously investigated by Pedley, Hill & Kessler (1988). The only qualitatively different result is that there no longer seem to be circumstances in which disturbances with a non-zero vertical wavenumber are more unstable than purely horizontal disturbances. On the way, it is demonstrated that the only significant contribution to sigma, other than the basic Newtonian stress, is that derived from the stresslets associated with the cells' intrinsic swimming motions.

Animals↗