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In infectious disease ecology and
epidemiology Epidemiology is the study and analysis of the distribution (who, when, and where), patterns and Risk factor (epidemiology), determinants of health and disease conditions in a defined population, and application of this knowledge to prevent dise ...
, a natural reservoir, also known as a disease reservoir or a reservoir of infection, is the
population Population is a set of humans or other organisms in a given region or area. Governments conduct a census to quantify the resident population size within a given jurisdiction. The term is also applied to non-human animals, microorganisms, and pl ...
of organisms or the specific environment in which an infectious
pathogen In biology, a pathogen (, "suffering", "passion" and , "producer of"), in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a Germ theory of d ...
naturally lives and reproduces, or upon which the pathogen primarily depends for its survival. A reservoir is usually a living
host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County * Host Island, in the Wilhelm Archipelago, Antarctica People * ...
of a certain species, such as an animal or a plant, inside of which a pathogen survives, often (though not always) without causing disease for the reservoir itself. By some definitions a reservoir may also be an environment external to an organism, such as a volume of contaminated air or water. Because of the enormous variety of infectious
microorganism A microorganism, or microbe, is an organism of microscopic scale, microscopic size, which may exist in its unicellular organism, single-celled form or as a Colony (biology)#Microbial colonies, colony of cells. The possible existence of unseen ...
s capable of causing disease, precise definitions for what constitutes a natural reservoir are numerous, various, and often conflicting. The reservoir concept applies only for pathogens capable of infecting more than one host population and only with respect to a defined target population – the population of organisms in which the pathogen causes disease. The reservoir is any population of organisms (or any environment) which harbors the pathogen and transmits it to the target population. Reservoirs may comprise one or more different species, may be the same or a different species as the target, and, in the broadest sense, may include vector species, which are otherwise distinct from natural reservoirs. Significantly, species considered reservoirs for a given pathogen may not experience symptoms of disease when infected by the pathogen. Identifying the natural reservoirs of infectious pathogens has proven useful in treating and preventing large outbreaks of disease in humans and domestic animals, especially those diseases for which no
vaccine A vaccine is a biological Dosage form, preparation that provides active acquired immunity to a particular infectious disease, infectious or cancer, malignant disease. The safety and effectiveness of vaccines has been widely studied and verifi ...
exists. In principle, zoonotic diseases can be controlled by isolating or destroying the pathogen's reservoirs of infection. The mass
culling Culling is the process of segregating organisms from a group according to desired or undesired characteristics. In animal breeding, it is removing or segregating animals from a breeding stock based on a specific trait. This is done to exagge ...
of animals confirmed or suspected as reservoirs for human pathogens, such as birds that harbor avian influenza, has been effective at containing possible
epidemic An epidemic (from Greek ἐπί ''epi'' "upon or above" and δῆμος ''demos'' "people") is the rapid spread of disease to a large number of hosts in a given population within a short period of time. For example, in meningococcal infection ...
s in many parts of the world; for other pathogens, such as the
ebolavirus The genus ''Ebolavirus'' (- or ; - or ) is a International Committee on Taxonomy of Viruses, virological taxon included in the family ''Filoviridae'' (filament-shaped viruses), order ''Mononegavirales''. The members of this genus are called ebo ...
es, the identity of the presumed natural reservoir remains obscure.


Definition and terminology

The great diversity of infectious pathogens, their possible hosts, and the ways in which their hosts respond to infection has resulted in multiple definitions for "natural reservoir", many of which are conflicting or incomplete. In a 2002 conceptual exploration published in the
CDC The Centers for Disease Control and Prevention (CDC) is the national public health agency of the United States. It is a United States federal agency under the Department of Health and Human Services (HHS), and is headquartered in Atlanta, ...
's '' Emerging Infectious Diseases'', the natural reservoir of a given pathogen is defined as "one or more epidemiologically connected populations or environments in which the pathogen can be permanently maintained and from which infection is transmitted to the defined target population." The ''target population'' is the population or species in which the pathogen causes disease; it is the population of interest because it has disease when infected by the pathogen (for example, humans are the target population in most medical epidemiological studies). A common criterion in other definitions distinguishes reservoirs from non-reservoirs by the degree to which the infected host shows symptoms of disease. By these definitions, a reservoir is a host that does not experience the symptoms of disease when infected by the pathogen, whereas non-reservoirs show symptoms of the disease. The pathogen still feeds, grows, and reproduces inside a reservoir host, but otherwise does not significantly affect its health; the relationship between pathogen and reservoir is more or less
commensal Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit f ...
, whereas in susceptible hosts that do develop disease caused by the pathogen, the pathogen is considered parasitic. What further defines a reservoir for a specific pathogen is where it can be maintained and from where it can be transmitted. A "multi-host" organism is capable of having more than one natural reservoir.


