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

Human sperm accumulation near surfaces: a simulation study

A hybrid boundary integral/slender body algorithm for modelling flagellar cell motility is presented. The algorithm uses the boundary element method to represent the ‘wedge-shaped’ head of the human sperm cell and a slender body theory representation of the flagellum. The head morphology is specified carefully due to its significant effect on the force and torque balance and hence movement of the free-swimming cell. The technique is used to investigate the mechanisms for the accumulation of human spermatozoa near surfaces. Sperm swimming in an infinite fluid, and near a plane boundary, with prescribed planar and three-dimensional flagellar waveforms are simulated. Both planar and ‘elliptical helicoid’ beating cells are predicted to accumulate at distances of approximately 8.5–22 μm from surfaces, for flagellar beating with angular wavenumber of 3π to 4π. Planar beating cells with wavenumber of approximately 2.4π or greater are predicted to accumulate at a finite distance, while cells with wavenumber of approximately 2π or less are predicted to escape from the surface, likely due to the breakdown of the stable swimming configuration. In the stable swimming trajectory the cell has a small angle of inclination away from the surface, no greater than approximately 0.5°. The trapping effect need not depend on specialized non-planar components of the flagellar beat but rather is a consequence of force and torque balance and the physical effect of the image systems in a no-slip plane boundary. The effect is relatively weak, so that a cell initially one body length from the surface and inclined at an angle of 4°–6° towards the surface will not be trapped but will rather be deflected from the surface. Cells performing rolling motility, where the flagellum sweeps out a ‘conical envelope’, are predicted to align with the surface provided that they approach with sufficiently steep angle. However simulation of cells swimming against a surface in such a configuration is not possible in the present framework. Simulated human sperm cells performing a planar beat with inclination between the beat plane and the plane-of-flattening of the head were not predicted to glide along surfaces, as has been observed in mouse sperm. Instead, cells initially with the head approximately 1.5–3 μm from the surface were predicted to turn away and escape. The simulation model was also used to examine rolling motility due to elliptical helicoid flagellar beating. The head was found to rotate by approximately 240° over one beat cycle and due to the time-varying torques associated with the flagellar beat was found to exhibit ‘looping’ as has been observed in cells swimming against coverslips

A Catalog of Absorption Lines in Eight HST/STIS E230M 1.0 < z < 1.7 Quasar Spectra

We have produced a catalog of line identifications and equivalent width measurements for all absorption features in eight ultraviolet echelle quasar spectra. These spectra were selected as having the highest signal-to-noise among the HST/STIS spectra obtained with the E230M grating. We identify 56 metal-line systems toward the eight quasars, and present plots of detected transitions, aligned in velocity-space. We found that about 1/4 - 1/3 of the features in the Lya forest region, redward of the incidence of the Lyb forest, are metal lines. High ionization transitions are common. We see both O VI and C IV in 88 - 90% of the metal-line systems for which the spectra cover the expected wavelength. Si III is seen in 58%, while low ionization absorption in C II, Si II, and/or Al II is detected in 50% of the systems for which they are covered. This catalog will facilitate future studies of the Lya forest and of metal-line systems of various types.Comment: 13 pages, 2 figures, submitted to Monthly Notices of the Royal Astronomical Society, a complete version with the appendix and all figures is available at http://www.astro.psu.edu/users/misawa/pub/Paper/qalcat.pdf.g

Nonlinear instability in flagellar dynamics: a notel modulation mechanism in sperm migration

Throughout biology, cells and organisms use flagella and cilia to propel fluid and achieve motility. The beating of these organelles, and the corresponding ability to sense, respond to and modulate this beat is central to many processes in health and disease. While the mechanics of flagellum–fluid interaction has been the subject of extensive mathematical studies, these models have been restricted to being geometrically linear or weakly nonlinear, despite the high curvatures observed physiologically. We study the effect of geometrical nonlinearity, focusing on the spermatozoon flagellum. For a wide range of physiologically relevant parameters, the nonlinear model predicts that flagellar compression by the internal forces initiates an effective buckling behaviour, leading to a symmetry-breaking bifurcation that causes profound and complicated changes in the waveform and swimming trajectory, as well as the breakdown of the linear theory. The emergent waveform also induces curved swimming in an otherwise symmetric system, with the swimming trajectory being sensitive to head shape—no signalling or asymmetric forces are required. We conclude that nonlinear models are essential in understanding the flagellar waveform in migratory human sperm; these models will also be invaluable in understanding motile flagella and cilia in other systems

