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Lyme Disease Studies
Tickborne Diseases in California
The abstracts for most of these articles relating to tickborne diseases in California are available by going to the National Library of Medicine website http://www.ncbi.nlm.nih.gov and typing in keywords. In some cases, free full text versions may be available to download.
People with an interest this subject are fortunate that UC Berkeley entomologist Robert Lane, PhD, a consummate researcher, has devoted his entire career to the study of ticks and Lyme disease, and has trained many other researchers who follow in his footsteps in California and other states. You will see his name on many of the papers in this list. Our knowledge about Lyme disease in California is immeasurably greater because of the work of Dr. Lane. CALDA thanks him for his leadership, dedication, professionalism, and generosity in sharing his time and expertise with us for many years.
The abstracts are listed in chronological order of publication. You may use the Find tool above to search for keywords. (Select Edit and then Find and type in keyword)
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Full text is available on PubMed as a scanned copy of the original print version.
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Full text is available on PubMed as a scanned copy of the original print version.
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Full text is available on PubMed as a scanned copy of the original print version.
Six species of rodents from Kern County, California, were inoculated subcutaneously with western equine encephalomyelitis (WEE) virus to determine their susceptibility to infection and their potential as natural hosts. Ammospermophilus nelsoni, Citellus beecheyi, Dipodomys heermanni, Dipodomys nitratoides, Peromyscus maniculatus, and Sciurus griseus were readily infected. Infection was usually fatal in Dipodomys species, C. beecheyi, and S. griseus, but was clinically inapparent in other species. Viremic responses varied greatly in magnitude and duration in different species and with different viral strains. Viremic animals that survived developed high titers of hemagglutination-inhibiting antibody. Hemagglutination-inhibiting and neutralizing antibodies persisted at high titers for at least 8 to 58 weeks after infection, except in P. maniculatus. If animals died during or shortly after the viremic phase of infection, the virus usually was recoverable from numerous organs. Long-term survival of virus could not be demonstrated in A. nelsoni and Dipodomys species. It is concluded that A. nelsoni and P. maniculatus are not important natural hosts of WEE virus; they are susceptible to infection and develop antibodies, but serological surveys of the same species rarely reveal evidence of infection. S. griseus, D. heermanni, D. nitratoides, and possibly C. beecheyi are aberrant hosts of WEE virus since most of them died when infected. Two species of ticks that are ectoparasitic on rodents in Kern County were evaluated as vectors of WEE virus. Dermacentor parumapertus failed to become infected after feeding on viremic hosts, and Ornithodorus parkeri became infected but failed to transmit virus.
Full text is available on PubMed as a scanned copy of the original print version.
Inoculation of susceptible calves confirmed that the modified card agglutination test accurately detected the anaplasmosis infection status of each of 35 Columbian black-tailed deer (Odocoileus hemionus columbianus). Anaplasma marginale, and specific antibodies, were demonstrated only in calves which received blood from deer that were positive by the card test. The modified card agglutination testing of deer serum was performed in the manner recommended for testing cattle serum with bovine-origin antigen and bovine serum factor.
Evidence is presented for an area of human risk to Rocky Mountain spotted fever (RMSF) in California extending beyond the range of its classic tick vector, Dermacentor andersoni. Geographic, seasonal, and host distributions of Dermacentor variabilis and Dermacentor occidentalis suggest these species as potential vectors of RMSF in California outside the range of Dermacentor andersoni.
Lyme disease, defined by erythema chronicum migrans and sometimes followed by neurologic, cardiac, or joint involvement, is known to have affected 512 patients in the United States. The disease seems to occur in three distinct foci: along the northeastern coast, in Wisconsin, and in California and Oregon, a distribution that correlates closely with that of Ixodes dammini in the first two areas and with Ixodes pacificus in the last. The implicated tick, saved by six patients in the Northeast, was identified as nymphal I. dammini. Residence in or travel to endemic areas and history of tick bite may be important clues to diagnosis.
An engorging female Ixodes pacificus was observed on a western harvest mouse (Reithrodontomys megalotis) in Humboldt County, CA. The mouse demonstrated a flaccid paraplegia, but it appeared to recover fully after the tick was removed.
Full text is available on PubMed as a scanned copy of the original print version.
A rickettsial survey of ixodid ticks known to bite man was conducted in 1979 in four coastal counties of California to obtain isolates from tick species that might be involved in the transmission of spotted fever-like illnesses, and to examine serologic characteristics of the rickettsiae relative to defined members of the spotted fever group (SFG). One hundred and seventy (19.4%) of 877 ticks comprising three species were shown by hemolymph test to harbor rickettsia-like organisms. A total of 85 SFG rickettsia isolates was obtained by Vero-cell culture; 82 were from Dermacentor occidentalis, two were from D. variabilis, and one was from Ixodes pacificus. As determined by microimmunofluorescence, the isolates comprised four distinct serotypes. Two serotypes were obtained only from D. occidentalis, and one each only from D. variabilis and I. pacificus, respectively. Most D. occidentalis isolates possessed the serologic characteristics of Rickettsia rhipicephali, but there were similar to yet distinguishable from, R. rickettsii and are members of an unclassified serotype referred to as 364D. The two isolates from D. variabilis resembled the unclassified 369C serotype previously shown to be associated with this species and D. andersoni elsewhere in the United States. The I. pacificus isolate was similar to strains of the unclassified Tillamook serotype isolated from this tick in several localities in western Oregon. Representative strains of the four serotypes could also be distinguished on the basis of pathogenicity for Vero cells, chick embryos, guinea pigs, and/or meadow voles. The significance of these findings relative to occurrence of tick-associated illnesses in western California is briefly discussed.
Lyme disease is a process of unknown etiology that has been linked to bites of ticks of the Ixodes ricinus complex. Central Minnesota is an area of known Ixodes dammini distribution. This case represents the first recognition of Lyme disease from this area. This case presented as a fluctuating meningoencephalitis with superimposed cranial neuropathy. The characteristic skin lesion, erythema chronicum migrans, was recognized by history, and oligoarticular arthritis subsequently developed. Physicians practicing in areas of known Ixodes tick distribution outside of the northeastern United States (coastal areas of Oregon and California, central Minnesota, northwestern Wisconsin, northwestern Utah, certain parts of Ontario) should be alert to this disease and aware of the variety of symptom complexes that are possible with it.
A virus very similar or identical to Colorado tick fever (CTF) virus was recovered from the blood clot of one of 104 black-tailed jack rabbits (Lepus californicus) examined during a survey for various zoonotic agents in mammals and ticks from the University of California, Hopland Field Station, Mendocino County, California, 1974--79. This is the first reported isolation of a CTF-like virus from L. californicus, and only the second time such a virus has been found in northwestern California. Mendocino County is located far outside the known distributional ranges of the most common mammalian hosts of CTF virus and of Dermacentor andersoni, the only proven tick vector for man. The viral isolate is very similar to a CTF-like virus previously recovered from the blood and spleen of a western gray squirrel (Sciurus griseus) from San Luis Obispo County, an area also outside of the previously-known CTF area. Virus was not isolated from 14 additional species of mammals (354 specimens) or from eight species of ticks (4,487 individuals), but CTF-neutralizing antibodies were detected in 28 of 771 (3.6%) sera from seven of 15 mammalian species including significant titers (greater than or equal to 1:8) in two species and one subspecies not previously reported as natural hosts, i.e., brush mouse (Peromyscus boylii), pinyon mouse (P. truei), and Columbian black-tailed deer (Odocoileus hemionus columbianus). CTF indirect immunofluorescent antibodies also were detected in 26 of 129 (20.2%) sera belonging to four of five mammalian species tested. Neutralizing antibodies were found in sera of deer from other localities in Mendocino County, from a deer mouse from Napa County, and from a brush rabbit from Monterey County as well. These findings suggest that a virus identical or similar to CTF virus is widespread in northwestern-westcentral California, and that surveillance for human cases of CTF or a similar disease should be extended to cover this region.
A strain of Rickettsia canada was recovered in 1980 an adult rabbit tick, Haemaphysalis leporispalustris, taken from a black-tailed jack rabbit, Lepus californicus, in Mendocino County, California. In all examined biologic characteristics, this isolate, CA410, is indistinguishable from the prototype, strain 2678, isolated in Ontario, Canada, in 1963. These similarities include serologic and immunologic reactivity in laboratory mice and guinea pigs, cultural characteristics in Vero cells, chick embryo cells and embryonated eggs, low pathogenicity for mice, meadow voles and guinea pigs, unusual resistance to streptomycin, morphology by electron microscopy, and molar percentages of guanine plus cytosine of the deoxyribonucleic acids. Recovery of this second strain in the same species of tick, but far removed in time and place from the origin of the prototype, provides evidence that R. canada is an established, ecologically stable, rickettsia in North America.
