Rabies in Sri Lanka: splendid isolation. (Dispatches).Rabies virus rabies virus n. A rather large, bullet-shaped virus of the genus Lyssavirus that causes rabies. exists in dogs on Sri Lanka Sri Lanka (srē läng`kə) [Sinhalese,=resplendent land], formerly Ceylon, ancient Taprobane, officially Democratic Socialist Republic of Sri Lanka, island republic (2005 est. pop. as a single, minimally divergent lineage only distantly related to other rabies virus lineages in Asia. Stable, geographically isolated virus populations are susceptible to local extinction Local extinction is where a species (or other taxon) ceases to exist in the chosen area of study, but still exists elsewhere. This phenomenon is also known as extirpation. Local extinctions are contrasted with global extinctions. . A fully implemented rabies-control campaign could make Sri Lanka the first Asian country Noun 1. Asian country - any one of the nations occupying the Asian continent Asian nation country, land, state - the territory occupied by a nation; "he returned to the land of his birth"; "he visited several European countries" in >30 years to become free of rabies virus. ********** 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 , an encephalomyelitis encephalomyelitis /en·ceph·a·lo·my·eli·tis/ (en-sef?ah-lo-mi?e-li´tis) inflammation of the brain and spinal cord. acute disseminated encephalomyelitis caused by infection with rabies virus or other lyssaviruses, is responsible for 40,000-50,000 human deaths each year in Asian countries (1). Human rabies is preventable, but the high cost of antirabies biologics limits their use. Because the source of almost all human rattles infections in Asia is a bite by a domestic dog, effective dog rabies control programs not only serve to reduce human deaths but also can reduce the overall costs associated with rabies prevention. Sri Lanka is an island with a land area of approximately 62,000 [km.sup.2] situated in the Indian Ocean Indian Ocean, third largest ocean, c.28,350,000 sq mi (73,427,000 sq km), extending from S Asia to Antarctica and from E Africa to SE Australia; it is c.4,000 mi (6,400 km) wide at the equator. It constitutes about 20% of the world's total ocean area. , 35 km from the southern end of the Indian Peninsula. More than 95% of approximately 100 human rabies deaths each year in Sri Lanka are the result of bites by unvaccinated dogs, and U.S.$1.5 million is spent each year for antirabies biologics (2). A national dog rabies elimination program begun 20 years ago has yet to reduce the number of bite exposures requiring rabies treatment or the number of human rabies deaths. Low vaccination coverage in the resident dog population and ineffective management of stray animals are the most likely reasons for the program's lack of success, but the abundant wild fauna in Sri Lanka could provide unrecognized reservoirs for rabies virus. Serologic se·rol·o·gy n. pl. se·rol·o·gies 1. The science that deals with the properties and reactions of serums, especially blood serum. 2. data also suggest that lyssaviruses other than rabies virus may be present in bats in Asia (3). Additionally, Sri Lanka's close proximity to India might allow frequent introduction of rabies virus--infected animals. Our research expands an earlier study of rabies in Sri Lanka (4) by using a panel of monoclonal antibodies This is a list of monoclonal antibodies, antibodies which are clones of a single parent cell. When used as medications, the generic names end in -mab (see "Nomenclature of monoclonal antibodies"). (MAbs) to survey Sri Lankan rabies samples for lyssaviruses other than rabies, by using genetic typing to identify viral lineages that might signify a wildlife reservoir for rabies virus, and by using 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. analysis to obtain estimates of the frequency of introductions of rabies virus from neighboring neigh·bor n. 1. One who lives near or next to another. 2. A person, place, or thing adjacent to or located near another. 3. A fellow human. 4. Used as a form of familiar address. v. Asian countries that might jeopardize a dog rabies--control program in Sri Lanka. The Study Most samples submitted for rabies testing Rabies testing is a test generally done on animals (predominantly wild animals) when a person has been bitten. Since the 1960's, the standard test for rabies has been Direct fluorescent antibody test (dFA test). in Sri Lanka originate from the area around the capital, Colombo. Our study included 44 samples: 36 from cases chosen randomly from the approximately 180 animals diagnosed with rabies at the Medical Research Institute, Colombo, from January to May 2001; 2 from elephants whose infections in 1998 and 1999 occurred in an area of Colombo with large active bat Active Bat is a low-power, wireless indoor location