BGD Biogeosciences Discussions BGD 1810-6285 Copernicus GmbH Göttingen, Germany 10.5194/bgd-5-1469-2008 Microbiology and atmospheric processes: biological, physical and chemical characterization of aerosol particles Georgakopoulos D. G. 1 Després V. 2 3 Fröhlich-Nowoisky J. 2 3 Psenner R. 4 Ariya P. A. 5 Pósfai M. 6 Ahern H. E. 7 Moffett B. F. 7 Hill T. C. J. 7 Agricultural University of Athens, Athens, Greece Max Planck Institute for Chemistry, Mainz, Germany Johannes Gutenberg University, Mainz, Germany University of Innsbruck, Innsbruck, Austria McGill University, W. Montreal, Canada University of Veszprem, Veszprem, Hungary University of East London, London, UK 08 04 2008 5 2 1469 1510 This is an open-access article ditributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. This article is available from http://www.biogeosciences-discuss.net/5/1469/2008/bgd-5-1469-2008.html The full text article is available as a PDF file from http://www.biogeosciences-discuss.net/5/1469/2008/bgd-5-1469-2008.pdf

The interest in bioaerosols has traditionally been linked to health hazards for humans, animals and plants. However, several components of bioaerosols exhibit physical properties of great significance for cloud processes, such as ice nucleation and cloud condensation. To gain a better understanding of their influence on climate, it is therefore important to determine the composition, concentration, seasonal fluctuation, regional diversity and evolution of bioaerosols. In this paper, we will review briefly the existing techniques for detection, quantification, physical and chemical analysis of biological particles, attempting to bridge physical, chemical and biological methods for analysis of biological particles and integrate them with aerosol sampling techniques. We will also explore some emerging spectroscopy techniques for bulk and single-particle analysis that have potential for in-situ physical and chemical analysis. Lastly, we will outline open questions and further desired capabilities (e.g., in-situ, sensitive, both broad and selective, on-line, time-resolved, rapid, versatile, cost-effective techniques) required prior to comprehensive understanding of chemical and physical characterization of bioaerosols.

 References Ahern, H. E.: Cloud Borne Bacteria: Community Composition and Potential Impact on Atmospheric Nucleation, PhD Thesis, University of East London, 2007. Ahern, H. E., Walsh, K. A., Hill, T. C. J., and Moffett, B. F.: Fluorescent pseudomonads isolated from Hebridean cloud and rain water produce biosurfactants but do not cause ice nucleation, Biogeosciences, 4, 115–124, 2007. Alfreider, A., Pernthaler, J., Amann, R., Sattler, B., Glockner, F. O., Wille, A., and Psenner, R.: Community analysis of the bacteria assemblages in winter cover and pelagic layers of a high mountain lake using in situ hybridization, Appl. Environ. Microb., 62, 2138–2144, 1996. Amann, R. and Ludwig, W.: Ribosomal RNA-targeted nucleic acid probes for studies in microbial ecology, FEMS Microbiol. Rev., 24, 555–565, 2000. Amato, P., Menager, M., Sancelme, M., Laj, P., Mailhot, G. D., and Delort, A.-M.: Microbial population in cloud water at the Puy de Dôme: implications for the chemistry of clouds, Atmos. Environ., 39, 4143–4153, 2005. Amato, P., Parazols, M., Sancelme, M., Laj, P., Mailhot, G., and Delort, A.-M.: Microorganisms isolated from the water phase of tropospheric clouds at the Puy de Dôme: major groups and growth abilities at low temperatures, FEMS Microbiol. Ecol., 59, 242–254, 2007. Amato, P., Parazols, M., Sancelme, M., Mailhot, G., Laj, P., and Delort, A.