Eighty-one 11-month-old, nonpregnant, Anaplasma marginale-seronegative beef heifers were allotted to 2 groups for evaluation of a modified live A marginale vaccine (n = 50 for vaccinated group and n = 31 for nonvaccinated group). The vaccine induced a mild form of anaplasmosis, as evidenced by a significant (P less than 0.01) decrease in the packed cell volume (PCV) between days 31 and 46 after vaccination. The lowest PCV was 11%, and 3 heifers had a PCV less than 20%. Slight lethargy was evident in some of the vaccinated heifers between days 30 and 45 after vaccination. All vaccinated heifers became seropositive to A marginale, as measured by the complement fixation test and the card test on days 35 and 42 after vaccination, respectively. All nonvaccinated heifers remained seronegative.
Lyme disease is a recently described clinical entity with cutaneous, neurologic, articular and cardiac manifestations. Since the original description of the disease in 1977, more than 500 cases have been reported. Although the vast majority of patients have been from the area near Lyme, Connecticut, we have seen four patients from northern California with various aspects of Lyme disease. This diagnosis should be considered in patients who have traveled to tick regions and who have a distinctive skin lesion (erythema chronicum migrans). Clinical and epidemiologic evidence suggests the disease is transmitted by a tick, lxodes dammini in the eastern United States, lxodes pacificus in the West and lxodes ricinus in Europe. The etiologic agent, a new spirochete, may have been recently discovered.
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To determine the significance of the western black-legged tick, Ixodes pacificus, as a vector of the Lyme disease spirochete, Borrelia burgdorferi, a tick/spirochete survey was conducted in northern California and southwestern Oregon from 1982 to 1984. Of 1,687 adult ticks collected off vegetation, 25 (1.48%) contained spirochetes. Of 715 ticks from Oregon, 14 (1.96%) were infected whereas 11 (1.13%) of 972 ticks from California harbored spirochetes. An isolate of one of the spirochetes reacted specifically when treated with monoclonal antibodies to B. burgdorferi. Polyacrylamide gel electrophoresis of a lysate of the isolate showed it to be nearly identical with two isolates of B. burgdorferi. Of the 25 infected I. pacificus, 17 had spirochetes in their midgut only; the remaining eight ticks showed a generalized infection of all the tissues, with midgut, central ganglion and ovary or testes showing heavy spirochetal infections. Decreased immunofluorescent staining reactivity of spirochetes in tissues other than midgut in six of eight I. pacificus with generalized infection may reflect adverse physiologic conditions for the development of spirochetes in the hemocele.
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The relationship of native Columbian Black-tailed Deer, two species of exotic deer (Axis and Fallow), and their ticks to the Lyme disease spirochete, Borrelia burgdorferi, was studied in coastal and inland areas of northern California, USA. Spirochetemias were detected in 27% of Black-tailed Deer, 50% of Axis Deer, and 56% of Fallow Deer collected in late fall and winter. Antibody prevalence was 38% in Black-tailed Deer, 24% in Fallow Deer, and 6% in Axis Deer. One to three tick species were collected from each species of deer, and two tick species were flagged from vegetation; of these, only the Western Black-legged Tick, Ixodes pacificus, was found to contain spirochetes. These findings suggest that all three deer species may be important hosts of spirochetes, possibly B. burgdorferi, and reconfirm that I. pacificus is the primary vector of the latter in California.
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Full text is available on PubMed as a scanned copy of the original print version.
One hundred and seventy five sera from cats in Ontario, Canada, were tested for hemagglutination inhibition (HI) antibodies to three arboviruses; namely, Powassan (POW) of the Flavivirus serogroup, and Snowshoe hare (SSH) and Jamestown Canyon (JC) viruses of the California (CAL) serogroup. All sera were negative for antibodies to POW virus. Twelve cats possessed CAL serogroup antibodies including three with antibodies to SSH alone, six with antibodies to JC alone, and three with antibodies to both SSH and JC antigens. POW virus was inoculated into seven cats, one intracerebrally and six intravenously. Neurologic signs were not detected in any of the cats. Histologic lesions of a nonsuppurative encephalitis and encephalomyelitis were observed in the intracerebrally inoculated cat and in one of the intravenously inoculated cats, respectively. POW virus was not isolated from the brain or spinal cord of either of these two cats. HI antibodies were detected in the sera of all inoculated animals. HI antibodies were not detected in the CSF of any animal.
During 1984 and 1985, blood samples were obtained from 271 dogs that were suspected of having borreliosis. The dogs lived in areas known to be infested with ticks and had been examined because of limb/joint disorders or for unknown illnesses marked by fever, anorexia, or fatigue. Lameness had been the most frequently reported clinical manifestation. Analyses of serum specimens, by an indirect fluorescent antibody (IFA) method or by an ELISA, detected antibodies to Borrelia burgdorferi, the etiologic agent of borreliosis in dogs and of Lyme disease in human beings. Antibody to B burgdorferi was detected in 76.3% of 114 specimens from dogs living in the lower Hudson Valley region of New York State (predominantly Westchester County), in 66.5% of 155 specimens from dogs from southern Connecticut, and in single specimens from dogs from Rhode Island and California. Geometric mean antibody titers peaked during the winter. Results of IFA tests and ELISA were in agreement, but the latter method yielded less variable results, had greater sensitivity, and was more easily standardized. Five dogs from New York State and Connecticut seropositive to B burgdorferi had developed kidney disorders during or after episodes of intermittent lameness. Application of murine monoclonal antibody in an IFA procedure verified the presence of B burgdorferi in renal cortical tissues from one dog.
In a survey of 1,714 adult Ixodes pacificus ticks collected in northern California, 24 (1.4%) were found to be infected with spirochetes that reacted with an anti-Borrelia burgdorferi polyvalent conjugate in direct immunofluorescence tests. Eleven isolates of B. burgdorferi from these ticks were characterized by monoclonal antibody, polyacrylamide gel electrophoresis, and Western blot (immunoblot) analyses. Ten of the isolates had molecular and antigenic characteristics similar to those of other U.S. isolates. One strain, cloned by limiting-dilution techniques, was different from any previously reported U.S. strain, but similar to reported European strains. The cloned strain, DN127-Cl9-2, did not react with monoclonal antibodies to Osp A and Osp B major proteins found in most of the U.S. strains. It exhibited an abundant protein with an apparent molecular weight of 25,000.
The relative abundance of, and spirochetal-infection rates in, adult ixodid ticks from eight Lyme borreliosis clinical-case areas and two comparison areas were investigated in northern California from late fall to early spring, 1984-87. The western black-legged tick (Ixodes pacificus) was the most abundant species at seven of nine sites yielding specimens as determined with a tick drag method. The Pacific Coast tick (Dermacentor occidentalis) was the most abundant species at two sites, and lesser numbers of this tick and the American dog tick (D. variabilis) were obtained from seven and two sites, respectively. Abundance of I. pacificus adults varied in clinical-case areas as well as in the comparison areas, and was not correlated significantly with spirochetal infection rates in this tick. Overall, spirochetes were detected in 1.4 and 1.0% of the adult I. pacificus collected from clinical case (n = 857) and comparison sites (n = 383), respectively, and in 0.8% of adult D. occidentalis (n = 253) from one comparison area. An additional 244 D. occidentalis adults from all other sites were tested with negative results. Five spirochetal isolates recovered from I. pacificus adults were identified as Borrelia burgdorferi with specific monoclonal antibodies. Seven of 10 patients interviewed reportedly contracted their infections in summer, and six presented with a history of tick bite. Nine patients owned or occasionally harbored one or more dogs, and at least one of the dogs contracted Lyme borreliosis at the same site as its owner. Clinical manifestations of the disease in human patients included erythema migrans (100%), arthritis/arthralgia (60%), neurologic abnormalities (60%), and cardiac involvement (20%). Four of nine patients whose sera were assayed by indirect immunofluorescence contained significant antibody titers to B. burgdorferi.