Doing more with less: the flagellar end piece enhances the propulsive effectiveness of human spermatozoa

Spermatozoa self-propel by propagating bending waves along a predominantly active elastic flagellum. The organized structure of the "9 + 2" axoneme is lost in the most-distal few microns of the flagellum, and therefore this region is unlikely to have the ability to generate active bending; as such it has been largely neglected in biophysical studies. Through elastohydrodynamic modeling of human-like sperm we show that an inactive distal region confers significant advantages, both in propulsive thrust and swimming efficiency, when compared with a fully active flagellum of the same total length. The beneficial effect of the inactive end piece on these statistics can be as small as a few percent but can be above 430%. The optimal inactive length, between 2-18% of the total length, depends on both wavenumber and viscous-elastic ratio, and therefore is likely to vary in different species. Potential implications in evolutionary biology and clinical assessment are discussed.Comment: To Appear, Physical Review Fluids. 25 pages, 14 figure

Hyaluronic Acid Binding Sperm Selection for assisted reproduction treatment (HABSelect): Study protocol for a multicentre randomised controlled trial

© Published by the BMJ Publishing Group Limited.Introduction The selection of a sperm with good genomic integrity is an important consideration for improving intracytoplasmic sperm injection (ICSI) outcome. Current convention selects sperm by vigour and morphology, but preliminary evidence suggests selection based on hyaluronic acid binding may be beneficial. The aim of the Hyaluronic Acid Binding Sperm Selection (HABSelect) trial is to determine the efficacy of hyaluronic acid (HA)-selection of sperm versus conventionally selected sperm prior to ICSI on live birth rate (LBR). The mechanistic aim is to assess whether and how the chromatin state of HA-selected sperm corresponds with clinical outcomes - clinical pregnancy rate (CPR), LBR and pregnancy loss (PL). Methods and analysis Couples attending UK Centres will be approached, eligibility screening performed and informed consent sought. Randomisation will occur within 24 hours prior to ICSI treatment. Participants will be randomly allocated 1:1 to the intervention arm (physiological intracytoplasmic sperm injection, PICSI) versus the control arm using conventional methods (ICSI). The primary clinical outcome is LBR ≥37 weeks' gestation with the mechanistic study determining LBR's relationship with sperm DNA integrity. Secondary outcomes will determine this for CPR and PL. Only embryologists performing the procedure will be aware of the treatment allocation. Steps will be taken to militate against biases arising from embryologists being non-blinded. Randomisation will use a minimisation algorithm to balance for key prognostic variables. The trial is powered to detect a 5% difference (24-29%: p=0.05) in LBR ≥37 weeks' gestation. Selected residual sperm samples will be tested by one or more assays of DNA integrity. Ethics and dissemination HABSelect is a UK NIHR-EME funded study (reg no 11/14/34; IRAS REF. 13/YH/0162). The trial was designed in partnership with patient and public involvement to help maximise patient benefits. Trial findings will be reported as per CONSORT guidelines and will be made available in lay language via the trial web site (http://www.habselect.org.uk/). Trial registration number ISRCTN99214271; Pre-results

