A field study was conducted to determine the effects of ambient conditions and burning practices of rice fields in Taiwan on the chemical and physical characteristics of the smoke aerosol. Rice straw was burned on an actual rice field under typical conditions and smoke particles were collected immediately downwind of the field over the full particle size spectrum. Here we present size distributions of levoglucosan, a common molecular tracer for biomass burning, as well as detailed concentration patterns of three anhydrosugars, including mannosan, and galactosan, in addition to smoke aerosol concentrations of inorganic ions and carbonaceous species. The generated smoke aerosol was characterized by a high OC/EC ratio (10) and a large fraction of potassium (K + ) and chloride (Cl -) ions at a Cl -/K + ratio of 2. Levoglucosan showed a distinct bimodal distribution in the smoke particles with a large fraction (up to 56%) of the total levoglucosan mass observed in very large particles (PM >10 ). The prevailing ambient conditions (such as relatively high humidity), atmospheric processes (e.g., particle coagulation, hygroscopic growth, and deposition), the specific burning practices of rice fields in Taiwan (slow burning of straw spread in thin layers on the ground), as well as the inherent properties of rice straw likely influenced the particle size characteristics of the smoke tracer. Moreover, the relative abundance of the three biomass burning tracers showed a unique pattern (in good agreement with previous chamber burn measurements): levoglucosanto-mannosan ratios were distinctly higher (with an average value of 27) than those observed for other types of biomass, such as softwood, hard wood, peat, or leaves, in previous studies. Such chemical fingerprint may be used in source apportionment studies for the assessment of contributions from the combustion of specific types of biomass.
Telomere maintenance is required for chromosome stability, and telomeres are typically replicated by the action of the reverse transcriptase telomerase. In both tumor and yeast cells that lack telomerase, telomeres are maintained by an alternative recombination mechanism. Genetic studies have led to the identification of DNA polymerases, cell cycle checkpoint proteins, and telomere binding proteins involved in the telomerase pathway. However, how these proteins affect telomere-telomere recombination has not been identified to date. Using an assay to trace the in vivo recombinational products throughout the course of survivor development, we show here that three major replicative polymerases, ␣, ␦, and , play roles in telomere-telomere recombination and that each causes different effects and phenotypes when they as well as the telomerase are defective. Polymerase ␦ appears to be the main activity for telomere extension, since neither type I nor type II survivors arising via telomere-telomere recombination were seen in its absence. The frequency of type I versus type II is altered in the polymerase ␣ and mutants relative to the wild type. Each prefers to develop a particular type of survivor. Moreover, type II recombination is mediated by the cell cycle checkpoint proteins Tel1 and Mec1, and telomere-telomere recombination is regulated by telomere binding protein Cdc13 and the Ku complex. Together, our results suggest that coordination between DNA replication machinery, DNA damage signaling, DNA recombination machinery, and the telomere protein-DNA complex allows telomere recombination to repair telomeric ends in the absence of telomerase.
Telomere maintenance is required for chromosome stability, and telomeres are typically replicated by the action of telomerase. In both mammalian tumor and yeast cells that lack telomerase, telomeres are maintained by an alternative (ALT) recombination mechanism. In yeast, Sgs1p and its associated type IA topoisomerase, Top3p, may work coordinately in removing Holliday junction intermediates from a crossover-producing recombination pathway. Previous studies have also indicated that Sgs1 helicase acts in a telomere recombination pathway. Here we show that topoisomerase III is involved in telomere-telomere recombination. The recovery of telomere recombination-dependent survivors in a telomerase-minus yeast strain was dependent on Top3p catalytic activity. Moreover, the RIF1 and RIF2 genes are required for the establishment of TOP3/SGS1-dependent telomeretelomere recombination. In human Saos-2 ALT cells, human topoisomerase III␣ (hTOP3␣) also contributes to telomere recombination. Strikingly, the telomerase activity is clearly enhanced in surviving si-hTOP3␣ Saos-2 ALT cells. Altogether, the present results suggest a potential role for hTOP3␣ in dissociating telomeric structures in telomerase-deficient cells, providing therapeutic implications in human tumors.
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