system accurate up to 3 cm. It is based on a principle of trilateration, and relies on multiple ultrasonic receivers embedded in the ceiling and measures time-of-flight to them. colonies; and 5 from human case-patients (1999-2001). A dog rabies virus collected in 1986 from Colombo served as a reference sample for mutational change in Sri Lankan rabies virus over time. Sri Lankan samples were submitted to virus typing by both antigenic and genetic methods. The antigenic profile of 31 Sri Lankan samples, displaying 3+ to 4+ antigen distribution, was determined by indirect immunofluorescence Noun 1. indirect immunofluorescence - a method of using fluorescence microscopy to detect the presence of an antigen indirectly fluorescence microscopy - light microscopy in which the specimen is irradiated at wavelengths that excite fluorochromes methods with a panel of 20 nucleoprotein-specific MAbs obtained from the 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. and the Wistar Institute The Wistar Institute, an independent nonprofit biomedical research institute in Philadelphia, Pennsylvania, United States, is dedicated to discovering the causes and cures for major diseases, including cancer, cardiovascular disease, autoimmune disorders, and infectious diseases. and compared with published reaction patterns for these MAbs (5). Only two antibodies were useful. C4, which is not known to react with any lyssavirus other than rabies virus, reacted with all samples from Sri Lanka, indicating the absence of any other lyssavirus among these samples. A negative reaction with MAb CR54 distinguished rabies virus from Sri Lanka from that from the Philippines. The reaction pattern for Sri Lankan virus was identical to that reported for rabies virus from Indonesia and Thailand. A phylogenetic analysis included the 44 Sri Lankan samples and nucleotide sequences for rabies virus and other lyssaviruses obtained from GenBank and from a sequence repository at CDC See Control Data, century date change and Back Orifice. CDC - Control Data Corporation (Table, Figure 1). Data analyzed included representatives of available sequence for mainland and island Asian countries and sequence data for samples representing a similarly broad geographic distribution of mainland and island countries of East Africa and the Arabian Peninsula Arabian Peninsula or Arabia Peninsular region, southwest Asia. With its offshore islands, it covers about 1 million sq mi (2.6 million sq km). Constituent countries are Bahrain, Kuwait, Oman, Qatar, United Arab Emirates, Yemen, and, the largest, Saudi Arabia. . With the use of standard methods (6), 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 the Sri Lankan rabies virus samples, reverse transcribed, and amplified by polymerase chain reaction polymerase chain reaction (pŏl`ĭmərās') (PCR), laboratory process in which a particular DNA segment from a mixture of DNA chains is rapidly replicated, producing a large, readily analyzed sample of a piece of DNA; the process is . Genetic typing was based on nucleotide sequence differences in complementary DNA complementary DNA n. cDNA. (cDNA) for the 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. gene as aligned with sequence for the Pasteur vaccine strain of rabies virus (7). [FIGURE 1 OMITTED] The 44 samples from Sri Lanka shared approximately 99% homology homology (hōmŏl`əjē), in biology, the correspondence between structures of different species that is attributable to their evolutionary descent from a common ancestor. over a 320-nucleotide region of the carboxy terminus of the N gene (bp 1157-1476). Only 10 nucleotide substitutions (all synonymous) were found when the entire protein encoding sequence of the N gene (1350 bp) was compared for isolates from 1986 and 2001 (1986 dog case no. 1294, GenBank accession no. AY138549; 2001 cow case no. 5657, GenBank accession no. AY138550), and the two samples were identical over an additional 56 bp of nontranslated sequence between the stop codon stop codon n. Any of three codons, UAA, UAG, or UGA, that signal the termination of the synthesis of a protein. Also called chain termination codon. and polyadenylation signal (bp 1421-1476). No evidence was found for a wildlife reservoir for rabies independent of rabies in domestic dogs. Mutations at nucleotide position 1231 and 1408 separated the virus samples into two clusters (Figure 1), but the clusters corresponded roughly to sample collection sites (Figure 2), and domestic dog samples were found in both clusters. Because the analysis included only two mongooses, we cannot exclude mongooses as a cryptic reservoir for rabies. Nevertheless, rabies cases in mongooses and other wild animals WILD ANIMALS. Animals in a state of nature; animals ferae naturae. Vide Animals; Ferae naturae. are only infrequently identified in