B. What Animals Are Not Susceptible To Infection With The Rabies Virus?
Vet Clin North Am Modest Anim Pract. Author manuscript; available in PMC 2009 Jul 1.
Published in last edited form as:
PMCID: PMC2518964
NIHMSID: NIHMS55023
Rabies in small animals
Introduction
Rabies is an aboriginal disease and its history tin can be traced dorsum more than than 5000 years ago24. Despite significant scientific progress, rabies remains an important global disease. Annually, more than 55,000 human being fatalities are reported, and millions of others require mail-exposure treatment38 , 39. Most of the human cases occur in the developing nations of Asia and Africa, where dog rabies remains owned or epizootic and thus is the main source for human exposure24. In developed countries, human rabies has dramatically declined during the past threescore years as a direct effect of routine vaccination of pet animals.
Rabies in the United States
In the United States, rabies was owned in small-scale animals, especially in dogs, and thus was a major public health problem in the beginning of the last century. Nigh x,000 rabies cases were reported annually in dogs and cats43. Massive immunizations in domestic dogs and cats were initiated in the 1940s and 1950s. As a outcome, rabies in dogs and cats declined dramatically and now only a few hundred cases are reported each year43 (Fig one). The rabies virus strains that used to be associated with dogs take disappeared during the concluding few years13. Viruses associated with small animals are derived from strains affecting wild fauna animals.
Currently wildlife rabies is enzootic in the United States. Seven to 8 thousand cases have been reported in wildlife annually during the past ii decadesviii , ix. Meantime there are a few rabies enzootics occurring in the U.s.a.. The distribution of the terrestrial animal rabies epizootics is shown in Fig. 2 45. Raccoon rabies has been spreading during the 1980s and 1990s along the eastern seaboard17 and spreading west in the new century7. Three different variants exist in striped skunks in longstanding reservoirs in California, the north key states, and the due south cardinal states16. Now skunks have been reported to be infected with raccoon and bat rabies variants in other states26 , 33. There are at least three fox rabies enzootics: arctic foxes in Alaska, forth with ruby and grayness foxes in the southeastfive , 12. Some of these terrestrial wildlife species may accept acquired rabies virus from dogs a long fourth dimension ago and have adapted to their ain species and locationseighteen. Others may have evolved from bat rabies variants33. Spillover from i species to some other occurs from time to fourth dimension25 , 33 and may lead to spreading in the new species. The distribution of these terrestrial rabies epizootics are depicted in Fig. 2A and the phylogenetic relationship of these rabies variants in the The states is summarized in Fig. 2B 45. In addition to terrestrial animal rabies, bat rabies has been detected in all the 48 face-to-face states and has been responsible for virtually of the human cases in the United States for the past 20 years8.
(A) Geographic distribution of the major terrestrial carnivore hosts of rabies virus variants. Each region is largely characterized by a unique rabies variant specific to a single carnivore host. (B) Neighbour-joining tree for nucleotide sequence of a 320-bp region of the nucleoprotein gene of selected RABV isolates from the United States, Mexico, and Canada. Each group of virus isolates that was sequenced to illustrate the unique RABV variants associated with terrestrial carnivores is boxed. The Polar Play a joke on variant (Artic and reddish fox) is no longer considered enzootic in the United States. Bootstrap values are shown at the branching indicate for clades recovered in .700/1000 iterations of the information. ABLV was used as the outgroup and to root the tree. Samples from a rabid trick in Ontario, Canada (CN OT FX 2/4), and from ii human rabies cases with exposures to rabid dogs in Mexico (MX/TX HM 1976 and 1979) are included to show variants of RABV shared across international boundaries. U.Due south. samples are identified by a two-letter of the alphabet abbreviation for the state and animal from which the sample originated, followed by the year the case occurred. With the exception of the Canadian sample (GenBank accretion {"blazon":"entrez-nucleotide","attrs":{"text":"U11735","term_id":"511234","term_text":"U11735"}}U11735), all RABV sequences were derived from samples in a virus repository at CDC. (Taken from Real et al45)
Wild animals rabies presents a health trouble to domestic pocket-size animals, which in turn accept a higher adventure of manual to humans due to their shut contact with people. Rabies variants found in domestic animals include variants establish in raccoons, due north cardinal skunks, south central skunks, Texas foxes, Texas canis familiaris-coyotes and California skunks35.
