Fluid mechanics of nodal flow due to embryonic primary cilia

Breaking of left–right symmetry is crucial in vertebrate development. The role of cilia-driven flow has been the subject of many recent publications, but the underlying mechanisms remain controversial. At approximately 8 days post-fertilization, after the establishment of the dorsal–ventral and anterior–posterior axes, a depressed structure is found on the ventral side of mouse embryos, termed the ventral node. Within the node, ‘whirling’ primary cilia, tilted towards the posterior, drive a flow implicated in the initial left–right signalling asymmetry. However, the underlying fluid mechanics have not been fully and correctly explained until recently and accurate characterization is required in determining how the flow triggers the downstream signalling cascades. Using the approximation of resistive force theory, we show how the flow is produced and calculate the optimal configuration to cause maximum flow, showing excellent agreement with in vitro measurements and numerical simulation, and paralleling recent analogue experiments. By calculating numerical solutions of the slender body theory equations, we present time-dependent physically based fluid dynamics simulations of particle pathlines in flows generated by large arrays of beating cilia, showing the far-field radial streamlines predicted by the theory

Mathematical modelling of cilia driven transport of biological fluids

Cilia-driven flow occurs in the airway surface liquid, in the female and male reproductive tracts and enables symmetry-breaking in the embryonic node. Viscoelastic rheology is found in healthy states in some systems, whereas in others may characterise disease, motivating the development of mathematical models that take this effect into account. We derive the fundamental solution for linear viscoelastic flow, which is subsequently used as a basis for slender-body theory. Our numerical algorithm allows efficient computation of three-dimensional time-dependent flow, bending moments, power and particle transport. We apply the model to the large-amplitude motion of a single cilium in a linear Maxwell liquid. A relatively short relaxation time of just 0.032 times the beat period significantly reduces forces, bending moments, power and particle transport, the last variable exhibiting exponential decay with relaxation time. A test particle is propelled approximately one-fifth as quickly along the direction of cilia beating for scaled relaxation time 0.032 as in the Newtonian case, and mean volume flow is abolished, emphasizing the sensitivity of cilia function to fluid rheology. These results may have implications for flow in the airways, where the transition from Newtonian to viscoelastic rheology in the peri-ciliary fluid may reduce clearance

A viscoelastic traction layer model of mucociliary flow

A new mathematical model of the transport of mucus and periciliary liquid (PCL) in the airways by cilia is presented. Mucus is represented by a linearly viscoelastic fluid, the mat of cilia is modelled as an ‘active porous medium.’ The propulsive effect of the cilia is modelled by a time-dependent force acting in a shear-thinned ‘traction layer’ between the mucus and the PCL. The effects of surface and interface tension are modelled by constraining the mucus free surface and mucus–PCL interface to be flat. It is assumed that the epithelium is impermeable to fluid. Using Fourier series, the system is converted into ODEs and solved numerically. We calculate values for mean mucus speed close to those observed by Matsui et~al. [{J. Clin. Invest.}, 102(6):1125’1131, 1998], (~40 μms−1). We obtain more detail regarding the dynamics of the flow and the nonlinear relationships between physical parameters in healthy and diseased states than in previously published models. Pressure gradients in the PCL caused by interface and surface tension are vital to ensuring efficient transport of mucus, and the role of the mucus–PCL interface appears to be to support such pressure gradients, ensuring efficient transport. Mean transport of PCL is found to be very small, consistent with previous analyses, providing insight into theories regarding the normal tonicity of PCL

Investigation of variables in turbine erosion, influence of aerodynamic and geometric parameters

Influence of aerodynamic and geometric parameters in turbine erosio

Measurements of PAN, alkyl nitrates, ozone, and hydrocarbons during spring in interior Alaska

Measurements of the atmospheric mixing ratios of ozone, peroxyacetylnitrate (PAN), hydrocarbons, and alkyl nitrates were made in a boreal forest ecosystem in the interior of Alaska from March 15 to May 14, 1993. During this period the mixing ratios of PAN, alkyl nitrates, and nonmethane hydrocarbons (NMHCs) generally decreased due to the influence of both meteorology and OH removal. Mean mixing ratios of ozone, PAN, C2 ‐ C6 alkyl nitrates, and total C2 ‐ C5 NMHC during southerly flow periods were 24.4 parts per billion (ppbv), 132.1 parts per trillion (pptv ), 34 pptv, and 8.2 ppbCv, respectively. During a short period of northerly flow, mixing ratios of PAN and total NMHC were approximately 2 times the southerly flow mixing ratios. PAN is correlated with ozone, and alkyl nitrates are correlated with alkanes. PAN and ozone mixing ratios exhibit similar diurnal variations on a number of days with an early morning minimum and afternoon maximum. This is likely due to a diurnal cycle in the boundary layer ‐ free troposphere exchange and loss processes in the boundary layer for both O3 and PAN. Higher molecular weight (mw) hydrocarbons and alkyl nitrates are observed to decrease more quickly than the lower mw hydrocarbons, consistent with removal by OH as the primary loss process