Types of reservoirs

Natural reservoirs can be divided into three main types: human, animal (non-human), and environmental.


Human reservoirs

''Human reservoirs'' are human beings infected by pathogens that exist on or within the human body. Infections like
poliomyelitis Poliomyelitis ( ), commonly shortened to polio, is an infectious disease caused by the poliovirus. Approximately 75% of cases are asymptomatic; mild symptoms which can occur include sore throat and fever; in a proportion of cases more severe ...
and
smallpox Smallpox was an infectious disease caused by Variola virus (often called Smallpox virus), which belongs to the genus '' Orthopoxvirus''. The last naturally occurring case was diagnosed in October 1977, and the World Health Organization (W ...
, which exist exclusively within a human reservoir, are sometimes known as ''anthroponoses''. Humans can act as reservoirs for
sexually transmitted disease A sexually transmitted infection (STI), also referred to as a sexually transmitted disease (STD) and the older term venereal disease (VD), is an infection that is spread by sexual activity, especially vaginal intercourse, anal sex, or ...
s,
measles Measles (probably from Middle Dutch or Middle High German ''masel(e)'', meaning "blemish, blood blister") is a highly contagious, Vaccine-preventable diseases, vaccine-preventable infectious disease caused by Measles morbillivirus, measles v ...
,
mumps MUMPS ("Massachusetts General Hospital Utility Multi-Programming System"), or M, is an imperative, high-level programming language with an integrated transaction processing key–value database. It was originally developed at Massachusetts Gen ...
, streptococcal infection, various respiratory pathogens, and the smallpox virus.


Animal reservoirs

Animal (non-human) reservoirs consist of domesticated and wild animals infected by pathogens. For example, the bacterium ''
Vibrio cholerae ''Vibrio cholerae'' is a species of Gram-negative bacteria, Gram-negative, Facultative anaerobic organism, facultative anaerobe and Vibrio, comma-shaped bacteria. The bacteria naturally live in Brackish water, brackish or saltwater where they att ...
'', which causes
cholera Cholera () is an infection of the small intestine by some Strain (biology), strains of the Bacteria, bacterium ''Vibrio cholerae''. Symptoms may range from none, to mild, to severe. The classic symptom is large amounts of watery diarrhea last ...
in humans, has natural reservoirs in
copepod Copepods (; meaning 'oar-feet') are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (living in the water column), some are benthos, benthic (living on the sedimen ...
s,
zooplankton Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...
, and
shellfish Shellfish, in colloquial and fisheries usage, are exoskeleton-bearing Aquatic animal, aquatic invertebrates used as Human food, food, including various species of Mollusca, molluscs, crustaceans, and echinoderms. Although most kinds of shellfish ...
. Parasitic blood-flukes of the genus ''
Schistosoma ''Schistosoma'' is a genus of trematodes, commonly known as blood flukes. They are Parasitism, parasitic flatworms responsible for a highly significant group of infections in humans termed ''schistosomiasis'', which is considered by the World H ...
'', responsible for
schistosomiasis Schistosomiasis, also known as snail fever, bilharzia, and Katayama fever is a neglected tropical helminthiasis, disease caused by parasitism, parasitic Schistosoma, flatworms called schistosomes. It affects both humans and animals. It affects ...
, spend part of their lives inside freshwater
snail A snail is a shelled gastropod. The name is most often applied to land snails, terrestrial molluscs, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gas ...
s before completing their life cycles in vertebrate hosts. Viruses of the taxon ''
Ebolavirus The genus ''Ebolavirus'' (- or ; - or ) is a International Committee on Taxonomy of Viruses, virological taxon included in the family ''Filoviridae'' (filament-shaped viruses), order ''Mononegavirales''. The members of this genus are called ebo ...
'', which causes
Ebola virus disease Ebola, also known as Ebola virus disease (EVD) and Ebola hemorrhagic fever (EHF), is a viral hemorrhagic fever in humans and other primates, caused by ebolaviruses. Symptoms typically start anywhere between two days and three weeks after infe ...
, are thought to have a natural reservoir in bats or other animals exposed to the virus. Other zoonotic diseases that have been transmitted from animals to humans include:
rabies Rabies is a viral disease that causes encephalitis in humans and other mammals. It was historically referred to as hydrophobia ("fear of water") because its victims panic when offered liquids to drink. Early symptoms can include fever and abn ...
, blastomycosis, psittacosis, trichinosis, cat-scratch disease, histoplasmosis,
coccidioidomycosis Coccidioidomycosis (, ) is a mammalian mycosis, fungal disease caused by ''Coccidioides immitis'' or ''Coccidioides posadasii''. It is commonly known as cocci, Valley fever, California fever, desert rheumatism, or San Joaquin Valley fever. Cocci ...
, and
salmonella ''Salmonella'' is a genus of bacillus (shape), rod-shaped, (bacillus) Gram-negative bacteria of the family Enterobacteriaceae. The two known species of ''Salmonella'' are ''Salmonella enterica'' and ''Salmonella bongori''. ''S. enterica'' ...
. Common animal reservoirs include: bats, rodents, cows, pigs, sheep, swine, rabbits, raccoons, dogs, and other mammals.


