Rabies virus maintained by dogs in humans and terrestrial wildlife, Ceara State, Brazil.Rabies viruses circulating in Ceara, Brazil, were identified by molecular analysis to be related to variants maintained by dogs, bats, and other wildlife. Most of these viruses are associated with human rabies rabies (rā`bēz, ră`–) or hydrophobia (hī'drəfō`bēə), acute viral infection of the central nervous system in dogs, foxes, raccoons, skunks, bats, and other animals, and in cases. We document the emergence of a rabies virus variant responsible for an independent epidemic cycle in the crab-eating fox Noun 1. crab-eating fox - wild dog of northern South America crab-eating dog, Dusicyon cancrivorus wild dog - any of various undomesticated mammals of the family Canidae that are thought to resemble domestic dogs as distinguished from jackals or wolves (Cerdocyon thous). ********** After dog rabies control programs were implemented in Ceara State, Brazil, a the number of human cases decreased (1,2). Thereafter, the epidemiologic importance of rabies in wildlife became evident. From 1990 through 2005, a total of 173 cases of rabies were reported in Cerdocyon thous (crab-eating fox), 25 in Callithrix. j. jacchus (common marmoset Common Marmoset (Callithrix (Callithrix) jacchus) is a New World monkey. It originally seemed to only live on the northeast coast of Brazil. It was recently found also in southeast Brazil. Description The fur of the Common Marmoset is grey. ) and 6 in Procyon cancrivorous (crab-eating raccoon raccoon, nocturnal New World mammal of the genus Procyon. The common raccoon of North America, Procyon lotor, also called coon, is found from S Canada to South America, except in parts of the Rocky Mts. and in deserts. ). During this period, in 13 of 40 human cases reported in Ceara, wildlife was the source of infection (2). In 1996, because of this new epidemiologic situation, public health authorities launched an educational program, and no human cases due to wildlife were recorded in 1999, despite 84 cases in wildlife registered that year (2). The objective of this study was to elucidate some of the epidemiologic events involved in rabies emergence among wildlife in Ceara. The Study We studied 22 samples, from dogs, cattle, wildlife, and humans in Ceara, obtained from 1997 to 2003 (Table). Samples were antigenically characterized by using a monoclonal antibody monoclonal antibody, an antibody that is mass produced in the laboratory from a single clone and that recognizes only one antigen. Monoclonal antibodies are typically made by fusing a normally short-lived, antibody-producing B cell (see immunity) to a fast-growing (MAb) panel against the viral nucleoprotein nucleoprotein Macromolecular complex consisting of a protein linked to a nucleic acid, either DNA or RNA. The proteins that combine with DNA are generally of characteristic types called histones and protamines. (3-5). Isolates were injected into the brains of suckling suckling In mammals, the drawing of milk into the mouth from the nipple of a mammary gland. In human beings, it is referred to as nursing or breast-feeding. The word also denotes an animal that has not yet been weaned—that is, whose access to milk has not yet been mice, and brain impressions were made for MAb typing (3,5). Characterization of the samples identified 4 antigenic variants. Antigenic variant-2 (AgV2), maintained by dogs, was found in all C. thous, P. crancrivourous, and human cases and in all dog isolates with the exception of brdg5360, which was positive with all the MAbs. Antigenic variant-3 (AgV3), epidemiologically associated with vampire bats, Desmodus rotundus Noun 1. Desmodus rotundus - mouse-sized bat of tropical Central America and South America having sharp incisor and canine teeth; feeds on the blood of birds and mammals , was identified in 3 bovine samples. A previously reported profile, representing an AgV that circulates in marmosets in Ceara (5), was detected in sample brsg5696. The Ceara viruses were analyzed genetically through a comparative phylogenetic phy·lo·ge·net·ic adj. 1. Of or relating to phylogeny or phylogenetics. 