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Summary, 1S Introduction, 1S Role of movement of populations in infectious diseases, 2S Historical events, 2S Current volume and range of movement, 2S Diversity of pathogens and mechanisms of transmission, 2S Trade, 3S Receptivity of places and populations, 3S Population size, density and vulnerability, 3S Location and physicochemical characteristics, 3S Competent vectors and intermediate hosts, 4S Socioeconomic, political factors; infrastructure and access to health care, 4S Populations and places affected, 4S Travel as a circuit, 4S En route transmission, 4S Process of travel; cruise ship as a new habitat, 5S Travellers, 6S Travellers as sentinels, 6S Travellers as couriers; key role of laboratory, 6S Travellers as transmitters/disseminators, 7S Time of consequences, 7S Specific disease examples, 7S Neisseria meningitidis, W135, 7S Dengue fever, 8S Microbial genetic material; resistance and virulence factors, 8S Conclusions, 8S Acknowledgements, 9S References, 9S The movement of populations shapes the patterns and distribution of infectious diseases globally. The consequences of travel are seen in the traveller and in places and populations visited and may persist long after travel. The traveller can be seen as an interactive biological unit who picks up, processes, carries and drops off microbial genetic material. A traveller can introduce potential pathogens in the absence of signs or symptoms of illness. Travellers can serve as a sentinel population; study of them can provide insights into the presence and level of risk of transmission of infections in other geographical regions. Travellers can also be seen as couriers who inadvertently ferry pathogens and microbial genetic material to regions where researchers can carry out detailed analyses that can help to map the location and movement of strains, genotypes and resistance patterns. The laboratory plays a key role in the identification and characterization of pathogens, which can inform management of individual patients and the public health response. The connectedness and mobility in the world today facilitate the emergence of infectious diseases in humans and also in animals and plants. Many traditional barriers have been breached by travel, roads and technology. Population size and density favour spread of many infections. The rapid generation time of microbes and their capacity to adapt to changes in the physico-chemical and immunological environment will pose continuing challenges. Travellers are interactive biological units that pick up, process and drop off microbial genetic material at different times and in different places. Travellers alter places and populations during and after travel. Changes in the biota may be sustained if a traveller carries a pathogen into a new environment where it is transmitted, and then survives, replicates, and persists in a new geographical area. Migrating humans and trade play a key role in the global dispersal of human pathogens and of microbial genetic material (Wilson 1995a). This paper will review the mechanisms through which the movement of populations affects the distribution and spread of infectious diseases and the factors that influence the vulnerability of places and populations to introductions of infections. The examples of dengue fever, influenza, meningococcus and multidrug-resistant pathogens will be used to illustrate key points. History books are full of vivid examples of the role of traders, travellers, pilgrims, missionaries, explorers, warriors and other migrating individuals or groups who introduced human pathogens into susceptible populations, often with devastating consequences. The New World was hard hit when European explorers brought with them measles, smallpox and tuberculosis, which caused high mortality in local non-immune populations (Crosby 1972). The major culprits were pathogens that were carried by humans and spread easily from person to person. Exploration of many parts of sub-Saharan Africa by Europeans was associated with high death rates from infections, such as malaria, yellow fever, and sleeping sickness – vector-borne infections that, in a sense, were embedded the landscape. In a book that explores the excess mortality or ‘relocation costs’ among European soldiers in the tropics between 1815 and 1914, Philip Curtin found that the death rates from disease were at least twice as high in soldiers during time spent in the tropics relative to that of soldiers who remained in temperate areas (Curtin 1989). Many of these vector-borne diseases persist today and pose a risk to local residents and visitors. Throughout most of history many more soldiers died from infectious diseases than directly from weapons of war. Population movements and displacements associated with war and violent conflicts also lead to outbreaks of infectious diseases that involve civilian populations. Infections, such as cholera, typhoid fever, and measles, in refugee and displaced populations are often a consequence of crowding, poor nutrition, lack of basic sanitation, and disruption of basic medical services, such as immunization programmes, during periods of social and political upheaval. Travellers form an important bridge to diverse geographical areas and populations. Infections that can be spread from person to person can be readily carried by travellers to any part of the globe. AIDS is a modern day example of an infection that, in a few short decades, has been carried to all parts of the world. The ease and volume of modern human traffic has facilitated its spread, assisted in many areas by sexual practices, sexual tourism, shared injection devices and the use of contaminated instruments or blood in the medical setting. An estimated 60 million persons have been infected since the mid-1980s and 40 million persons are estimated to be living with HIV today. The speed, volume and reach of travel today are unprecedented. The world has more and faster links than at any time in history. Today it is possible to travel to virtually any part of the world in a time period that is less than the incubation for many infectious diseases. The World Tourism Organization (WTO) estimated that world tourism grew by 7·4% in 2000 and the total number of international arrivals reached 699 million (World Tourism Organization 2001). More than 1·4 million persons cross-international borders on flights everyday. In the 1990s more than 5000 airports had regularly scheduled international flights, which means that dense urban centres and their surrounding periurban areas all over the world are linked. Of course, air travel is only one means of travel, as humans travel