Sri Lanka and occur in areas where dog rabies cases are common. Arai et al. (4) found similar results for wildlife samples in their study. Given that >95% of human rabies cases report a dog bite dog bite Public health The clamping of skin and subjacent soft tissues between the upper and lower mandible of a canine, which may cause infections, acting as a disease vector or even death. See Dog. as the source of infection, expanding rabies-control programs to include wild animals in Sri Lanka without more specific evidence of a reservoir status for these animals would not be helpful. [FIGURE 2 OMITTED] In contrast to the limited sequence diversity among samples from Sri Lanka, extensive sequence divergence was found when samples from Sri Lanka were compared with samples from other Asian countries, and genetic change was much more evident in the nontranslated sequence (17.2% to 31.2% divergence) as compared with the protein-encoding sequence (11.8% to 21.5% divergence). Identity for nontranslated sequence in some intercountry comparisons was as low as 50% (e.g., 50% to 58% identity over the nontranslated sequence for comparison of viruses from Indonesia and India). Additionally, samples from Vietnam, Indonesia, Laos, the Philippines, and Thailand contained an extra nucleotide in the nontranslated region as compared with the Pasteur vaccine strain of rabies virus. The exact position of this extra nucleotide could not be determined because it varied in relation to different sample groups, and the single extra nucleotide likely represents multiple insertion and deletion events. Molecular aspects of the epidemiology of rabies within Sri Lanka and between Sri Lanka and other countries were determined by constructing phylogenetic trees (Figure 1). Because the possibility of multiple insertions and deletions prevented an exact alignment of the nontranslated sequence, phylogenetic analysis was conducted with only the protein-encoding sequence. The phylogenetic analysis showed that all samples positive for rabies in diagnostic tests from Sri Lanka were rabies virus. Samples from Sri Lanka clustered with the Pasteur vaccine strain of rabies virus and other samples identified as genotype genotype (jēn`ətīp'): see genetics. genotype Genetic makeup of an organism. The genotype determines the hereditary potentials and limitations of an individual. 1 lyssavirus (8) and were monophyletic monophyletic /mono·phy·let·ic/ (mon?o-fi-let´ik) descended from a common ancestor or stem cell. mon·o·phy·let·ic adj. 1. Descended or derived from one original stock or source. with respect to other lyssavirus genotypes (not shown). Rabies virus samples from Asia were segregated into six clades on the basis of the geographic origin of the samples. Despite the exclusion of the highly variable nontranslated sequence from the analysis, no evidence was found for a common ancestry for all six Asian lineages (Figure 1). Three lineages were composed of samples from a single country (Philippines, Vietnam, and Indonesia). The remaining three lineages comprised samples from geographically proximate proximate /prox·i·mate/ (prok´si-mit) immediate or nearest. prox·i·mate adj. Closely related in space, time, or order; very near; proximal. proximate immediate; nearest. countries (Sri Lanka and the sample from Madras Madras. 1 State and former province, India: see Tamil Nadu. 2 City, India: see Chennai. , India; Thailand and Laos; and Pakistan, India, and Nepal). Similar findings were achieved with a more limited sample set encompassing the entire nucleoprotein gene (inset in Figure 1). In contrast to the genetic diversity of Asian rabies virus samples, samples of African 1 rabies virus (8) collected from the Arabian Peninsula and mainland and island countries of East Africa shared much higher sequence identity (approximately 93%) and could be resolved as a single lineage sharing a common ancestry with European vaccine strains of the virus dating from the 19th century (Pasteur rabies virus). Molecular clock data suggest that this virus emerged 300-500 years ago (9) and support historical data for rabies virus dissemination during early European exploration and colonization colonization, extension of political and economic control over an area by a state whose nationals have occupied the area and usually possess organizational or technological superiority over the native population. of Africa (8-10). The genetic divergence Genetic divergence is the process of one species diverging over time into more than one species. Passing small random advantages characteristic changes over time from one generation to the next generations. between samples from Sri Lanka and the most closely related Asian sample (Madras, India; >6.3% difference) suggests a similarly distant common ancestor for these two rabies virus populations. Conclusions