Rabies in Dogs
Dogs are the natural host for rabies. At that place are two forms of rabies—the excitatory or "furious" form, and the paralytic or "impaired" gradethree , 28. At that place are several overlapping phases during the progression of the illness: the prodromal menstruation, the furious period, and the paralytic periodthree , 28. Withal the clinical signs of rabies may vary among animals. The first stage lasts 2 to iii days in dogs. During this phase, infected animals always show different behavior. The excitement phase may last up to a week, simply sometimes animals progress direct from the prodromal phase to the paralytic stage. In the second flow, animals suddenly become barbarous and behave erratic. Within several days, the illness progresses to the paralytic period. In the last menses, animals show paralysis kickoff in the wound limb, and then in the neck and head. Affliction in animals ends in respiratory failure and death3 , 28. The grade of rabies typically lasts 3 to viii days in dogs.
Recently a report described rabies symptoms in a 6 month quondam mixed breed female person dog in Florida, which provides valuable insight into clinical presentation of rabies meningoencephalomyelitis. At presentation, the domestic dog has a three day history of acute paraplegia, including areflexia, hyperesthesia, and non-painful swelling of the left 2nd and 3rd digits of the affected limb, somewhen progressing to flaccid paralysis of the correct pelvic limb. Analysis of lumbar CSF showed abnormally high protein, RBC, and WBC counts. Cytopathology revealed 78% lymphocytes, 21% mononuclear phagocytes, and ane% neutrophils. Serum testing for rabies neutralization antibodies using RFFIT assay was negative. EMG of left pelvic limb revealed moderate fibrillations and positive abrupt waves suggestive of denervation/myopathy. No Thousand wave could be generated for the left sciatic nerve, indicating a lack of axonal or neuromuscular transmission. F waves were also absent on the left sciatic, tibial and ulnar nerves. Results for the correct limb, paravertebral muscles and thoracic limb muscles, and right sciatic, tibial and ulnar nerves were normal. Dementia, salivation and development of bilateral ventrolateral strabismus, focal and facial limb seizure and aggression occurred upon recovery from anesthesia. After euthanasia, the beast tested positive for raccoon rabies. Intracytoplasmic inclusion bodies could exist seen in the encephalon stem and spinal cord. Degenerate and necrotic neurons were seen within thoracic and lumbar spinal cord6.
It is interesting to note that at that place have been cases of cerebral cysticercosis caused past the larval Taenia solium which mimics rabies virus infection in dogs51. Additionally, there have been cases of cutaneous vasculitis associated with rabies vaccine administration in dogs, all with a like inflammatory pattern of mononuclear cells (nonleukocytoclstic)40.
Rabies in Cats
Cats are the domestic animals most frequently reported rabid in the Us, and 200 to 300 cases are reported annually31. In ane report in Pennsylvania, 44% of human being mail exposure prophylaxis (PEP) were due to exposure to a potentially rabid cat25. Factors influencing the increased incidence of rabies in cats include community tolerance of free ranging felines, also as less frequent rabies vaccination due to more lenient country laws for cats equally compared to dogs. Additionally, communities of feral cats exist, and people who treat these feral animals are at run a risk for coming into contact with rabies virus. Cats are predominantly affected by the variant of rabies virus endemic to the region in which they reside. For example, forth the North American East declension, cats are ordinarily infected with raccoon rabies virus variant. Withal, cats can also contract bat rabies virus variants, as both cats and bats are nocturnal, and cats trap minor animals such as bats27. Rabid cats display symptoms similar to those in dogs, simply have a trend to hide in secluded places and are often more barbarous. Like to recommendations for dogs, it is a common recommendation to confine and find a cat involved in a human bite, in club to rule out rabies exposure27 , 53.
Rabies in other small animals
In addition to dogs and cats, rabies has been reported in other domestic small animals such equally ferrets and rabbits. Two species of ferrets are mutual in the United states of america, the common ferret (Mustela putorius) and the black-footed ferret (Mustela nigripes). Ferrets have become popular companion animals in the United States. Ferrets were originally used to hunt small-scale game and suckling animals, and may be attracted by the olfactory property of milk49. Although rare, rabid pet ferrets take been reported in the United States13. Therefore, it has become increasingly of import to be enlightened of clinical signs of rabies in domestic ferrets, to avoid potentially harmful interactions with their human being owners. Clinical signs of paralytic rabies in ferrets included lethargy, ataxia, paresis, paraperesis, paralysis, bladder atony, constipation, hypothermia, inappetance and anorexia, abnormal or frequent vocalization, sneezing, parasthesia, and ptyalism (moist or disordered fur around the mouth). But about ten% of rabid ferrets in experimental infection showed aggressive beliefs with rapid attack and destruction towards a paper applicator, most had no to mild interest in the applicator. It has been recommended to vaccinate all pet ferrets confronting rabies, and to consider rabies in the differential diagnosis of ferrets with acute personality modify and/or paralysis41 , 42.
