Rabies in animals
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Rabies Rabies is a viral disease that causes encephalitis in humans and other mammals. Early symptoms can include fever and tingling at the site of exposure. These symptoms are followed by one or more of the following symptoms: nausea, vomiting, ...
is a viral
zoonotic A zoonosis (; plural zoonoses) or zoonotic disease is an infectious disease of humans caused by a pathogen (an infectious agent, such as a bacterium, virus, parasite or prion) that has jumped from a non-human (usually a vertebrate) to a human. ...
neuroinvasive disease which causes inflammation in the brain and is usually fatal. Rabies, caused by the
rabies virus Rabies virus, scientific name ''Rabies lyssavirus'', is a neurotropic virus that causes rabies in humans and animals. Rabies transmission can occur through the saliva of animals and less commonly through contact with human saliva. ''Rabies lys ...
, primarily infects mammals. In the laboratory it has been found that birds can be infected, as well as cell cultures from birds, reptiles and insects. The brains of animals with rabies deteriorate. As a result, they tend to behave bizarrely and often aggressively, increasing the chances that they will bite another animal or a person and transmit the disease. Most cases of humans contracting the disease from infected animals are in developing nations. In 2010, an estimated 26,000 people died from rabies, down from 54,000 in 1990.


Stages of disease

Three stages of rabies are recognized in dogs and other animals. # The first stage is a one- to three-day period characterized by behavioral changes and is known as the prodromal stage. # The second stage is the excitative stage, which lasts three to four days. It is this stage that is often known as ''furious rabies'' due to the tendency of the affected animal to be hyperreactive to external stimuli and bite at anything near. # The third stage is the paralytic or dumb stage and is caused by damage to motor neurons. Incoordination is seen due to rear limb paralysis and drooling and difficulty swallowing is caused by paralysis of facial and throat muscles. This disables the host's ability to swallow, which causes saliva to pour from the mouth. This causes bites to be the most common way for the infection to spread, as the virus is most concentrated in the throat and cheeks, causing major contamination to saliva. Death is usually caused by respiratory arrest.


Mammals


Bats

Bat-transmitted rabies occurs throughout North and South America but it was first closely studied in
Trinidad Trinidad is the larger and more populous of the two major islands of Trinidad and Tobago. The island lies off the northeastern coast of Venezuela and sits on the continental shelf of South America. It is often referred to as the southernmos ...
in the
West Indies The West Indies is a subregion of North America, surrounded by the North Atlantic Ocean and the Caribbean Sea that includes 13 independent island countries and 18 dependencies and other territories in three major archipelagos: the Greate ...
. This island was experiencing a significant toll of livestock and humans alike to rabid bats. In the 10 years from 1925 and 1935, 89 people and thousands of livestock had died from it—"the highest human mortality from rabies-infected bats thus far recorded anywhere." In 1931, Dr.
Joseph Lennox Pawan Joseph is a common male given name, derived from the Hebrew Yosef (יוֹסֵף). "Joseph" is used, along with "Josef", mostly in English, French and partially German languages. This spelling is also found as a variant in the languages of the mo ...
of
Trinidad Trinidad is the larger and more populous of the two major islands of Trinidad and Tobago. The island lies off the northeastern coast of Venezuela and sits on the continental shelf of South America. It is often referred to as the southernmos ...
in the
West Indies The West Indies is a subregion of North America, surrounded by the North Atlantic Ocean and the Caribbean Sea that includes 13 independent island countries and 18 dependencies and other territories in three major archipelagos: the Greate ...
, a government bacteriologist, found
Negri bodies Negri bodies are eosinophilic, sharply outlined, pathognomonic inclusion bodies (2–10 μm in diameter) found in the cytoplasm of certain nerve cells containing the virus of rabies, especially in pyramidal cells within Ammon's horn of t ...
in the brain of a bat with unusual habits. In 1932, Dr. Pawan discovered that infected
vampire bats Vampire bats, species of the subfamily Desmodontinae, are leaf-nosed bats found in Central and South America. Their food source is blood of other animals, a dietary trait called hematophagy. Three extant bat species feed solely on blood: the com ...
could transmit rabies to humans and other animals. In 1934, the Trinidad and Tobago government began a program of eradicating vampire bats, while encouraging the screening off of livestock buildings and offering free vaccination programs for exposed livestock. After the opening of the Trinidad Regional Virus Laboratory in 1953, Arthur Greenhall demonstrated that at least eight species of bats in Trinidad had been infected with rabies; including the
common vampire bat The common vampire bat (''Desmodus rotundus'') is a small, leaf-nosed bat native to Latin America. It is one of three extant species of vampire bat, the other two being the hairy-legged and the white-winged vampire bats. The common vampire bat ...
, the rare white-winged vampire bat, as well as two abundant species of fruit bats: the
Seba's short-tailed bat Seba's short-tailed bat (''Carollia perspicillata'') is a common and widespread bat species in the family Phyllostomidae. They are found in Central America, the northern parts of South America, and in the Antilles islands. Description ''C. pe ...
and the
Jamaican fruit bat The Jamaican, common or Mexican fruit bat (''Artibeus jamaicensis'') is a fruit-eating bat native to Mexico, through Central America to northwestern South America, as well as the Greater and many of the Lesser Antilles. It is also an uncommon re ...
. Recent data sequencing suggests recombination events in an American bat led the modern rabies virus to gain the head of a G-protein
ectodomain An ectodomain is the domain of a membrane protein that extends into the extracellular space (the space outside a cell). Ectodomains are usually the parts of proteins that initiate contact with surfaces, which leads to signal transduction.A notable ...
thousands of years ago. This change occurred in an organism that had both rabies and a separate carnivore virus. The recombination resulted in a cross-over that gave rabies a new success rate across hosts since the G-protein ectodomain, which controls binding and pH receptors, was now suited for carnivore hosts as well.