In northern California, antibodies to Borrelia burgdorferi were detected in 58 of 73 (79%), and spirochetemias in one of 26 (4%) black-tailed jackrabbits (Lepus californicus californicus), by indirect and direct immunofluorescence, respectively. Five species of ticks (Dermacentor occidentalis, D. parumapertus, Ixodes neotomae, I. pacificus, and Haemaphysalis leporispalustris) were collected from rabbits. Two of these species of ticks were found to contain spirochetes; two of 10 (20%) I. neotomae and two of 174 (1%) H. leporispalustris. A strain of B. burgdorferi was recovered from I. neotomae. One infected H. leporispalustris female passed spirochetes via eggs to about 67% of her progeny. The widespread distribution of the black-tailed jackrabbit, its infestation by at least four ticks (D. occidentalis, D. parumapertus, I. neotomae, and I. pacificus) known to be infected naturally with B. burgdorferi, and the high prevalence of spirochetal antibody in this lagomorph suggest that it might be useful as a sentinel for surveillance of Lyme borreliosis. Spirochetes were detected in 15% of 40 Columbian black-tailed deer (Odocoileus hemionus columbianus) by direct immunofluorescence bound with a Borrelia-specific monoclonal antibody (H9724), but not with a monoclonal antibody (H5332) specific for B. burgdorferi. The geographical overlap of different borreliae in ticks that bite wildlife such as deer may confound spirochetal serosurveys, and underscores the need for more specific serologic tests than those currently available.
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The relationship of immature western black-legged ticks, Ixodes pacificus Cooley and Kohls, to the western fence lizard, Sceloporus occidentalis Baird and Girard, and to the Lyme disease spirochete, Borrelia burgdorferi, was investigated in chaparral and woodland-grass habitats in northern California from 1984 to 1986. Immature ticks were found on lizards in spring and summer, but the prevalence and abundance of ticks on this host were considerably greater in spring. The peak of larval abundance preceded that of nymphs by several weeks, but there was considerable seasonal overlap between these parasitic stages. Larvae and nymphs attached primarily to the lateral nuchal pockets of lizards in chaparral (99.5%) and woodland-grass (91.8%). The numbers of larvae infesting lizards in spring fit the negative binomial distribution in woodland-grass but not in chaparral; insufficient data precluded similar analyses for nymphs. Tick loads did not differ significantly with respect to age or gender of the lizard. Spirochetal infection rates (range, 0-3.7%) in I. pacificus immatures were comparable in both habitats and were similar to those reported previously for adults of this tick. Overall, 1 (0.9%) of 117 larvae and 10 (1.8%) of 552 nymphs were infected with spirochetes resembling B. burgdorferi. Spirochetes were not observed in blood smears prepared from 261 wild-caught lizards, including five lizards fed upon by infected ticks at the time of collection. These and other findings suggest that S. occidentalis, although an important host of I. pacificus immatures, may be less important as a source for infecting ticks with B. burgdorferi.
Twenty (1.4%) of 1,421 adult Ixodes pacificus ticks and two (20%) of 10 adult Ixodes neotomae ticks collected in five counties of northern California were found to contain spirochetes by direct immunofluorescence examination of their tissues with a polyvalent conjugate. Borreliae isolated from the tissues of nine of these ticks (I. pacificus, 8; I. neotomae, 1) were identified as Borrelia burgdorferi with specific monoclonal antibodies and characterized further by polyacrylamide gel electrophoresis and Western blot (immunoblot) analyses. The isolate from I. neotomae was the first to be characterized from a tick other than I. pacificus in western North America. All strains were relatively homogeneous with respect to the kind of OspA proteins they produced, whereas they were heterogeneous with regard to their OspB proteins and to several low-molecular-weight proteins in the 21,500-to-24,000 region. Significant phenotypic variation was observed among isolates obtained within and between populations of I. pacificus. This investigation nearly doubles the number of isolates of B. burgdorferi that have been characterized from ixodid ticks in the far western United States.
Nine isolates of Borrelia burgdorferi from ixodid ticks collected in northern California were characterized. Restriction endonuclease analysis, pulsed-field gel electrophoresis, and Western blot (immunoblot) analysis were used in this study. Four isolates were very similar to each other. The others shared some similarities but were classified as having unique genotypes. A strain from an Ixodes neotomae tick displayed the greatest genetic and antigenic diversity when compared to the isolates collected from Ixodes pacificus ticks. A computerized library based on DNA banding patterns of the isolates by restriction enzyme analysis is also reported. This library was created by using a scanning laser densitometer.
Diurnal host-seeking by adults of the western black-legged tick, Ixodes pacificus Cooley & Kohls in relation to vegetative habitat type and meterological factors was investigated at a coastal and an inland site in northern California. Host-seeking behavior and relative tick abundance were determined with a tick drag method from late fall to midwinter 1987-1988. At the inland site, tick abundance usually was significantly greater in chaparral-grassland ecotones than in adjoining dense chaparral on the south-facing slope of a mountaintop, whereas both of these vegetative types produced significantly fewer ticks on a north slope compared with a contiguous south-facing slope. There was no evidence for an association between tick abundance and plant species within ecotonal chaparral. Multiple regression analyses revealed that tick abundance in ecotonal chaparral at the inland site and in grassland at the coastal site was not associated consistently with either ambient temperature or relative humidity. Compared with the inland site, ticks were considerably more abundant at the coastal site where sampling occasionally yielded more than 100 ticks per 100 drag samples. The incidence of infection with Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner, the etiologic agent of Lyme borreliosis, in adult ticks collected in the morning (1.5%) versus afternoon (2.2%), and in male (1.3%) versus female ticks (2.5%), at the coastal site in winter was similar.
An unusual strain of Borrelia burgdorferi (DN 127 cl 9-2) that was isolated from an Ixodes pacificus tick did not react with monoclonal antibodies (MAbs) to OspA and OspB surface proteins, which are found in most U.S. strains. The strain exhibited an abundant protein with an apparent molecular weight of 25,000 (25K protein). A MAb, 86 DN-1, that was prepared to the 25K protein was used in studies on the effect of proteases on the intact spirochetes, immune electron microscopy, and Western blot (immunoblot) analyses; the results indicated that the low-molecular-weight protein was an apparent surface protein that was loosely attached to the spirochete. Five tick isolates from California possessed low-molecular-weight proteins in the 20,000- to 25,000-molecular-weight range that reacted with the 86 DN-1 MAb. The 25K protein of DN 127 cl 9-2 was unaffected by prolonged in vitro passage of cultures in BSK II medium, while the low-molecular-weight proteins of the other strains of B. burgdorferi from California either decreased in quantity or became undetectable on long-term in vitro passage.
A 35-year-old woman presented with a bilateral palpebral follicular conjunctivitis. Subsequently, she developed a bilateral keratitis and, on a separate occasion, an episcleritis that was associated with a recrudescence of Lyme disease and poor compliance with the antibiotic regimen. Both the keratitis and episcleritis cleared completely after topical corticosteroid therapy and reinstitution of appropriate antibiotic treatment. This report emphasizes the importance of collaboration between internal medicine and ophthalmologic specialists during the long-term management of Lyme disease.
A conventional indirect immunofluorescence assay (IFA) and an anticomplement indirect immunofluorescence assay (ACIF) for detecting serum antibodies to Borrelia burgdorferi in humans were evaluated during a prevalence survey in northern California. Sera obtained from 119 current or former residents of an area in which Lyme disease is endemic were split and tested by the IFA in two laboratories and the ACIF in a third. The seropositivity rate ranged from 15 to 20% with 88 to 93% agreement among laboratories. Interlaboratory agreement was statistically highly significant in each of the three pairwise comparisons and was positively associated with clinical manifestations of Lyme disease. Intralaboratory agreement ranged from 93 to 96% in two laboratories and was also statistically highly significant. Immunoblotting confirmed 100 of 101 of the nondiscrepant immunofluorescence test results and likewise was positively correlated with the degree of interlaboratory agreement. The ACIF was found to be a highly specific test (100% specificity) with a much lower cutoff titer (1:8) than the conventional IFA (determined to be 1:128 or 1:256 in two laboratories) for detecting antibodies to B. burgdorferi. It also appeared to be more sensitive (80 versus 68%) than the IFA as determined by comparative immunoblotting, though the absolute sensitivity of the ACIF for serodiagnosis of early Lyme disease has yet to be determined. Significant serologic cross-reactivity was demonstrated between B. burgdorferi, Borrelia coriaceae, and Borrelia hermsii by the IFA, which may confound spirochetal serosurveys in California where all three spirochetes are known to coexist.
