Survival and Growth of Francisella tularensis in = Acanthamoeba castellanii

Journal = List >=20 Appl=20 Environ Microbiol > v.69(1); = Jan=20 2003

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Appl Environ Microbiol. 2003 January; = 69(1):=20 600=E2=80=93606.

doi: 10.1128/AEM.69.1.600-606.2003.

Survival and Growth of Francisella = tularensis=20 in Acanthamoeba castellanii

Hadi Abd,^1,² Thorsten=20 Johansson,¹ Igor Golovliov,³ Gunnar=20 Sandstr=C3=B6m,² and Mats = Forsman¹^*

Swedish Defence Research Agency, SE 901 82=20 Ume=C3=A5,¹ Clinical Bacteriology, Department of = Clinical=20 Microbiology, Ume=C3=A5 University, SE 901 85 = Ume=C3=A5,³ Division of=20 Clinical Bacteriology, Department of Microbiology, Pathology and=20 Immunology, Karolinska Institute, Huddinge University Hospital, SE = 141 86=20 Stockholm, Sweden²

^*Corresponding author. Mailing address: = Swedish=20 Defense Research Agency, SE 901 82, Ume=C3=A5, Sweden. Phone: 46 = 90 106669.=20 Fax: 46 90 106800. E-mail: mats.forsman@foi.se. = =E2=80=A0 3D"

Present address: Division of Clinical Bacteriology, = Department of=20 Microbiology, Pathology and Immunology, Huddinge University = Hospital, SE=20 141 86 Stockholm, Sweden.

Received June 10, 2002; Accepted October 15, 2002.

= This=20 article has been cited=20 by other articles in PMC.

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MATERIALS AND=20 METHODS

RESULTS

DISCUSSION

REFERENCES

Abstract

Francisella tularensis is a highly infectious, = facultative=20 intracellular bacterium which causes epidemics of tularemia in = both humans=20 and mammals at regular intervals. The natural reservoir of the = bacterium=20 is largely unknown, although it has been speculated that protozoa = may=20 harbor it. To test this hypothesis, Acanthamoeba = castellanii was=20 cocultured with a strain of F. tularensis engineered to = produce=20 green fluorescent protein (GFP) in a nutrient-rich medium. GFP=20 fluorescence within A. castellanii was then monitored by = flow=20 cytometry and fluorescence microscopy. In addition, extracellular = bacteria=20 were distinguished from intracellular bacteria by targeting with=20 monoclonal antibodies. Electron microscopy was used to determine = the=20 intracellular location of F. tularensis in A. = castellanii,=20 and viable counts were obtained for both extracellular and = intracellular=20 bacteria. The results showed that many F. tularensis cells = were=20 located intracellularly in A. castellanii cells. The = bacteria=20 multiplied within intracellular vacuoles and eventually killed = many of the=20 host cells. F. tularensis was found in intact trophozoites, = excreted vesicles, and cysts. Furthermore, F. tularensis = grew=20 faster in cocultures with A. castellanii than it did when = grown=20 alone in the same medium. This increase in growth was accompanied = by a=20 decrease in the number of A. castellanii cells. The = interaction=20 between F. tularensis and amoebae demonstrated in this = study=20 indicates that ubiquitous protozoa might be an important = environmental=20 reservoir for F. tularensis.

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Abstract

MATERIALS AND=20 METHODS

RESULTS

DISCUSSION

REFERENCES

Francisella tularensis, a gram-negative, nonsporulating, = nonmotile, and capsulated coccobacillus, is a facultative = intracellular=20 bacterium which causes tularemia in humans and animals (37).=20 Inoculation or inhalation of as few as 10 F. tularensis = organisms=20 may cause disease in humans (35,=20 36),=20 and because of its high virulence and contagiousness, the = bacterium is=20 considered a possible bioterrorism agent (7).=20 It has been isolated from approximately 250 wildlife species (37),=20 but its principal natural reservoir is largely unknown, although = studies=20 have shown that the organism may persist for more than a year in = water or=20 mud (30).

Virtually nothing is known about the virulence mechanisms of = F.=20 tularensis or about how it survives within host cells. Even = the=20 avirulent live vaccine strain (LVS), which has been used as a = human=20 vaccine (34),=20 is strongly pathogenic for certain animals and causes a lethal = infection=20 in mice that is indistinguishable from human disease (8).=20 It has been shown that LVS is very well adapted to the = intracellular=20 environment of macrophages, survives in phagosomes (3,=20 13),=20 and exerts a cytopathogenic effect on murine macrophages (3a,=20 5,=20 13)=20 Furthermore, it has been found that LVS localizes within an acidic = vesicle, which facilitates its iron uptake (11),=20 and that LVS releases an acid phosphatase that inhibits the = respiratory=20 burst in neutrophils (33).=20 Recently, it has also been demonstrated that infection with the = LVS strain=20 induces apoptosis in murine macrophages (24).

