Acidosis Is a Key Regulator of Osteoblast Ecto-Nucleotidase Pyrophosphatase/Phosphodiesterase 1 (NPP1) Expression and Activity

Previous work has shown that acidosis prevents bone nodule formation by osteoblasts in vitro by inhibiting mineralisation of the collagenous matrix. The ratio of phosphate (Pi) to pyrophosphate (PPi) in the bone microenvironment is a fundamental regulator of bone mineralisation. Both Pi and PPi, a potent inhibitor of mineralisation, are generated from extracellular nucleotides by the actions of ecto‐nucleotidases. This study investigated the expression and activity of ecto‐nucleotidases by osteoblasts under normal and acid conditions. We found that osteoblasts express mRNA for a number of ecto‐nucleotidases including NTPdase 1–6 (ecto‐nucleoside triphosphate diphosphohydrolase) and NPP1‐3 (ecto‐nucleotide pyrophosphatase/phosphodiesterase). The rank order of mRNA expression in differentiating rat osteoblasts (day 7) was Enpp1 > NTPdase 4 > NTPdase 6 > NTPdase 5 > alkaline phosphatase > ecto‐5‐nucleotidase > Enpp3 > NTPdase 1 > NTPdase 3 > Enpp2 > NTPdase 2. Acidosis (pH 6.9) upregulated NPP1 mRNA (2.8‐fold) and protein expression at all stages of osteoblast differentiation compared to physiological pH (pH 7.4); expression of other ecto‐nucleotidases was unaffected. Furthermore, total NPP activity was increased up to 53% in osteoblasts cultured in acid conditions (P < 0.001). Release of ATP, one of the key substrates for NPP1, from osteoblasts, was unaffected by acidosis. Further studies showed that mineralised bone formation by osteoblasts cultured from NPP1 knockout mice was increased compared with wildtypes (2.5‐fold, P < 0.001) and was partially resistant to the inhibitory effect of acidosis. These results indicate that increased NPP1 expression and activity might contribute to the decreased mineralisation observed when osteoblasts are exposed to acid conditions

Activation of the P2Y2 receptor regulates bone cell function by enhancing ATP release

Bone cells constitutively release ATP into the extracellular environment where it acts locally via P2 receptors to regulate bone cell function. Whilst P2Y2 receptor stimulation regulates bone mineralisation, the functional effects of this receptor in osteoclasts remain unknown. This investigation used the P2Y2 receptor knockout (P2Y2R−/−) mouse model to investigate the role of this receptor in bone. MicroCT analysis of P2Y2R−/− mice demonstrated age-related increases in trabecular bone volume (≤48%), number (≤30%) and thickness (≤17%). In vitro P2Y2R−/− osteoblasts displayed a 3-fold increase in bone formation and alkaline phosphatase activity, whilst P2Y2R−/− osteoclasts exhibited a 65% reduction in resorptive activity. Serum cross-linked C-telopeptide levels (CTX, resorption marker) were also decreased (≤35%). The resorption defect in P2Y2R−/− osteoclasts was rescued by the addition of exogenous ATP, suggesting that an ATP deficit could be a key factor in the reduced function of these cells. In agreement, we found that basal ATP release was reduced up to 53% in P2Y2R−/− osteoclasts. The P2Y2 receptor agonists, UTP and 2-thioUTP, increased osteoclast activity and ATP release in wild-type but not in P2Y2R−/− cells. This indicates that the P2Y2 receptor may regulate osteoclast function indirectly by promoting ATP release. UTP and 2-thioUTP also stimulate ATP release from osteoblasts suggesting that the P2Y2 receptor exerts a similar function in these cells. Taken together, our findings are consistent with the notion that the primary action of P2Y2 receptor signalling in bone is to regulate extracellular ATP levels

Stellar Hydrodynamics in Radiative Regions

We present an analysis of the response of a radiative region to waves generated by a convective region of the star; this wave treatment of the classical problem of ``overshooting'' gives extra mixing relative to the treatment traditionally used in stellar evolutionary codes. The interface between convectively stable and unstable regions is dynamic and nonspherical, so that the nonturbulent material is driven into motion, even in the absence of ``penetrative overshoot.'' These motions may be described by the theory of nonspherical stellar pulsations, and are related to motion measured by helioseismology. Multi-dimensional numerical simulations of convective flow show puzzling features which we explain by this simplified physical model. Gravity waves generated at the interface are dissipated, resulting in slow circulation and mixing seen outside the formal convection zone. The approach may be extended to deal with rotation and composition gradients. Tests of this description in the stellar evolution code TYCHO produce carbon stars on the asymptotic giant branch (AGB), an isochrone age for the Hyades and three young clusters with lithium depletion ages from brown dwarfs, and lithium and beryllium depletion consistent with observations of the Hyades and Pleiades, all without tuning parameters. The insight into the different contributions of rotational and hydrodynamic mixing processes could have important implications for realistic simulation of supernovae and other questions in stellar evolution.Comment: 27 pages, 5 figures, accepted to the Astrophysical Journa