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A total of 138 birds (24 species) was captured in an oak woodland between December 1988 and June 1989 at the University of California, Sierra Foothill Range Field Station, Yuba County, Calif. Ticks were not found on 71 birds captured between December 1988 and March 1989. Five subadult Ixodes pacificus Cooley & Kohls were removed from 3 of 67 birds caught between April and June 1989. These three birds, an orange-crowned warbler (Vermivora celata (Say, a lazuli bunting (Passerina amoena (Say], and a chipping sparrow (Spizella passerina (Bechstein], represent new host records for I. pacificus in California. Tissues from two ticks and thick blood films prepared from 126 birds tested negative for spirochetes by direct immunofluorescence (DI). A total of 172 larval and 197 nymphal I. pacificus was removed from 15 of 16 western fence lizards (Sceloporus occidentalis Baird & Girard) caught between April and June 1989 in the same location as were birds. Thick blood films prepared from all 16 lizards and tissue smears from 334 of the ticks (143 larvae and 191 nymphs) were DI test-negative for spirochetes. One (1.1%) of 93 adult I. pacificus collected at the bird-lizard capture site in February 1989 was infected with spirochetes that resembled B. burgdorferi.
The etiologic agent of Lyme disease, Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner, was isolated repeatedly from dusky-footed wood rats, Neotoma fuscipes Baird, and California kangaroo rats, Dipodomys californicus Merriam, in northern California. All animals were collected in a region endemic for Lyme disease but for which the natural reservoir of B. burgdorferi was unknown. Similar attempts to isolate spirochetes from lizards, other species of rodents, jack rabbits, and deer between 1987 and 1991 were unsuccessful. Spirochetes isolated from wood rats and kangaroo rats were antigenically similar to strains of B. burgdorferi that had been isolated previously from the western black-legged tick, Ixodes pacificus Cooley & Kohls, in California. Similar enzootic cycles involving wood rats or kangaroo rats should be sought in other regions of the United States where the reservoirs of this spirochete are unknown.
The association of immature ixodid ticks, several species of rodents, and the Lyme disease spirochete, Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner, was studied in two habitats in northern California in spring and summer 1985 and year-round in 1986. A total of 428 rodents were collected from ecotonal chaparral and a woodland-grass-rock outcrop; the former habitat yielded six species, the latter three species. The deer mouse, Peromyscus maniculatus (Wagner), and the piñon mouse, P. truei (Shufeldt), were the dominant species year-round and collectively comprised 78% of rodents captured within chaparral and 87% from the rock outcrop in 1986. In both habitats, rodents were trapped most frequently in winter and spring, and least often in summer and fall. A total of 306 rodent blood films from all six species were assayed for spirochetes by direct immunofluorescence; of these, only one film prepared from P. truei (n = 123 films from 53 individual mice) was found to contain spirochetes. Immature western black-legged ticks, Ixodes pacificus Cooley & Kohls, and Pacific Coast ticks, Dermacentor occidentalis Marx, were collected from each species of rodent. Larvae of I. pacificus infested P. maniculatus and P. truei in low numbers year-round, but nymphs of this tick rarely parasitized these rodents. D. occidentalis larvae infested P. maniculatus and P. truei in spring and particularly in summer; nymphal ticks infested these mice primarily in summer. The efficiency of visual inspection for collecting immatures of these ticks from P. maniculatus ranged from 45 to 69% in spring and summer, whereas the efficiency of a drop-off technique appeared to be 100%. Spirochetes were detected in <1% of D. occidentalis larvae (n = 310) and nymphs (n = 120), and in approximately 4% of I. pacificus larvae (n = 75) derived from these hosts. The potential significance of these findings in the enzootiology of B. burgdorferi is discussed.
Babesia gibsoni caused severe hemolytic anemia in 11 dogs from southern California. The most common clinical signs of B gibsoni infection were lethargy, anorexia, anemia, and thrombocytopenia. Acute infection with B gibsoni may be misdiagnosed as autoimmune hemolytic anemia. Diagnosis was most reliably determined by identification of the intraerythrocytic parasites on Giemsa-stained blood smears. The pathogenicity of B gibsoni, difficulties in diagnosis, the parasite's resistance to treatment with available drugs, and frequent interstate movement of dogs indicate that this disease may be a serious threat to dogs throughout the United States.
Duration of attachment was determined for the mites Neotrombicula californica and Geckobiella texana and the tick Ixodes pacificus on the iguanid lizards Sceloporus graciosus and Uta stansburiana from southern California. Neotrombicula californica infestations lasted approximately one wk. Attachment of Ixodes pacificus larvae and nymphs lasted approximately eight days on S. graciosus and 16 days on U. stansburiana. Geckobiella texana remained attached to S. graciosus for five days and to U. stansburiana for 28 days.
Mass screening ELISA methods were developed for testing cattle serum for antibodies against 14 common livestock diseases simultaneously. The absorbance values were transformed to a %ELISA (spectrophotometric antibody end point) by a computer interfaced with a microplate reader. A histogram indicating a cutoff point and a report for the veterinarian also was generated. The computer program produced a print-out of the antibody profile for each animal tested, the antibody concentration against each disease, and a histogram (antibody profile) showing the prevalence of each disease in the herd. Serum samples were obtained from 1,953 cattle, including 880 dairy cattle from 10 herds and 1,073 beef cattle from 20 herds. These samples were obtained from June 1988 through June 1989. The highest antibody prevalence was against bluetongue virus. Of the 1,953 cattle tested, 1,223 (63%) were seropositive for bluetongue virus, including 502 (57%) of the dairy cattle and 721 (67%) beef cattle. Other antibody prevalences, in descending order, were: rotavirus (44%), Pasteurella spp (25%), Leptospira spp and Haemophilus spp (22%), Mycoplasma spp (18%), parainfluenza virus (17%), Campylobacter spp (16%), Anaplasma marginale (15%), bovine leukosis virus (13%), Brucella spp (8%), Mycobacterium paratuberculosis (8%), bovine viral diarrhea virus (3%), and infectious bovine rhinotracheitis virus (3%). Major differences in antibody prevalence between dairy and beef cattle were that only 4% of the dairy cattle were seropositive for A marginale, compared with 25% of the beef cattle, and conversely, 29% of the dairy cattle were seropositive for bovine leukosis virus, compared with 1% of the beef cattle.
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Argas (Argas) monolakensis, n. sp., is described from adults, nymphs, and larvae collected from under and around nests of California gulls, Larus californicus Lawrence, on islands in Mono Lake, Mono County, Calif., and from specimens reared in the laboratory. This species is closely related to A. cooleyi Kohls & Hoogstraal, a parasite of cliff swallows, Hirundo pyrrhonota Vieillot, but is easily distinguished by hypostome dentition and roof of Haller's organ in all stages and chaetotaxy of the larvae. This tick was successfully reared and maintained in the laboratory by feeding them on domestic chickens. Larvae require 5-8 d to feed, whereas all postlarval stages feed rapidly within 9-62 min. At Mono Lake, ticks are above ground and seek hosts only at night. The number of nymphal stages varies from 2 to 5 depending on the developmental temperature and sex of the tick. Ticks over winter at Mono Lake as second- to fifth-stage nymphs and adults. Ovarian diapause is common with preoviposition periods in extreme cases lasting up to 20 months. This tick will readily feed on humans and has the potential to transmit Mono Lake virus, which has been isolated from an estimated 2-8% of ticks on various islands. To date, A. monolakensis is known only from islands in Mono Lake, Calif.
The Lyme disease spirochete, Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner, was isolated from the blood of a dusky-footed wood rat, Neotoma fuscipes Baird, in the San Bernardino Mountains of southern California. Antigenic, protein, and molecular analyses demonstrated that the isolate varied slightly from most isolates of B. burgdorferi from northern California and was clearly distinct from other species of Borrelia that are endemic to the state. This is the first reported isolate of B. burgdorferi from southern California and demonstrates that the Lyme disease spirochete is enzootic in mountains near the major human population center of the state.
Knowledge of zoonotic transmission cycles is essential for the development of effective strategies for disease prevention. The enzootiology of Lyme disease in California differs fundamentally from that reported from the eastern United States. Woodrats, not mice, serve as reservoir hosts, and Ixodes neotomae, a nonhuman-biting tick, maintains the agent of Lyme disease, Borrelia burgdorferi, in enzootic cycles. The western black-legged tick, Ixodes pacificus, is the primary vector to humans, but it appears to be an inefficient maintenance vector. Isolates of B. burgdorferi from California exhibit considerable antigenic heterogeneity, and some isolates differ strikingly from isolates recovered from this and other geographic regions.