2. Relating to or based on evolutionary development or history. study based on a 320-bp fragment of the nucleoprotein gene, from position 1157 to 1476, as compared with SADB SADB Security Association Database SADB Syrian Agricultural Database SADB Subject Area Database SADB Small and Disadvantaged Business SADB Single Asset Database SADB Security Audit Database SADB Security Audit Data Base 19 (5-7,8). These isolates were also compared with rabies virus variants circulating among domestic animals and wildlife from the Americas. The viral RNA RNA: see nucleic acid. RNA in full ribonucleic acid One of the two main types of nucleic acid (the other being DNA), which functions in cellular protein synthesis in all living cells and replaces DNA as the carrier of genetic was extracted from infected tissues, and the cDNA was obtained by reverse transcription-PCR techniques, using primers 21 g and 304, and was sequenced with primer 304 (7,9). The phylogenetic analyses were made by using the PileUp program of the Wisconsin Package Version 10.1 (10) and the programs DNADIST, NEIGHBOR, SEQBOOT, and CONSENSE of the PHYLIP PHYLIP Phylogeny Inference Package (genetics software) package (11). The expressed percentages of identity refer to the nucleotide sequences. The trees were obtained with the TREEVIEW program (12). The phylogenetic analyses showed segregation in 5 lineages, A-E A-E, AE above-elbow; see under amputation. (Figure 1), which was statistically supported by high bootstrap See boot. (operating system, compiler) bootstrap - To load and initialise the operating system on a computer. Normally abbreviated to "boot". From the curious expression "to pull oneself up by one's bootstraps", one of the legendary feats of Baron von Munchhausen. values. [FIGURE 1 OMITTED] Lineage A was represented by a sample from a dog from Maranguape, which was obtained in 2001. This virus showed its closest genetic relationship with lineage B (identity 92.4%-94.2%). Lineage B was formed by all the C. thous isolates, a sample from a human bitten by a P. cancrivorous raccoon in Fortaleza in 1997, and a virus from a P. crancrivorous raccoon collected in Maranguape during 2001 (intrinsic identity 96.5%-100%). This lineage showed its highest percentage of identity with lineage C (intrinsic identity 90.6%-92.8%). Lineage C consisted of 9 human samples collected in 5 different counties from 2000 to 2003 and an isolate obtained from a dog in 2000. The samples were highly homologous homologous /ho·mol·o·gous/ (ho-mol´ah-gus) 1. corresponding in structure, position, origin, etc. 2. allogeneic. ho·mol·o·gous adj. 1. (intrinsic identity 99.1%-100%). Lineage D included 3 bovines collected in 3 geographically distant counties during 2000 and 2001 (intrinsic identity 97.5%-98.4%). Lineage E was represented by the only sample collected from a C. j jacchus marmoset marmoset (mär`məzĕt'), name for many of the small, squirrellike New World monkeys of the family Callithricidae. Members of this family are all found in tropical South America, with one species found also in Central America. , These last 2 lineages were related distantly to all the others. When compared with representatives of rabies variants maintained by terrestrial and bat species in the Americas (Figure 2), lineages A, B, and C continue to segregate seg·re·gate v. seg·re·gat·ed, seg·re·gat·ing, seg·re·gates v.tr. 1. To separate or isolate from others or from a main body or group. See Synonyms at isolate. 2. as independent lineages with high statistical support. Ceara bovine samples representing lineage D clustered with D. rotundus and D. rotundus-related cases from Latin America Latin America, the Spanish-speaking, Portuguese-speaking, and French-speaking countries (except Canada) of North America, South America, Central America, and the West Indies. (intrinsic identity 94%-97.8%). [FIGURE 2 OMITTED] The only sample representative of lineage E segregated with 2 isolates from humans bitten by C. j. jacchus and a sample collected from a marmoset kept as a pet (5). The isolates were highly homologous to each other (intrinsic