Molecular properties of (U)LIRGs: CO, HCN, HNC and HCO+

The observed molecular properties of a sample of FIR-luminous and OH megamaser (OH-MM) galaxies have been investigated. The ratio of high and low-density tracer lines is found to be determined by the progression of the star formation in the system. The HCO+/HCN and HCO+/HNC line ratios are good proxies for the density of the gas, and PDR and XDR sources can be distinguished using the HNC/HCN line ratio. The properties of the OH-MM sources in the sample can be explained by PDR chemistry in gas with densities higher than 10^5.5 cm^-3, confirming the classical OH-MM model of IR pumped amplification with (variable) low gains.Comment: 5 pages, 2 figures, to appear in: IAU Symposium 242 Astrophysical Masers and their Environment

Evolution of the ISM in Luminous IR Galaxies

Molecules that trace the high-density regions of the interstellar medium may be used to evaluate the changing physical and chemical environment during the ongoing nuclear activity in (Ultra-)Luminous Infrared Galaxies. The changing ratios of the HCN(1-0), HNC(1-0), HCO+(1-0), CN(1-0) and CN(2-1), and CS(3-2) transitions were compared with the HCN(1-0)/CO(1-0) ratio, which is proposed to represent the progression time scale of the starburst. These diagnostic diagrams were interpreted using the results of theoretical modeling using a large physical and chemical network to describe the state of the nuclear ISM in the evolving galaxies. Systematic changes are seen in the line ratios as the sources evolve from early stage for the nuclear starburst (ULIRGs) to later stages. These changes result from changing environmental conditions and particularly from the lowering of the average density of the medium. A temperature rise due to mechanical heating of the medium by feedback explains the lowering of the ratios at later evolutionary stages. Infrared pumping may affect the CN and HNC line ratios during early evolutionary stages. Molecular transitions display a behavior that relates to changes of the environment during an evolving nuclear starburst. Molecular properties may be used to designate the evolutionary stage of the nuclear starburst. The HCN(1-0)/CO(1-0) and HCO+(1-0)/HCN(1-0) ratios serve as indicators of the time evolution of the outburst.Comment: To be published in Astronomy and Astrophysics - 11 pages, 9 figures, 1 tabl

The irradiated ISM of ULIRGs

The nuclei of ULIRGs harbor massive young stars, an accreting central black hole, or both. Results are presented for molecular gas that is exposed to X-rays (1-100 keV, XDRs) and far-ultraviolet radiation (6-13.6 eV, PDRs). Attention is paid to species like HCO+, HCN, HNC, OH, H2O and CO. Line ratios of HCN/HCO+ and HNC/HCN discriminate between PDRs and XDRs. Very high J (>10) CO lines, observable with HIFI/Herschel, discriminate very well between XDRs and PDRs. In XDRs, it is easy to produce large abundances of warm (T>100 K) H2O and OH. In PDRs, only OH is produced similarly well.Comment: 5 pages, 6 figures, to appear in: IAU Symposium 242 Astrophysical Masers and their Environment

Diagnostics of active galaxies: I. Modeling the infrared properties of dusty cores of starburst galaxies

An evolutionary model of star formation in ULIRGs was created using existing star formation and radiative transfer codes (STARBURST99, RADMC and RADICAL) as building blocks. The results of the simulations are compared to data from two IRAS catalogs. From the simulations it is found that the dust surrounding the starburst region is made up from two components. There is a low optical depth (tau=0.1, which corresponds to 0.1% of the total dust mass), hot (T~400K) non-grey component close to the starburst (scale size 10pc) and a large scale, colder grey component (100pc, 75K) with a much larger column (tau=10). The simulations also show that starburst galaxies can be powered by massive star formation. The parameters for this star forming region are difficult to determine, since the IR continuum luminosity is only sensitive to the total UV input. Therefore, there is a degeneracy between the total starburst mass and the initial mass function (IMF) slope. A less massive star formation with a shallower IMF will produce the same amount of OB stars and therefore the same amount of irradiating UV flux. Assuming the stars are formed according to a Salpeter IMF (Psi(M) ~ M^-2.35), the star formation region should produce 10^9 Msun of stars (either in one instantaneous burst, or in a continuous process) in order to produce enough IR radiation.Comment: 37 pages, 32 figures, accepted by Astronomy and Astrophysics. Version with Appendices can be obtained from http://www.astro.rug.nl/~loenen/Loenen_et_al_2006.pd