The role of the Western fence lizard Sceloporus occidentalis in the enzootiology of the Lyme disease spirochete Borrelia burgdorferi was evaluated in the Hopland and Ukiah areas of Mendocino County, California. In 1989, half of 74 lizards collected monthly from April to October at Hopland were infested by the immature western black-legged tick Ixodes pacificus at a mean intensity of 6.0 ticks per lizard. The prevalence of infestation of lizards by immature I. pacificus (36 of 73) at Ukiah was similar, but the mean intensity (12.9) was approximately twice as great. Overall, zero of 223 larvae and two (0.6%) of 330 nymphs from both sites were found to contain spirochetes by direct immunofluorescence. Larval and nymphal I. pacificus fit the negative binomial distribution in spring, and the prevalence and abundance of these stages were significantly greater in spring than in summer at both sites. Spirochetes were not visualized in thick blood films prepared from 133 lizards from both localities. Plasma antibodies against B. burgdorferi were detected in seven of 10 experimentally inoculated lizards, in five (8%) of 63 lizards from Hopland, and in 10 (14%) of 70 lizards from Ukiah. Adult lizards had a significantly greater tick burden and seropositivity rate than juvenile lizards only at Ukiah. In 1991, efforts to detect and culture spirochetes from the blood of 21 wild-caught lizards and from the tissues of 189 associated ticks that fed xenodiagnostically on them were unsuccessful.(ABSTRACT TRUNCATED AT 250 WORDS)
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A 1-year prospective study of risk factors for seropositivity to and contraction of Lyme disease among members of a small rural community (population, approximately 150) was conducted in northwestern California in 1988-1989. The initial rate of seropositivity for Borrelia burgdorferi for 119 current or former residents ranged from 15 to 20% among three laboratories, with statistically significant interlaboratory agreement. Questionnaires were completed by 93 current residents at entry and 80 residents a year later to evaluate the association of serologic status with 20 categorical and 47 continuous variables. Seropositive subjects had resided in the study area about 2 years longer, were bitten by unspecified biting flies more often, and were less likely to have engaged in hiking than seronegative subjects. One of 59 seronegative subjects seroconverted a year later (annual incidence = 1.7%). The cumulative frequency of seropositivity for Lyme disease in the study population was > or = 24%. Of 83 subjects examined physically, 13 were diagnosed as having definite and 18 as having probable Lyme disease. The seropositivity rate was significantly higher (38.7%) among individuals with definite/probable Lyme disease than in asymptomatic subjects (13.5%). Subjects who were seronegative or free of Lyme disease reported nearly as many tick bites as subjects who were seropositive or had a diagnosis of the disease. Age, time spent outdoors in the fall multiplied by a clothing index, and woodcutting were significantly associated with Lyme disease in logistic regression analyses.
The efficiency of transovarial transmission of Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner was evaluated in Ixodes pacificus Cooley & Kohls collected from two areas of northern California where Lyme disease is endemic. In total, 132 (8.8%) of 1,499 replete females examined by direct immunofluorescence were demonstrated to be infected with B. burgdorferi. Larvae or eggs from 119 of these females were examined for the presence of spirochetes by direct immunofluorescence, placing them in culture, or both; none was found to contain B. burgdorferi. The fecundity of 20 midgut-infected (mean = 874.2) and 20 uninfected (mean = 1,048.3) I. pacificus females did not differ statistically. Likewise, the fertility of infected (mean = 87.0%) and uninfected (mean = 89.9%) females and the mean engorged weights of both groups (infected, 120.8 mg versus uninfected, 132.7 mg), were comparable. The fecundity, fertility, and mean weights of six replete females having ovarian infections, six females having midgut-restricted infections, and six uninfected females were also similar. We conclude that transovarial transmission is not efficient for maintaining B. burgdorferi in populations of I. pacificus, a known vector of that pathogen. Infection with the spirochete does not appear to affect either feeding or reproductive success adversely in females of this tick.
Reciprocal crosses between Ixodes dammini Spielman, Clifford, Piesman & Corwin from Massachusetts and Ixodes scapularis Say from Georgia produced offspring through the F3 generation when the experiment was discontinued. Reciprocal I. dammini x Ixodes pacificus Cooley & Kohls (California) and I. scapularis x I. pacificus crosses produced F1 progeny; however, all progeny were sterile. Assortative mating experiments between I. dammini and I. scapularis indicated that males and females of both species mated with the opposite sex of heterospecific or conspecific ticks when there was a choice. Conventional discriminant analysis of morphometric measurements of ticks from Georgia, North Carolina, Maryland, Massachusetts, and two populations of F1 hybrids indicated that there were recognizable differences. However, size-free (sheared) discriminant analysis indicated that these differences were largely size-dependent, with much overlap of the four eastern and two hybrid populations but no overlap with I. pacificus from California. Analysis of chromosomes (morphology and C band) indicated no differences between the Georgia and Massachusetts populations but showed a difference between them and the California population of I. pacificus. Analysis of isozymes showed that the genetic identity value for the Georgia and Massachusetts populations was within the normal range for conspecific populations, whereas the California population indicated congeneric but not conspecific relatedness to the Georgia and Massachusetts populations. Life cycle data collected under similar laboratory conditions showed no differences in length of feeding and molting periods among Georgia, Massachusetts, and California populations. These data and results of the work of other authors on tick host preferences and vector competence indicate that I. dammini is not a valid species separate from I. scapularis. Because the name Ixodes scapularis Say, 1821, has priority over the name Ixodes dammini Spielman, Clifford, Piesman & Corwin, 1979, I. dammini is relegated to a junior subjective synonym of I. scapularis (based on Article 23 of the International Code of Zoological Nomenclature).
A standard system for infecting ticks with the Lyme disease spirochete Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner is defined. Rodents infected via tick feeding or inoculation of tick homogenates were more infectious to ticks than rodents infected with culture-derived spirochetes. White laboratory mice were more infectious than hamsters. Three strains of B. burgdorferi (JD1, B31, and WI210) produced batches of infected ticks with > 80% infection when mice were infected with tick-derived material. Ixodes dammini Spielman, Clifford, Piesman & Corwin were 3.6x more efficient than I. pacificus Cooley & Kohls in acquiring and maintaining infection with two California strains of B. burgdorferi, originally isolated from I. pacificus.
Protozoal parasites of the genus Babesia were isolated for the first time from free-ranging desert bighorn sheep (Ovis canadensis nelsoni) and mule deer (Odocoileus hemionus) populations in California by in vitro culture of host blood. These naturally infected animals did not have microscopically detectable parasitemia at the time blood was collected for parasite cultivation. Three isolates of small Babesia parasites were cultured from different sample groups of bighorn sheep, and two isolates of large Babesia parasites were cultured from a group of bighorn sheep and a group of mule deer, respectively. The size and structure of the various forms of piroplasms from each isolate remained consistent throughout the period of cultivation. Statistical comparison of the sizes of the piroplasms among the isolates indicated that there were at least two distinct morphotypes. Four of the five isolates were maintained with continuous growth in cultures containing erythrocytes from uninfected donor bighorn sheep, mule deer, and domestic sheep. Cryopreservation or storage of cultures at 4 C for seven days did not affect viability of the isolates. These results demonstrate the potential for use of in vitro cultivation methods for the isolation of Babesia parasites from free-ranging artiodactylids.
The seasonal activity and spatial distribution of adult and immature Ixodes pacificus Cooley & Kohls and Dermacentor occidentalis Marx were determined along trails and on hillsides in two parks in Contra Costa County, CA. I. pacificus and D. occidentalis adults were most numerous in January and May, respectively. Adult ticks were significantly more abundant along heavily vegetated trails than on open grassy hillsides, and on the uphill versus the downhill side of trails. Five species of rodents were captured, and numbers of I. pacificus and D. occidentalis larvae per rodent were highest in May-June and July, respectively. Few nymphs were recovered either by flagging or from captured rodents. An average of 2.2 and 2.8% of the I. pacificus adults collected from the two parks were infected with the Lyme disease spirochete, Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner. The greatest risk of contracting Lyme disease from adult I. pacificus in these two Contra Costa County parks is during the winter months, especially while hiking near the uphill side of trails.
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This article reviews epizootiology, public health considerations, antibody testing, and molecular biology of Lyme borreliosis. Correlation of clinical signs with titer response is discussed.
Three phyletic groups of Borrelia associated with Lyme disease, B. burgdorferi, B. garinii and group VS461 can be distinguished from each other and other species of Borrelia by BfaI restriction site polymorphisms in PCR amplified 16S rRNA genes. One strain isolated from an Ixodes pacificus tick in California that was previously unclassifiable was distinguishable from B. burgdorferi by an Mn/I restriction site polymorphism.