Observational Tests and Predictive Stellar Evolution II: Non-standard Models

We examine contributions of second order physical processes to results of stellar evolution calculations amenable to direct observational testing. In the first paper in the series (Young et al. 2001) we established baseline results using only physics which are common to modern stellar evolution codes. In the current paper we establish how much of the discrepancy between observations and baseline models is due to particular elements of new physics. We then consider the impact of the observational uncertainties on the maximum predictive accuracy achievable by a stellar evolution code. The sun is an optimal case because of the precise and abundant observations and the relative simplicity of the underlying stellar physics. The Standard Model is capable of matching the structure of the sun as determined by helioseismology and gross surface observables to better than a percent. Given an initial mass and surface composition within the observational errors, and no additional constraints for which the models can be optimized, it is not possible to predict the sun's current state to better than ~7%. Convectively induced mixing in radiative regions, seen in multidimensional hydrodynamic simulations, dramatically improves the predictions for radii, luminosity, and apsidal motions of eclipsing binaries while simultaneously maintaining consistency with observed light element depletion and turnoff ages in young clusters (Young et al. 2003). Systematic errors in core size for models of massive binaries disappear with more complete mixing physics, and acceptable fits are achieved for all of the binaries without calibration of free parameters. The lack of accurate abundance determinations for binaries is now the main obstacle to improving stellar models using this type of test.Comment: 33 pages, 8 figures, accepted for publication in the Astrophysical Journa

Neutrino-driven Explosions

The question why and how core-collapse supernovae (SNe) explode is one of the central and most long-standing riddles of stellar astrophysics. A solution is crucial for deciphering the SN phenomenon, for predicting observable signals such as light curves and spectra, nucleosynthesis, neutrinos, and gravitational waves, for defining the role of SNe in the evolution of galaxies, and for explaining the birth conditions and properties of neutron stars (NSs) and stellar-mass black holes. Since the formation of such compact remnants releases over hundred times more energy in neutrinos than the SN in the explosion, neutrinos can be the decisive agents for powering the SN outburst. According to the standard paradigm of the neutrino-driven mechanism, the energy transfer by the intense neutrino flux to the medium behind the stagnating core-bounce shock, assisted by violent hydrodynamic mass motions (sometimes subsumed by the term "turbulence"), revives the outward shock motion and thus initiates the SN blast. Because of the weak coupling of neutrinos in the region of this energy deposition, detailed, multidimensional hydrodynamic models including neutrino transport and a wide variety of physics are needed to assess the viability of the mechanism. Owing to advanced numerical codes and increasing supercomputer power, considerable progress has been achieved in our understanding of the physical processes that have to act in concert for the success of neutrino-driven explosions. First studies begin to reveal observational implications and avenues to test the theoretical picture by data from individual SNe and SN remnants but also from population-integrated observables. While models will be further refined, a real breakthrough is expected through the next Galactic core-collapse SN, when neutrinos and gravitational waves can be used to probe the conditions deep inside the dying star. (abridged)Comment: Author version of chapter for 'Handbook of Supernovae,' edited by A. Alsabti and P. Murdin, Springer. 54 pages, 13 figure

A dynamical model of surrogate reactions

A new dynamical model is developed to describe the whole process of surrogate reactions; transfer of several nucleons at an initial stage, thermal equilibration of residues leading to washing out of shell effects and decay of populated compound nuclei are treated in a unified framework. Multi-dimensional Langevin equations are employed to describe time-evolution of collective coordinates with a time-dependent potential energy surface corresponding to different stages of surrogate reactions. The new model is capable of calculating spin distributions of the compound nuclei, one of the most important quantity in the surrogate technique. Furthermore, various observables of surrogate reactions can be calculated, e.g., energy and angular distribution of ejectile, and mass distributions of fission fragments. These features are important to assess validity of the proposed model itself, to understand mechanisms of the surrogate reactions and to determine unknown parameters of the model. It is found that spin distributions of compound nuclei produced in $^{18}$O+$^{238}$U $\rightarrow ^{16}$O+$^{240*}$U and $^{18}$O+$^{236}$U $\rightarrow ^{16}$O+$^{238*}$U reactions are equivalent and much less than 10$\hbar$, therefore satisfy conditions proposed by Chiba and Iwamoto (PRC 81, 044604(2010)) if they are used as a pair in the surrogate ratio method.Comment: 17 pages, 5 figure