Cats

In the United States, domestic cats are the most commonly reported rabid animal. In the United States, , between 200 and 300 cases are reported annually; in 2017, 276 cats with rabies were reported. , in every year since 1990, reported cases of rabies in cats outnumbered cases of rabies in dogs. Cats that have not been vaccinated and are allowed access to the outdoors have the most risk for contracting rabies, as they may come in contact with rabid animals. The virus is often passed on during fights between cats or other animals and is transmitted by bites, saliva or through mucous membranes and fresh wounds. The virus can incubate from one day up to over a year before any symptoms begin to show. Symptoms have a rapid onset and can include unusual aggression, restlessness, lethargy, anorexia, weakness, disorientation, paralysis and seizures. Vaccination of felines (including boosters) by a veterinarian is recommended to prevent rabies infection in outdoor cats.


Cattle

In cattle-raising areas where vampire bats are common, fenced-in cows often become a primary target for the bats (along with horses), due to their easy accessibility compared to wild mammals. In Latin America, vampire bats are the primary reservoir of the rabies virus, and in Peru, for instance, researchers have calculated that over 500 cattle per year die of bat-transmitted rabies. Vampire bats have been extinct in the United States for thousands of years (a situation that may reverse due to climate change, as the range of vampire bats in northern Mexico has recently been creeping northward with warmer weather), thus United States cattle are not currently susceptible to rabies from this vector. However, cases of rabies in dairy cows in the United States has occurred (perhaps transmitted by bites from
canines Canine may refer to: Zoology and anatomy * a dog-like Canid animal in the subfamily Caninae ** ''Canis'', a genus including dogs, wolves, coyotes, and jackals ** Dog, the domestic dog * Canine tooth, in mammalian oral anatomy People with the surn ...
), leading to concerns that humans consuming
unpasteurized Pasteurization or pasteurisation is a process of food preservation in which packaged and non-packaged foods (such as milk and fruit juices) are treated with mild heat, usually to less than , to eliminate pathogens and extend shelf life. Th ...
dairy products from these cows could be exposed to the virus. Vaccination programs in Latin America have been effective at protecting cattle from rabies, along with other approaches such as the culling of vampire bat populations.


Coyotes

Rabies is common in coyotes, and can be a cause for concern if they interact with humans.