Rabies cases take too been reported in both rodents and lagomorphs, including a rabid pet guinea pig in 2003 which chip its possessor in the clavicle. The guinea squealer was later institute to exist infected with raccoon rabies virus. Between 1991 and 2001, the Wadsworth Center Rabies Lab received 7 lagomorphs, all pet domestics, three of which were exposed to a raccoon and i to a skunk. All 7 lagomorphs were infected with raccoon rabies virus23. Still, rodents and lagomorphs should exist considered "spillover" species, non reservoirs. Unfortunately often times no clinical signs are obvious in rabies-infected rodents. In 1972, a written report on rabid squirrels showed that ½ of infected animals that died of rabies showed no clinical signs23 , 57.
Ii cases of rabies in domestic rabbits (Oryctolagus cuniculus) in Maryland in 1999 are worth note hither. In both of these cases rabbits were sent home with owners afterwards examination, and owners were instructed to hand or forcefulness feed rabbits which later died and were found to be rabid. Clinical signs of illness in these rabbits upon test included weakness in forelimbs, palpable subcutaneous crepitus, slight intermittent caput tremors, ear infection, nasal discharge, and anorexia. Upon readmission one rabbit exhibited heavy wheezing, inability to stand, caput tilt, and bilateral conjunctivitis. The disease class culminated in a recumbent and non-responsive state. The case history of this rabbit included an attack by a raccoon in a rabies owned area resulting in a wound to the ear and the rabbit being covered in saliva29. Information technology is critical that rabies be considered in the differential diagnosis of any rabbit coming into contact with raccoons, peculiarly those displaying neurological signs. Furthermore, discharging an animal that has been exposed to potentially rabid wild animals should exist avoided, equally should recommending owners to forcefulness feed these animals, bringing them in closer contact to a potentially rabid pet.
Despite natural infection of rabbits being rare, it is imperative to remember that rabbits are used for rabies diagnostic testing, and were used for creation of the fist rabies vaccine by Louis Pasteur in the 1880s. Rabbits are highly susceptible to rabies virus infection, have incubation periods between 2 and 3 weeks later intracerebral inoculation, and normally develop paralytic rabies. Experimentally infected rabbits brandish anorexia, fever, restlessness, weight loss, and neurological signs such as teeth grinding, caput tremors, poor coordination of the hind limbs, and ascending paralysis. The afflicted rabbit usually dies inside iii–iv days29. Veterinarians should advise patients that no rabies vaccine is available for rabbits, and then prevention is essential. Rabbits should be kept indoors or kept in elevated hutches without exposed wire mesh floors, and rabbits should exist supervised at all times when exercising outdoors29.
Laboratory diagnosis for animal rabies
Clinical signs are good indication for rabies in minor animals. Still rapid and accurate laboratory diagnosis for animal rabies is of import for confirmation. In addition, many animals may not testify typical signs of rabies. Normally rabid or suspected rabid wild fauna are road kill or otherwise deceased when brought into diagnostic laboratories. Laboratory diagnosis is very important considering it provides not only data for epidemiological investigation of animal rabies, but also guidance for initiation of PEP in affected people14.
Direct florescent antibiotic assay (dFA)
The most oftentimes used method for rabies diagnosis in the laboratory is dFAiv xx , 21. Usually, brain smears or encephalon imprints from rabid or suspected rabid animals are reacted with fluorescein isothiocyanate (FITC)-conjugated-anti-rabies N antibodies4 , 55. When observed under a fluorescent microscope, the green-fluorescent foci will show the rabies virus antigen (Fig. 3A). dFA is rapid, economic, and sensitive for laboratory diagnosis of animal rabies. Rabies antigens tin be detected by the specific antibiotic; however, they should be differentiated from the not-specific groundwork.
Detection of rabies virus antigens past dFA (A) and immunohistochemistry (B). The virus foci prove positive stains with green-fluorescent color (A). Courtesy of http://world wide web.cdc.gov/rabies/diagnosis.html. The paraffin embedded slide was stained past anti-RV nucleoprotein monoclonal antibody 802-2. Rabies antigens in the cytoplasm and inclusions are shown in brown (using diaminobenzidine as the substance) and the cell nuclei are shown in bluish (B).
Direct rapid immunohistochemistry exam (dRIT)
Recently, the CDC developed the dRIT32, which is similar to dFA. Brain smears or imprints on glass slides are stock-still with ten% buffered formalin32. According to standard immunohistochemical staining, the virus antigen tin be detected by anti-rabies N monoclonal antibody and examined under a lite microscope. The sensitivity and specificity of dRIT is equivalent to that of the dFA32.
