Star Formation in Atomic Gas

Observations of nearby galaxies have firmly established, over a broad range of galactic environments and metallicities, that star formation occurs exclusively in the molecular phase of the interstellar medium (ISM). Theoretical models show that this association results from the correlation between chemical phase, shielding, and temperature. Interstellar gas converts from atomic to molecular only in regions that are well shielded from interstellar ultraviolet (UV) photons, and since UV photons are also the dominant source of interstellar heating, only in these shielded regions does the gas become cold enough to be subject to Jeans instability. However, while the equilibrium temperature and chemical state of interstellar gas are well-correlated, the time scale required to reach chemical equilibrium is much longer than that required to reach thermal equilibrium, and both timescales are metallicity-dependent. Here I show that the difference in time scales implies that, at metallicities below a few percent of the Solar value, well-shielded gas will reach low temperatures and proceed to star formation before the bulk of it is able to convert from atomic to molecular. As a result, at extremely low metallicities, star formation will occur in a cold atomic phase of the ISM rather than a molecular phase. I calculate the observable consequences of this result for star formation in low metallicity galaxies, and I discuss how some current numerical models for H2-regulated star-formation may need to be modified.Comment: 9 pages, 8 figures, ApJ in press; very minor changes from previous versio

In-Situ Survival Mechanisms of U and Tc Reducing Bacteria in Contaminated Sediments

Desulfovibrio desulfuricans G20 and Shewanella oneidensis MR-1 are model subsurface organisms for studying genes involving in situ radionuclide transformation and sediment survival. Our research objective for this project has been to develop a signature-tagged mutagenesis (STM) procedure and use it to identify mutants in genes of these subsurface bacteria involved in sediment survival and radionuclide reduction. The mutant genes identified in these studies allow us for the first time to describe at the genetic level microbial processes that are actually being used by environmental bacteria while growing in their natural ecosystems. Identification of these genes revealed facets of microbial physiology and ecology that are not accessible through laboratory studies. Ultimately, this information may be used to optimize bioremediation or other engineered microbial processes. Furthermore, the identification of a mutant in a gene conferring multidrug resistance in strain MR-1 shows that this widespread mechanism of antibiotic resistance, likely has its origins as a mechanism of bacterial defense against naturally occurring toxins. Studies with D. desulfuricans G20: The STM procedure first involved generating a library of 5760 G20 mutants and screening for potential non-survivors in subsurface sediment microcosms. After two rounds of screening, a total of 117 mutants were confirmed to be true non-survivors. 97 transposon insertion regions have been sequenced to date. Upon further analysis of these mutants, we classified the sediment survival genes into COG functional categories. STM mutant insertions were located in genes encoding proteins related to metabolism (33%), cellular processes (42%), and information storage and processing (17%). We also noted 8% of STM mutants identified had insertions in genes for hypothetical proteins or unknown functions. Interestingly, at least 64 of these genes encode cytoplasmic proteins, 46 encode inner membrane proteins, and only 7 encode periplasmic space and outer membrane associated proteins. Through blast search analysis, we also showed that 81 out of 94 proteins shown to be important in sediment survival have homologs in D. vulgaris, 70 have homologs in Geobacter metallireducens, and 69 have homologs in Geobacter sulfurreducens PCA. Some interesting proteins include ribonucleotide reductase and chemotaxis related proteins. Ribonucleotide reductase catalyzes the reductive synthesis of deoxyribonucleotides from their corresponding ribonucleotides, providing the precursors necessary for DNA synthesis. Two ribonucleotide reductase genes (nrdE, nrdD) were found to be essential for G20 survival in the sediment, but not essential for growth in the lactate-sulfate medium. Bacterial methyl-accepting chemotaxis proteins (MCP) respond to changes in the concentration of attractants and repellents in the environment

Testing models for molecular gas formation in galaxies: hydrostatic pressure or gas and dust shielding?