Dogs

Rabies has a long history of association with
dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is derived from the extinct Pleistocene wolf, and the modern wolf is the dog's nearest living relative. Do ...
s. The first written record of rabies is in the Codex of Eshnunna (), which dictates that the owner of a dog showing symptoms of rabies should take preventive measure against bites. If a person was bitten by a rabid dog and later died, the owner was fined heavily. Almost all of the human deaths attributed to rabies are due to rabies transmitted by dogs in countries where dog vaccination programs are not sufficiently developed to stop the spread of the virus.


Horses

Rabies can be contracted in horses if they interact with rabid animals in their pasture, usually through being bitten (e.g. by vampire bats) on the muzzle or lower limbs. Signs include aggression, incoordination, head-pressing, circling, lameness, muscle tremors, convulsions, colic and fever. Horses that experience the paralytic form of rabies have difficulty swallowing, and drooping of the lower jaw due to paralysis of the throat and jaw muscles. Incubation of the virus may range from 2–9 weeks. Death often occurs within 4–5 days of infection of the virus. There are no effective treatments for rabies in horses. Veterinarians recommend an initial vaccination as a foal at three months of age, repeated at one year and given an annual booster.


Monkeys

Monkeys, like humans, can get rabies; however, they do not tend to be a common source of rabies. Monkeys with rabies tend to die more quickly than humans. In one study, 9 of 10 monkeys developed severe symptoms or died within 20 days of infection. Rabies is often a concern for individuals travelling to developing countries as monkeys are the most common source of rabies after dogs in these places.


Rabbits

Despite natural infection of rabbits being rare, they are particularly vulnerable to the rabies virus; rabbits were used to develop the first rabies vaccine by Louis Pasteur in the 1880s, and continue to be used for rabies diagnostic testing. The virus is often contracted when attacked by other rabid animals and can incubate within a rabbit for up to 2–3 weeks. Symptoms include weakness in limbs, head tremors, low appetite, nasal discharge, and death within 3–4 days. There are currently no vaccines available for rabbits. The
National Institutes of Health The National Institutes of Health, commonly referred to as NIH (with each letter pronounced individually), is the primary agency of the United States government responsible for biomedical and public health research. It was founded in the late ...
recommends that rabbits be kept indoors or enclosed in hutches outside that do not allow other animals to come in contact with them.


Red Pandas

Although rare, cases of rabies in red pandas have been recorded.


Skunks

In the United States, there is currently no
USDA The United States Department of Agriculture (USDA) is the federal executive department responsible for developing and executing federal laws related to farming, forestry, rural economic development, and food. It aims to meet the needs of com ...
-approved vaccine for the strain of rabies that afflicts skunks. When cases are reported of pet skunks biting a human, the animals are frequently killed in order to be tested for rabies. It has been reported that three different variants of rabies exist in striped skunks in the north and south central states. Humans exposed to the rabies virus must begin post-exposure prophylaxis before the disease can progress to the central nervous system. For this reason, it is necessary to determine whether the animal, in fact, has rabies as quickly as possible. Without a definitive quarantine period in place for skunks, quarantining the animals is not advised as there is no way of knowing how long it may take the animal to show symptoms. Destruction of the skunk is recommended and the brain is then tested for presence of rabies virus. Skunk owners have recently organized to campaign for USDA approval of both a vaccine and an officially recommended quarantine period for skunks in the United States.


Wolves

Under normal circumstances, wild
wolves The wolf (''Canis lupus''; : wolves), also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, and gray wolves, as popularly un ...
are generally timid around humans, though there are several reported circumstances in which wolves have been recorded to act aggressively toward humans. The majority of fatal wolf attacks have historically involved rabies, which was first recorded in wolves in the 13th century. The earliest recorded case of an actual rabid wolf attack comes from Germany in 1557. Though wolves are not reservoirs for the disease, they can catch it from other species. Wolves develop an exceptionally severe aggressive state when infected and can bite numerous people in a single attack. Before a vaccine was developed, bites were almost always fatal. Today, wolf bites can be treated, but the severity of rabid wolf attacks can sometimes result in outright death, or a bite near the head will make the disease act too fast for the treatment to take effect. Rabid attacks tend to cluster in winter and spring. With the reduction of rabies in Europe and North America, few rabid wolf attacks have been recorded, though some still occur annually in the Middle East. Rabid attacks can be distinguished from predatory attacks by the fact that rabid wolves limit themselves to biting their victims rather than consuming them. Plus, the timespan of predatory attacks can sometimes last for months or years, as opposed to rabid attacks which end usually after a fortnight. Victims of rabid wolves are usually attacked around the head and neck in a sustained manner.