-M.: An important source of micro-organisms for cloud water at the Puy de Dôme (France), Atmos. Environ., 41, 8253–8263, 2007. Anderson, J. R., Aggett, F. J., Buseck, P. R., Germani, M. S., and Shattuck, T. W.: Chemistry of individual aerosol particles from Chandler, Arizona, an arid urban environment, Environ. Sci. Technol., 22(7), 811–818, 1988. Andreeva, I. S., Belan, B. D., Borodulin, A. I., Buryak, G. A., Zhukov, V. A., Panchenko, M. V., Penenko, V. V., Petrishchenko, V. A., and Safatov, A. S.: Variability of the content of live microorganisms in the atmospheric aerosol in southern regions of Western Siberia, Doklady Biol. Sci., 381, 530–534, 2001. Ariya, P. A., Nepotchatykh, O., Igntova, O., and Amyot, M.: Microbiological degradation of organic compounds in the atmosphere, Geophys. Res. Lett., 29, 341–344, 2002. Barkay, Z., Teller, A., Ganor, E., Levin, Z., and Shapira, Y.: Atomic force and scanning electron microscopy of atmospheric particles, Microsc. Res. Techniq., 68(2), 107–114, 2005. Bauer, H., Kasper-Giebl, A., Loflund, H., Giebl, H., Hitzenberger, R., Zibuschka, F., and Puxbaum, H.: The contribution of bacteria and fungal spores to the organic carbon content of cloud water, precipitation and aerosols, Atmos. Res., 64, 100–119, 2002. Beddows, D. C. S. and Telle, H. H.: Prospects of real-time single-particle biological aerosol analysis: A comparison between laser-induced breakdown spectroscopy and aerosol time-of-flight mass spectrometry, Spectrochim. Acta B, 60(7–8), 1040–1059, 2005. Bent, S. J., Pierson, J. D., and Forney, L. J.: Measuring species richness based on microbial community fingerprints: the emperor has no clothes, Appl. Environ. Microb., 73, 2399–401, 2007. Birenszvige, A., Eversole, J., Seaver, M., Francesconi, S., Valdes, E., and Kulaga, H.: Aerosol characteristics in a subway environment, Aerosol Sci. Tech., 37, 210–220, 2003. Blando, J. D., Porcja, R. J., Li, T. H., Bowman, D., Lioy, P. J., and Turpin, B. J.: Secondary formation and Smokey Mountain organic aerosol: an examination of aerosol polarity and functional group composition during SEAVS, Environ. Sci. Technol., 32, 604–613, 1998. Brodie, E. L., DeSantis, T. Z., Parker, J. P. M., Zubietta, I. X., Piceno, Y. M., and Andersen, G. L.: Urban aerosols harbor diverse and dynamic bacterial populations, Proc. Natl. Acad. Sci. USA, 104, 299–304, 2007. Burge, H. A.: Bioaerosol investigations, in: Bioaerosols, edited by: Burge, H. A., CRC Press, Boca Raton, FL, 1–23, 1995. Castrillo, L. A., Lee Jr., R. E., Lee, M. R., and Rutherford, S. T.: Identification of ice-nucleating active Pseudomonas fluorescens strains for biological control of overwintering Colorado potato beetles (Coleoptera: Chrysomelidae), J. Econ. Entomol., 93(2), 226–233, 2000. Chapman, M. D.: Analytical methods: immunoassays, in: Bioaerosols, edited by: Burge, H. A., CRC Press, Boca Raton, FL, 235–248, 1995. Chen, Y. Z., Shah, N., Huggins, F. E., and Huffman, G. P.: Characterization of ambient airborne particles by energy-filtered transmission electron microscopy, Aerosol Sci. Tech., 39(6), 509–518, 2005. Chi, M.-C. and Li, C.-S.: Fluorochrome and Fluorescent In Situ Hybridization to monitor bioaerosols in swine buildings, Aerosol Sci. Tech., 39, 1101–1110, 2005. Christner, B. C., Mosley-Thompson, E., Thompson, L. G., and Reeve, J. N.: Isolation of bacteria and 16S rDNAs from Lake Vostok accretion ice, Environ. Microb., 3, 570–577, 2001. Cote, V., Kos, G., Mortazavi, A., and Ariya, P. A.: Microbial and "de novo" transformation of dicarboxylic acids by three airborne fungi, Sci. Total Environ., 390(2–3), 530–537, 2008. Danovaro, R., Luna, G. M., Dell'Anno, A., and Pietrangeli, B.: Comparison of two fingerprinting techniques, Terminal Restriction Fragment Length Polymorphism and Automated Ribosomal Intergenic Spacer Analysis, for determination of bacterial diversity in aquatic environments, Appl. Environ. Microb., 72, 5982–5989, 2006. de Hoog, J., Osan, J., Szaloki, I., Eyckmans, K., Worobiec, A., Ro, C. U., and Van Grieken, R.: Thin-window electron probe X-ray microanalysis of individual atmospheric particles above the North Sea, Atmos. Environ., 39(18), 3231–3242, 2005. Decesari, S., Facchini, M. C., Fuzzi, S., and Tagliavini, E.: Characterization of Water-Soluble Organic Compounds in Atmospheric Aerosol: A New Approach, J. Geophys. Res., 105(D1), 1481–1489, 2000. Decesari, S., Facchini, M. C., Matta, E., Lettini, F., Mircea, M., Fuzzi, S., Tagliavini, E., and Putaud, J. P.: Chemical features and seasonal trend of water soluble organic compounds in the Po Valley fine aerosol, Atmos. Environ., 35, 3691–3699, 2001. Decesari, S., Mircea, M., Cavalli, F., Fuzzi, S., Moretti, F., Tagliavini, E., and Facchini, M. C.: Source Attribution of Water-Soluble Organic Aerosol by Nuclear Magnetic Resonance Spectroscopy, Environ. Sci. Technol., 41, 2479–2484, 2007. Després, V. R., Nowoisky, J. F., Klose, M., Conrad, R., Andreae, M. O., and Pöschl, U.: Characterization of primary biogenic aerosol particles in urban, rural, and high-alpine air by DNA sequence and restriction fragment analysis of ribosomal RNA genes, Biogeosciences, 4, 1127–1141, 2007. Dimos, D., Kallimanis, A., Perisynakis, A., Chatziloukas, E., and Drainas, C.: A contribution to the analysis of the genetic diversity of bacterial ice nucleation genes in Hellenic niches, HSBMB Newsletter, 53, 66–67, 2006. Ebert, M., Inerle-Hof, M., and Weinbruch, S.: Environmental scanning electron microscopy as a new technique to determine the hygroscopic behaviour of individual aerosol particles, Atmos. Environ., 36(39–40), 5909–5916, 2002. Ebert, M., Weinbruch, S., Hoffmann, P., and Ortner, H. M.: The chemical composition and complex refractive index of rural and urban influenced aerosols determined by individual particle analysis, Atmos. Environ., 38(38), 6531–6545, 2004. Fogel, G., Collins, C., Li, J., and Brunk, C.: Prokaryotic genome size and SSU rDNA copy number: estimation of relative abundance from a mixed population, Microbial Ecol., 38, 93–113, 1999. Friedbacher, G., Grasserbauer, M., Meslmani, Y., Klaus, N., and Higatsberger, M. J.: Investigation of environmental aerosol by atomic force microscopy, Anal. Chem., 67, 1749–1754, 1995. Gilardoni, S., Russell, L. M., Sorooshian, A., Flagan, R. C., Seinfeld, J. H., Bates, T. S., Quinn, P. K., Allan, J. D., Williams, B., Goldstein, A. H., Onasch, T. B., and Worsnop, D. R.: Regional variation of organic functional groups in aerosol particles on four US east coast platforms during the International Consortium for Atmospheric Research on Transport and Transformation 2004 campaign, J. Geophys. Res., 112, D10S27, doi:10.1029/2006JD007737, 2007. Grüntzig, V., Nold, S. C., Zhou, Z., and Tiedje, J. M.: Pseudomonas stutzeri nitrite reductase gene abundance in environmental samples measured by real-time PCR, Appl. Environ. Microb., 67, 760–768, 2001. Guilbaud, C., Ahern, H., Dominguez, H., Glaux C., Moffett, B., Hill, T., and Morris, C.: Detecting ice nucleating bacteria in environmental samples using PCR of the gene conferring ice nucleation activity. Paper presented at the IUGG's Earth: Our Changing Planet Symposium, Perugia, July, 2007. Jaenicke, R.: Abundance of cellular material and proteins in the atmosphere, Science, 308, p. 73, 2005. Jansson, J. and Leser, T.: Quantitative PCR of