To understand the role of bighorn sheep (Ovis canadensis) in the epidemiology of anaplasmosis, we recovered a field isolate from a suspected enzootic area in southern California (USA). Whole blood was collected from three desert bighorn sheep (Ovis canadensis nelsoni) and inoculated into a susceptible splenectomized domestic sheep, calf and a susceptible spleen-intact bighorn sheep. No infection occurred in the calf, but a detectable infection developed in both the domestic sheep and bighorn sheep 24 days after inoculation. The infection in both domestic and bighorn sheep resulted in severe clinical disease but was resolved with the use of tetracycline. Using monoclonal antibodies and DNA probes, we confirmed that the isolate was Anaplasma ovis.
Using an indirect immunofluorescence assay, we determined the prevalence of Anaplasma-reactive antibody in three herds of bighorn sheep, each a different subspecies and occupying a different habitat in California (USA). Antibodies to Anaplasma spp. were identified in none of twenty California bighorn (Ovis canadensis californiana) sampled from the Mt. Baxter herd, 11 of 17 peninsular bighorn (O. canadensis cremnobates) sampled in the Santa Rosa Mountains, and all 20 desert bighorn (O. canadensis nelsoni) sampled at Old Dad Peak/Kelso Mountains. Based on an assay and an adsorption technique, the titers most likely were due to Anaplasma ovis. The presence and species of tick vectors in each of the habitats, and the presence or absence of deer or livestock were identified as factors potentially influencing seroprevalence of antibodies.
The effect of biweekly flagging on adult Ixodes pacificus Cooley & Kohls abundance was determined by using mark-recapture techniques. Ticks were recaptured up to three times during the 12-wk study, and increasingly higher proportions of marked ticks were recaptured as the study progressed. By week 6 of the study, 46% of the collected ticks had been marked, and on the final sampling date, all ticks had been captured previously. Removal sampling would have substantially reduced estimates of the abundance of the questing I. pacificus adult population. For ecological studies that require the repeated sampling of a given habitat, the most representative data are obtained by returning flagged ticks to their collection site.
Previous studies describing the occurrence and molecular characteristics of Lyme disease spirochetes, Borrelia burgdorferi, from California have been restricted primarily to isolates obtained from the north coastal region of this large and ecologically diverse state. Our objective was to look for and examine B. burdorferi organisms isolated from Ixodes pacificus ticks collected from numerous regions spanning most parts of California where this tick is found. Thirty-one isolates of B. burgdorferi were examined from individual or pooled I. pacificus ticks collected from 25 counties throughout the state. One isolate was obtained from ticks collected at Wawona Campground in Yosemite National Park, documenting the occurrence of the Lyme disease spirochete in an area of intensive human recreational use. One isolate from an Ixodes neotomae tick from an additional county was also examined. Sodium dodecyl sulfate-polyacrylamide gel electrophoresis, immunoblot analysis, agarose gel electrophoresis, Southern blot analysis, and the polymerase chain reaction were used to examine the molecular and genetic determinants of these uncloned, low-passage-number isolates. All of the isolates were identified as B. burgdorferi by their protein profiles and reactivities with monoclonal and polyclonal antibodies, and all the isolates were typed by the polymerase chain reaction as North American-type spirochetes (B. burgdorferi sensu stricto). Although products of the ospAB locus were identified in protein analyses in all of the isolates, several isolates contained deleted forms of this locus that would result in the expression of chimeric OspA-OspB proteins. The analysis of OspC demonstrated that this protein was widely conserved among the isolates but was also quite variable in its molecular mass and the amount of it that was expressed.
We tested 276 sera from 18 free-ranging black-tailed and mule deer (Odocoileus hemionus) herds in California (USA) collected from 1987 to 1991 in five biogeographical habitat types, for antibodies against eight infectious disease agents. Overall antibody prevalence was 56% for Anaplasma marginale, 31% for Borrelia burgdorferi, 16% for bluetongue virus serotype 17, 15% for epizootic hemorrhagic disease virus, 7% for Coxiella burnetii and Toxoplasma gondii, respectively, and 0% for bovine leukosis virus and caprine arthritis/encephalitis virus, respectively. Antibodies against Lyme borreliosis and anaplasmosis were found in deer throughout California, but antibodies against bluetongue and epizootic hemorrhagic disease were most prevalent in deer from southern California.
OBJECTIVE--To determine the reason(s) why serologic tests for Lyme disease are performed, who initiates the test requests, and how the test results are used in a clinical setting. DESIGN--Retrospective cohort. SETTING--Prepaid health plan in northern California. PATIENTS--Consecutive sample of 117 patients for whom at least one serologic test for Lyme disease was performed during a 3-month period. MAIN OUTCOME MEASURES--Reason for ordering and result of the serologic test, differential diagnoses, and treatment. RESULTS--One of 117 patients had antibodies to Borrelia burgdorferi. Fifty-six percent of test requests were initiated by the physician and 35% by the patient. Of 66 tests ordered by the physician, 20% were performed because of suspected early Lyme disease, 6% as follow-up of a tick bite, 2% to confirm a prior history of Lyme disease, 14% as a workup for arthritis, and 60% as one of a battery of laboratory tests for vague symptoms. Of 41 tests initiated by the patient, 51% were performed because of a history of a tick bite. The reasons for ordering 10 tests were undetermined. CONCLUSION--Only 19% of all serologic tests for Lyme disease were performed because the physician suspected Lyme disease in the patient. Particularly in light of the low probability of contracting Lyme disease in California, it appears that this serologic test is being overused. Indiscriminate testing increases health care costs and does not appear to affect treatment decisions. Education is needed regarding the limitations of this serologic test.
The vector competence of the western black-legged tick, Ixodes pacificus Cooley & Kohls, and the Pacific Coast tick, Dermacentor occidentalis Marx, for the Lyme disease spirochete (Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner) was compared. Rabbits, hamsters, and the deer mouse, Peromyscus maniculatus (Wagner), were injected with cultured spirochetes or infected tick-suspensions, or were fed upon by spirochete-infected ticks. Five of seven isolates used as inocula were reisolated from vertebrates with the ear-punch biopsy technique. Three isolates (CA4, 5, 7) that were infectious for both vertebrates and ticks possessed prominent low-molecular-weight protein bands that had relative mobilities of approximately 24-26 kd. The ability of ticks to acquire and maintain various inocula of B. burgdorferi was evaluated by feeding uninfected larvae xenodiagnostically on all three hosts 0-63 d postinjection. Low percentages (0-10.6%) of the I. pacificus and none of the D. occidentalis became infected. By contrast, 33% of I. pacificus and 40% of Ixodes scapularis Say (= I. dammini Spielman, Clifford, Piesman & Corwin) that fed on hamsters infected by tick-bite acquired and transstadially passed spirochetes; 10% of D. occidentalis fed on infected hamsters similarly acquired but did not maintain spirochetes. Ixodes pacificus nymphs efficiently transmitted B. burgdorferi to deer mice and a hamster. Feeding by one spirochete-infected nymph was sufficient to produce patent infections in each of five mice.
To determine the incidence of physician-diagnosed Lyme disease in an endemic area of California, an active surveillance program was implemented in Lake, Mendocino, Sonoma, and southern Humboldt counties. More than 200 medical care providers were called monthly for their list of suspected cases of Lyme disease. Pertinent information was abstracted from the medical record of each patient. Of 153 cases of possible early Lyme disease ascertained from July 1991 to December 1992, 37% consisted of physician-diagnosed erythema migrans. Only 58% of erythema migrans rashes were at least 5 cm in diameter. An additional 43 patients had suspicious rashes not classified as erythema migrans. Of 166 patients with possible late-stage Lyme disease, 31% had specific clinical symptoms and 75% had a positive serologic test. With an incident case defined as physician-diagnosed erythema migrans of at least 5 cm in diameter, the annual incidence of Lyme disease in northwestern coastal California according to active surveillance only was 5.5 per 100,000. The rate of Lyme disease in California is substantially lower than that in the Atlantic northeastern United States. Many suspected cases of Lyme disease in this endemic area do not meet surveillance criteria, which are intentionally restrictive. Although some of the illnesses not meeting surveillance criteria may be due to infection with Borrelia burgdorferi, it appears that Lyme disease is being overdiagnosed in this area.