As depicted by the antigenic and genetic analysis of samples in this report and the earlier work by Arai et al. (4), rabies in Sri Lanka is associated with a single lineage of rabies virus, which shares only a distant common ancestry with rabies viruses from any other Asian country. Genetic diversity between rabies virus lineages from mainland or island Asian countries was much greater than among samples collected over a similar geographic expanse in Africa. Differences between Asia and Africa in the partitioning and degree of spread of rabies virus in host populations might be explained by agricultural practices that evolve in areas of high rainfall versus the agricultural practices necessary in areas where rain is seasonal and scarce. For example, nomadic See nomadic computing. herding, a common agricultural practice in Africa, would spread an introduced rabies virus over great distances, whereas the primarily small-scale farming operations in Sri Lanka and other Asian countries, maintained locally for generations, could restrict virus spread. These observations suggest that restrictions to domestic animal movement can be exploited in rabies-control efforts in Asia. Rabies viruses in Sri Lanka and India may be more closely related than described in this and previous studies (4). Rabies virus more closely related to Indian viruses may be present in the northern and eastern parts of Sri Lanka, where civil unrest prevented sample collection. The Sri Lankan variant of the virus may still be actively transmitted by animals in India; only limited virus typing has been conducted on Indian rabies samples. However, although considerable genetic diversity was found in the four available sequences for rabies virus from the Indian subcontinent Indian subcontinent, region, S central Asia, comprising the countries of Pakistan, India, and Bangladesh and the Himalayan states of Nepal, and Bhutan. Sri Lanka, an island off the southeastern tip of the Indian peninsula, is often considered a part of the subcontinent. , only one Indian lineage shared a common ancestry with Sri Lankan samples. The degree of divergence between this Indian sample and the Sri Lankan samples and the lack of heterogeneity in Sri Lankan samples collected over a 16-year period suggest that the introduction of the ancestral virus occurred only once and in the distant past. While the data reported here and in the work by Arai et al. (4) represent only a fraction of all rabies cases that occur each year in Sri Lanka, the island is small and heavily populated pop·u·late tr.v. pop·u·lat·ed, pop·u·lat·ing, pop·u·lates 1. To supply with inhabitants, as by colonization; people. 2. . If other rabies virus lineages are present, or if introductions from India are more frequent than our data indicate, future virus typing should detect them easily. Antigenic and genetic typing of samples collected for rabies testing in Sri Lanka found no lyssavirus other than rabies virus, no evidence for cycles of rabies virus transmission in wild species independent of endemic rabies virus in domestic dogs, no evidence of a recent introduction of the virus to Sri Lanka through the translocation translocation /trans·lo·ca·tion/ (trans?lo-ka´shun) the attachment of a fragment of one chromosome to a nonhomologous chromosome. Abbreviated t. of animals from other areas of Asia, and (surprisingly, given the close proximity to mainland Asia) no evidence for outside introduction of the virus to Sri Lanka in the recent past. Historically, rabies of this type has been susceptible to control with a fully implemented vaccination campaign and an animal-control effort that addresses local cultural and religious attitudes toward management of stray animals. The economic impact of rabies on Sri Lanka argues for national and international cooperation in a strong rabies-control program for this island nation. The prospects are good that such an effort could eliminate rabies virus from the island and create the first rabies virus--free country of the millennium.
Table. Virus samples from countries other than
Sri Lanka included in phylogenetic analysis
Sample ID Sample history and GenBank accession numbers
Australian bat Australian bat lyssavirus from insectivorous
lysssavirus bat (Saccolaimus flaviventris); 1996.
Lyssavirus most closely related to rabies
virus. Used as outgroup to root phylogenetic
analysis. AF081020.
Duvenhage virus Duvenhage virus from human bitten by bat,
South Africa, 1970. Bat lyssavirus used as
outgroup to root phylogenetic analysis.
U22848.
Nepal U.S. citizen bitten by dog
in Nepal, 1996. AY138578.
Pakistan Dog, 1990. AY138565.
India Resident of U. K., bitten by dog
in India, circa 1988. AY138566.
Kenya U.S. Peace Corps volunteer bitten
by dog in Kenya, 1983. AY138567.
Tanzania Goat, 1992, Zanzibar, Tanzania. AY 138568.