Other placental mammals

The most commonly infected terrestrial animals in the United States are
raccoon The raccoon ( or , ''Procyon lotor''), sometimes called the common raccoon to distinguish it from other species, is a mammal native to North America. It is the largest of the procyonid family, having a body length of , and a body weight of ...
s, skunks,
fox Foxes are small to medium-sized, omnivorous mammals belonging to several genera of the family Canidae. They have a flattened skull, upright, triangular ears, a pointed, slightly upturned snout, and a long bushy tail (or ''brush''). Twelve sp ...
es, and coyotes. Any bites by such wild animals must be considered a possible exposure to the rabies virus. Most cases of rabies in rodents reported to the
Centers for Disease Control and Prevention 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, and is headquartered in Atlanta, Georgi ...
in the United States have been found among
groundhog The groundhog (''Marmota monax''), also known as a woodchuck, is a rodent of the family Sciuridae, belonging to the group of large ground squirrels known as marmots. The groundhog is a lowland creature of North America; it is found through mu ...
s (woodchucks). Small rodents such as
squirrels Squirrels are members of the family Sciuridae, a family that includes small or medium-size rodents. The squirrel family includes tree squirrels, ground squirrels (including chipmunks and prairie dogs, among others), and flying squirrels. Squ ...
,
hamsters Hamsters are rodents (order Rodentia) belonging to the subfamily Cricetinae, which contains 19 species classified in seven genera.Fox, Sue. 2006. ''Hamsters''. T.F.H. Publications Inc. They have become established as popular small pets. The ...
,
guinea pigs The guinea pig or domestic guinea pig (''Cavia porcellus''), also known as the cavy or domestic cavy (), is a species of rodent belonging to the genus '' Cavia'' in the family Caviidae. Breeders tend to use the word ''cavy'' to describe the a ...
,
gerbils Gerbillinae is one of the subfamilies of the rodent family Muridae and includes the gerbils, jirds, and sand rats. Once known as desert rats, the subfamily includes about 110 species of African, Indian, and Asian rodents, including sand rats an ...
,
chipmunks Chipmunks are small, striped rodents of the family Sciuridae. Chipmunks are found in North America, with the exception of the Siberian chipmunk which is found primarily in Asia. Taxonomy and systematics Chipmunks may be classified either as ...
,
rats Rats are various medium-sized, long-tailed rodents. Species of rats are found throughout the order Rodentia, but stereotypical rats are found in the genus ''Rattus''. Other rat genera include ''Neotoma'' ( pack rats), ''Bandicota'' (bandicoot ...
, mice, and lagomorphs like rabbits and
hares Hares and jackrabbits are mammals belonging to the genus ''Lepus''. They are herbivores, and live solitarily or in pairs. They nest in slight depressions called forms, and their young are able to fend for themselves shortly after birth. The gen ...
are almost never found to be infected with rabies, and are not known to transmit rabies to humans.


Marsupial and monotreme mammals

The
Virginia opossum The Virginia opossum (''Didelphis virginiana''), also known as the North American opossum, is the only opossum living north of Mexico, its range extending south into Central America. It is the northernmost marsupial in the world. In the United S ...
(a marsupial, unlike the other mammals named above, which are all eutherians/
placental Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distinguishe ...
), has a lower internal body temperature than the rabies virus prefers and therefore is resistant but not immune to rabies. Marsupials, along with
monotremes Monotremes () are prototherian mammals of the order Monotremata. They are one of the three groups of living mammals, along with placentals (Eutheria), and marsupials ( Metatheria). Monotremes are typified by structural differences in their br ...
(
platypuses The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal endemic to eastern Australia, including Tasmania. The platypus is the sole living representative or monotyp ...
and
echidnas Echidnas (), sometimes known as spiny anteaters, are quill-covered monotremes (egg-laying mammals) belonging to the family Tachyglossidae . The four extant species of echidnas and the platypus are the only living mammals that lay eggs and the ...
), typically have lower body temperatures than similarly sized eutherians.