Effective one-body approach to general relativistic two-body dynamics

We map the general relativistic two-body problem onto that of a test particle moving in an effective external metric. This effective-one-body approach defines, in a non-perturbative manner, the late dynamical evolution of a coalescing binary system of compact objects. The transition from the adiabatic inspiral, driven by gravitational radiation damping, to an unstable plunge, induced by strong spacetime curvature, is predicted to occur for orbits more tightly bound than the innermost stable circular orbit in a Schwarzschild metric of mass M = m1 + m2. The binding energy, angular momentum and orbital frequency of the innermost stable circular orbit for the time-symmetric two-body problem are determined as a function of the mass ratio.Comment: 52 pages, RevTex, epsfig, 8 figure

Observing Supernova 1987A with the Refurbished Hubble Space Telescope

Observations with the Hubble Space Telescope (HST), conducted since 1990, now offer an unprecedented glimpse into fast astrophysical shocks in the young remnant of supernova 1987A. Comparing observations taken in 2010 using the refurbished instruments on HST with data taken in 2004, just before the Space Telescope Imaging Spectrograph failed, we find that the Ly-a and H-a lines from shock emission continue to brighten, while their maximum velocities continue to decrease. We observe broad blueshifted Ly-a, which we attribute to resonant scattering of photons emitted from hotspots on the equatorial ring. We also detect NV~\lambda\lambda 1239,1243 A line emission, but only to the red of Ly-A. The profiles of the NV lines differ markedly from that of H-a, suggesting that the N^{4+} ions are scattered and accelerated by turbulent electromagnetic fields that isotropize the ions in the collisionless shock.Comment: Science, accepted. Science Express, 02 Sept 2010. 5 figures. Supporting online material can be found at http://www.sciencemag.org/cgi/content/full/sci;science.1192134/DC

Explosive Nucleosynthesis: What we learned and what we still do not understand

This review touches on historical aspects, going back to the early days of nuclear astrophysics, initiated by B$^2$FH and Cameron, discusses (i) the required nuclear input from reaction rates and decay properties up to the nuclear equation of state, continues (ii) with the tools to perform nucleosynthesis calculations and (iii) early parametrized nucleosynthesis studies, before (iv) reliable stellar models became available for the late stages of stellar evolution. It passes then through (v) explosive environments from core-collapse supernovae to explosive events in binary systems (including type Ia supernovae and compact binary mergers), and finally (vi) discusses the role of all these nucleosynthesis production sites in the evolution of galaxies. The focus is put on the comparison of early ideas and present, very recent, understanding.Comment: 11 pages, to appear in Springer Proceedings in Physics (Proc. of Intl. Conf. "Nuclei in the Cosmos XV", LNGS Assergi, Italy, June 2018

Post-Newtonian Models of Binary Neutron Stars

Using an energy variational method, we calculate quasi-equilibrium configurations of binary neutron stars modeled as compressible triaxial ellipsoids obeying a polytropic equation of state. Our energy functional includes terms both for the internal hydrodynamics of the stars and for the external orbital motion. We add the leading post-Newtonian (PN) corrections to the internal and gravitational energies of the stars, and adopt hybrid orbital terms which are fully relativistic in the test-mass limit and always accurate to PN order. The total energy functional is varied to find quasi-equilibrium sequences for both corotating and irrotational binaries in circular orbits. We examine how the orbital frequency at the innermost stable circular orbit depends on the polytropic index n and the compactness parameter GM/Rc^2. We find that, for a given GM/Rc^2, the innermost stable circular orbit along an irrotational sequence is about 17% larger than the innermost secularly stable circular orbit along the corotating sequence when n=0.5, and 20% larger when n=1. We also examine the dependence of the maximum neutron star mass on the orbital frequency and find that, if PN tidal effects can be neglected, the maximum equilibrium mass increases as the orbital separation decreases.Comment: 53 pages, LaTex, 9 figures as 10 postscript files, accepted by Phys. Rev. D, replaced version contains updated reference