Virus Isolation
Mouse inoculation is a globe health organization (WHO) recommended method to confirm dFA when the issue is negativethirty 56. Commonly, brain suspension or spinal fluid from rabid or suspected rabid animals is intracerebrally inoculated into mouse brain. Two mice are killed every 2 days post infection until day 20 and brain smears are subjected to dFA. The 50% mouse intracerebral lethal dose (MICLD50) tin be calculated46. Virus isolation can besides be performed in cell civilisation, commonly on neuroblastoma cells59. The 50% tissue civilization infective dose (TCID50) can be calculated46. Jail cell civilisation inoculation is as sensitive as the mouse inoculation test47 and it requires less fourth dimension to obtain results.
Reverse transcriptase Polymerase chain reaction (RT-PCR)
RT-PCR is a newly developed method for rabies diagnosis4 , l. RT-PCR is very useful when sample size is small such as saliva and spinal fluid. Viral RNA is amplified by RT-PCR with primers usually designed from the N gene, the most conserved gene in rabies virus. RT-PCR for rabies diagnosis is as rapid as dFA and is every bit sensitive equally mouse inoculation test34. RT-PCR is also widely used in epidemiological investigation and outbreak studies. When combined with sequencing, this method can besides be used to differentiate rabies virus variants from multiple species of animals19 , 35 , 50 , 54. Viral variants tin also be differentiated with dissimilar monoclonal antibodies in an indirect fluorescent antibody assay22 , 35.
Histopathology and Immunohistochemistry
Rabies diagnosis in small animals tin besides be performed on brain tissues by histopathology and immunohistochemistry44. Histopathology may prove lymphocytic inflammation, perivascular cuffing, gliosis, and neurodegeneration52. Inflammation is diffuse in neuraxis. The parenchymal glial response is at first microglial just later mixed with astrocytes. Neurondegeneration is often not severe48. The severity of inflammation may vary between animal species. Sometimes, a spongiform encephalopathy with vacuolation in the gray matter can be observed15. Negri bodies which are ovoid eosinophilic intracytoplasmic inclusions11 , 58 are a hallmark for rabies diagnosis. Nonetheless Negri bodies are not plant in all rabies cases52.
In fixed encephalon tissue, immunohistochemistry tin can be used to confirm diagnosis (Fig. 3B). By using the rabies specific antibody and avidin-biotin colorimetric detection arrangement, the virus tin be detected. Antigen-positive neurons can be establish in the brain and spinal cord.
Detection of rabies virus-specific antibodies
Detection of specific antibodies can be used as diagnostic tools for rabies. In that location are many methods that have been adult to observe rabies-specific antibodies. Rapid fluorescent focus inhibition test (RFFIT) is the method used most ofttimes to detect virus neutralizing antibodies4 , x. ELISA has besides been used to detect virus-specific antibodies when the ELISA plate is coated with rabies virus antigens36 , 37. Since antibodies have several days to develop, this method is rarely used in diagnosis of animal rabies. Rather detection of virus-specific antibodies is often used in vaccination studies.
Rabies control in domestic pocket-sized animals
Rabies control in small animals is by routine immunization with inactivated rabies virus vaccines, which have been approved for dogs, cats, and ferrets. Offset vaccination is carried out at iii months of age and followed past a booster i yr later. Subsequent immunization is carried out annually or triennially depending on the type of vaccines usedi. Recently a recombinant canarypox vaccine is licensed for cats with a similar immunization schedule2. Currently, it is required by police force that dogs and cats be vaccinated against rabies.
Conclusion
Rabies in small-scale animals has been dramatically reduced in the Us since the introduction of rabies vaccination of domestic animals in the 1940s. As a consequence, the number of man rabies cases has declined to but a couple per yr. During the last several years, dog rabies variant has almost disappeared completely. Nonetheless, rabies in wildlife has skyrocketed. At the present, there are many concurrent rabies epizootics in wild fauna in the The states: raccoon rabies along the eastern seaboard, skunk rabies in the central states and California, artic fox rabies in Alaska, and red and gray fox rabies in the southwestern states. In addition, bat rabies is endemic in the 48 face-to-face states. Each wild animals species carries its own rabies variant(south). These wild fauna epizootics present a constant public health threat as well as the danger of reintroducing rabies to domestic animals. Vaccination is the key to prevent both rabies in small animals and rabies transmission to humans.
Footnotes
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