Birds

Bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s were first artificially infected with rabies in 1884; however, infected birds are largely, if not wholly, asymptomatic, and recover. Other bird species have been known to develop rabies antibodies, a sign of infection, after feeding on rabies-infected mammals.


Transport of pet animals between countries

Rabies is
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...
to many parts of the world, and one of the reasons given for
quarantine A quarantine is a restriction on the movement of people, animals and goods which is intended to prevent the spread of disease or pests. It is often used in connection to disease and illness, preventing the movement of those who may have been ...
periods in international animal transport has been to try to keep the disease out of uninfected regions. However, most developed countries, pioneered by Sweden, now allow unencumbered travel between their territories for pet animals that have demonstrated an adequate immune response to rabies vaccination. Such countries may limit movement to animals from countries where rabies is considered to be under control in pet animals. There are various lists of such countries. The United Kingdom has developed a list, and France has a rather different list, said to be based on a list of the ''Office International des Epizooties'' (OIE). The European Union has a harmonised list. No list of rabies-free countries is readily available from OIE. In recent years, canine rabies has been practically eliminated in North America and
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located entirel ...
due to extensive and often mandatory
vaccination Vaccination is the administration of a vaccine to help the immune system develop immunity from a disease. Vaccines contain a microorganism or virus in a weakened, live or killed state, or proteins or toxins from the organism. In stimulating ...
requirements. However it is still a significant problem in parts of
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
, parts of the
Middle East The Middle East ( ar, الشرق الأوسط, ISO 233: ) is a geopolitical region commonly encompassing Arabia (including the Arabian Peninsula and Bahrain), Asia Minor (Asian part of Turkey except Hatay Province), East Thrace (Europ ...
, parts of
Latin America Latin America or * french: Amérique Latine, link=no * ht, Amerik Latin, link=no * pt, América Latina, link=no, name=a, sometimes referred to as LatAm is a large cultural region in the Americas where Romance languages — languages derived f ...
, and parts of
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
. Dogs are considered to be the main
reservoir A reservoir (; from French ''réservoir'' ) is an enlarged lake behind a dam. Such a dam may be either artificial, built to store fresh water or it may be a natural formation. Reservoirs can be created in a number of ways, including contro ...
for rabies in
developing countries A developing country is a sovereign state with a lesser developed industrial base and a lower Human Development Index (HDI) relative to other countries. However, this definition is not universally agreed upon. There is also no clear agreem ...
. However, the recent spread of rabies in the northeastern United States and further may cause a restrengthening of precautions against movement of possibly rabid animals between developed countries.


See also

* Prevalence of rabies * Rabies transmission *
Rabies vaccine The rabies vaccine is a vaccine used to prevent rabies. There are a number of rabies vaccines available that are both safe and effective. They can be used to prevent rabies before, and, for a period of time, after exposure to the rabies virus, ...


Footnotes


References

* Baynard, Ashley C. ''et al.'' (2011). "Bats and Lyssaviruses." In: ''Advances in VIRUS RESEARCH VOLUME 79. Research Advances in Rabies''. Edited by Alan C. Jackson. Elsevier. . * Goodwin G. G., and A. M. Greenhall. 1961. "A review of the bats of
Trinidad and Tobago Trinidad and Tobago (, ), officially the Republic of Trinidad and Tobago, is the southernmost island country in the Caribbean. Consisting of the main islands Trinidad and Tobago, and numerous much smaller islands, it is situated south of ...
." ''Bulletin of the American Museum of Natural History'', 122. *
Joseph Lennox Pawan Joseph is a common male given name, derived from the Hebrew Yosef (יוֹסֵף). "Joseph" is used, along with "Josef", mostly in English, French and partially German languages. This spelling is also found as a variant in the languages of the mo ...
(1936). "Transmission of the Paralytic Rabies in Trinidad of the Vampire Bat: ''Desmodus rotundus murinus'' Wagner, 1840." ''Annual Tropical Medicine and Parasitol'', 30, April 8, 1936, pp. 137–156. {{DEFAULTSORT:Rabies In Animals in animals