Free-living amoebae such as Acanthamoeba species are = commonly=20 found in natural aquatic systems (25).=20 As a part of biofilms in aquatic environments, free-living amoebae = and=20 bacteria are involved in complex interactions. Francisella = species=20 are also known to be connected with natural water systems (18,=20 30),=20 and it is well known that Acanthamoeba species are = environmental=20 hosts of several intracellular pathogens, such as = Legionella,=20 Chlamydia, and Mycobacterium (2,=20 26,=20 38,=20 39).=20 Legionella cells interact with their protozoan hosts and = mammalian=20 cells in a similar way. The known genetic factors required by=20 Legionella to infect protozoa are also required for the = infection=20 process in mammalian cells (16,=20 17).=20 Previous studies of the interaction between F. tularensis = and=20 Acanthamoeba castellanii have shown that growth of F.=20 tularensis is enhanced in media preconditioned by amoebae (15).=20 Here, we have tested the hypothesis that protozoa may comprise a=20 significant environmental reservoir for F. tularensis.

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RESULTS

DISCUSSION

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MATERIALS AND METHODS

Microorganisms and plasmid construction. = A.=20 castellanii (ATCC 30234) was obtained from the American Type = Culture=20 Collection (ATCC), Manassas, Va., and F. tularensis LVS = (type B)=20 was obtained from the U.S. Army Medical Research Institute of = Infectious=20 Diseases, Fort Detrick, Md. F. tularensis LVS carrying a=20 destabilized form of green fluorescent protein (GFP) was = constructed as=20 follows. First, the promoter for GroEL from F. tularensis = LVS (9)=20 was amplified by PCR, cloned into the SmaI-PstI = sites=20 upstream of the chloramphenicol acetyltransferase (CAT) gene in = pKK214 (23,=20 29),=20 and then introduced into F. tularensis LVS by = cryotransformation=20 (31).=20 The resulting plasmid was designated pKK214CAT and was used as the = basic=20 construct in the present work. A mutant form (M2) of wild-type GFP = (6)=20 was amplified by PCR and cloned into the PstI-EcoRI = sites of=20 the pKK214CAT plasmid under the control of the GroEL promoter. = Because the=20 EcoRI site is inside the CAT gene, this procedure led to = the=20 partial removal of the CAT gene. The resulting plasmid was = designated=20 pKK214GFP. Next, a destabilized variant (ASV) of GFP (Clontech, = Palo Alto,=20 Calif.) was amplified by PCR and cut with = Bsp1407I-EcoRI,=20 and the resulting Bsp1407I-EcoRI fragment was cloned = into=20 pKK214GFP cut with the same enzymes. This resulted in substitution = of the=20 C-terminal part of GFP (M2 form) for the destabilized C-terminal = form=20 (ASV), which contains sequences encoding protease targets = responsible for=20 the rapid degradation of the expressed protein. This construct was = designated pKK214GFP/ASV. To determine the half-life of this = GFP/ASV=20 construction in F. tularensis LVS, samples containing 2.0 = =C3=97=20 10⁹ CFU/ml were inactivated by treatment with 250 = =CE=BCg of=20 gentamicin (Sigma, St. Louis, Mo.) per ml for 1 h at room = temperature in=20 darkness. After confirming that the cells were unable to grow, as=20 determined by viable counts, following this treatment, viable = count and=20 flow cytometry analyses were carried out in parallel for 7 days. = No growth=20 on modified Thayer-Martin agar was observed, and the fluorescence=20 gradually decreased.

Culture media and growth conditions. = A.=20 castellanii was grown without shaking at 30=C2=B0C to a final = concentration=20 of 10⁶/ml in ATCC medium no. 712 (ATCC). Modified = Thayer-Martin=20 agar plates containing 36 g of GC base medium (Difco Laboratories, = Detroit, Mich.) per liter, 10 g of hemoglobin (Difco) per liter, = 10 mg of=20 IsoVitaleX (BBL Microbiology Systems, Cockeysville, Md.) per = liter, and 10=20 mg of tetracycline (Sigma) per liter were used for viable counts = and=20 growth of F. tularensis LVS/GFP/ASV. After the inoculum was = spread=20 as serial dilutions, the agar plates were incubated at 37=C2=B0C = in 5%=20 CO₂ for at least 2 days if not otherwise stated. = Cocultures of=20 F. tularensis LVS/GFP/ASV and A. castellanii were = incubated=20 in Falcon tissue culture flasks (75 cm²) filled with 50 = ml of=20 ATCC medium 712 containing 20 =CE=BCg of tetracycline per ml and = an initial=20 concentration of 10⁵ A. castellanii cells and=20 10⁶ F. tularensis cells per ml. Control flasks = with the=20 microorganisms cultured separately were prepared in the same way = and with=20 the same initial concentrations. The flasks were incubated at = 30=C2=B0C without=20 shaking. Samples were withdrawn regularly for electron microscopy, = flow=20 cytometry, and fluorescence microscopy analysis and to obtain = viable=20 counts.