Sequestration of spirochetes and concurrent histopathologic lesions were evaluated in tissues of Borrelia burgdorferi-infected dusky-footed woodrats (Neotoma fuscipes) and deer mice (Peromyscus maniculatus). Rodents were either wild-caught and naturally infected or were experimentally infected by tick bite, by intramuscular (i.m.) injection of cultured spirochetes, or by i.m. injection of tick suspensions. Samples of host tissues, including skin, blood, ear, brain, eye, heart, lung, liver, spleen, kidney, and urinary bladder, were removed from up to 21 woodrats and four deer mice and cultured in BSK II medium. Borreliae-positive cultures of ear punch biopsies were obtained from 10 of 11 woodrats and from all of four deer mice. Additionally, positive cultures were obtained from three of 36 skin biopsies of woodrats, and from one of 36 cultures of woodrat blood. In contrast, spirochetes were not observed in 505 cultures of internal organs or whole blood. Samples of tissues from seven naturally infected woodrats, four experimentally infected woodrats, and nine experimentally infected deer mice also were examined for histopathologic lesions. Nonsuppurative cellular infiltrates were recognized in samples from most tissue types from woodrats, but few lesions were observed in tissues from deer mice. Recognized lesions in woodrats that were consistent with infections of Lyme borreliosis in other species included synovitis, myositis, and myocarditis. Such lesions were more common in woodrats than in deer mice. Inflammatory lesions, especially synovitis, were more common in woodrats with long-term infections than in woodrats with relatively short-term infections. No clinical signs of disease were observed in either species of rodent.
A 4-year-old male mixed-breed dog from the Sierra Nevada mountains in California was referred because of epistaxis and signs of cervical pain. Dermacentor variabilis ticks were found on the dog at the time of physical examination. Clinicopathologic abnormalities included nonregenerative anemia, thrombocytopenia, and rare intracytoplasmic morulae within circulating neutrophils. Abnormalities of the CSF included pleocytosis and intracytoplasmic morulae in approximately 9% of neutrophils. Serum antibody titers for Ehrlichia canis (40,960) and Rickettsia rickettsii (5,120) were high, and titer for E equi (40) was moderate. Treatment included administration of tetracycline, chloramphenicol, doxycycline, and prednisone. The dog had several relapses, but long-term remission was eventually achieved. Granulocytic ehrlichiosis has previously been associated with anemia, thrombocytopenia, and polyarthritis in dogs. This case suggests that granulocytic ehrlichiosis may be associated with meningitis and that the organisms that cause granulocytic ehrlichiosis may have the same vector as do the spotted fever-group rickettsiae.
BACKGROUND. Human babesiosis is a tick-transmitted zoonosis associated with two protozoa of the family Piroplasmorida: Babesia microti (in the United States) and B. divergens (in Europe). Recently, infection with an unusual babesia-like piroplasm (designated WA1) was described in a patient from Washington State. We studied four patients in California who were identified as being infected with a similar protozoal parasite. All four patients had undergone splenectomy, three because of trauma and one because of Hodgkin's disease. Two of the patients had complicated courses, and one died. METHODS. Piroplasm-specific nuclear small-subunit ribosomal DNA was recovered from the blood of the four patients by amplification with the polymerase chain reaction. The genetic sequences were compared with those of other known piroplasm species. Indirect immunofluorescent-antibody testing of serum from the four patients and from other potentially exposed persons was performed with WA1 and babesia antigens. RESULTS. Genetic sequence analysis showed that the organisms from all four patients were nearly identical. Phylogenic analysis showed that this strain is more closely related to a known canine pathogen (B. gibsoni) and to theileria species than to some members of the genus babesia. Serum from three of the patients was reactive to WA1 but not to B. microti antigen. Serologic testing showed WA1-antibody seroprevalence rates of 16 percent (8 of 51 persons at risk) and 3.5 percent (4 of 115) in two geographically distinct areas of northern California. CONCLUSIONS. A newly identified babesia-like organism causes infections in humans in the western United States. The clinical spectrum associated with infection with this protozoan ranges from asymptomatic infection or influenza-like illness to fulminant, fatal disease.
The transmission of Borrelia burgdorferi to deer mice (Peromyscus maniculatus) by Ixodes pacificus nymphs was investigated experimentally. Deer mice were exposed to infected nymphs for 24, 48, or 72 hr, or until ticks had fed to repletion (> or = 96 hr). Infection status of hosts was assessed 4 weeks later by culture of ear-punch biopsies in BSK II medium and by indirect immunofluorescence. Eight mice exposed to ticks for 24 hr did not become infected. In contrast, infection was acquired by one of nine (11%), two of eight (25%), and eight of 10 (80%) mice exposed for 48, 72, and > or = 96 hr, respectively. Eight weeks after exposure to infected nymphs, the infectivity of five deer mice for I. pacificus larvae was assessed. Overall, 33% of I. pacificus larvae fed on these mice acquired and transstadially passed spirochetes. We conclude that most I. pacificus nymphs require four days or longer to transmit spirochetes to deer mice, and that larvae efficiently acquire and maintain spirochetes from mice that have been infected by tick-bite.
To clarify the role of nymphal versus adult western black-legged ticks (Ixodes pacificus) in the epidemiology of Lyme disease, the seasonal distribution, abundance, and spirochetal infection rates in these stages, and the seasonal occurrence of ticks biting humans and of incident cases of Lyme disease were determined in northern California. Although their seasonal activity periods overlapped for about one-third of the year, nymphs and adults predominated in different seasons, the former from late spring to summer and the latter from fall to early spring. At one site, four (4%) of 100 adults from low vegetation bordering a hardwood forest and 44 (13.6%) of 324 nymphs from leaf litter in the forest were found to contain Borrelia burgdorferi. Biting-collection records revealed that nymphs attach to people more commonly than recognized previously; I. pacificus nymphs comprised 12.5% of 967 ticks of various species and stages and 42% of all nymphs submitted for identification. Attachments by nymphs occurred primarily between April and August, which coincided with the seasonal occurrence of most incident cases of Lyme disease. Collectively, these findings strongly implicate the nymphal stage of I. pacificus as the primary vector of B. burgdorferi to humans in this region.
Sera from 111 bighorn sheep (Ovis canadensis) and 95 mule deer (Odocoileus hemionus) were tested using an indirect immunofluorescence assay for antibodies to two isolates of Babesia spp. recently obtained from these hosts in California (USA). The study populations were from six locations: three areas of real or potential sympatry of bighorn sheep and deer, one area with deer only, and two areas with bighorn sheep only. Antibody titers from seroreactive individuals were similar with both babesial isolate antigens (P < 0.05), and seroprevalence was highest in the areas of host sympatry. A moderate to high seroprevalence (> or = 30%) in some of the study populations was evidence that babesial parasites may be common in bighorn sheep and mule deer in some areas of California.
Aspects of the reservoir competence of four rodents for the Lyme disease spirochete, Borrelia burgdorferi, were evaluated in California. Rodents were live-trapped and ear-punch biopsies were cultured during each season. A second set of biopsies was cultured from representative individuals after 2-3 weeks of captivity and the results of culturing biopsies taken on both dates were compared with the results of feeding Ixodes pacificus larvae on hosts xenodiagnostically. The prevalence of infections did not differ significantly between dusky-footed woodrats (Neotoma fuscipes) and California kangaroo rats (Dipodomys californicus) nor among seasons. Combined results of the three tests showed that 85.7% of dusky-footed woodrats (n = 21) and 78.6% of California kangaroo rats (n = 14) were infected with B. burgdorferi. In contrast, only 22.2% of brush mice (Peromyscus boylii) (n = 14) and 7.1% of pinyon mice (P. truei) (n = 9) were infected. The sensitivity of culturing ear-punch biopsies as an assay for borrelial infection was significantly greater when biopsies were taken after a short period of captivity (0.89) rather than on the day of capture (0.52). Tick xenodiagnosis, in which I. pacificus was used as the vector, revealed borrelial infections in 90.3% of infected rodents. Spirochetes were observed in 37.7% of 239, 45.2% of 155, 60.0% of 10, and 7.1% of 14 cultures of nymphal I. pacificus fed as larvae on naturally infected woodrats, kangaroo rats, brush mice, and a pinyon mouse, respectively. The mean prevalence of infection in xenodiagnostic ticks varied significantly among host species with a greater proportion of ticks infected while feeding on woodrats and kangaroo rats than on mice. This study reconfirms previous reports that implicated woodrats and kangaroo rats as reservoirs of B. burgdorferi in California.