Mammalian Sperm Motility: Observation and Theory

Mammalian spermatozoa motility is a subject of growing importance because of rising human infertility and the possibility of improving animal breeding. We highlight opportunities for fluid and continuum dynamics to provide novel insights concerning the mechanics of these specialized cells, especially during their remarkable journey to the egg. The biological structure of the motile sperm appendage, the flagellum, is described and placed in the context of the mechanics underlying the migration of mammalian sperm through the numerous environments of the female reproductive tract. This process demands certain specific changes to flagellar movement and motility for which further mechanical insight would be valuable, although this requires improved modeling capabilities, particularly to increase our understanding of sperm progression in vivo. We summarize current theoretical studies, highlighting the synergistic combination of imaging and theory in exploring sperm motility, and discuss the challenges for future observational and theoretical studies in understanding the underlying mechanics.\ud Acronyms and Definitions\ud Acrosome: the cap of the sperm head containing enzymes allowing penetration of the zona pellucida via the acrosome reaction\ud Adenosine triphosphate (ATP): the currency unit of chemical energy transfer in living cells\ud Axoneme: a phylogenetically conserved structure within the eukaryotic flagellum consisting of a ring of nine microtubule doublets and a central pair, frequently referred to as 9 + 2\ud Bending moment density: the moment per unit length associated with flagellar bending; it can be divided into a hydrodynamic moment, an elastic moment (from the flagellar bending stiffness), an active moment (generated by dyneins exerting forces between adjacent microtubule doublets), and a passive moment resisting shear\ud Capacitation: the physiological state of a sperm required for fertilization, which is accompanied by the motility patterns associated with hyperactivation, characterized in saline by high-amplitude asymmetric beating\ud Central pair: a pair of microtubules along the length of the axoneme, symmetrically and slightly offset from the axoneme centerline\ud Cumulus oophorus: the outer vestment of the mammalian egg consisting of hundreds of cells radiating out from the egg embedded within a non-Newtonian hyaluronic acid gel\ud Dynein: a molecular motor within the axoneme, attached between adjacent microtubule doublets, that exerts a shearing force to induce axonemal bending\ud Flagellum: a motile cellular appendage that drives the swimming of sperm and other cells; this article focuses on the eukaryotic flagellum\ud Microtubule doublet: a pair of proteinaceous filament structures running the length of the axoneme; dyneins drive their bending, which induces flagellar motion\ud Mid-piece: the region of a sperm flagellum with a mitochondrial sheath, where ATP is generated\ud Oocyte: the egg\ud Outer dense fibers and fibrous sheath: accessory structures reinforcing the mammalian sperm flagellum; the combined axoneme and accessory structures are referred to as 9+9+2\ud Resistive-force theory: an approximation for the local drag of a slender filament element in Stokes flow (or a viscoelastic generalization thereof)\ud Rheotaxis: directed motility in response to the influence of fluid flow\ud Shear: in the context of the flagellum, the relative movement of adjacent microtubule doublets\ud Slender-body theory: an improved approximation for the local drag on a slender filament element in Stokes flow (or a viscoelastic generalization thereof)\ud Zona pellucida: a tough glycoprotein coat between the human egg and the cumulus oophorus, which a sperm must penetrate for successful fertilizatio

Cultural Resources Survey of the Leander Rehabilitation Center, Williamson County, Texas