by boat, bus, train, car, truck and by foot. An estimated 400 million international travellers entered the US by air, land or ship in 1999. Travelling humans also carry or facilitate the transport of plants, animals, seeds and all matter of biological material. The movement of biota (including humans, animals and plants) impacts on the traveller and also on populations and places en route, at the destination, and where the traveller returns after travel. The traveller may be at risk of acquiring unfamiliar infections in a new environment, but the impact of travel encompasses a range of events that derive from microbes and other biota carried by the traveller, but also the traveller's behaviour, often based on cultural or societal traditions that may influence risk of exposure. The process of travel also changes the environment in ways that may make other populations more or less vulnerable to infections. The idea of emerging infections for many people conjures images of exotic or remote viruses that cause haemorrhagic fever, viruses such as Ebola, Marburg or Lassa fever. Although these remain a serious concern in some regions of the world, agents causing emerging microbial threats include common, familiar infections, such as influenza and tuberculosis. They come from all classes of organisms – viruses, bacteria, protozoa, fungi and helminths. They include infections spread from person to person as well as vector-borne infections and those acquired from contact with animals, soil or water. Although the focus in this paper is on emerging infections in humans, emerging infections also affect plants and animals, and these can have direct and indirect health and economic impacts on humans. Prominent examples include bovine spongiform encephalopathy (BSE), foot-and-mouth disease, avian influenza and West Nile virus infection. Among the agents causing emerging infections are microbes previously well characterized that have spread geographically (e.g. dengue and West Nile virus), increased in incidence (e.g. diphtheria, HIV and tuberculosis), changed in virulence (e.g. Neisseria meningitidis clones), or become more resistant to treatment (e.g. Mycobacterium tuberculosis, Streptococcus pneumoniae, Salmonella species, falciparum and vivax malaria, Staphylococcus aureus). For several established pathogens, the presence of HIV infection increases the risk of infection, alters the clinical presentation, or reduces the efficacy of standard treatment. Most of the infections perceived as being ‘new’ are not caused by novel organisms but by microbes long present but not well characterized or not known previously to cause human disease. Examples include Helicobacter pylori, ehrlichiae, cyclospora and Sin Nombre virus (one cause of hantavirus pulmonary syndrome). In some instances, humans on or changed a in ways that increased contact between humans and a This is of some of the viruses, such as the and the cause of haemorrhagic fever. New laboratory have it possible to at to of and that were causing infections in the US the disease was and the pathogen of of from and that from the that disease, was present in the US at least as as trade is dispersal that carries human pathogens, such as and intermediate the world. The is and may be and of from where it is and distribution also make it possible for a in one location and in to be to or For Salmonella outbreaks in in and the US in US and in were to from seeds from a are regularly carried the world by or other some can become established in a new environment and may spread to of these have the capacity to serve as vectors of human infections, to organisms from to to be an for the of to new regions. In was introduced into the in used from The the was found in in and had spread to in The dispersal and The of this more than the presence of In the laboratory is a for at least many of them human virus and dengue viruses have been from in is the in an of dengue fever, caused by dengue that in in of human found on of the had been laboratory all geographical regions and populations are to the and spread of new infections. Infections that are spread directly from person to person (e.g. direct or by the route may be by such factors as living (e.g. crowding, and level of size and of the from infections or size of the (e.g. of sexual and number of of the among In for a pathogen to in a it have a basic it is than it will may if it is an infection, such as tuberculosis, which can persist in a form and can or after the infection. The physicochemical environment has direct and indirect on infectious diseases through its impact on pathogens, vectors and (Wilson changes can influence all of organisms and all of that a (e.g. or or intermediate may have a For be introduced into an are present that can part of the of the The presence of the of is but not for the transmission of the or with reach with other humans have contact with (e.g. in in for the to the human and and in the human The as more than a to carry a pathogen from one to The pathogen from one to and The period of time in the for the incubation is to for the of a in a is than the time period for the to in the transmission will not in that if a that is to that also affects time for from to it can affect the of species, such as Many areas are by vectors that are known to be to a infection may not The of presence of and and other factors may influence the contact between and humans, the of infection An or can be resistant to the and spread of a pathogen through several means – some are disease and are The may be of from infection (e.g. or immunization (e.g. measles, of and spread of many and infections (e.g. cholera, typhoid presence of and air and may vector-borne infections. The physicochemical environment may the of a pathogen or intermediate in a events (e.g. and other (e.g. can influence risk of infectious diseases through among them of human populations, of and and or of (Wilson of and medical care, and and may the impact and the spread of infections. travel to focus on the destination, but the of is The process of travel often the contact with of For the that will provide transport for most of the the traveller may have by and spent time in a with of people from often in with a environment (e.g. bus, train, with shared air The traveller shared Many outbreaks of infections to on include outbreaks to on the (e.g. among For infections with short incubation may the of a long as with an of on a from to the US for and and The provide for air during is during and is on in the range of and this may and to infections and In one influenza A was to of on a A person on the was in the of influenza, and the was not during a in In a different during a long on a a with multidrug-resistant infection to of persons and to of persons the of the infections that have spread on include smallpox and infections, spread by the route also have the potential to spread to new populations the The from of to may be and include many the A of of of were in in in the of 2000 2001). 