A B S T R A C T Atmospheric aerosol particles of biological origin are a very diverse group of biological materials and structures, including microorganisms, dispersal units, fragments and excretions of biological organisms. In recent years, the impact of biological aerosol particles on atmospheric processes has been studied with increasing intensity, and a wealth of new information and insights has been gained. This review outlines the current knowledge on major categories of primary biological aerosol particles (PBAP): bacteria and archaea, fungal spores and fragments, pollen, viruses, algae and cyanobacteria, biological crusts and lichens and others like plant or animal fragments and detritus. We give an overview of sampling methods and physical, chemical and biological techniques for PBAP analysis (cultivation, microscopy, DNA/RNA analysis, chemical tracers, optical and mass spectrometry, etc.). Moreover, we address and summarise the current understanding and open questions concerning the influence of PBAP on the atmosphere and climate, i.e. their optical properties and their ability to act as ice nuclei (IN) or cloud condensation nuclei (CCN). We suggest that the following research activities should be pursued in future studies of atmospheric biological aerosol particles: (1) develop efficient and reliable analytical techniques for the identification and quantification of PBAP; (2) apply advanced and standardised techniques to determine the abundance and diversity of PBAP and their seasonal variation at regional and global scales (atmospheric biogeography); (3) determine the emission rates, optical properties, IN and CCN activity of PBAP in field measurements and laboratory experiments; (4) use field and laboratory data to constrain numerical models of atmospheric transport, transformation and climate effects of PBAP.

Aerosols of biological origin play a vital role in the Earth system, particularly in the interactions between atmosphere, biosphere, climate, and public health. Airborne bacteria, fungal spores, pollen, and other bioparticles are essential for the reproduction and spread of organisms across various ecosystems, and they can cause or enhance human, animal, and plant diseases. Moreover, they can serve as nuclei for cloud droplets, ice crystals, and precipitation, thus influencing the hydrological cycle and climate. The sources, abundance, composition, and effects of biological aerosols and the atmospheric microbiome are, however, not yet well characterized and constitute a large gap in the scientific understanding of the interaction and co-evolution of life and climate in the Earth system. This review presents an overview of the state of bioaerosol research, highlights recent advances, and outlines future perspectives in terms o fbioaerosolidentification, characterization, transport, and transforma- tion processes, as well as their interactions with climate, health, and ecosystems, focusing on the role bioaerosols play in the Earth system

Fungal spores can account for large proportions of air particulate matter, and they may potentially influence the hydrological cycle and climate as nuclei for water droplets and ice crystals in clouds, fog, and precipitation. Moreover, some fungi are major pathogens and allergens. The diversity of airborne fungi is, however, not well-known. By DNA analysis we found pronounced differences in the relative abundance and seasonal cycles of various groups of fungi in coarse and fine particulate matter, with more plant pathogens in the coarse fraction and more human pathogens and allergens in the respirable fine particle fraction (<3 m). Moreover, the ratio of Basidiomycota to Ascomycota was found to be much higher than previously assumed, which might also apply to the biosphere.atmospheric aerosol ͉ bioaerosol ͉ DNA analysis ͉ fungal spores R ecent studies have shown that fungal spores and other biological particles can account for large proportions of aerosol particle mass in pristine rainforest air as well as in rural and urban environments (1-5). For example, fungal spores were found to account for up to 45% of coarse particle mass (Ͼ1 m) in tropical rainforest air and up to 4-11% of fine particle mass (Յ2.5 m) in urban and rural air (3). Fungi have also been found in clouds, fog, and precipitation where these and other biological particles can act as nuclei for water droplets and ice crystals and can influence precipitation patterns and the Earth's energy budget (e.g., refs. 6-17). On average, the number and mass concentrations of fungal spores in continental boundary layer air are on the order of 10 3 -10 4 m Ϫ3 and approximately 1 g m Ϫ3 , respectively, and the estimated global emissions of approximately 50 Tg yr Ϫ1 are among the largest sources of organic aerosol (1).Some fungi are major pathogens or allergens for humans, animals, and plants, and air is the primary medium for their dispersal (18)(19)(20), but the diversity of fungi in air particulate matter is not well-known. The traditional cultivation, microscopy, and chemical tracer techniques applied in earlier studies were insufficient for a broad coverage of different fungal species, and recently reported first applications of molecular genetic techniques were very limited in scope and methodology as discussed below.In this study, we investigated and characterized the diversity and frequency of occurrence of fungi in air particulate matter by DNA extraction and sequence analysis of the internal transcribed spacer region (ITS).

