Differentiated Smooth Muscle Cells Generate a Subpopulation of Resident Vascular Progenitor Cells in the Adventitia Regulated by Klf4

RATIONALE: The vascular adventitia is a complex layer of the vessel wall consisting of vasa vasorum microvessels, nerves, fibroblasts, immune cells, and resident progenitor cells. Adventitial progenitors express the stem cell markers, Sca1 and CD34 (adventitial sca1-positive progenitor cells [AdvSca1]), have the potential to differentiate in vitro into multiple lineages, and potentially contribute to intimal lesions in vivo. OBJECTIVE: Although emerging data support the existence of AdvSca1 cells, the goal of this study was to determine their origin, degree of multipotency and heterogeneity, and contribution to vessel remodeling. METHODS AND RESULTS: Using 2 in vivo fate-mapping approaches combined with a smooth muscle cell (SMC) epigenetic lineage mark, we report that a subpopulation of AdvSca1 cells is generated in situ from differentiated SMCs. Our data establish that the vascular adventitia contains phenotypically distinct subpopulations of progenitor cells expressing SMC, myeloid, and hematopoietic progenitor-like properties and that differentiated SMCs are a source to varying degrees of each subpopulation. SMC-derived AdvSca1 cells exhibit a multipotent phenotype capable of differentiating in vivo into mature SMCs, resident macrophages, and endothelial-like cells. After vascular injury, SMC-derived AdvSca1 cells expand in number and are major contributors to adventitial remodeling. Induction of the transcription factor Klf4 in differentiated SMCs is essential for SMC reprogramming in vivo, whereas in vitro approaches demonstrate that Klf4 is essential for the maintenance of the AdvSca1 progenitor phenotype. CONCLUSIONS: We propose that generation of resident vascular progenitor cells from differentiated SMCs is a normal physiological process that contributes to the vascular stem cell pool and plays important roles in arterial homeostasis and disease

On the evolution of a star cluster and its multiple stellar systems following gas dispersal

We investigate the evolution, following gas dispersal, of a star cluster produced from a hydrodynamical calculation. We find that when the gas, initially comprising 60% of the mass, is removed, the system settles into a bound cluster containing 30-40% of the stellar mass surrounding by an expanding halo of ejected stars. The bound cluster expands from an initial radius of <0.05 pc to 1-2 pc over 4-10 Myr, depending on how quickly the gas is removed, implying that stellar clusters may begin with far higher stellar densities than usually assumed. With rapid gas dispersal the most massive stars are found to be mass segregated for the first ~1 Myr of evolution, but classical mass segregation only develops for cases with long gas removal timescales. Eventually, many of the most massive stars are expelled from the bound cluster. Despite the high initial stellar density and the extensive dynamical evolution of the system, we find that the stellar multiplicity is almost constant during the 10 Myr of evolution. This is because the primordial multiple systems are formed in a clustered environment and, thus, by their nature are already resistant to further evolution. The majority of multiple system evolution is confined to the decay of high-order systems and the formation of a significant population of very wide (10^4-10^5 AU) multiple systems in the expanding halo. This formation mechanism for wide binaries potentially solves the problem of how most stars apparently form in clusters and yet a substantial population of wide binaries exist in the field. Many of these wide binaries and the binaries produced by the decay of high-order multiple systems have unequal mass components, potentially solving the problem that hydrodynamical simulations of star formation are found to under-produce unequal-mass solar-type binaries.Comment: Accepted by MNRAS, 18 pages, 13 figure

A parametric physical model for the intracluster medium and its use in joint SZ/X-ray analyses of galaxy clusters