Flow cytometry analysis. The flow = cytometer=20 (FACSort; Becton Dickinson Immuno Systems, San Jose, Calif.), = equipped=20 with an argon laser giving a 488-nm primary emission line, was = calibrated=20 with unlabeled and labeled beads (Becton Dickinson) and FACSComp = software=20 (Becton Dickinson). Unlabeled cells were adjusted for forward = scatter=20 (relative size), side scatter (relative granularity), FL1 (green = color),=20 FL2 (red color), and FL3 (deep red color). The measuring time per = sample=20 was 15 to 50 s, with a medium flow rate of 60 =CE=BCl/min. From = each sample=20 10,000 events were registered, and data were analyzed with Cell = Quest=20 software (Becton Dickinson). Samples (3 ml) of cell suspension = from the=20 coculture flasks were centrifuged for 10 min at 300 =C3=97 = g in a model=20 TJ6 centrifuge (Beckman Instruments, Palo Alto, Calif.) and washed = six=20 times with FacsWash solution (Becton Dickinson) prior to the = analyses.

Microscopy analysis. A. = castellanii=20 cells were counted in a B=C3=BCrker counting chamber (Merck = Eurolab, Stockholm,=20 Sweden) under a light microscope (Leica Microscopy Systems, = Heerburgg,=20 Switzerland), while the abundance and distribution of F. = tularensis=20 LVS/GFP/ASV were analyzed by fluorescence microscopy (Leica = Microscopy=20 Systems). Prior to analysis, using an oil immersion lens and a = green=20 filter, 2-ml samples of cell suspension from the coculture flask = were=20 centrifuged for 10 min at 300 =C3=97 g in a Beckman model = TJ6=20 centrifuge, and the resulting pellets were washed six times with = FacsWash=20 solution.

Antibody labeling of extracellular F. tularensis = LVS/GFP/ASV was=20 performed by adding 10 =CE=BCl of biotin-labeled antibodies, = specific for F.=20 tularensis LVS (German Armed Forces Medical Academy, Munich, = Germany),=20 to 1-ml samples of cell suspension from coculture flasks. The = samples were=20 incubated for 20 min at room temperature, washed, reincubated for = 20 min=20 with 10 =CE=BCl of R-phycoerythrin-conjugated Streptoavidin (DAKO, = Glostrup,=20 Denmark), washed again, and examined under the fluorescence=20 microscope.

For electron microscopy, 2-ml samples of cell suspension from = the=20 coculture flasks were centrifuged for 10 min at 300 =C3=97 = g in a=20 Beckman model TJ6 centrifuge, and the resulting pellets were = washed six=20 times with FacsWash solution. Each pellet of infected amoebae was = fixed=20 with 2.5% glutaraldehyde in 0.1 M sodium cacodylate buffer (pH = 7.3) for 2=20 h at room temperature. The samples were then washed twice in the = same=20 buffer and fixed in 1% reduced osmium tetroxide for 1 h. The = specimens=20 were centrifuged into pellets, which were embedded in a 2% = solution of=20 agarose at 45=C2=B0C. The solidified agarose containing the = amoebae was cut=20 into small cubes, which were dehydrated in graded acetone and = embedded in=20 an Epon-Araldite mixture. Ultrathin sections were stained with = uranyl=20 acetate and lead citrate and examined with a transmission electron = microscope (EM 900; Carl Zeiss, Oberkochen, Germany).

Analysis of viable F. tularensis = within A.=20 castellanii. Samples (3 ml) of cell suspensions from = coculture=20 flasks were centrifuged for 10 min at 300 =C3=97 g in a = Beckman model=20 TJ6 centrifuge and washed six times with FacsWash solution to = minimize=20 extracellular F. tularensis contamination. The pellets were = resuspended in 500 =CE=BCl of FacsWash solution and incubated with = 250 =CE=BCg of=20 gentamicin per ml for 1 h at room temperature. Next, the samples = were=20 centrifuged for 10 min at 300 =C3=97 g in a Beckman model = TJ6=20 centrifuge, and 100-=CE=BCl portions of the supernatant were = spread on modified=20 Thayer-Martin agar plates. Each pellet was diluted twofold with = 0.5%=20 sodium deoxycholate and incubated for 5 min at room temperature, = after=20 which series of 10-fold dilutions from 10¹ to = 10⁴=20 were prepared and spread on modified Thayer-Martin agar.