environmental samples, in: Molecular Microbiology Manual, 2nd Edition, Kluwer Academic Publishers, The Nethlands, 445–464, 2004. Kaegi, R. and Holzer, L.: Transfer of a single particle for combined ESEM and TEM analyses, Atmos. Environ., 37(31), 4353–4359, 2003. Katrinak, K. A., Rez, P., and Buseck, P. R.: Structural variations in individual carbonaceous particles from an urban aerosol, Environ. Sci. Technol., 26, 1967–1976, 1992. Kellogg, C. A. and Griffin, D. W.: Aerobiology and the global transport of desert dust, Trends Ecol. Evol., 21, 638–644, 2006. Kemp, B. M., Monroe, C., and Smith, D. G.: Repeat silica extraction: a simple technique for the removal of PCR inhibitors from DNA extracts, J. Archaeol. Sci., 33, 1680–1689, 2006. Kis, V. K., Pósfai, M., and Labar, J. L.: Nanostructure of atmospheric soot particles, Atmos. Environ., 40(29), 5533–5542, 2006. Knut, R., Moen, B., Drømtorp, S. M., and Holck, A. L.: Use of ethidium monoazide and PCR in combination for quantification of viable and dead cells in complex samples, Appl. Environ. Microb., 71, 1018–1024, 2004. Köllensperger, G., Friedbacher, G., Kotzick, R., Niessner, R., and Grasserbauer, M.: In-situ atomic force microscopy investigation of aerosols exposed to different humidities, Fresen. J. Anal. Chem., 364, 296–304, 1999. Krueger, B. J., Grassian, V. H., Iedema, M. J., Cowin, J. P., and Laskin, A.: Probing heterogeneous chemistry of individual atmospheric particles using scanning electron microscopy and energy-dispersive X-ray analysis, Anal. Chem., 75(19), 5170–5179, 2003. Laskin, A., Cowin, J. P., and Iedema, M. J.: Analysis of individual environmental particles using modern methods of electron microscopy and X-ray microanalysis, J. Electron. Spectrosc., 150(2–3), 260–274, 2006. Liu, X. D., Zhu, J., Van Espen, P., Adams, F., Xiao, R., Dong, S. P., and Li, Y. W.: Single particle characterization of spring and summer aerosols in Beijing: Formation of composite sulfate of calcium and potassium, Atmos. Environ., 39(36), 6909–6918, 2005. Lighthart, B.: The ecology of bacteria in the alfresco atmosphere, FEMS Microbiol. Ecol., 23, 263–274, 1997. Margulies, M., Egholm, M., Altman, W. E., Attiya, S., Bader, J. S., Bemben, L. A., Berka, J., Braverman, M. S., Chen, Y.-J., Chen, Z., Dewell, S. B., Du, L., Fierro, J. M., Gomes, X. V., Godwin, B. C., He, W., Helgesen, S., Ho, C. H., Irzyk, G. P., Jando, S. C., Alenquer, M. L. I., Jarvie, T. P., Jirage, K. B., Kim, J.-B., Knight, J. R., Lanza, J. R., Leamon, J. H., Lefkowitz, S. M., Lei, M., Li, J., Lohman, K. L., Lu, H., Makhijani, V. B., McDade, K. E., McKenna, M. P., Myers, E. W., Nickerson, E., Nobile, J. R., Plant, R., Puc, B. P., Ronan, M. T., Roth, G. T., Sarkis, G. J., Simons, J. F., Simpson, J. W., Srinivasan, M., Tartaro, K. R., Tomasz, A., Vogt, K. A., Volkmer, G. A., Wang, S. H., Wang, Y., Weiner, M. P., Yu, P., Begley, R. F., and Rothberg, J. M.: Genome sequencing in microfabricated high-density picolitre reactors, Nature, 437, 376–380, 2005. Maria, S. F., Russell, L. M., Turpin, B. J., Porcja, R. J., Campos, T. L., Weber, R. J., and Huebert, B. J.: Source Signatures of Carbon Monoxide and Organic Functional Groups in Asian Pacific Regional Aerosol Characterization Experiment (ACE-Asia) Submicron Aerosol Types, J. Geophys. Res., 108, 8637, doi:10.1029/2003JD003703, 2003. Maria, S. F., Russell, L. M., Turpin, B. J., and Porcja, R. J.: FTIR measurements of functional groups and organic mass in aerosol samples over the Caribbean, Atmos. Environ., 36, 5185–5196, 2002. Maron, P., Lejon, D., Carvalho, E., Bizet, K., Lemanceau, P., Ranjard, L., and Mougel, C.: Assessing genetic structure and