We present a parameterized model of the intra-cluster medium that is suitable for jointly analysing pointed observations of the Sunyaev-Zel'dovich (SZ) effect and X-ray emission in galaxy clusters. The model is based on assumptions of hydrostatic equilibrium, the Navarro, Frenk and White (NFW) model for the dark matter, and a softened power law profile for the gas entropy. We test this entropy-based model against high and low signal-to-noise mock observations of a relaxed and recently-merged cluster from N-body/hydrodynamic simulations, using Bayesian hyper-parameters to optimise the relative statistical weighting of the mock SZ and X-ray data. We find that it accurately reproduces both the global values of the cluster temperature, total mass and gas mass fraction (fgas), as well as the radial dependencies of these quantities outside of the core (r > kpc). For reference we also provide a comparison with results from the single isothermal beta model. We confirm previous results that the single isothermal beta model can result in significant biases in derived cluster properties.Comment: Published in MNRAS. 20 pages. 9 figure

Importance of the Initial Conditions for Star Formation - III: Statistical Properties of Embedded Protostellar Clusters

We investigate the formation of protostellar clusters during the collapse of dense molecular cloud cores with a focus on the evolution of potential and kinetic energy, the degree of substructure, and the early phase of mass segregation. Our study is based on a series of hydrodynamic simulations of dense cores, where we vary the initial density profile and the initial turbulent velocity. In the three-dimensional adaptive mesh refinement simulations, we follow the dynamical formation of filaments and protostars until a star formation efficiency of 20%. Despite the different initial configurations, the global ensemble of all protostars in a setup shows a similar energy evolution and forms sub-virial clusters with an energy ratio $E_\mathrm{kin}/|E_\mathrm{pot}|\sim0.2$. Concentrating on the innermost central region, the clusters show a roughly virialised energy balance. However, the region of virial balance only covers the innermost $\sim10-30$ of all the protostars. In all simulations with multiple protostars, the total kinetic energy of the protostars is higher than the kinetic energy of the gas cloud, although the protostars only contain 20% of the total mass. The clusters vary significantly in size, mass, and number of protostars, and show different degrees of substructure and mass segregation. Flat density profiles and compressive turbulent modes produce more subclusters then centrally concentrated profiles and solenoidal turbulence. We find that dynamical relaxation and hence dynamical mass segregation is very efficient in all cases from the very beginning of the nascent cluster, i.e., during a phase when protostars are constantly forming and accreting.Comment: 19 pages, MNRAS accepte

21 cm fluctuations from inhomogeneous X-ray heating before reionization

Many models of early structure formation predict a period of heating immediately preceding reionization, when X-rays raise the gas temperature above that of the cosmic microwave background. These X-rays are often assumed to heat the intergalactic medium (IGM) uniformly, but in reality they will heat the gas more strongly closer to the sources. We develop a framework for calculating fluctuations in the 21 cm brightness temperature that originate from this spatial variation in the heating rate. High-redshift sources are highly clustered, leading to significant gas temperature fluctuations (with fractional variations ~40%, peaking on k~0.1 Mpc^{-1} scales). This induces a distinctive peak-trough structure in the angle-averaged 21 cm power spectrum, which may be accessible to the proposed Square Kilometre Array. This signal reaches the ~10 mK level, and is stronger than that induced by Lyman alpha flux fluctuations. As well as probing the thermal evolution of the IGM before reionization, this 21 cm signal contains information about the spectra of the first X-ray sources. Finally, we consider disentangling temperature, density and Lyman alpha flux fluctuations as functions of redshift.Comment: 14 pages, 12 figures, accepted by MNRAS, subsection and figure added, text reworked for clarit

Cosmology at Low Frequencies: The 21 cm Transition and the High-Redshift Universe

