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1f4ec41f723e758522faa99829a52f00ea45a9e2.json
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{
"paper_id": "1f4ec41f723e758522faa99829a52f00ea45a9e2",
"metadata": {
"title": "Aerobiology and Its Role in the Transmission of Infectious Diseases",
"authors": [
{
"first": "Aaron",
"middle": [],
"last": "Fernstrom",
"suffix": "",
"affiliation": {
"laboratory": "",
"institution": "Mid-Atlantic Venture Investment Company, LLC",
"location": {
"postCode": "20009",
"settlement": "Washington",
"region": "DC",
"country": "USA"
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"email": ""
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{
"first": "Michael",
"middle": [],
"last": "Goldblatt",
"suffix": "",
"affiliation": {
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"institution": "Functional Genetics, Inc",
"location": {
"postCode": "20818",
"settlement": "Gaithersburg",
"region": "MD",
"country": "USA"
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"email": ""
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"text": "Aerobiology plays a fundamental role in the transmission of infectious diseases. As infectious disease and infection control practitioners continue employing contemporary techniques (e.g., computational \ufffduid dynamics to study particle \ufffdow, polymerase chain reaction methodologies to quantify particle concentrations in various settings, and epidemiology to track the spread of disease), the central variables affecting the airborne transmission of pathogens are becoming better known. \ue062is paper reviews many of these aerobiological variables (e.g., particle size, particle type, the duration that particles can remain airborne, the distance that particles can travel, and meteorological and environmental factors), as well as the common origins of these infectious particles. We then review several real-world settings with known difficulties controlling the airborne transmission of infectious particles (e.g., office buildings, healthcare facilities, and commercial airplanes), while detailing the respective measures each of these industries is undertaking in its effort to ameliorate the transmission of airborne infectious diseases.",
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"text": "Exposure to airborne pathogens is a common denominator of all human life [1] . With the improvement of research methods for studying airborne pathogens has come evidence indicating that microorganisms (e.g., viruses, bacteria, and fungal spores) from an infectious source may disperse over very great distances by air currents and ultimately be inhaled, ingested, or come into contact with individuals who have had no contact with the infectious source [2] [3] [4] [5] . Airborne pathogens present a unique challenge in infectious disease and infection control, for a small percentage of infectious individuals appear to be responsible for disseminating the majority of infectious particles [6] . \ue062is paper begins by reviewing the crucial elements of aerobiology and physics that allow infectious particles to be transmitted via airborne and droplet means. Building on the basics of aerobiology, we then explore the common origins of droplet and airborne infections, as these are factors critical to understanding the epidemiology of diverse airborne pathogens. We then discuss several environmental considerations that in\ufffduence the airborne transmission of disease, for these greatly impact particular environments in which airborne pathogens are commonly believed to be problematic. Finally, we discuss airborne pathogens in the context of several speci\ufffdc examples: healthcare facilities, office buildings, and travel and leisure settings (e.g., commercial airplanes, cruise ships, and hotels).",
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"text": "Aerobiology is the study of the processes involved in the movement of microorganisms in the atmosphere from one geographical location to another [7] , including the aerosolized transmission of disease. \ue062e aerosolized transmission of disease occurs through both \"droplet\" and \"airborne\" means. Droplet transmission is de\ufffdned as the transmission of diseases by expelled particles that are likely to settle to a surface quickly, typically within three feet of the source [8] [9] [10] . \ue062us, for example, in order for an infection to be caused by droplet transmission, a susceptible individual must be close enough to the source of the infection (e.g., an infected individual) in order for the droplet (containing the infectious microorganism) to make contact with the susceptible individual's respiratory tract, eyes, mouth, nasal passages, and so forth [11] . In contrast, airborne transmission is de\ufffdned as the transmission of infection by expelled particles that are comparatively smaller in size and thus can remain suspended in air for long periods of time. Airborne particles are particularly