Ehrlichia equi, a rickettsia described from horses in California 30 years ago, causes equine granulocytic ehrlichiosis throughout the Americas and possibly Europe. Here, we report experimental transmission of E. equi from infected to susceptible horses through bites of western blacklegged ticks, Ixodes pacificus (Cooley & Kohls). In preliminary field studies, only I. pacificus consistently infested horses and vegetation at three locations with contemporary cases of equine ehrlichosis, and in particular, I. pacificus was the only species found attached to all of the infected horses. Exposure to bites of ticks in the genus Ixodes poses previously unrecognized and serious health risks to humans and animals.
The infectivity and dissemination to the skin of six isolates of Borrelia burgdorferi were evaluated by inoculating them into groups of deer mice (Peromyscus maniculatus), hamsters, and Swiss Webster mice. Rodent infection was assayed by culture of ear punch biopsy specimens taken at 4, 8, and 12 weeks postinoculation (p.i.). Spirochetes were detected in biopsy specimens from individuals of all three host species that had been inoculated with four isolates (CA3, CA4, CA7, and CA8). Ear punch biopsy specimens taken from Swiss Webster mice at 12 weeks p.i. yielded an additional reisolate (CA2), even though these animals did not seroconvert. The remaining isolate (CA9) was not recovered from any host. However, two deer mice and all hamsters and Swiss Webster mice inoculated with CA9 seroconverted. All six isolates were of low infectivity to ticks when inoculated intramuscularly into hosts. Only 4 (1.6%) of 250 Ixodes pacificus larvae acquired and transstadially maintained infection from hosts inoculated intramuscularly. Infectivity of three isolates for ticks also was tested in Swiss Webster mice injected intradermally. The mean prevalences of infection in xenodiagnostic ticks fed on these mice at 4 weeks p.i. were 47.9, 1.2, and 2.2% for isolates CA4, CA7, and CA8, respectively. The mean prevalences of infection for ticks fed on the same mice at 12 weeks p.i. were 36.4, 11.8, and 20.4%, respectively. Such differences in the infectivity and rate of dissemination of individual isolates of B. burgdorferi should be considered during studies of reservoir and vector competence.
Ehrlichiae are small, gram-negative, obligately intracellular bacteria that reside within a phagosome. The first human ehrlichial infection was recognized in the United States in 1987. It was later shown to be caused by a new species, Ehrlichia chaffeensis. In 1994, an ehrlichial pathogen within neutrophils that is closely related to the known veterinary pathogens E. equi and E. phagocytophila was found to infect humans. Molecular methods were required to detect, characterize, and identify these fastidious and uncultivated bacteria. Subsequently, E. chaffeensis infection was documented in more than 400 patients in 30 states, Europe, and Africa. Likewise, approximately 170 cases of human granulocytic ehrlichiosis have been diagnosed, most since 1994, predominantly in the upper midwestern and northeastern states, but also in northern California. The disease caused by ehrlichiae is generally undifferentiated but is often associated with leukopenia, thrombocytopenia, and elevated serum hepatic transaminase levels in tick-exposed patients. Infection ranges from subclinical to fatal; tetracycline appears to be an effective therapy. The emergence of these two newly recognized tickborne infections as threats to human health is probably due to increased clinical cognizance, but as in other emerging tickborne infections, it is likely that the rapid increase in identified cases signals a true emergence of disease associated with a changing vector-host ecology.
Nymphal and larval stages of Ixodes (Ixodes) jellisoni Cooley & Kohls and I. (I.) neotomae Cooley are described for the first time. These two tick species occur only in the western United States, predominantly in California. The primary host for I. jellisoni is the California kangaroo rat, Dipodomys californicus (Merriam); that for I. neotomae is the dusky-footed woodrat, Neotoma fuscipes Baird. The etiologic agent of Lyme disease Borrelia burgdorferi Johnson, Schmidt, Hyde, Steigerwalt & Brenner has recently been isolated from both tick species, and I. neotomae was proven a competent enzootic vector of the Lyme disease spirochete.
The dusky-footed woodrat, Neotoma fuscipes Baird is a natural reservoir of the Lyme disease spirochete, Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner, in California. To investigate the potential of host-targeted insecticide to control the tick vectors of B. burgdorferi, permethrin-impregnated or untreated cotton balls were distributed in metal cylinders as potential nesting material adjacent to 95 woodrat houses in chaparral-covered rangeland. Laboratory experiments demonstrated that adult woodrats would enter the cylinders and construct nests from permethrin-treated or untreated cotton. The residual concentration of permethrin did not vary significantly during an 11-mo period and remained > 60% of the registered insecticidal formulation (7.5% [AI] by cotton weight). The abundance of four species of ticks (Ixodes neotomae Cooley; the western blacklegged tick I. pacificus Cooley & Kohls; I. woodi Bishopp; and the Pacific Coast tick, Dermacentor occidentalis Marx) infesting woodrats was similar in the treatment and control areas. Although > 90% of the cotton disappeared from the metal cylinders in both areas, examination of eight active woodrat houses revealed that small amounts of cotton had been incorporated into the nest cups of only 25%. In contrast, the abundance of the flea Orchopeas sexdentatus (Baker) decreased significantly in the treatment area only. Spirochetes were not detected in 168 adult O. sexdentatus fleas that had fed on spirochetemic woodrats, which demonstrates that this flea is an inefficient host of B. burgdorferi. We conclude that the use of permethrin-impregnated cotton as potential nesting material is ineffective for controlling ticks associated with the dusky-footed woodrat in brushlands, but this methodology may be useful for reducing populations of sylvatic fleas.
Babesiosis is a malaria-like, tick-transmitted zoonosis caused by protozoa of the family Piroplasmorida, which includes Babesia and Theileria species. In the United States, the infection is endemic in the Northeast and upper Midwest, although cases have recently been described in Northern California and Washington State. We report a case of babesiosis in a patient infected with HIV who presented with a prolonged fever of unknown origin; the patient had not undergone splenectomy. Parasitemia persisted despite initial clinical improvement after treatment with quinine and clindamycin. Babesiosis was controlled with a maintenance regimen consisting of clindamycin, doxycycline, and high-dose azithromycin, but the infection was not eradicated. Babesiosis should be considered in the differential diagnosis of HIV-infected patients with fevers and/or anemia in areas where the infection is endemic. HIV-infected patients who are severely immunosuppressed, even those without a history of splenectomy, may present with severe manifestations of babesiosis and develop a chronic infection, which may require therapy to prevent relapse of disease.
No abstract available.
The risk of humans encountering vector ticks along hiking trails or in picnic grounds in a Lyme disease-endemic area was evaluated in a multipurpose recreational area (Tilden Regional Park) in the populous San Francisco Bay region of California. Four hillside hiking trails (two high-use, two low-use) were sampled by dragging and walking through low vegetation biweekly for one year; four heavily used picnic areas were sampled concurrently by dragging. Adults of three human-biting ticks were enumerated (n = 1,911) along all trials: Dermacentor occidentalis (63.6% of total), Ixodes pacificus (26.2%), and D. variabilis (0.2%). Subadults (n = 1,669) of D. occidentalis (0.06% of total) and two nonhuman-biters, D. albipictus (70.3%) and Haemaphysalis leporispalustris (29.7%), also were collected. Dragging yielded many more adult ticks than walking year-round for all trials. These methods were significantly correlated during periods of peak tick abundance, but the associations were not sufficiently strong or consistent to allow prediction of captures for either method based on the other. Adult ticks were distributed largely in clusters along the uphill sides of trails. Several adult ticks collected adjacent to trails were found to contain spirochetes identified with polyclonal antibodies as Borrelia burgdorferi (D. occidentalis, 0 of 861; D. variabilis, two of 126 [1.6%]; I. pacificus, 1 of 609 [0.2%]). Picnic areas produced low numbers of adult D. occidentalis and I. pacificus, which prohibited testing them for spirochetes. Two measures for evaluating risk were calculated, the encounter distance (= mean number of meters traveled before encountering a vector tick by either dragging or walking) and the mean number of spirochete-infected ticks encountered by these methods per kilometer of trial. These measures revealed that the risk of exposure to spirochete-infected adult ticks along trails was low year-round irrespective of usage, and risk was even lower in picnic areas. Future studies evaluating human exposure to vector ticks in recreational areas should incorporate, whenever possible, testing for multiple pathogens because most ixodid ticks that commonly bite people in the United States are capable of transmitting two or more microbial disease agents.
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Arthropod-borne viruses are important causes of diseases of the central nervous system. In addition to the tick-borne encephalitis viruses, other arboviruses in Europe are known to cause neurological disorders. Among them are West Nile, California group, Bhanja, Erve, Kemerovo group, Eyach, and Thogoto viruses. The ecologies and epidemiologies of these viruses are presented and their medical importance as travel-related diseases is discussed.