identity 98.7%-100%). Conclusions A thorough description of rabies epidemiology depends on a comprehensive surveillance program and application of accurate molecular methods to discriminate among different variants and the emergence of new foci. Antigenic and limited sequencing analyses were used to better understand the emergent epidemiologic events in wildlife in Ceara, Brazil. These analyses allowed identification of 5 potential cycles in this region, despite antigenic homogeneity. Lack of antigenic and genetic relationships of sample brdg5693, representing lineage A, with the rest of the isolates from Ceara and the known terrestrial rabies vectors from the Americas shows that this virus is a variant not previously described. This virus was geographically and temporarily associated with samples brpcr5698 and brcth5692, obtained in Maranguape during 2001. These circumstances demonstrated the existence of at least 2 overlapping endemic cycles in this area. Lineage B was formed mainly by isolates from C. thous, which indicates the existence of an emerging rabies cycle in this species. The epidemiologic situation in Ceara was complicated because of overlapping distributions of dog and C. thous rabies cases (Figure 1). Tree topology and genetic relationships between dog and C. thous variants suggested that the canine virus was introduced in C. thous populations because of spillover spill·o·ver n. 1. The act or an instance of spilling over. 2. An amount or quantity spilled over. 3. A side effect arising from or as if from an unpredicted source: events, which gave rise to an emergent cycle. A similar event was described between domestic dogs and Canis adustus (jackal jackal, name for several Old World carnivorous mammals of the genus Canis, which also includes the dog and the wolf. Jackals are found in Africa and S Asia, where they inhabit deserts, grasslands, and brush country. ) in Zimbawe. In this case, the variant circulating in dogs was introduced into the C. adustus population by spillover events, with the consequent emergence of an independent cycle (13). Recently, the hoary fox
Inclusion in lineage B of an isolate obtained from a human bitten by a P. cancrivorous raccoon and another sample collected from this species suggested the risk of establishing C. thous variant in P. cancrivorous. The niches of these 2 species overlap, which facilitates their encounters. Additional surveillance is necessary to clarify this situation. Epidemiologic data which indicates that humans had been exposed to dog bites, results of molecular characterization, and inclusion of a dog isolate in the C lineage strongly incriminate To charge with a crime; to expose to an accusation or a charge of crime; to involve oneself or another in a criminal prosecution or the danger thereof; as in the rule that a witness is not bound to give testimony that would tend to incriminate him or her. the dog as the reservoir of this variant. Identification of the source of infection by using classic surveillance alone is complicated by the presence of multiple cycles of transmission. Genetic comparison of samples from lineage D with viruses representing bats viruses from the Americas helped to identify D. rotundus as the source of livestock infection. The close genetic relationship of sample brsg5696 with rabies isolates obtained from C. j. jacchus and human cases bitten by marmosets further supported C. j. jaccuss as the most important vector of this variant. This finding indicates that this species plays an important role for disease maintenance in nature. Methods for antigenic and genetic identification of rabies samples isolated in the Americas have contributed effectively to the development of health programs, as well as recognition of possible wild reservoirs of urban rabies. The emergence of new cycles in Latin American wildlife indicates the need to strengthen surveillance programs in these species and research development for the evaluation of the feasibility of oral vaccination interventions. Acknowledgments We thank Debora R. V. Sacramento for her technical collaboration, Fabiana L. A. da