Observations of the high-redshift Universe with the 21 cm hyperfine line of neutral hydrogen promise to open an entirely new window onto the early phases of cosmic structure formation. Here we review the physics of the 21 cm transition, focusing on processes relevant at high redshifts, and describe the insights to be gained from such observations. These include measuring the matter power spectrum at z~50, observing the formation of the cosmic web and the first luminous sources, and mapping the reionization of the intergalactic medium. The epoch of reionization is of particular interest, because large HII regions will seed substantial fluctuations in the 21 cm background. We also discuss the experimental challenges involved in detecting this signal, with an emphasis on the Galactic and extragalactic foregrounds. These increase rapidly toward low frequencies and are especially severe for the highest redshift applications. Assuming that these difficulties can be overcome, the redshifted 21 cm line will offer unique insight into the high-redshift Universe, complementing other probes but providing the only direct, three-dimensional view of structure formation from z~200 to z~6.Comment: extended review accepted by Physics Reports, 207 pages, 44 figures (some low resolution); version with high resolution figures available at http://pantheon.yale.edu/~srf28/21cm/index.htm; minor changes to match published versio

The micro-structure of the intergalactic medium I: the 21cm signature from dynamical minihaloes

A unified description is provided for the 21cm signatures arising from minihaloes against a bright background radio source and against the Cosmic Microwave Background (CMB), within the context of a dynamical collapsing cosmological spherical halo model. The equivalent width distribution of the resulting 21cm forest is computed for LCDM cosmologies, along with the brightness temperature differential relative to the CMB. The effects of an ambient Ly-alpha radiation field and heating of the IGM on the signatures are included. It is shown that the dynamical effects of heating substantially suppress absorption features in the 21cm forest with observed equivalent widths exceeding 0.15 kHz. It is demonstrated how measurements of excess fluctuations beyond detector noise could make a statistical detection of the weaker absorption features against a bright background radio source. It is also shown that a Ly-alpha radiation field only a few percent the thermalisation rate is sufficient to render the minihalo signal against the CMB negligible compared with the signal from the diffuse IGM component. The 21cm signals are found to be very sensitive to the amount of small scale power in the primordial density fluctuation spectrum. The effects of gas cooling via radiative atomic and molecular processes and of star formation on setting the maximum mass of the minihaloes giving rise to a 21cm signal are included, with allowance made for the suppression of molecular hydrogen formation by an ambient UV radiation field.Comment: Substantial revision: paper restructured; hydrodynamical model with heating included; discussion of statistical detection of absorption fluctuations from 21cm forest added. 34 pages; 34 figures. Definitive version published in Monthly Notice

Properties of the dense core population in Orion B as seen by the Herschel Gould Belt survey

We present a detailed study of the Orion B molecular cloud complex (d ~ 400 pc), which was imaged with the PACS and SPIRE photometric cameras at wavelengths from 70 to 500 μm as part of the Herschel Gould Belt survey (HGBS). We release new high-resolution maps of column density and dust temperature for the whole complex, derived in the same consistent manner as for other HGBS regions. In the filamentary subregions NGC 2023 and 2024, NGC 2068 and 2071, and L1622, a total of 1768 starless dense cores were identified based on Herschel data, 490–804 (~28−45%) of which are self-gravitating prestellar cores that will likely form stars in the future. A total of 76 protostellar dense cores were also found. The typical lifetime of the prestellar cores was estimated to be tpreOrionB = 1.7−0.6+0.8Myr. The prestellar core mass function (CMF) derived for the whole sample of prestellar cores peaks at ~0.5 M⊙ (in dN/dlogM format) and is consistent with a power-law with logarithmic slope −1.27 ± 0.24 at the high-mass end, compared to the Salpeter slope of − 1.35. In the Orion B region, we confirm the existence of a transition in prestellar core formation efficiency (CFE) around a fiducial value AVbg ~ 7 mag in background visual extinction, which is similar to the trend observed with Herschel in other regions, such as the Aquila cloud. This is not a sharp threshold, however, but a smooth transition between a regime with very low prestellar CFE at AVbg 90% when a more complete sample of filamentary structures is considered. Interestingly, the median separation observed between nearest core neighbors corresponds to the typical inner filament width of ~0.1 pc, which is commonly observed in nearby molecular clouds, including Orion B. Analysis of the CMF observed as a function of background cloud column density shows that the most massive prestellar cores are spatially segregated in the highest column density areas, and suggests that both higher- and lower-mass prestellar cores may form in denser filaments