Common animal reservoirs


= Bats

= Numerous zoonotic diseases have been traced back to bats. There are a couple of theories that serve as possible explanations as to why bats carry so many viruses. One proposed theory is that there exist so many bat-borne illnesses because there exist a large number of bat species and individuals. The second possibility is that something about bats' physiology makes them especially good reservoir hosts. Perhaps bats' "food choices, population structure, ability to fly, seasonal migration and daily movement patterns, torpor and hibernation, life span, and roosting behaviors" are responsible for making them especially suitable reservoir hosts.
Lyssavirus ''Lyssavirus'' (from the Greek ''lyssa'' "rage, fury, rabies" and the Latin '' vīrus'') is a genus of RNA viruses in the family ''Rhabdoviridae'', order '' Mononegavirales''. Mammals, including humans, can serve as natural hosts. The genus ...
es (including the
Rabies virus Rabies virus (''Lyssavirus rabies'') is a neurotropic virus that causes rabies in animals, including humans. It can cause violence, hydrophobia, and fever. Rabies transmission can also occur through the saliva of animals and less commonly throu ...
), Henipaviruses, Menangle and
Tioman Tioman Island () is , off the east coast of Peninsular Malaysia, more specifically off the east coast of Rompin District, Pahang, Malaysia. It is long and wide and has seven villages, the largest and most populous being Kampung Tekek on the ...
viruses, SARS-CoV-Like Viruses, and
Ebola Ebola, also known as Ebola virus disease (EVD) and Ebola hemorrhagic fever (EHF), is a viral hemorrhagic fever in humans and other primates, caused by ebolaviruses. Symptoms typically start anywhere between two days and three weeks after in ...
viruses have all been traced back to different species of bats. Fruit bats in particular serve as the reservoir host for Nipah virus (NiV).


= Rats

= Rats are known to be the reservoir hosts for several zoonotic diseases. Norway rats were found to be infested with the
Lyme disease Lyme disease, also known as Lyme borreliosis, is a tick-borne disease caused by species of ''Borrelia'' bacteria, Disease vector, transmitted by blood-feeding ticks in the genus ''Ixodes''. It is the most common disease spread by ticks in th ...
spirochetes. In Mexico rats are known carriers of ''
Trypanosoma cruzi ''Trypanosoma cruzi'' is a species of parasitic euglenoids. Among the protozoa, the trypanosomes characteristically bore tissue in another organism and feed on blood (primarily) and also lymph. This behaviour causes disease or the likelihood ...
'', which causes
Chagas disease Chagas disease, also known as American trypanosomiasis, is a tropical parasitic disease caused by ''Trypanosoma cruzi''. It is spread mostly by insects in the subfamily Triatominae, known as "kissing bugs". The symptoms change throughout the ...
.


= Mice

= White-footed mice ('' Peromyscus leucopus'') are one of the most important animal reservoirs for the Lyme disease spirochete (''
Borrelia burgdorferi ''Borrelia burgdorferi'' is a bacterial species of the spirochete class in the genus '' Borrelia'', and is one of the causative agents of Lyme disease in humans. Along with a few similar genospecies, some of which also cause Lyme disease, it m ...
''). Deer mice serve as reservoir hosts for Sin Nombre virus, which causes hantavirus pulmonary syndrome (HPS).


= Monkeys

= The
Zika virus Zika virus (ZIKV; pronounced or ) is a member of the virus family ''Flaviviridae''. It is spread by daytime-active ''Aedes'' mosquitoes, such as '' A. aegypti'' and '' A. albopictus''. Its name comes from the Ziika Forest of Uganda, where ...
originated from monkeys in Africa. In São José do Rio Preto and Belo Horizonte, Brazil the Zika virus has been found in dead monkeys. Genome sequencing has revealed the virus to be very similar to the type that infects humans.