diversity of airborne bacterial communities by DNA fingerprinting and 16S rDNA clone library, Atmos. Environ., 39, 3687–3695, 2005. Matthias-Maser, S. and Jaenicke, R.: Examination of atmospheric bioaerosol particles with radii >0.2 $\mu $m. J. Aerosol Sci., 25, 1605–1613, 1994. Maynard, A. D.: The application of electron-energy-loss spectroscopy to the analysis of ultrafine aerosol-particles, J. Aerosol Sci., 26(5), 757–777, 1995. Maynard, A. D., Ito, Y., Arslan, I., Zimmer, A. T., Browning, N., and Nicholls, A.: Examining elemental surface enrichment in ultrafine aerosol particles using analytical scanning transmission electron microscopy, Aerosol Sci. Tech., 38(4), 365–381, 2004. Moffett, B. F., Nicholson, F. A., Uwakwe, N. C., Chambers, B. J., Harris, J. A., and Hill, T. C. J.: Zinc contamination decreases the bacterial diversity of agricultural soil, FEMS Microbiol. Ecol., 43, 13–19, 2003. Moyer, C. L., Tiedje, J. M., Dobbs, F. C., and Karl, D. M.: A computer-simulated restriction fragment length polymorphism analysis of bacterial small-subunit rRNA genes: efficacy of selected tetrameric restriction enzymes for studies of microbial diversity in nature, Appl. Environ. Microb., 62, 2501–2507, 1996. Murphy, D. M., Cziczo, D. J., Froyd, K. D., Hudson, P. K., Matthew, B. M., Middlebrook, A. M., Peltier, R. E., Sullivan, A., Thomson, D. S., and Weber, R. J.: Single-particle mass spectrometry of tropospheric aerosol particles, J. Geophys. Res., 111(D23), D23S32, doi:10.1029/2006JD007340, 2006. Muyzer, G. and Smalla, K.: Application of denaturing gradient gel electrophoresis and temperature gradient gel electrophoresis in microbial molecular ecology, A. Van Leeuw J. Microb., 73, 127–141, 1998. Myers, M. R., Fischer, S. G., Maniatis, T., and Lerman, L. S.: Modification of the melting properties of duplex DNA by attachment of a GC-rich DNA sequence as determined by denaturing gradient gel electrophoresis, Nucl. Acids Res., 13, 3111–3129, 1985. Niemi, J. V., Saarikoski, S., Tervahattu, H., Makela, T., Hillamo, R., Vehkamaki, H., Sogacheva, L., and Kulmala, M.: Changes in background aerosol composition in Finland during polluted and clean periods studied by TEM/EDX individual particle analysis, Atmos. Chem. Phys., 6, 5049–5066, 2006. Osan, J., Szaloki, I., Ro, C. U., and Van Grieken, R.: Light element analysis of individual microparticles using thin-window EPMA, Mikrochim. Acta, 132(2–4), 349–355, 2000. Peccia, J. and Hernandez, M.: Incorporating polymerase chain reaction-based identification, population characterization, and quantification of microorganisms into aerosol science: a review, Atmos. Environ., 40, 3491–3961, 2006. Pernthaler, A., Pernthaler, J., and Amann, R.: Fluorescent in situ hybridisation and catalyzed reported deposition for the identification of marine bacteria, Appl. Environ. Microb., 68, 3094–3101, 2002. Pernthaler, J., Glöckner, F.-O., Unterholzner, S., Alfreider, A., Psenner, R., and Amann, R.: Seasonal community and population dynamics of pelagic bacteria and Archaea in a high mountain lake, Appl. Environ. Microb., 64, 4299–4306, 1998. Pósfai, M., Anderson, J. R., Buseck, P. R., Shattuck, T. W., and Tindale, N. W.: Constituents of a Remote Pacific Marine Aerosol: A TEM Study, Atmos. Environ., 28, 1747–1756, 1994. Pósfai, M., Anderson, J. R., Buseck, P. R., and Sievering, H.: Compositional variations of sea-salt-mode aerosol particles from the North Atlantic, J. Geophys. Res., 100, 23 063–23 074, 1995. Pósfai, M., Gelencsér, A., Simonics, R., Arató, K., Li, J., Hobbs, P. V., and Buseck, P. R.: Atmospheric tar balls: Particles from biomass and biofuel burning, J. Geophys. Res., 