In August-September 1996, personnel from Prewitt and Associates, Inc., conducted a cultural resources survey of ca. 725 acres of the former Leander Rehabilitation Center. The project area lies adjacent to U.S. Highway 183 and FM 620 in southern Williamson County, Texas. The survey resulted in additional documentation of one previously recorded prehistoric archeological site (41 WM452), the identification and recording of four historic archeological sites (41WM892, 41WM893, 41WM896, and 41WM897), and reconnaissance-level documentation of 45 historic buildings and structures. Site 41WM452 is an extensive upland lithic scatter and lithic procurement site which lacks subsurface deposits, features, and datable materials. Site 41WM892 is a wood-chopper camp that contains a number of rock alignments and limited artifact deposits dating to the first decade of the twentieth century. Site 41WM893 is a remnant of a railroad spur used during the 1937-1941 construction of Marshall Ford Dam (now Mansfield Dam). Site 41WM896 contains a small number of features and sparse artifact deposits associated with the 1937-1945 Rhodes farmstead. Site 41WM897 is an isolated historic well with unknown associations. None of these archeological sites contains important information, and it is recommended that they be considered not eligible for listing in the National Register of Historic Places or for designation as State Archeological Landmarks. The 45 buildings and structures, at 36 locations, are associated with the former State Dairy and Hog Farm. This farm was established in 1942, expanded after 1945, and reached its peak years of production as a hog farm between 1950 and the late 1960s, Created to serve the needs of the State Board of Control and the State Hospital, the facility is significant for its success in food production for eleemosynary institutions in Austin and throughout Texas, as well as for its role in the application of modern psychiatric treatment based on the therapeutic value of manual labor. Among the surveyed resources are dwellings, an office and warehouse building, a dormitory, a variety of agricultural buildings and structures, and infrastructural elements, all built between 1943 and 1955. Twenty-one of the 45 surveyed resources are recommended as being eligible for listing in the National Register of Historic Places under Criteria A and C as Contributing resources in a historic district and for designation as State Archeological Landmarks

Ground-based searches for interstellar H_(2)D^+

We present ground-based searches for the 1_(10) - 1_(11) line of interstellar H_(2)D^(+) at 372 GHz which are more sensitive than those obtained from the Kuiper Airborne Observatory by factors of 3-4 for extended sources and by more than two orders of magnitude for compact sources. The line was not detected in a variety of interstellar clouds, including NGC 2264 toward which a possible detection had been suggested previously. The inferred H_(2)D^(+) abundance limits of 10^(-10) - 10^(-11) are still consistent with, but approach the abundances predicted by chemical models. Simultaneous observations of the DCO^(+) 3-2 and N_(2)H^(+) 4-3 lines have been used to place additional limits on the H_(3)^(+) abundance, and suggest 10^(-11) < x(H_(3)^(+))< 10^(-9). The N_(2)H^(+) data also indicate that for NGC 2264, but perhaps not for the other sources, gas-phase N_2 contains a substantial fraction of the available nitrogen in the cloud

Constraints on the age and dilution of Pacific Exploratory Mission-Tropics biomass burning plumes from the natural radionuclide tracer 210Pb

During the NASA Global Troposphere Experiment Pacific Exploratory Mission-Tropics (PEM-Tropics) airborne sampling campaign we found unexpectedly high concentrations of aerosol-associated 210Pb throughout the free troposphere over the South Pacific. Because of the remoteness of the study region, we expected specific activities to be generally less than 35 μBq m−3 but found an average in the free troposphere of 107 μBq m−3. This average was elevated by a large number of very active (up to 405 μBq m−3) samples that were associated with biomass burning plumes encountered on nearly every PEM-Tropics flight in the southern hemisphere. We use a simple aging and dilution model, which assumes that 222Rn and primary combustion products are pumped into the free troposphere in wet convective systems over fire regions (most likely in Africa), to explain the elevated 210Pb activities. This model reproduces the observed 210Pb activities very well, and predicts the ratios of four hydrocarbon species (emitted by combustion) to CO to better than 20% in most cases. Plume ages calculated by the model depend strongly on the assumed 222Rn activities in the initial plume, but using values plausible for continental boundary layer air yields ages that are consistent with travel times from Africa to the South Pacific calculated with a back trajectory model. The model also shows that despite being easily recognized through the large enhancements of biomass burning tracers, these plumes must have entrained large fractions of the surrounding ambient air during transport

Revisiting two-step Forbush decreases

Interplanetary coronal mass ejections (ICMEs) and their shocks can sweep out galactic cosmic rays (GCRs), thus creating Forbush decreases (FDs). The traditional model of FDs predicts that an ICME and its shock decrease the GCR intensity in a two-step profile. This model, however, has been the focus of little testing. Thus, our goal is to discover whether a passing ICME and its shock inevitably lead to a two-step FD, as predicted by the model. We use cosmic ray data from 14 neutron monitors and, when possible, high time resolution GCR data from the spacecraft International Gamma Ray Astrophysical Laboratory (INTEGRAL). We analyze 233 ICMEs that should have created two-step FDs. Of these, only 80 created FDs, and only 13 created two-step FDs. FDs are thus less common than predicted by the model. The majority of events indicates that profiles of FDs are more complicated, particularly within the ICME sheath, than predicted by the model. We conclude that the traditional model of FDs as having one or two steps should be discarded. We also conclude that generally ignored small-scale interplanetary magnetic field structure can contribute to the observed variety of FD profiles