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

Assessing Honors Internationalization: A Case Study of Lloyd International Honors College at UNC Greensboro

Lloyd International Honors College (LIHC) of the University of North Carolina at Greensboro (UNC Greensboro) is a useful example of the reimagining of a traditional honors program into an honors college with an international focus.1 The process of becoming an internationally focused honors college, which began in 2006, was part of the university’s strategic goal of internationalizing its curriculum, student body, faculty, and culture. It has involved an extended process of program development; campus-wide partnership building, specifically in conjunction with the university’s International Programs Center (IPC) and Global Engagement Quality Enhancement Plan (QEP); and iterative assessment. This chapter outlines the internationalization of the honors college as it is embedded in an iterative assessment process. In doing so, it highlights the implementation of international programs and structures at the university and in the honors college, defines the assessment framework the university and honors used to guide their internationalization efforts, discusses specific assessment measures and outcomes, and considers future directions

The biology of Australian seagrasses

Seagrasses are angiosperms which have ventured into the marine environment, where they produce totally submerged flowers which are pollinated under water. They occur in many regions of the world,but the following account is concerned primarily with studies on the biology of these plants in Australian waters. It also attempts to contrast some of the features displayed by the seagrasses with those of more familiar terrestrial plants

The Deuterium Abundance in the z=0.7 absorber towards QSO PG1718+4807

We report a further analysis of the ratio of deuterium to hydrogen (D/H) using HST spectra of the z=0.701 Lyman limit system towards the QSO PG1718+481. Initial analyses of this absorber found it gave a high D/H value, 1.8 - 3.1 \times 10^{-4} (Webb et al. 1998), inconsistent with several higher redshift measurements. It is thus important to critically examine this measurement. By analysing the velocity widths of the DI, HI and metal lines present in this system, Kirkman et al. (2001) report that the additional absorption in the blue wing of the lya line can not be DI, with a confidence level of 98%. Here we present a more detailed analysis, taking into account possible wavelength shifts between the three sets of HST spectra used in the analysis. We find that the constraints on this system are not as strong as those claimed by Kirkman et al. The discrepancy between the parameters of the blue wing absorption and the parameters expected for DI is marginally worse than 1 sigma. Tytler et al.(1999) commented on the first analysis of Webb et al.(1997,1998), reporting the presence of a contaminating lower redshift Lyman limit system, with log[N(HI)] = 16.7 at z=0.602, which biases the N(HI) estimate for the main system. Here we show that this absorber actually has log[N(HI)] < 14.6 and does not impact on the estimate of N(HI) in the system of interest at z = 0.701. The purpose of the present paper is to highlight important aspects of the analysis which were not explored in previous studies, and hence help refine the methods used in future analyses of D/H in quasar spectra.Comment: 11 pages, 9 figures. Accepted by MNRA

Intrinsic Properties of the <z>=2.7 Lyman Alpha Forest from Keck Spectra of QSO HS 1946+7658

We present the highest quality Lyman Alpha forest spectra published to date, from the QSO HS 1946+7658. The distribution of H I column densities is a power law of slope -1.5 from Log N = 12.1 - 14. This power law can extend to N = 0, because lines weaker than Log N = 12.1 do not have a large H I optical depth. Low column lines with Log N > 9 could account for all observed He II absorption, but lines with Log N > 12 alone are unlikely to do so. The b distribution between 20 and 60 km/sec is a Gaussian with a mean of 23 km/sec (less than reported in past at this z), and a sigma b of 14 km/sec. We report no evolution in the Lyman alpha forest (except the number of lines), because Lu et al. (1997) found the same N and b distributions at = 3.7. We see lines with 14 80 km/sec that cannot be accounted for by noise or blending effects. We discover that the lower cutoff in the b distribution varies with N, from b = 14 km/sec at Log N = 12.5 to b = 22 km/sec at Log N = 14.0, but otherwise b and N are not correlated. We see no Lyman Alpha line clustering above 50 \kms, in disagreement with previous results from lower signal to noise data, but we do see a 3 sigma clustering signal at 25 - 50 km/sec among lines with Log N > 13.6Comment: (Minor changes including new identifications for two weak lines) 46 pages including 16 Figures, Latex Table 1 Available at http://nately.ucsd.edu/~david . To appear in Ap