Stars in galaxies form in giant molecular clouds that coalesce when the atomic hydrogen is converted into molecules. There are currently two dominant models for what property of the galactic disk determines its molecular fraction: either hydrostatic pressure driven by the gravity of gas and stars, or a combination of gas column density and metallicity. To assess the validity of these models, we compare theoretical predictions to the observed atomic gas content of low-metallicity dwarf galaxies with high stellar densities. The extreme conditions found in these systems are optimal to distinguish the two models, otherwise degenerate in nearby spirals. Locally, on scales <100 pc, we find that the state of the interstellar medium is mostly sensitive to the gas column density and metallicity rather than hydrostatic pressure. On larger scales where the average stellar density is considerably lower, both pressure and shielding models reproduce the observations, even at low metallicity. We conclude that models based on gas and dust shielding more closely describe the process of molecular formation, especially at the high resolution that can be achieved in modern galaxy simulations or with future radio/millimeter arrays.Comment: 20 pages, 12 figures. Accepted for publication in Ap

What Phase of the Interstellar Medium Correlates with the Star Formation Rate?

Nearby spiral galaxies show an extremely tight correlation between tracers of molecular hydrogen (H_2) in the interstellar medium (ISM) and tracers of recent star formation, but it is unclear whether this correlation is fundamental or accidental. In the galaxies that have been surveyed to date, H_2 resides predominantly in gravitationally bound clouds cooled by carbon monoxide (CO) molecules, but in galaxies of low metal content the correlations between bound clouds, CO, and H_2 break down, and it is unclear if the star formation rate will then correlate with H_2 or with some other quantity. Here we show that star formation will continue to follow H_2 independent of metallicity. This is not because H_2 is directly important for cooling, but instead because the transition from predominantly atomic hydrogen (HI) to H_2 occurs under the same conditions as a dramatic drop in gas temperature and Bonnor-Ebert mass that destabilizes clouds and initiates collapse. We use this model to compute how star formation rate will correlate with total gas mass, with mass of gas where the hydrogen is H_2, and with mass of gas where the carbon is CO in galaxies of varying metallicity, and show that preliminary observations match the trend we predict.Comment: 13 pages, 7 figures, emulateapj format, accepted to ApJ; minor revisions to discussion, and a minor error in figures 4 and 5 fixed. No other change

Metallicity and the Universality of the IMF

The stellar initial mass function (IMF), along with the star formation rate, is one of the fundamental properties that any theory of star formation must explain. An interesting feature of the IMF is that it appears to be remarkably universal across a wide range of environments. Particularly, there appears to be little variation in either the characteristic mass of the IMF or its high-mass tail between clusters with different metallicities. Previous attempts to understand this apparent independence of metallicity have not accounted for radiation feedback from high-mass protostars, which can dominate the energy balance of the gas in star-forming regions. We extend this work, showing that the fragmentation of molecular gas should depend only weakly on the amount of dust present, even when the primary heating source is radiation from massive protostars. First, we report a series of core collapse simulations using the ORION AMR code that systematically vary the dust opacity and show explicitly that this has little effect on the temperature or fragmentation of the gas. Then, we provide an analytic argument for why the IMF varies so little in observed star clusters, even as the metallicity varies by a factor of 100.Comment: 11 pages, 6 figures, emulateapj format, accepted to ApJ. Typos removed, references added, and discussion revised in section 3.2. Conclusions unchange

Metallicity-Dependent quenching of Star Formation at High Redshift in Small Galaxies