Mechanical feedback in the molecular ISM of luminous IR galaxies

Aims: Molecular emission lines originating in the nuclei of luminous infra-red galaxies are used to determine the physical properties of the nuclear ISM in these systems. Methods: A large observational database of molecular emission lines is compared with model predictions that include heating by UV and X-ray radiation, mechanical heating, and the effects of cosmic rays. Results: The observed line ratios and model predictions imply a separation of the observedsystems into three groups: XDRs, UV-dominated high-density (n>=10^5 cm-3) PDRs, and lower-density (n=10^4.5 cm-3) PDRs that are dominated by mechanical feedback. Conclusions: The division of the two types of PDRs follows naturally from the evolution of the star formation cycle of these sources, which evolves from deeply embedded young stars, resulting in high-density (n>=10^5 cm-3) PDRs, to a stage where the gas density has decreased (n=10^4.5 cm-3) and mechanical feedback from supernova shocks dominates the heating budget.Comment: 4 pages, 3 figures, published as Letter to the Editor in A&A (see http://www.aanda.org/articles/aa/abs/2008/34/aa10327-08/aa10327-08.html

Molecules as tracers of galaxy evolution: an EMIR survey. I. Presentation of the data and first results

We investigate the molecular gas properties of a sample of 23 galaxies in order to find and test chemical signatures of galaxy evolution and to compare them to IR evolutionary tracers. Observation at 3 mm wavelengths were obtained with the EMIR broadband receiver, mounted on the IRAM 30 m telescope on Pico Veleta, Spain. We compare the emission of the main molecular species with existing models of chemical evolution by means of line intensity ratios diagrams and principal component analysis. We detect molecular emission in 19 galaxies in two 8 GHz-wide bands centred at 88 and 112 GHz. The main detected transitions are the J=1-0 lines of CO, 13CO, HCN, HNC, HCO+, CN, and C2H. We also detect HC3N J=10-9 in the galaxies IRAS 17208, IC 860, NGC 4418, NGC 7771, and NGC 1068. The only HC3N detections are in objects with HCO+/HCN<1 and warm IRAS colours. Galaxies with the highest HC3N/HCN ratios have warm IRAS colours (60/100 {\mu}m>0.8). The brightest HC3N emission is found in IC 860, where we also detect the molecule in its vibrationally excited state.We find low HNC/HCN line ratios (<0.5), that cannot be explained by existing PDR or XDR chemical models. Bright HC3N emission in HCO+-faint objects may imply that these are not dominated by X-ray chemistry. Thus the HCN/HCO+ line ratio is not, by itself, a reliable tracer of XDRs. Bright HC3N and faint HCO+ could be signatures of embedded starformation, instead of AGN activity

Insights into the regulation of DMSP synthesis in the diatom Thalassiosira pseudonana through APR activity, proteomics and gene expression analyses on cells acclimating to changes in salinity, light and nitrogen

Despite the importance of dimethylsulphoniopropionate (DMSP) in the global sulphur cycle and climate regulation, the biological pathways underpinning its synthesis in marine phytoplankton remain poorly understood. The intracellular concentration of DMSP increases with increased salinity, increased light intensity and nitrogen starvation in the diatom Thalassiosira pseudonana. We used these conditions to investigate DMSP synthesis at the cellular level via analysis of enzyme activity, gene expression and proteome comparison. The activity of the key sulphur assimilatory enzyme, adenosine 5′- phosphosulphate reductase was not coordinated with increasing intracellular DMSP concentration. Under all three treatments coordination in the expression of sulphur assimilation genes was limited to increases in sulphite reductase transcripts. Similarly, proteomic 2D gel analysis only revealed an increase in phosphoenolpyruvate carboxylase following increases in DMSP concentration. Our findings suggest that increased sulphur assimilation might not be required for increased DMSP synthesis, instead the availability of carbon and nitrogen substrates may be important in the regulation of this pathway. This contrasts with the regulation of sulphur metabolism in higher plants, which generally involves upregulation of several sulphur assimilatory enzymes. In T. pseudonana changes relating to sulphur metabolism were specific to the individual treatments and, given that little coordination was seen in transcript and protein responses across the three growth conditions, different patterns of regulation might be responsible for the increase in DMSP concentration seen under each treatment

Gravitational lensing reveals extreme dust-obscured star formation in quasar host galaxies