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

QSOs and Absorption Line Systems Surrounding the Hubble Deep Field

We have imaged a 45x45 sq. arcmin. area centered on the Hubble Deep Field (HDF) in UBVRI passbands, down to respective limiting magnitudes of approximately 21.5, 22.5, 22.2, 22.2, and 21.2. The principal goals of the survey are to identify QSOs and to map structure traced by luminous galaxies and QSO absorption line systems in a wide volume containing the HDF. We have selected QSO candidates from color space, and identified 4 QSOs and 2 narrow emission-line galaxies (NELGs) which have not previously been discovered, bringing the total number of known QSOs in the area to 19. The bright z=1.305 QSO only 12 arcmin. away from the HDF raises the northern HDF to nearly the same status as the HDF-S, which was selected to be proximate to a bright QSO. About half of the QSO candidates remain for spectroscopic verification. Absorption line spectroscopy has been obtained for 3 bright QSOs in the field, using the Keck 10m, ARC 3.5m, and MDM 2.4m telescopes. Five heavy-element absorption line systems have been identified, 4 of which overlap the well-explored redshift range covered by deep galaxy redshift surveys towards the HDF. The two absorbers at z=0.5565 and z=0.5621 occur at the same redshift as the second most populated redshift peak in the galaxy distribution, but each is more than 7Mpc/h (comoving, Omega_M=1, Omega_L=0) away from the HDF line of sight in the transverse dimension. This supports more indirect evidence that the galaxy redshift peaks are contained within large sheet-like structures which traverse the HDF, and may be precursors to large-scale ``pancake'' structures seen in the present-day galaxy distribution.Comment: 36 pages, including 9 figures and 8 tables. Accepted for publication in the Astronomical Journa

Crossing the Suspension Bridge: Navigating the Road from School Suspension to College Success – How Some Students Have Overcome the Negative Implications of School Suspension to Bridge the Road to College

Out-of-school suspensions for middle and high school students can have negative, long-lasting consequences. Researchers have documented that suspensions have a negative impact on academic development, increase likelihood of dropping out of school, and are associated with a stronger likelihood that students will be involved in the legal system. However, there are students who overcome these negative statistics and matriculate to post-secondary education successfully. This study examines the lived experiences and personal attributes in students’ lives that enabled them to overcome a history of suspension to enter and succeed in higher education. Using Bronfenbrenner’s ecological model, the study’s researchers interviewed individuals who have a history of middle and/or high school suspensions and matriculated to higher education. Findings suggest that sense of belonging; family, home,/school support; and strength of relationships helped participants neutralize the impact and mitigate the negative aspects of suspension

Intergalactic HeII absorption towards QSO 1157+3143

We report the discovery of a further line of sight allowing detection of HeII Ly alpha absorption by the intergalactic medium. A HST/STIS survey of 32 bright z ~ 3 quasars selected from the Hamburg Quasar Surveys yielded one detection toward QSO 1157+3143 (z ~ 3, B ~ 17). A 10 orbit follow-up spectrum reveals a UV spectrum significantly supressed by two intervening Lyman limit systems at z=2.77 and 2.94, but with the continuum flux recovering sufficiently shortward of ~ 1700 A to allow study of the HeII absorption spectrum in the redshift range 2.75 < z < 2.97. The absorption is characterized by alternating voids and dense filament structures seen in both HeII and HI. Attempts to model the HeII opacity in terms of HI Ly alpha forest absorption are not successful in the voids, suggesting that HeII reionization is not complete between z=2.77 and 2.97 or that an optically thin Lyman limit system with z ~ 0.3 is responsible for the additional opacity.Comment: 6 pages, 4 figures, accepted by A&

Spectroscopic Analysis of H I Absorption Line Systems in 40 HIRES Quasars

We list and analyze H I absorption lines at redshifts 2 < z < 4 with column density (12 < log(N_HI) < 19) in 40 high-resolutional (FWHM = 8.0 km/s) quasar spectra obtained with the Keck+HIRES. We de-blend and fit all H I lines within 1,000 km/s of 86 strong H I lines whose column densities are log(N_HI/[cm^-2]) > 15. Unlike most prior studies, we use not only Lya but also all visible higher Lyman series lines to improve the fitting accuracy. This reveals components near to higher column density systems that can not be seen in Lya. We list the Voigt profile fits to the 1339 H I components that we found. We examined physical properties of H I lines after separating them into several sub-samples according to their velocity separation from the quasars, their redshift, column density and the S/N ratio of the spectrum. We found two interesting trends for lines with 12 < log(N_HI) < 15 which are within 200-1000 km/s of systems with log(N_HI) > 15. First, their column density distribution becomes steeper, meaning relatively fewer high column density lines, at z < 2.9. Second, their column density distribution also becomes steeper and their line width becomes broader by about 2-3 km/s when they are within 5,000 km/s of their quasar.Comment: 32 pages, 14 figures, accepted for publication in the Astronomical Journal. A complete version with all tables and figures is available at http://www.astro.psu.edu/users/misawa/pub/Paper/40hires.ps.g