Environmental reservoirs

Environmental reservoirs include living and non-living reservoirs that harbor infectious pathogens outside the bodies of animals. These reservoirs may exist on land (plants and soil), in water, or the air. Pathogens in these reservoirs are sometimes free-living. The bacteria ''
Legionella pneumophila ''Legionella pneumophila'', the primary causative agent for Legionnaires' disease, Legionnaire's disease, is an Aerobic organism, aerobic, pleomorphic, Flagellum, flagellated, non-spore-forming, Gram-negative bacteria, Gram-negative bacterium. ' ...
'', a
facultative intracellular parasite Intracellular parasites are microparasites that are capable of growing and reproducing inside the cells of a Host (biology), host. They are also called intracellular pathogens. Types There are two main types of intracellular parasites: Facultat ...
which causes
Legionnaires' disease Legionnaires' disease is a form of atypical pneumonia caused by any species of ''Legionella'' bacteria, quite often ''Legionella pneumophila''. Signs and symptoms include cough, shortness of breath, high fever, myalgia, muscle pains, and headach ...
, and ''
Vibrio cholerae ''Vibrio cholerae'' is a species of Gram-negative bacteria, Gram-negative, Facultative anaerobic organism, facultative anaerobe and Vibrio, comma-shaped bacteria. The bacteria naturally live in Brackish water, brackish or saltwater where they att ...
'', which causes
cholera Cholera () is an infection of the small intestine by some Strain (biology), strains of the Bacteria, bacterium ''Vibrio cholerae''. Symptoms may range from none, to mild, to severe. The classic symptom is large amounts of watery diarrhea last ...
, can both exist as free-living parasites in certain water sources as well as in invertebrate animal hosts.


Disease transmission

A disease reservoir acts as a transmission point between a pathogen and a susceptible host. Transmission can occur directly or indirectly.


Direct transmission

Direct transmission can occur from direct contact or direct
droplet A drop or droplet is a small column of liquid, bounded completely or almost completely by free surfaces. A drop may form when liquid accumulates at the end of a tube or other surface boundary, producing a hanging drop called a pendant drop. Dro ...
spread. Direct contact transmission between two people can happen through skin contact, kissing, and sexual contact. Humans serving as disease reservoirs can be symptomatic (showing illness) or asymptomatic (not showing illness), act as disease carriers, and often spread illness unknowingly. Human carriers commonly transmit disease because they do not realize they are infected and take no special precautions to prevent transmission. Symptomatic persons aware of their illness are not as likely to transmit infection because they take precautions to reduce possible transmission of the disease and/or seek out treatment to prevent the spread of the disease. Direct droplet spread is due to solid particles or liquid droplets suspended in the air for some time. Droplet spread is considered the transmission of the pathogen to a susceptible host within a meter of distance; said droplet spread can occur from coughing, sneezing, and/or just talking. * ''
Neisseria gonorrhoeae ''Neisseria gonorrhoeae'', also known as ''gonococcus'' (singular) or ''gonococci'' (plural), is a species of Gram-negative diplococci bacteria first isolated by Albert Ludwig Sigesmund Neisser, Albert Neisser in 1879. An obligate human pathog ...
'' (
Gonorrhea Gonorrhoea or gonorrhea, colloquially known as the clap, is a sexually transmitted infection (STI) caused by the bacterium ''Neisseria gonorrhoeae''. Infection may involve the genitals, mouth, or rectum. Gonorrhea is spread through sexual c ...
) is transmitted by sexual contact involving the penis, vagina, mouth, and anus through direct contact transmission. * '' Bordetella pertussis'' (
Pertussis Whooping cough ( or ), also known as pertussis or the 100-day cough, is a highly contagious, vaccine-preventable bacterial disease. Initial symptoms are usually similar to those of the common cold with a runny nose, fever, and mild cough, bu ...
) is transmitted by cough from the human reservoir to the susceptible host through direct droplet spread. *