[abridged] The star formation rates (SFR) of low-metallicity galaxies depend sensitively on the gas metallicity, because metals are crucial to mediating the transition from intermediate-temperature atomic gas to cold molecular gas, a necessary precursor to star formation. We study the impact of this effect on the star formation history of galaxies. We incorporate metallicity-dependent star formation and metal enrichment in a simple model that follows the evolution of a halo main progenitor. Our model shows that including the effect of metallicity leads to suppression of star formation at redshift z>2 in dark halos with masses <~ 10^11 Msun, with the suppression becoming near total for halos below ~10^9.5-10 Msun. We find that at high redshift the SFR cannot catch up with the gas inflow rate (IR), because the SFR is limited by the free-fall time, and because it is suppressed further by a lack of metals. As a result, in each galaxy the SFR is growing in time faster than the IR, and the integrated cosmic SFR density is rising with time. The suppressed in situ SFR at high z makes the growth of stellar mass dominated by ex situ SFR which implies that the specific SFR (sSFR) remains constant with time. The intensely accreted gas at high z is accumulating as an atomic gas reservoir. This provides additional fuel for star formation in 10^10 - 10^12 Msun halos at z ~ 1-3, which allows the SFR to exceed the instantaneous IR, and may enable an even higher outflow rate. At z<1, following the natural decline in IR with time due to the universal expansion, the SFR and sSFR are expected to drop. We specify the expected dependence of sSFR and metallicity on stellar mass and redshift. At a given z, and below a critical mass, these relations are predicted to be flat and rising respectively. Our model predictions qualitatively match some of the puzzling features in the observed star formation history.Comment: 25 pages, 18 figures, emulateapj format, accepted to ApJ. Resolution of some figures degraded to fit within arXiv size limits. A full resolution version is available at http://www.ucolick.org/~krumholz/publications.htm

Interconnection of Key Microbial Functional Genes for Enhanced Benzo[a]pyrene Biodegradation in Sediments by Microbial Electrochemistry.

Sediment microbial fuel cells (SMFCs) can stimulate the degradation of polycyclic aromatic hydrocarbons in sediments, but the mechanism of this process is poorly understood at the microbial functional gene level. Here, the use of SMFC resulted in 92% benzo[a]pyrene (BaP) removal over 970 days relative to 54% in the controls. Sediment functions, microbial community structure, and network interactions were dramatically altered by the SMFC employment. Functional gene analysis showed that c-type cytochrome genes for electron transfer, aromatic degradation genes, and extracellular ligninolytic enzymes involved in lignin degradation were significantly enriched in bulk sediments during SMFC operation. Correspondingly, chemical analysis of the system showed that these genetic changes resulted in increases in the levels of easily oxidizable organic carbon and humic acids which may have resulted in increased BaP bioavailability and increased degradation rates. Tracking microbial functional genes and corresponding organic matter responses should aid mechanistic understanding of BaP enhanced biodegradation by microbial electrochemistry and development of sustainable bioremediation strategies

Exposure of Soil Microbial Communities to Chromium and Arsenic Alters Their Diversity and Structure

Extensive use of chromium (Cr) and arsenic (As) based preservatives from the leather tanning industry in Pakistan has had a deleterious effect on the soils surrounding production facilities. Bacteria have been shown to be an active component in the geochemical cycling of both Cr and As, but it is unknown how these compounds affect microbial community composition or the prevalence and form of metal resistance. Therefore, we sought to understand the effects that long-term exposure to As and Cr had on the diversity and structure of soil microbial communities. Soils from three spatially isolated tanning facilities in the Punjab province of Pakistan were analyzed. The structure, diversity and abundance of microbial 16S rRNA genes were highly influenced by the concentration and presence of hexavalent chromium (Cr (VI)) and arsenic. When compared to control soils, contaminated soils were dominated by Proteobacteria while Actinobacteria and Acidobacteria (which are generally abundant in pristine soils) were minor components of the bacterial community. Shifts in community composition were significant and revealed that Cr (VI)-containing soils were more similar to each other than to As contaminated soils lacking Cr (VI). Diversity of the arsenic resistance genes, arsB and ACR3 were also determined. Results showed that ACR3 becomes less diverse as arsenic concentrations increase with a single OTU dominating at the highest concentration. Chronic exposure to either Cr or As not only alters the composition of the soil bacterial community in general, but affects the arsenic resistant individuals in different ways