109(D6), D06213, doi:10.1029/2003JD004169, 2004. Pósfai, M., Li, J., Anderson, J. R., and Buseck, P. R.: Aerosol bacteria over the Southern Ocean during ACE-1, Atmos. Res., 66, 231–240, 2003. Pósfai, M. and Molnár, A.: Aerosol particles in the troposphere: A mineralogical introduction, edited by: Vaughan, D. J. and Wogelius, R. A., Environmental Mineralogy, 2, 197–252, 2000. Pósfai, M., Xu, H., Anderson, J. R., and Buseck, P. R.: Wet and dry sizes of atmospheric aerosol particles: A combined AFM-TEM study, Geophys. Res. Lett., 25, 1907–1910, 1998. Radosevich, J., Wilson, W., Shinn, J., DeSantis, T., and Anderse, G.: Development of a high-volume aerosol collection system for the identification of air-borne micro-organisms, Lett. Appl. Microbiol., 34, 162–167, 2002. Ramirez-Aguilar, K. A., Lehmpuhl, D. W., Michel, A. E., Birks, J. W., and Rowlen, K. L.: Atomic force microscopy for the analysis of environmental particles, Ultramicroscopy, 77, 187–194, 1999. Ranjard, L., Poly, F., and Nazaret, S.: Monitoring complex bacterial communities using culture-independent molecular techniques: application to soil environment, Res. Microbiol., 151, 167–177, 2000. Read, S.: Recovery efficiencies of nucleic acid extraction kits as measured by quantitative LightCycler$^\mboxTM$ PCR, J. Clin. Pathol., 54, 86–90, 2001. Russell, S. C., Czerwieniec, G., Lebrilla, C., Steele, P., Riot, V., Coffee, K., Frank, M., and Gard, E. E.: Achieving high detection sensitivity (14 zmol) of biomolecular ions in bioaerosol mass spectrometry, Anal. Chem., 77(15), 4734–4741, 2005. Russell, S. C., Czerwieniec, G., Lebrilla, C., Tobias, H., Fergenson, D. P., Steele, P., Pitesky, M., Horn, J., Srivastava, A., Frank, M., and Gard, E. E.: Toward understanding the ionization of biomarkers from micrometer particles by bio-aerosol mass spectrometry, J. Am. Soc. Mass Spectr., 15(6), 900–909, 2004. Rutledge, R. G.: Sigmoidal curve-fitting quantitative real-time PCR with the prospective of developing automated high-throughput applications, Nucl. Acids Res., 32(22), p. e178, 2004. Samburova, V., Didenko, T., Kunenkov, E., Emmenegger, C., Zenobi, R., and Kalberer, M.: Functional Group Analysis of High Molecular Weight Comounds in the Water-soluble Fraction of Organic Aerosols, Atmos. Environ., 41, 4703–4710, 2007. Sannigrahi, P., Sullivan, A. P., Weber, R. J., and Ingall, E. D.: Characterization of water-soluble organic carbon in urban atmospheric aerosols using solid-state 13C NMR spectroscopy, Environ. Sci. Technol., 40, 666–672, 2006. Sattler, B., Puxbaum, H., and Psenner, R.: Bacterial growth in supercooled cloud droplets, Geophys. Res. Lett., 28, 239–243, 2001. Scheinert, P., Krausse, R., Ullmann, U., Söller, R., and Krupp, G.: Molecular differentiation of bacteria by PCR amplification of the 16S-23SrRNA spacer, J. Microbiol. Meth., 26, 103–117, 1996. Schwieger, F. and Tebbe, C. C.: A new approach to utilize PCR-single strand conformation polymorphism for 16S rRNA gene-based microbial community analysis, Appl. Environ. Microbiol., 64, 4870–4876, 1998. Smalla, K., Oros-Sichler, M., Milling, A., Heuer, H., Baumgarte, S., Becker, R., Neuber, G., Kropf, S., Ulrich, A., and Tebbe, C. C.: Bacterial diversity of soils assessed by DGGE, T-RFLP and SSCP fingerprints of PCR-amplified 16S gene fragments: Do the different methods provide similar results?, J. Microbiol. Meth., 69, 470–479, 2007. Sobanska, S., Coeur, C., Maenhaut, W., and Adams, F.: SEM-EDX Characterisation of Tropospheric Aerosols in the Negev Desert (Israel), J. Atmos. Chem., 44, 299–322, 2003. Stetzenbach, L. D., Buttner, M. P., and Cruz, P.: Detection and enumeration of airborne