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AIRBIOTA-CM Program (S2013/MAE-2874, Community 34 of Madrid, Spain) supported the Microbiome analysis. J. Oteros was partially supported 35 by the Postdoctoral Fellowship Program of the Helmholtz Zentrum München. 36 37 Disclosure of potential conflict of interest: All the authors declare that they have no 38 relevant conflicts of interest. 39 - 2 - 40 Abstract: 41 Background: Endotoxin (lipopolysaccharides, LPS) released from Gram-negative 42 bacteria causes strong immunologic and inflammatory effects and when airborne may 43 contribute to respiratory conditions such as allergic asthma. 44 Objectives: To identify the source of airborne endotoxin and the effect of this endotoxin 45 on allergic sensitization. 46 Methods: We determined LPS in outdoor air on a daily basis for 4 consecutive years in 47 Munich (Germany) and Davos (Switzerland). Air was sampled as Particulate Matter 48 PM>10μm and 10>PM>2.5. LPS was determined using the recombinant Factor C (rFC) 49 assay. 50 Results: Over 60% of the annual endotoxin exposure was detected in the PM>10 fraction 51 showing that bacteria do not aerosolize as independent units or aggregates, but adhered 52 to large particles. In Munich 70% of the annual exposure was detected between June 53 12th and August 28th. Multivariate modelling showed that endotoxin levels could be 54 explained by phenological parameters i.e. plant growth. Indeed, days with high airborne 55 endotoxin levels correlated well with the amount of Artemisia pollen in the air. Pollen 56 collected from plants across Europe (100 locations) showed that the highest levels of 57 endotoxin were detected on Artemisia vulgaris (mugwort) pollen, with little on other 58 pollen. Microbiome analysis showed that LPS concentrations on mugwort pollen were 59 related to the presence of Pseudomonas spp. and Pantoea spp. communities. In a mouse 60 model of allergic disease, the presence of LPS on mugwort pollen was needed for allergic 61 sensitization. 62 Conclusions: The majority of airborne endotoxin stems from bacteria dispersed with 63 pollen of only one plant: mugwort. This LPS was essential for inducing inflammation of 64 the lung and allergic sensitization. 65 66 Key Messages: 67 70% of airborne endotoxin was dispersed with only one specific pollen type: Artemisia 68 pollen. 69 In an animal model, the endotoxin was essential for inducing allergic sensitization and 70 lung inflammation. - 3 - 71 The microbial load of pollen could enhance their allergenic impact. 72 73 Capsule Summary: Artemisia pollen is the main vector for airborne endotoxin. This 74 endotoxin (LPS) was essential for allergic sensitization against Artemisia pollen. 75 76 Keywords: Endotoxin; ambient; bacteria; pollen; Gram-negative; recombinant Factor c; 77 Davos; Munich; Artemisia; microbiome; mouse model; allergy; lipopolysaccharide; LPS; 78 source; PM10 79 80 Abbreviations: 81 LPS: Lipopolysaccharides. 82 NGS: Next Generation Sequencing. 83 rFC: recombinant Factor C 84 PM: Particulate Matter 85 - 4 - 86 Introduction 87 Endotoxins (lipopolysaccharides, LPS) are macromolecules from the outer 88 cellular wall of all Gram-negative bacteria, and are essential for their viability. These 89 molecules consist of an O-polysaccharide, a Core-oligosaccharide and Lipid A. LPS is one 90 of the most potent activators of the immune system, including in humans. There are 91 many LPS types in nature, but all of them have a phosphorylated diglucosamine 92 backbone substituted with several acyl chains and one or more Kdo (2-Keto-3-desoxy- 93 octonate) residues 1. 94 Endotoxin exposure can influence human health. Airborne endotoxin is a known 95 immunotoxin causing inflammatory reactions of the respiratory system, the main 96 symptoms being: fever, chest tightness, bronchospasm, pyrexia and ultimately chronic 97 neutrophilic airway inflammation. An excessive exposure to LPS results in a systemic 98 inflammatory reaction, leading to multiple organ failure, shock and potentially death 1. 99 A positive association has been found between indoor LPS exposure and wheezing 2. 100 Upon long-term exposure, lung inflammation caused by LPS is a determinant for the 101 progression of chronic respiratory diseases 3. Inhaled endotoxin causes asthma 102 intensification and adverse respiratory symptoms and is also a risk factor for increased 103 asthma prevalence 4, but not all studies show a consensus 2. Although high LPS levels 104 increase asthmatic symptoms, epidemiological data shows that low LPS exposure is 105 related with less allergy and atopic sensitization 5. The possibility of a protective effect 106 of exposure to endotoxin in the development of allergy resulted in the hygiene 107 hypothesis. The exposure to this pro-inflammatory agent may activate the immune 108 system towards Th1 responses. Th1 responses suppress the development of 109 immunoglobulin E antibodies. 110 Endotoxin concentrations in the workplace and other indoor environments, 111 where populations spend most of their time, have been extensively studied. There is, 112 however, a large gap in our knowledge about exposure to endotoxins in outdoor 113 environments. The consequences of prolonged exposure to a constant outdoor LPS are 114 still unknown. Although indoor LPS concentrations are affected by specific indoor factors 115 (e.g. pets in the house, the age of the building, building usage, number of occupants, 116 smoking, etc.) 6, they are also affected by conditions outdoors 7. Furthermore, indoor - 5 - 117 bacterial communities show seasonality over the year 8, which could be driven by 118 outdoor variations 9. Indoor LPS concentrations can be either higher or lower than 119 outside concentrations 10, 11. 120 The identity and source of bacteria producing airborne LPS outdoors is also not 121 well known. Airborne bacteria are ubiquitous, but their communities vary depending on 122 the surrounding environment and are much higher over terrestrial areas than over 123 oceans 12. Bacteria in outdoor air mostly originate from natural rather than anthropic 124 sources 13. Soil dust is thought to be one of the main natural sources
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