We have observed 104 gravitationally-lensed quasars at $z\sim1-4$ with Herschel/SPIRE, the largest such sample ever studied. By targeting gravitational lenses, we probe intrinsic far-infrared (FIR) luminosities and star formation rates (SFRs) more typical of the population than the extremely luminous sources that are otherwise accessible. We detect 72 objects with Herschel/SPIRE and find 66 percent (69 sources) of the sample have spectral energy distributions (SEDs) characteristic of dust emission. For 53 objects with sufficiently constrained SEDs, we find a median effective dust temperature of $38^{+12}_{-5}$ K. By applying the radio-infrared correlation, we find no evidence for an FIR excess which is consistent with star-formation-heated dust. We derive a median magnification-corrected FIR luminosity of $3.6^{+4.8}_{-2.4}~\times 10^{11}~{\rm L_{\odot}}$ and median SFR of $120^{+160}_{-80}~{\rm M_{\odot}~yr^{-1}}$ for 94 quasars with redshifts. We find $\sim10$ percent of our sample have FIR properties similar to typical dusty star-forming galaxies at $z\sim2-3$ and a range of SFRs $<20-10000~{\rm M_{\odot}~yr^{-1}}$ for our sample as a whole. These results are in line with current models of quasar evolution and suggests a coexistence of dust-obscured star formation and AGN activity is typical of most quasars. We do not find a statistically-significant difference in the FIR luminosities of quasars in our sample with a radio excess relative to the radio-infrared correlation. Synchrotron emission is found to dominate at FIR wavelengths for $<15$ percent of those sources classified as powerful radio galaxies.Comment: 47 pages, 89 figures, accepted for publication in MNRA

HSV Neutralization by the Microbicidal Candidate C5A

Genital herpes is a major risk factor in acquiring human immunodeficiency virus type-1 (HIV-1) infection and is caused by both Herpes Simplex virus type 1 (HSV-1) and HSV-2. The amphipathic peptide C5A, derived from the non-structural hepatitis C virus (HCV) protein 5A, was shown to prevent HIV-1 infection but neither influenza nor vesicular stomatitis virus infections. Here we investigated the antiviral function of C5A on HSV infections. C5A efficiently inhibited both HSV-1 and HSV-2 infection in epithelial cells in vitro as well as in an ex vivo epidermal infection model. C5A destabilized the integrity of the viral HSV membrane. Furthermore, drug resistant HSV strains were inhibited by this peptide. Notably, C5A-mediated neutralization of HSV-1 prevented HIV-1 transmission. An in vitro HIV-1 transmigration assay was developed using primary genital epithelial cells and HSV infection increased HIV-1 transmigration. Treatment with C5A abolished HIV-1 transmigration by preventing HSV infection and by preserving the integrity of the genital epithelium that was severely compromised by HSV infection. In conclusion, this study demonstrates that C5A represents a multipurpose microbicide candidate, which neutralizes both HIV-1 and HSV, and which may interfere with HIV-1 transmission through the genital epithelium

Signal Peptide-Dependent Inhibition of MHC Class I Heavy Chain Translation by Rhesus Cytomegalovirus

The US2-11 region of human and rhesus cytomegalovirus encodes a conserved family of glycoproteins that inhibit MHC-I assembly with viral peptides, thus preventing cytotoxic T cell recognition. Since HCMV lacking US2-11 is no longer able to block assembly and transport of MHC-I, we examined whether this is also observed for RhCMV lacking the corresponding region. Unexpectedly, recombinant RhCMV lacking US2-11 was still able to inhibit MHC-I expression in infected fibroblasts, suggesting the presence of an additional MHC-I evasion mechanism. Progressive deletion analysis of RhCMV-specific genomic regions revealed that MHC-I expression is fully restored upon additional deletion of rh178. The protein encoded by this RhCMV-specific open reading frame is anchored in the endoplasmic reticulum membrane. In the presence of rh178, RhCMV prevented MHC-I heavy chain (HC) expression, but did not inhibit mRNA transcription or association of HC mRNA with translating ribosomes. Proteasome inhibitors stabilized a HC degradation intermediate in the absence of rh178, but not in its presence, suggesting that rh178 prevents completion of HC translation. This interference was signal sequence-dependent since replacing the signal peptide with that of CD4 or murine HC rendered human HCs resistant to rh178. We have identified an inhibitor of antigen presentation encoded by rhesus cytomegalovirus unique in both its lack of homology to any other known protein and in its mechanism of action. By preventing signal sequence-dependent HC translocation, rh178 acts prior to US2, US3 and US11 which attack MHC-I proteins after protein synthesis is completed. Rh178 is the first viral protein known to interfere at this step of the MHC-I pathway, thus taking advantage of the conserved nature of HC leader peptides, and represents a new mechanism of translational interference