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In the for million persons are Many have and in an with these regions. Travellers can also be as couriers who pick to microbial genetic material and carry it Travellers provide a of an area. from travellers may the of an or and other characterization that is through clinical and and This researchers to of a disease to characterization and the in some instances, its and route of The laboratory an part of the the and reach of global travel in be to agents where to that are found the world. is in the of the individual – and in when may be Many agents pose public health but such as Lassa or multidrug-resistant typhoid fever, have the potential for to or into the In those instances, pathogen identification and is key in of and for of the and also in if public health be and identification of a pathogen in a traveller can provide insights into is in other parts of the world. in may not have the capacity to and local of microbes by travellers to areas where are can populations in regions and help to inform a global This can help inform public health in other if is in a and if it those with a capacity to of resistance patterns among organisms such as falciparum can influence of of resistance of influence of for typhoid and affect the at which is travel. of the of meningitidis has of of and or may have as new become of laboratory can be in if the individual The important role of travellers in the spread of HIV has been Travellers can carry and microbes or microbial genetic material in the absence of Many infections in travellers are not or that in travellers only a of microbial of A person who a pathogen or resistance into a new may be and may be For travellers may carry bacteria, such as or that may carry virulence or resistance that can a new 2001). may be consequences may not be or In the an of caused by which after the had become infected in 1989). Infections that can remain for periods (e.g. or can in a form (e.g. may have consequences or after the time of which may have been HIV and can be from person to person. infections may be only after blood or A from with infection with were into several was the a of of disease in the in the US Neisseria meningitidis the of people but also can cause and and Most human infection is caused by and with and a time to time that are more or to spread more affect part of the which all or part of sub-Saharan of the by air by with other pathogens may also vulnerability to infection that A and have been used in Africa to In carried an of A meningitidis to 1989). with the and introduced it into sub-Saharan where it caused a of used to the spread of the to several other In a major in sub-Saharan Africa in more than of A infection were with a of of outbreaks among to the and their of for all Many in and Africa have used a and a is and has been used in the US and some other The have efficacy in the range of but this by and In that have the impact of on the of but not of the million persons from different come in for the have contact with persons from the world and then In the of 2000 meningitidis caused an of among from and their This was the time had been found in a were in were in other used and other to that from the and from had and were from found in infections in In were from from New on direct flights to were also on carried at the time of carried on In at least persons with disease had the of to the were in Africa the associated In between and researchers an in the of caused by and the of strains, which for of of in in the and have the of after to A and with the since 2000 have of infection in many in pilgrims, of pilgrims, and more in persons travel or contact with pilgrims, that this has been introduced and is causing sustained transmission in some areas and In the the of from the was than the mortality for other disease in and from to 2000 Among the from 2000 to were pilgrims, were of pilgrims, but had contact with the Dengue is caused by a by a that in to humans. Dengue is the most infection in the world and is and in and areas of dengue and humans. with one that but the individual for more disease if infected in the with a different The of disease, dengue haemorrhagic and dengue can be mortality can be by In a of dengue in of were in persons with a dengue infection. The of in with dengue infection who were infected with a different was Dengue viruses also to in virus as well as factors may influence disease humans, travellers, or carry the virus into new geographical which can lead to if a the new area. of the world today make many populations more to outbreaks of dengue and Travel has as The is more and the most rapid is in More urban areas in and areas have reached the size, estimated at to to the of dengue of a in areas where many have had infection increases the and mortality from infection. for have in many areas and vectors are resistant to also for and and used The global distribution of the vectors to a to populations in which viruses may also more of and human with more than one of virus This the for events that lead to the emergence of more or have that the number of dengue has been in with the size of the human over the of and of and air may also influence potential contact with the parts of and transmission of dengue in the US in has been transmission in has been of dengue have been the in In the major in and Travellers regularly and borders and and 2001). viruses, such as can also be carried in the world and then and in from persons and after travel to They that resistance in increased to and in with travel. This in persons who had A multidrug-resistant of is to have spread from to carried by one or more humans who were or infected A resistance may on a and be carried to where it may to A to have been spread on a resistant carried by travellers can also spread after in a factors influence the and distribution of infectious diseases. changes and are the and will to the for emerging infections, caused by previously or populations also the world are through movement of people and animals is a of microbial – and to – through a range of in to in an environment with and other and in a human that has been a of the microbes that have been changes and continuing in the between humans and This paper was the was a at the for in the for the by The and
Mary Wilson (2003) studied this question.