Poor air quality is globally the largest environmental health risk. Epidemiological studies have uncovered clear relationships of gaseous pollutants and particulate matter (PM) with adverse health outcomes, including mortality by cardiovascular and respiratory diseases. Studies of health impacts by aerosols are highly multidisciplinary with a broad range of scales in space and time. We assess recent advances and future challenges regarding aerosol effects on health from molecular to global scales through epidemiological studies, field measurements, health-related properties of PM, and multiphase interactions of oxidants and PM upon respiratory deposition. Global modeling combined with epidemiological exposure-response functions indicates that ambient air pollution causes more than four million premature deaths per year. Epidemiological studies usually refer to PM mass concentrations, but some health effects may relate to specific constituents such as bioaerosols, polycyclic aromatic compounds, and transition metals. Various analytical techniques and cellular and molecular assays are applied to assess the redox activity of PM and the formation of reactive oxygen species. Multiphase chemical interactions of lung antioxidants with atmospheric pollutants are crucial to the mechanistic and molecular understanding of oxidative stress upon respiratory deposition. The role of distinct PM components in health impacts and mortality needs to be clarified by integrated research on various spatiotemporal scales for better evaluation and mitigation of aerosol effects on public health in the Anthropocene.

Cloud glaciation is critically important for the global radiation budget (albedo) and for initiation of precipitation. But the freezing of pure water droplets requires cooling to temperatures as low as 235 K. Freezing at higher temperatures requires the presence of an ice nucleator, which serves as a template for arranging water molecules in an ice-like manner. It is often assumed that these ice nucleators have to be insoluble particles. We point out that also free macromolecules which are dissolved in water can efficiently induce ice nucleation: the size of such ice nucleating macromolecules (INMs) is in the range of nanometers, corresponding to the size of the critical ice embryo. As the latter is temperature-dependent, we see a correlation between the size of INMs and the ice nucleation temperature as predicted by classical nucleation theory. Different types of INMs have been found in a wide range of biological species and comprise a variety of chemical structures including proteins, saccharides, and lipids. Our investigation of the fungal species Acremonium implicatum, Isaria farinosa, and Mortierella alpina shows that their ice nucleation activity is caused by proteinaceous water-soluble INMs. We combine these new results and literature data on INMs from fungi, bacteria, and pollen with theoretical calculations to develop a chemical in-terpretation of ice nucleation and water-soluble INMs. This has atmospheric implications since many of these INMs can be released by fragmentation of the carrier cell and subsequently may be distributed independently. Up to now, this process has not been accounted for in atmospheric models

Abstract. Soil organic matter (SOM) may be a significant source of atmospheric ice nucleating particles (INPs), especially of those active > −15 • C. However, due to both a lack of investigations and the complexity of the SOM itself, the identities of these INPs remain unknown. To more comprehensively characterize organic INPs we tested locally representative soils in Wyoming and Colorado for total organic By contrast, lysozyme, which digests bacterial cell walls, only reduced INPs active at ≥ −7.5 or ≥ −6 • C, depending on the soil. The known IN bacteria were not detected in any soil, using PCR for the ina gene that codes for the active protein. We directly isolated and photographed two INPs from soil, using repeated cycles of freeze testing and subdivision of droplets of dilute soil suspensions; they were complex and apparently organic entities. Ice nucleation activity was not affected by digestion of Proteinase K-susceptible proteins or the removal of entities composed of fulvic and humic acids, sterols, or aliphatic alcohol monolayers. Organic INPs active colder than −10 to −12 • C were resistant to all investigations other than heat, oxidation with H 2 O 2 , and, for some, digestion with papain. They may originate from decomposing plant material, microbial biomass, and/or the humin component of the SOM. In the case of the latter then they are most likely to be a carbohydrate. Reflecting the diversity of the SOM itself, soil INPs have a range of sources which occur with differing relative abundances.

Specialized bacteria trigger ice formation by controlling the molecular structure and energy transfer in interfacial water.

Abstract. Biological residues in soil dust are a potentially strong source of atmospheric ice nuclei (IN). So far, however, the abundance, diversity, sources, seasonality, and role of biological – in particular, fungal – IN in soil dust have not been characterized. By analysis of the culturable fungi in topsoils, from a range of different land use and ecosystem types in southeast Wyoming, we found ice-nucleation-active (INA) fungi to be both widespread and abundant, particularly in soils with recent inputs of decomposable organic matter. Across all investigated soils, 8% of fungal isolates were INA. All INA isolates initiated freezing at −5 to −6 °C, and belonged to a single zygomycotic species, Mortierella alpina (Mortierellales, Mortierellomycotina). To our knowledge this is the first report of ice nucleation activity in a zygomycotic fungi because the few known INA fungi all belong to the phyla Ascomycota and Basidiomycota. M. alpina is known to be saprobic and widespread in soil, and Mortierella spores are present in air and rain. Sequencing of the ITS region and the gene for γ-linolenic elongase revealed four distinct clades, affiliated to different soil types. The IN produced by M. alpina seem to be proteinaceous, < 300 kDa in size, and can be easily washed off the mycelium. Ice nucleating fungal mycelium will ramify topsoils and probably also release cell-free IN into it. If these IN survive decomposition or are adsorbed onto mineral surfaces, their contribution might accumulate over time, perhaps to be transported with soil dust and influencing its ice nucleating properties.