Indirect transmission

Indirect transmission can occur by airborne transmission, by vehicles (including fomites), and by vectors. Airborne transmission is different from direct droplet spread as it is defined as disease transmission that takes place over a distance larger than a meter. Pathogens that can be transmitted through airborne sources are carried by particles such as dust or dried residue (referred to as droplet nuclei). Vehicles such as food, water, blood and fomites can act as passive transmission points between reservoirs and susceptible hosts. Fomites are inanimate objects (doorknobs, medical equipment, etc.) that become contaminated by a reservoir source or someone/something that is a carrier. A vehicle, like a reservoir, may also be a favorable environment for the growth of an infectious agent, as coming into contact with a vehicle leads to its transmission. Vector transmission occurs most often from insect bites from mosquitoes, flies, fleas, and ticks. There are two sub-categories of vectors: ''mechanical'' (an insect transmits the pathogen to a host without the insect itself being affected) and ''biological'' (reproduction of the pathogen occurs within the vector before the pathogen is transmitted to a host). To give a few examples, '' Morbillivirus'' (
measles Measles (probably from Middle Dutch or Middle High German ''masel(e)'', meaning "blemish, blood blister") is a highly contagious, Vaccine-preventable diseases, vaccine-preventable infectious disease caused by Measles morbillivirus, measles v ...
) is transmitted from an infected human host to a susceptible host as they are transmitted by respiration through airborne transmission. '' Campylobacter'' ( campylobacteriosis) is a common bacterial infection that is spread from human or non-human reservoirs by vehicles such as contaminated food and water. ''
Plasmodium falciparum ''Plasmodium falciparum'' is a Unicellular organism, unicellular protozoan parasite of humans and is the deadliest species of ''Plasmodium'' that causes malaria in humans. The parasite is transmitted through the bite of a female ''Anopheles'' mos ...
'' (
malaria Malaria is a Mosquito-borne disease, mosquito-borne infectious disease that affects vertebrates and ''Anopheles'' mosquitoes. Human malaria causes Signs and symptoms, symptoms that typically include fever, Fatigue (medical), fatigue, vomitin ...
) can be transmitted from an infected mosquito, an animal (non-human) reservoir, to a human host by biological vector transmission.


Implications for public health

LH Taylor found that 61% of all human pathogens are classified as zoonotic. Thus, the identification of the natural reservoirs of pathogens before zoonosis would be incredibly useful from a public health standpoint. Preventive measures can be taken to lessen the frequency of outbreaks, such as vaccinating the animal sources of disease or preventing contact with reservoir host animals. To predict and prevent future outbreaks of zoonotic diseases, the U.S. Agency for International Development started the Emerging Pandemic Threats initiative in 2009. In alliance with University of California-Davis, EcoHealth Alliance, Metabiota Inc.,
Smithsonian Institution The Smithsonian Institution ( ), or simply the Smithsonian, is a group of museums, Education center, education and Research institute, research centers, created by the Federal government of the United States, U.S. government "for the increase a ...
, and
Wildlife Conservation Society The Wildlife Conservation Society (WCS) is a global 501(c)(3) organization, 501(c)(3) non-governmental organization, headquartered at the Bronx Zoo in New York City, with a mission to save "wildlife and wild places across the globe". Founded in ...
with support from Columbia and Harvard universities, the members of the PREDICT project are focusing on the "detection and discovery of zoonotic diseases at the wildlife-human interface." There are numerous other organizations around the world experimenting with different methods to predict and identify reservoir hosts. Researchers at the University of Glasgow created a
machine learning Machine learning (ML) is a field of study in artificial intelligence concerned with the development and study of Computational statistics, statistical algorithms that can learn from data and generalise to unseen data, and thus perform Task ( ...
algorithm that is designed to use "viral genome sequences to predict the likely natural host for a broad spectrum of RNA viruses, the viral group that most often jumps from animals to humans."


See also

*
Fomite A fomite () or fomes () is any inanimate object that, when contaminated with or exposed to infectious agents (such as pathogenic bacteria, viruses or fungi), can transfer disease to a new host. Transfer of pathogens by fomites A fomite is any ...
*
Host (biology) In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasite, parasitic, a mutualism (biology), mutualistic, or a commensalism, commensalist ''guest'' (symbiont). The guest is typically provided with ...
* Refuge (ecology) *
Vector (epidemiology) In epidemiology, a disease vector is any living agent that carries and transmits an infectious pathogen such as a parasite or microbe, to another living organism. Agents regarded as vectors are mostly blood-sucking (Hematophagy, hematophagous) art ...
*
Zoonosis A zoonosis (; plural zoonoses) or zoonotic disease is an infectious disease of humans caused by a pathogen (an infectious agent, such as a virus, bacterium, parasite, fungi, or prion) that can jump from a non-human vertebrate to a human. When ...


References

{{Concepts in infectious disease Disease ecology Epidemiology