Unravelling the bacterial diversity in the atmosphere.

The study of airborne biological particles ('bioaerosols') has gained interest in recent years, due to an increasing amount of evidence suggesting that this fraction of airborne particulate matter may play a critical role in the negative effects of aerosols on biological systems. Pioneer investigations demonstrated that bacteria do exist in the atmosphere and can be metabolically active, although studies have not proved whether they actually form ecological communities or are merely assemblages of organisms passively transported from different sources. For a long time, cultivation-based methods have been the gold standard to describe and quantify airborne microorganisms. However, the use of culture-independent techniques and, more recently, of the next-generation sequencing-based methods, has improved the ability of the scientific community to investigate bioaerosols in detail and to address further research questions, such as the temporal and spatial variability of airborne bacterial assemblages, the environmental factors affecting this variability and the potential sources of atmospheric bacteria. This paper provides a systematic review of the state-of-the-art methodologies used in the study of airborne bacteria to achieve each of the aforementioned research objectives, as well as the main results obtained so far. Critical evaluations of the current state of the knowledge and suggestions for further researches are provided.