Human sperm swimming in a high viscosity mucus analogue

Remarkably, mammalian sperm maintain a substantive proportion of their progressive swimming speed within highly viscous fluids, including those of the female reproductive tract. Here, we analyse the digital microscopy of a human sperm swimming in a highly viscous, weakly elastic mucus analogue. We exploit principal component analysis to simplify its flagellar beat pattern, from which boundary element calculations are used to determine the time-dependent flow field around the sperm cell. The sperm flow field is further approximated in terms of regularized point forces, and estimates of the mechanical power consumption are determined, for comparison with analogous low viscosity media studies. This highlights extensive differences in the structure of the flows surrounding human sperm in different media, indicating how the cell-cell and cell- boundary hydrodynamic interactions significantly differ with the physical microenvironment. The regularized point force decomposition also provides cell-level information that may ulti- mately be incorporated into sperm population models. We further observe indications that the core feature in explaining the effectiveness of sperm swimming in high viscosity media is the loss of cell yawing, which is related with a greater density of regularized point force singularities along the axis of symmetry of the flagellar beat to represent the flow field. In turn this implicates a reduction of the wavelength of the distal beat pattern – and hence dynamical wavelength selection of the flagellar beat – as the dominant feature governing the effectiveness of sperm swimming in highly viscous media

Bottom sediments of Lake Rotoma

Lake Rotoma is a deep (70-80 m), oligotrophic, warm monomictic lake of volcanic origin with insignificant stream inflow and no clearly defined outflow. For at least 60 years up to 1972 the lake level fluctuated markedly about an overall rising trend of some 6-10 m. Nearshore profiles are related to the prevailing wave climate superimposed upon the overall rising lake level, shelves being wider, less steep, and deeper about the more exposed eastern and southern shorelines. The outer portions of shelves extending well below modern storm wave base into waters as deep as 15-25 m are relict features from lower lake level stands. Sediments fine from sand-gravel mixtures nearshore to silts in basinal areas. Their composition reflects a composite provenance involving the lavas and tephras about the lake, as well as intralake diatom frustules and organic matter. The distribution pattern of surficial bottom sediments is an interplay between grains of both biological and terrigenous origin, supplied presently and in the past by a variety of processes, that have been dispersed either by the modern hydrodynamic regime or by former ones associated with lower lake levels. These interrelationships are structured by erecting 5 process-age sediment classes in the lake, namely neoteric, amphoteric, proteric, palimpsest, and relict sediments, analogous to categories postulated for sediments on oceanic continental shelves. Short-core stratigraphy includes the Kaharoa (A.D. -1020) and Tarawera (A.D. 1886) tephras. The rates of sedimentation of diatomaceous silts in basinal areas have more than doubled since the Tarawera eruption, indicating an overall increase in the fertility level of lake waters associated, perhaps, with recent farm development in the catchment

Fall 1956

Greetings from the Club President (page 2) Winter School and Turf Conference Cancelled for 1957 (2) Value of Turf Clippings (3) Dedication--Minor J. Markuson (4) Interrelations between Design, Construction and Maintenance (6) Golf Course Design (6) Relationship of Golf Course Design to Maintenance Costs (9) Influence of Construction upon maintenance (12) Snow Blue - Snow White (14) Post Emergence Control of Crabgrass (15) Third Generation in Greenkeeping (16) Hand Greens Mowers (17) Equipment (18) Pelham Country Club vs New England Thruway (18) Should Junior Memberships be Allowed (19) Meet the Staff (20) Turf Club News (23) Chairman-superintendent Relationship: Quote by Dickinson (24) Letter To Dean Sieling from New England Golf Course Superintendent\u27s Association (24) Associate Members of Stockbridge Turf Management Club (25) Membership Application Form for Turf Management Club (26

Proactive and politically skilled professionals: What is the relationship with affective occupational commitment?

The aim of this study is to extend research on employee affective commitment in three ways: (1) instead of organizational commitment the focus is on occupational commitment; (2) the role of proactive personality on affective occupational commitment is examined; and (3) occupational satisfaction is examined as a mediator and political skills as moderator in the relationship between proactive personality and affective occupational commitment. Two connected studies, one in a hospital located in the private sector and one in a university located in the public sector, are carried out in Pakistan, drawing on a total sample of over 400 employees. The results show that proactive personality is positively related to affective occupational commitment, and that occupational satisfaction partly mediates the relationship between proactive personality and affective occupational commitment. No effect is found for a moderator effect of political skills in the relationship between proactive personality and affective occupational commitment. Political skills however moderate the relationship between proactive personality and affective organizational commitment