worrisome simply because they can remain suspended in the air for extended periods of time. Seminal studies from the 1930s and 1940s [8, 12, 13] demonstrated that airborne particles can remain airborne for as long as one week a\ue09der initial aerosolization, and suggested further [13] that these particles likely remained airborne for much longer. \ue062ey thus potentially expose a much higher number of susceptible individuals at a much greater distance from the source of infection [10, 11, 14, 15] . Depending on environmental factors (e.g., meteorological conditions outdoors and \ufffduid dynamic effects and pressure differentials indoors), airborne particles are easily measured 20 m from their source [16] . \ue062ese factors would be of no concern but for the fact that airborne bacterial, viral, and fungal particles are o\ue09den infectious [17] .",
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"text": "A complicating factor is the heterogeneous nature of droplet and airborne releases, which generally consist of mixtures of both single and multiple cells, spores, and viruses carried by both respiratory secretions and inert particles (e.g., dust) [17] . \ue062e origins of droplet or airborne infectious microorganisms are also heterogeneous: infectious particles may be generated from, for example, infectious persons, heating, ventilation, and air conditioning (HVAC) systems, and cooling tower water in hospitals [17] . All of these sources can produce airborne infectious particles [17] . Furthermore, Aspergillus fumigatus spores are common in dusts during outdoor and indoor construction, in air conditioners, ceiling tile, carpet, and other infectious aerosol carriers generated from dry sources; they may absorb water in the airborne state but still measure in the infectious particle size range [17] . Also, droplet and airborne transmission are not mutually exclusive. \ue062at is, independent of origin, particles carrying infectious microorganisms do not exclusively disperse by airborne or droplet transmission, but by both methods simultaneously [11] .",
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"text": "Transmission of infectious disease by the airborne route is dependent on the interplay of several critical factors, primarily particle size (i.e., the diameter of the particle) and the extent of desiccation [17] . \ue062e literature suggests that a particle's size is of central importance in determining whether it becomes and remains airborne and infectious [18] [19] [20] [21] [22] [23] . Simply illustrated, large particles fall out of the air and small particles remain airborne. \ue062e World Health Organization uses a particle diameter of 5 m to delineate between airborne (\u22645 m) and droplet (>5 m) transmission [17, 24, 25] . How particle size affects spatial distribution in the human respiratory tract has been studied extensively. Some studies suggest that particles over 6 m tend to mainly deposit in the upper airway, while particles under 2 m deposit mainly in the alveolar region [26] . Other studies conclude that particles under 10 m can penetrate deeper into the respiratory tract, and particles over 10 m are more likely to deposit on the surfaces of the upper airways and are less likely to penetrate into the lower pulmonary region [27] [28] [29] [30] [31] [32] [33] [34] [35] .",
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"text": "One of the challenges facing practitioners, particularly in an enclosed building, is that even large-sized droplets can remain suspended in air for long periods [17] . \ue062e reason is that droplets settle out of air onto a surface at a velocity dictated by their mass [17] . If the upward velocity of the air in which they circulate exceeds this velocity, they remain airborne. Hence, droplet aerosols up to 100 m diameter have been shown to remain suspended in air for prolonged periods when the velocity of air moving throughout a room exceeds the terminal settling velocity of the particle [17] .",
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"text": "Another critical variable is the rate at which particles desiccate. Even large, moisture laden droplet particles desiccate rapidly. In his seminal paper, Wells showed that particles begin desiccating immediately upon expulsion into the air and do so rapidly: particles up to 50 m can desiccate completely within 0.5 seconds [8] . Rapid desiccation is a concern since the smaller and lighter the infectious particle, the longer it will remain airborne. Hence, even when infectious agents are expelled from the respiratory tract in a matrix of mucus and other secretions, causing large, heavy particles, rapid desiccation can lengthen the time they remain airborne (the dried residuals of these large aerosols, termed droplet nuclei, are typically 0.5-12 m in diameter [17] ). Of further concern, very large aerosol particles may initially fall out of the air only to become airborne again once they have desiccated [17] .",