biocontaminants, Curr. Opin. Biotech., 15, 170–174, 2004. Sullivan, R. C. and Prather, K. A.: Recent advances in our understanding of atmospheric chemistry and climate made possible by on-line aerosol analysis instrumentation, Anal. Chem., 77(12), 3861–3885, 2005. Tobias, H. J., Schafer, M. P., Pitesky, M., Fergenson, D. P., Horn, J., Frank, M., and Gard, E. E.: Bioaerosol mass spectrometry for rapid detection of individual airborne Mycobacterium tuberculosis H37Ra particles, Appl. Environ. Microb., 71(10), 6086–6095, 2005. Tong, Y. and Lighthart, B.: The annual bacterial particle concentration and size distribution in the ambient atmosphere in a rural area of the Willamette Valley, Oregon, Aerosol. Sci. Tech., 32, 393–403, 2000. van Borm, W. A., Adams, F. C., and Maenhaut, W.: Characterization of individual particles in the Antwerp aerosol, Atmos. Environ., 23(5), 1139–1151, 1989. van Poppel, L. H., Friedrich, H., Spinsby, J., Chung, S. H., Seinfeld, J. H., and Buseck, P. R.: Electron tomography of nanoparticle clusters: Implications for atmospheric lifetimes and radiative forcing of soot, Geophys. Res. Lett., 32(24), L24811, doi:10.1029/2005GL024461, 2005. van Wuijckhuijse, A. L., Stowers, M. A., Kleefsman, W. A., van Baar, B. L. M., Kientz, C. E., and Marijnissen, J. C. M.: Matrix-assisted laser desorption/ionisation aerosol time-of-flight mass spectrometry for the analysis of bioaerosols: development of a fast detector for airborne biological pathogens, J. Aerosol Sci., 36(5–6), 677–687, 2005. von Wintzingerode, F., Gobel, U. B., and Stackebrandt, E.: Determination of microbial diversity in environmental samples: pitfalls of PCR-based rRNA analysis, FEMS Microbiol. Rev., 21, 213–29, 1997. Warren, G. J.: Identification and analysis of ina genes and proteins, in: Biological Ice Nucleation and its Applications, edited by: Lee Jr., R. E., Warren, G. J., and Gusta, L. V., APS Press, St. Paul, MN, 85–100, 1995. Warren, G., Corotto, L., and Wolber, P.: Conserved repeats in diverged ice nucleation structural genes from two species of Pseudomonas, Nucl. Acids Res., 14, 8047–8060, 1986. Wilson, K. H., Wilson, W. J., Radosevich, J. L., Desantis, T. Z., Viswanathan, V. S., Kuczmarski, T. A., and Andersen, G. L.: High-density micro-array of small subunit ribosomal DNA probes, Appl. Environ. Microbiol., 68, 2535–2541, 2002. Wise, M. E., Biskos, G., Martin, S. T., Russell, L. M., and Buseck, P. R.: Phase transitions of single salt particles studied using a transmission electron microscope with an environmental cell, Aerosol Sci. Tech., 39(9), 849–856, 2005. Worobiec, A., de Hoog, J., Osan, J., Szaloki, I., Ro, C. U., and Van Grieken, R.: Thermal stability of beam sensitive atmospheric aerosol particles in electron probe microanalysis at liquid nitrogen temperature, Spectrochim. Acta B, 58(3), 479–496, 2003. Yamamoto, S., Kasai, H., Arnold, D., Jackson, R., Vivian, A., and Harayama, S.: Phylogeny of the genus Pseudomonas: intrageneric structure reconstructed from the nucleotide sequences of gyrB and rpoD genes, Microbiology, 146, 2385–2394, 2000. Zappoli, S., Andracchio, A., Fuzzi, S., Facchini, M. C., Gelencser, A., Kiss, G., Krivacsy, Z., Molnar, A., Meszaros, E., Hansson, H. C., Rosman, K., and Zebuhr, Y.: Inorganic, organic and macromolecular components of fine aerosol in different areas of Europe in relation to their water solubility, Atmos. Environ., 33, 2733–2743, 1999. Zeng, Q.-Y., Westermark, S.-O., Rasmuson-Lestander, Å., and Wang, X.-R.: Detection and quantification of Wallemia sebi in aerosols by real-time PCR, conventional PCR and cultivation, Appl. Environ. Microbiol., 70, 7295–7302, 2004.