Saudi Arabia Arabian-American Oil Company employee
infected in Saudi Arabia, circa 1981,
AY138569; fox, 1987. U22481.
Oman Red fox, 1990. U22480.
Iran Dog, 1986. U22482.
Israel Dog, 1993. AY138570.
Madagascar Human, 1980. AY138571.
Pasteur rabies virus Cow bitten by dog, Paris, 1882, multiple
laboratory passages. M13215.
Namibia Jackal, 1992. U22649.
Algeria Dog, 1982. U22643.
Ethiopia Hyena, 1987. U22637.
India-Madras Unpublished, 2001. AF374721.
Sri Lanka, reference Dog case sample collected by Dr. Alex
dog sample, 1986 Wandeler, Colombo, 1986. AY138549.
Sri Lanka, human cases Five human case-patients with diagnosed
rabies, 1999-2001. Cluster A samples
originated from Kandy (n=2) and Ratnapura.
Cluster B samples originated from Colombo
and Karapitiya. Two cluster A human case
samples with nucleotide substitutions not
found in consensus sequence were submitted
to GenBank. AY138554 and AY138558.
Sri Lanka, dog cases 24 dogs with diagnosed rabies, January-May
2001. Cluster A samples originated from
Ragama, Polgasowita, Katunayake, Puttalam,
Meerigama, Panadura, Dewulapitiya. Cluster B
samples originated from Ragama, Polgasowita,
Mirihana, Dehiwela, Moratuwa, Chillaw,
Katana, Mount Lavinia, Balapitiya, Wattala
(n=2), Payagala (n=2), Colombo (n=3),
Nugegoda. Dog samples with nucleotide
substitutions not found in consensus
sequence for cluster A (AY138553, AY138555,
AY138557, AY138559) and cluster B (AY138551,
AY138552) were submitted to GenBank.
Sri Lanka, goat cases Two goats with diagnosed rabies,
January-May 2001. Cluster A sample originated
from Seeduwa. AY138561. Cluster B sample
originated from Kandy. AY138556.
Sri Lanka, mongoose Two mongooses (cluster B) with diagnosed
cases rabies, January-May 2001, Panadura and
Moratuwa. Sample with nucleotide
substitutions not found in consensus
sequence for cluster B was submitted
to GenBank. AY138562.
Sri Lanka, cow cases Four cows (cluster B) with diagnosed rabies,
January-May 2001, Kandy, Kaduwela, Horana,
Polgasowita. Samples with nucleotide
substitutions not found in consensus
sequence for cluster were submitted to
GenBank. AY138550 and AY138560.
Sri Lanka, elephant Two elephants (cluster B) with diagnosed
cases rabies in 1998 and 1999, in an area of
Colombo with large active bat colonies.
Sample with nucleotide substitutions not
found in consensus sequence for cluster B
was submitted to GenBank. AY138564.
Sri Lanka, cat cases Four cats (cluster B) with diagnosed rabies,
January-May 2001, Kelaniya, Nittambuwa,
Horana, Kaduwela. Sample with nucleotide
substitutions not found in consensus
sequence for cluster B was submitted to
GenBank. AY138563.
Philippines Resident of California, 1987, bitten by dog
in the Philippines, AY138575; resident of
California, 1972, bitten by dog in the
Philippines. AY138576.
Viet Nam Vietnamese refugee in Sidney,
Australia, 1991. AY138579.
Thailand Human, 1983, AY138572; dog, 1995, U22653.
Laos Laotian immigrant, Texas, 1984. AY 138577.
Indonesia Human, 1989, Jakarta, Java; AY 138573.
dog, 1989, Java; AY138574.