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"text": "One reason why particle size is such an important variable in airborne and droplet disease transmission is that the ability of an infectious disease to cause an infection depends on the concentration of the microorganism, the human infectious dose, and the virulence of the organism [17] . Humans can acquire devastating infectious diseases through exposure to very low levels of infectious particles. For example, In\ufffduenza A is believed to transmit via airborne and droplet means, and the infectious dose of In\ufffduenza A for humans is very low [62] . Additionally, the infectious dose for Francisella tularensis is reported to be a single organism [17] . Only a few cells of Mycobacterium tuberculosis are required to overcome normal lung clearance and inactivation mechanisms in a susceptible host [17] .",
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"text": "Airborne Infections \ue062e origins of infections resulting from droplet and airborne transmission are at the intersection of the clinical manifestation of disease, the site of infection, the presence of a pathogen, and the type of pathogen [11] . \ue062us, when investigating the origins of droplet and airborne infections, there are several well-known primary sources of infectious particles (see Table 1 ): vomiting, toilet \ufffdushing (i.e., toilet water aerosolization), sneezing, coughing, and talking. Moreover, toilet bowls, the water in them, and toilet seats may harbor infectious particles a\ue09der the initial \ufffdush, making additional aerosolization of infectious particles possible with additional \ufffdushes for as long as 30 minutes a\ue09der the initial \ufffdush [63] . Particle desiccation, discussed above, is important in this context. A single sneeze, for example, generates as many as 40,000 large droplet particles; most will desiccate Journal of Pathogens 3 T\uf761\uf762\uf76c\uf765 1: Droplet or airborne microorganisms released from various activities.",
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"text": "Approximate particle count Units Sneezing [36] 40,000 Per sneeze Bowel evacuation [37] 20,000 Per event Vomiting [38] 1,000 Per event Coughing [36] 710 Per cough Talking [36] 36 Per 100 words immediately into small, infectious droplet nuclei [17] , with 80% of the particles being smaller than 100 m [64] . \ue062e transmission of infectious diseases via airborne or droplet routes may also depend on the frequency of the initiating activity. For example, while a single sneeze may produce more total infectious particles than a cough [11, 28, 65, 66] , Couch et al. reported that coughing is more frequent than sneezing during infection with Coxsackievirus A [67] . \ue062is \ufffdnding suggests that coughing is a more likely method of airborne transmission for this disease than sneezing [67] . As coughing is also a common symptom of in\ufffduenza infection [68, 69] , it may also contribute to the airborne transmission of this pathogen.",
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"text": "Finally, infectious individuals are not always the immediate source of airborne infectious particles. Many people spend considerable time in office buildings, for example, and as a result become exposed to airborne pathogens that originate from nonhuman sources (e.g., molds, toxins produced by molds, pollen, pet dander, and pest droppings) [70] [71] [72] [73] [74] [75] [76] [77] . \ue062e health effects associated with naturally occurring indoor biological air pollutants include disease, toxicoses, and hypersensitivity (i.e., allergic) diseases [70] [71] [72] [73] [74] [75] [76] [77] . In addition, exposure to indoor biological air pollutants has been associated with \"sick building syndrome, \" a set of nonspeci\ufffdc symptoms that may include upper-respiratory symptoms, headaches, fatigue, and rash and \"appear to be linked to time spent in a building, but no speci\ufffdc illness or cause can be identi\ufffded. \" [78] .",
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"text": "While the airborne transmission of disease depends on several physical variables endemic to the infectious particle, environmental factors substantially in\ufffduence the efficacy of airborne disease transmission. \ue062e environmental factors most o\ue09den cited as modifying the airborne transmission of disease are temperature and relative humidity [17] . Together, they help determine whether or not an airborne particle can remain infectious [17] . For example, the size of infectious particles can change depending on relative humidity and temperature (i.e., factors that in\ufffduence desiccation or hygroscopicity). An added complication is the fact that temperature and humidity in\ufffduence viral, bacterial, and fungal particles differently [17] .",