Silveira Lebrun for structural support, Maria das Gracas Silva for help with the bibliography and illustrations, and all staff at Instituto Pasteur of Sao Paulo, Brazil, and Laboratory of Rabies Diagnosis of Ceara, Brazil, for collaboration and incalculable incentives. Dr Favoretto is a scientific researcher and specialist in public health at the Instituto Pasteur of Sao Paulo. Her research interests include diagnosis and molecular epidemiology molecular epidemiology Molecular medicine An evolving field that combines the tools of standard epidemiology–case studies, questionnaires and monitoring of exposure to external factors with the tools of molecular biology–eg, restriction endonucleases, of rabies virus. References (1.) Pan-American Health Organization/World Health Organization. Prevention and control of diseases. Veterinary Public Health Unit. Elimination of the human rabies transmitted by dogs in Latin America: analysis of the situation, 2004. Washington: Pan American Health Organization The Pan American Health Organization (PAHO) is an international public health agency with 100 years of experience in working to improve health and living standards of the countries of the Americas. It serves as the specialized organization for health of the Inter-American System. ; 2005. vol. 37, p. 10-29. (2.) Nucleo de Controle das Endemias Transmissiveis por vetores--NUEND, Coordenadoria de Apoio ao Desenvolvimento da Atencao a Saude--CODAS, 1998-2003. Boletim da Sccretaria da Saude do Estado do Ceara, Brasil, 2004. (3.) Favoretto SR, Carrieri ML, Cunha EMS, Aguiar EAC EAC an abbreviation used in studies of complement, in which E represents erythrocyte, A antibody, and C complement. , Silva LHQ LHQ Local Headquarters LHQ Life History Questionnaire , Sodre M, et al. Antigenic typing of Brazilian rabies virus samples isolated from animals and humans, 1989-2000. Rev Inst Med Trop Sao Paulo. 2002;44:91-5. (4.) Pan-American Health Organization/World Health Organization. Los anticuerpos monoclonales en la caracterizacidn y vigilancia de los virus de la rabia en America Latina y el Caribe. Pan American Journal of Public Health The American Journal of Public Health (AJPH) is a peer reviewed monthly journal of the American Public Health Association (APHA). The Journal also regularly publishes authoritative editorials and commentaries and serves as a forum for the analysis of health policy. . 2000;8:214-7. (5.) Favoretto SR, Mattos CC, Morais NB, Araujo FAA, Mattos CA. Rabies in marmosets (Callithrix jacchus) from the state of Ceara, Brazil. Emerg Infect Dis. 2001;7:1062-5. (6.) Conzelmann KK, Cox JK, Schneider LG, Thiel HJ. Molecular cloning and complete sequence of the attenuated Attenuated Alive but weakened; an attenuated microorganism can no longer produce disease. Mentioned in: Tuberculin Skin Test attenuated having undergone a process of attenuation. rabies virus SABB SABB Saudi British Bank SABB Storage Array Building Block SABB Solar Array Blanket Box(es) 19. Virology virology, study of viruses and their role in disease. Many viruses, such as animal RNA viruses and viruses that infect bacteria, or bacteriophages, have become useful laboratory tools in genetic studies and in work on the cellular metabolic control of gene expression . 1990;175:485-99. (7.) Smith JS. Rabies virus, In: Murray PR, Baron EJ, Pfaller MA, Tenover FC, Yolken R, editors. Manual of clinical microbiology, 6th ed. Washington: American Society for Microbiology The American Society for Microbiology (ASM) is a scientific organization, based in the United States although with over 43,000 members throughout the world. It is the largest single life science professional organization and its members include those whose interests encompass basic Press; 1995. p. 997-1003. (8.) de Mattos CC, de Mattos CA, Loza-Rubio E, Aguilar-Setien A, Orciari LA, Smith JS. Molecular characterization of rabies virus isolates from Mexico: implications for transmission dynamics and human risk. Am J Trop Med Hyg. 1999;61:587-97. (9.) de Mattos CA, de Mattos CC, Smith JS, Miller E, Papo S, Utrera A, et al. Genetic characterization of rabies field isolates from Venezuela. J Clin Microbiol. 