Acknowledgments We thank Carlos and Cecilia de Mattos and the staff of the Rabies Section of the Centers for Disease Control and Prevention for their assistance and Omala Wimalaratne for technical cooperation. References (1.) Rabies, Asia. Wkly Epidemiol Rec 2001;76:320-3. (2.) Wimalaratne O. Rabies in humans and animals in Sri Lanka. In: Dodet B, Meslin F-X, editors. Rabies control in Asia. Paris: John Libbey Eurotext; 2001. p. 248-51. (3.) Arguin PM, Murray-Lillibridge K, Miranda ME, Smith JS, Calaor AB, Rupprecht CE. Serologic evidence of Lyssavirus infections among bats, the Philippines. Emerg Infect Dis 2002;8:258-62. (4.) Arai YT, Takahashi H, Kameoka Y, Shiino T, Wimalaratne O, Lodmell DL. Characterization of Sri Lanka rabies virus isolates using nucleotide sequence analysis of nucleoprotein gene. Acta Virol 2001;45:327-33. (5.) Smith JS, Fishbein DB, Rupprecht CE, Clark K. Unexplained rabies in three immigrants in the United States United States, officially United States of America, republic (2005 est. pop. 295,734,000), 3,539,227 sq mi (9,166,598 sq km), North America. The United States is the world's third largest country in population and the fourth largest country in area. : a virologic investigation. N Engl J Med 1991;324:205-11. (6.) de Mattos CA, Favi M, Yung V, Pavletic C, de Mattos CC. Bat rabies in urban centers in Chile. J Wildl Dis 2000;36:231-40. (7.) Tordo N, Poch O, Ermine ermine, name for a number of northern species of weasel having white coats in winter, and highly prized for their white fur. It most commonly refers to the white phase of Mustela erminea, called short-tailed weasel in North America and stoat in the Old World. A, Keith G. Primary structure of leader RNA and nucleoprotein genes of the rabies genome: segmented homology with VSV VSV Vesicular Stomatitis Virus VSV Verband Schweizerischer Vermögensverwalter VSV Vacuum Switching Valve (car part) VSV Variable Stator Vanes VSV Vliegtuigbouwkundige StudieVereniging (Dutch) . Nucleic Acids Nucleic acids The cellular molecules DNA and RNA that act as coded instructions for the production of proteins and are copied for transmission of inherited traits. Res 1986;14:2671-3. (8.) Kissi B, Tordo N, Bourhy H. Genetic polymorphism polymorphism, of minerals, property of crystallizing in two or more distinct forms. Calcium carbonate is dimorphous (two forms), crystallizing as calcite or aragonite. Titanium dioxide is trimorphous; its three forms are brookite, anatase (or octahedrite), and rutile. in the rabies virus nucleoprotein gene. 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 1995;209:526-37. (9.) Badrane H, Tordo N. Host switching in Lyssavirus history from the Chiroptera to the Carnivora orders. J Virol 2001;75:8096-104. (10.) Smith JS, Orciari LA, Yager PA, Seidel sei·del n. A beer mug. [German, from Middle High German s  del, from Latin situla, bucket.]Noun 1. H, Warner CK. Epidemiologic and historical relationships among 97 rabies virus isolates as determined by limited sequence analysis. J Infect Dis 1992;166:296-307. Address for correspondence: Jean S. Smith, Viral and Rickettsial rickettsial /rick·ett·si·al/ (ri-ket´se-al) pertaining to or caused by rickettsiae. rick·ett·si·al adj. Relating to, or caused by a member of the genus Rickettsia. Zoonosis Zoonosis Definition Zoonosis, also called zoonotic disease refers to diseases that can be passed from animals, whether wild or domesticated, to humans. Branch, Division of Viral and Rickettsial Diseases, National Center for Infectious Diseases infectious diseases: see communicable diseases. , Centers for Disease Control and Prevention, Mailstop G33, 1600 Clifton Road Clifton Road is main street in Clifton neighborhood of Saddar Town in Karachi, Sindh, Pakistan. Its name dates from the British Colonial rule, and its market is posh areas of Karachi. , Atlanta, GA 30333, USA: fax: 404-639-1564; e-mail:jeansmithga@earthlink.net Susilakanthi Nanayakkara,* ([dagger]) Jean S. Smith, * and Charles E. Rupprecht * * Centers for Disease Control and Prevention, Atlanta, Georgia, USA; and ([dagger]) Medical Research Institute, Colombo, Sri Lanka Dr. Susilakanthi Nanayakkara is the assistant microbiologist of the Rabies Diagnosis and Research Laboratory of the Medical Research Institute of Sri Lanka. She obtained her post-M.D, overseas training on new techniques of rabies diagnosis at the Rabies Section of the Division of Viral and Rickettsial Diseases, National Center for Infectious Diseases, Centers for Disease Control and Prevention (CDC), for a 1-year period from September 2001 to August 2002. This work was performed while she was visiting CDC. |
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