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"text": "Temperature is an important factor affecting virus survival [79, 80] . Generally, as temperature rises, virus survival decreases [79] . For example, low temperatures (i.e., 44.6 \u2218 F-46.4 \u2218 F) have been suggested to be ideal for airborne in\ufffduenza survival, with survival decreasing progressively at moderate (i.e., 68.9 \u2218 F-75.2 \u2218 F) and high temperatures (i.e., >86 \u2218 F). \ue062is relationship holds across a range of relative humidities (i.e., 23%-81%) [81] . In\ufffduenza has also been shown to be transmissible via airborne vector under cold, dry conditions [82] . While relative humidity is recognized to be a factor in the viability of airborne and droplet viral transmissions [79, 80] , the exact relationship is presently not well understood. For example, the report of Arundel et al. that minimal survival for both lipid-enveloped and non-lipidenveloped viruses occurs at relative humidities between 40% and 70% [82] contrasts with that for in\ufffduenza noted above.",
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"text": "In general, bacteria are more resistant to temperature than viruses [83, 84] . Temperatures above 75.2 \u2218 F are required to reduce airborne bacterial survival [83, 84] . \ue062is temperature relationship has been found with gram-negative, gram-positive, and intracellular bacteria: Pseudomonas sp. [83, 84] , Pasteurella sp. [85] , Salmonella sp. [86] , Serratia sp. [87] , Escherichia sp. [87] [88] [89] , Bacillus sp. [87] , Bordetella sp. [90] , Chlamydia sp. [91] , and Mycoplasma sp. [92] . \ue062e survival of aerosolized gram-negative bacteria (including Pseudomonas sp., Enterobacter sp., and Klebsiella sp.) has been reported to be greatest at high relative humidity and low temperature [93] . However, available data on the effects of relative humidity on the survival of airborne bacteria are thus far inconsistent. For example, airborne gram-negative bacteria (e.g., E. coli, Salmonella sp., etc.) are reported not to survive well at increased relative humidity [94, 95] , while some airborne gram-positive bacteria (Staphylococcus albus, Streptococcus haemolyticus, Bacillus subtilis, and Streptococcus pneumoniae (type 1)) survive poorly at intermediate relative humidities [94] [95] [96] . Determining the rates of survival of airborne bacteria appears to be more complicated than with viruses [97, 98] . Even bacteria within the same structural classi\ufffdcation (e.g., gram-negative) may vary in how they respond to different changes in temperature and relative humidity [79] .",
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"text": "For fungi, extensive studies have characterized the levels of both indoor and outdoor airborne fungi and their spores [99, 100] . More than viruses or bacteria, airborne fungi and their spores have been suggested to have the potential to enter a building that uses natural ventilation. Certain species (e.g., Aspergillus sp.) are also well-known, potentially life-threatening airborne contaminants when introduced to immunocompromised patients (such as in a healthcare facility) [99] . Other fungi hazardous to the immunocompromised include Blastomyces sp., Coccidioides sp., Cryptococcus sp., and Histoplasma sp. [100] . Even in healthy people, individuals working consistently in indoor environments (such as an office or school) have shown hypersensitivity reactions such as rhinitis, sinusitis, or asthma in response to fungi exposure [79] . Relatively few laboratory studies have examined the airborne transmission of fungi and their spores in relation to temperature and relative humidity. Most data relating these variables to airborne fungi viability have been obtained in their natural environments [79] . Nonetheless, the results of such studies suggest a seasonal variation in airborne fungal and spore concentrations associated with common environmental conditions, including ambient temperature, relative humidity, precipitation, and wind speed [97, 98, 101] . Generally, fungi and their spores appear to be more resilient than viruses and bacteria, being able to withstand greater stresses due to dehydration and rehydration, as well as UV radiation [97, 98, 101] .",
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"text": "Given the diversity of viruses and bacteria that can spread via airborne or droplet means (see Table 2 ), an understanding of aerobiology, typical origins of droplet and airborne infections, and how different environmental factors affect airborne and droplet particles is critical to any discussion of the amelioration or mitigation of infectious airborne and droplet particle transmission.",
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"text": "\ue062ere are two principal challenges when working to ameliorate or mitigate the airborne transmission of infectious particles indoors: preventing in\ufffdltration and preventing transmission. We discuss the \ufffdrst in the context of office buildings and the latter in the context of healthcare facilities (both below).",
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