1996;34:1553-8. (10.) Version WR 10.1 Madison (WI): Genetics Computer Group; 2000. (11.) Felsenstein, J. Phylip: Phylogeny inference pakage. Version 3,5c. Seattle (WA): University of Washington;, 1993. (12.) Page, Ram. D. TREEVIEW: An application to display phylogenetic trees on personal computers. Computer Applied Biocience. 1996;12:357-8. (13.) Bingham J, Foggim CM, Wandeler AI, Hill FW. The epidemiology of rabies in Zimbabwe. 2. Rabies in jackals (Canis adustus and Canis mesomelas). Onderstepoort J Vet Res. 1999;l:11-23. (14.) Beruardi F, Nadin Davis SA, Wandeler AI, Armstrong J, Gomes A, Lima FS, et al. Antigenic and genetic characterization of rabies viruses isolated from domestic and wild animals WILD ANIMALS. Animals in a state of nature; animals ferae naturae. Vide Animals; Ferae naturae. of Brazil identifies the hoary fox as a rabies reservoir. J Gen Virol. 2005;86:3153-62. Silvana R. Favoretto, * ([dagger]) Cecilia C. de Mattos, ([double dagger]) Nelio B. de Morals, ([section]) Maria Luiza Carrieri, * Benedito N. Rolim, ([section]) Lucia M. Silva, ([section]) Charles E. Rupprecht, ([double dagger]) Edison L. Durigon, ([dagger]) and Carlos A. de Mattos ([double dagger]) * Instituto Pasteur de Sao Paulo, Sao Paulo, Brazil; ([dagger]) Universidade de Sao Paulo, Sao Paulo, Brazil; ([double dagger]) Centers for Disease Control and Prevention Centers for Disease Control and Prevention (CDC), agency of the U.S. Public Health Service since 1973, with headquarters in Atlanta; it was established in 1946 as the Communicable Disease Center. , Atlanta, Georgia, USA; and ([section]) Secretaria Estadual de Saude do Ceara, Fortaleza, Brazil Address for correspondence: Silvana R. Favoretto, Instituto Pasteur of Sao Paulo, Brazil, Av: Paulista, 393, Paraiso, Silo silo, watertight and airtight structure for making and storing silage. Silos vary in form from a covered pit, such as was used by the early Romans, to the modern storage tower, dating from the 19th cent. Paulo, SP--Brazil, CEP CEP congenital erythropoietic porphyria. CEP abbr. congenital erythropoietic porphyria : 01311000; email: srfavoretto@ig.com.br The opinions expressed by authors contributing to this journal do not necessarily reflect the opinions of the Centers for Disease Control and Prevention or the institutions with which the authors are affiliated.
Table. Identification, origin, and antiqenic and genetic variant of
22 rabies virus samples isolated from Ceara State, Brazil
Year of
Identification Animal species isolation Origin
Brhm4531 Human 1997 Fortaleza
Brcth4122 Cerdocyon thous 1998 --
Brhm5325 Human 2000 Caucaia
Brdg5360 Dog 2000 Caucaia
Brcth5361 C. thous 2000 Paracuru
Brbv5339 Bovine 2000 Antonina do Norte
Brbv5374 Bovine 2000 Quixere
Brhm5691 Human 2001 Caucaia
Brdg5693 Dog 2001 Maranguape
Brcth5695 C. thous 2001 Barroquinha
Brcth5697 C. thous 2001 Caninde
Brcth5692 C. thous 2001 Maranguape
Brpcr5698 P. cancrivorous 2001 Maranguape
Brbv5694 Bovine 2001 Aquiraz
Brsg5696 C. j. jacchus 2001 Caucaia
Brhmu138 Human 2002 Fortaleza
Brhmu142 Human 2003 Fortaleza
Brhmu130 Human 2003 Umirim
Brhmu146 Human 2003 Fortaleza
Brhmu145 Human 2003 Tururu
Brhmu129 Human 2003 Fortaleza
Brhmu131 Human 2003 Maracanau
Antigenic GenBank
Identification variant Genetic variant Group accession no.
Brhm4531 AgV2 Dog B DQ447947
Brcth4122 AgV2 Dog B DQ447948
Brhm5325 AgV2 Dog C DQ447949
Brdg5360 All+ Dog C DQ447950
Brcth5361 AgV2 Dog B DQ447951
Brbv5339 AgV3 Vampire bat D DQ447952
Brbv5374 AgV3 Vampire bat D DQ447953
Brhm5691 AgV2 Dog C DQ447954
Brdg5693 AgV2 Dog A DQ447955
Brcth5695 AgV2 Dog B DQ447956
Brcth5697 AgV2 Dog B DQ447957
Brcth5692 AgV2 Dog B DQ447958
Brpcr5698 AgV2 Dog B DQ447959
Brbv5694 AgV3 Vampire bat D DQ447960
Brsg5696 AgV new * Marmoset E DQ447961
Brhmu138 AgV2 Dog C DQ447962
Brhmu142 AgV2 Dog C DQ447963
Brhmu130 AgV2 Dog C DQ447964
Brhmu146 AgV2 Dog C DQ447965
Brhmu145 AgV2 Dog C DQ447966
Brhmu129 AgV2 Dog C DQ447967
Brhmu131 AgV2 Dog C DQ447968
* Pattern related to isolates from marmosets.
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