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MATERIALS AND=20 METHODS

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DISCUSSION

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RESULTS

GFP fluorescence from F. tularensis = in A.=20 castellanii. Prior to the analyses of GFP fluorescence = from=20 F. tularensis in A. castellanii, we determined the = half-life=20 of the destabilized form (ASV) of the GFP expressed in F.=20 tularensis LVS/GFP/ASV. The half-life was found to be = approximately 34=20 h. This ensured that viable cells were measured, since dying cells = would=20 show lower fluorescence intensity with time.

Flow cytometer analysis of samples taken from the flasks = containing=20 both F. tularensis LVS/GFP/ASV and A. castellanii = showed=20 that the fluorescence intensity increased with time in the A.=20 castellanii population, from 0% at day 0 to 54% at day 15 = (Fig. 1).

Intracellular location of F. = tularensis in=20 A. castellanii. Samples taken from cocultures of = F.=20 tularensis LVS/GFP/ASV and A. castellanii showed an = increase in=20 the numbers of intracellular bacteria with time (Fig. 2).=20 Moreover, different stages of infection were observed, including = growth in=20 intracellular-membrane-limited vacuoles, release of vesicles, and = cysts=20 containing bacteria. Samples taken at day 18 from the coculture = were mixed=20 with R-phycoerythrin-labeled antibodies specific for F. = tularensis,=20 and the antibody-directed staining was visualized by fluorescence=20 microscopy. The results showed that GFP-labeled bacteria within = A.=20 castellanii were not accessible to the antibodies and hence = exhibited=20 green fluorescence. In contrast, extracellular bacteria were bound = by the=20 antibodies and thus exhibited red fluorescence. This demonstrated = that=20 many of the F. tularensis cells were located = intracellularly (Fig.=20 3).=20 The fluorescence microscopy also showed that F. = tularensis-infected=20 amoebae disintegrated as vesicles inside them filled with F.=20 tularensis and that bacteria occurred within cysts (Fig. 2=20 and 3).

Electron micrographs confirmed that many F. tularensis = cells=20 were located within vacuoles in A. castellanii (Fig. 4).=20 The vacuoles containing bacteria seemed to attract amoebal = organelles such=20 as mitochondria and rough endoplasmic reticulum (Fig. 4C=20 and D) As shown in Fig. 4E,=20 F. tularensis cells could be seen lining up between an = emerging=20 double wall, which is a sign of encystation. However, another = outcome was=20 also observed in some cases (Fig. 2F,=20 3C,=20 and 4F),=20 in which all the bacteria seemed to be located inside the double = wall of=20 the cyst.

Growth of cocultures of F. = tularensis=20 LVS/GFP/ASV and A. castellanii. Viable counts of=20 extracellular F. tularensis LVS/GFP/ASV increased in the = presence=20 of A. castellanii from 2 =C3=97 10⁶ CFU/ml at = day 0 to 2 =C3=97=20 10⁹ CFU/ml at day 15. In contrast, viable counts of = F.=20 tularensis LVS/GFP/ASV cultured alone decreased from 2 =C3=97=20 10⁶ CFU/ml at day 0 to nondetectable levels at day 15 = (Fig. 5).=20 Differences between the numbers of A. castellanii cells in = cultures=20 with or without F. tularensis LVS/GFP/ASV were also = apparent. The=20 total counts, according to B=C3=BCrker chamber determinations, = showed that the=20 numbers of A. castellanii cells increased from 4 =C3=97 = 10⁵=20 cells/ml at day 0 to 8 =C3=97 10⁵ cells/ml at day 20 = when cultured=20 alone and decreased from 4 =C3=97 10⁵ cells/ml at day 0 = to 3 =C3=97=20 10⁵ cells/ml at day 20 when cocultured with F.=20 tularensis LVS/GFP/ASV (Fig. 6).

The viability of the F. tularensis cells within A.=20 castellanii was analyzed by taking viable counts of the = bacteria after=20 gentamicin treatment to kill extracellular bacteria followed by=20 deoxycholate treatment of the A. castellanii cells to = release the=20 intracellular bacteria. No bacteria were recovered from = gentamicin-treated=20 medium, but in A. castellanii cells, an increase in F.=20 tularensis LVS/GFP/ASV viable counts was observed over time, = from 0=20 cells/ml at days 0, 2, 4, and 8 to 2 =C3=97 10⁶ = cells/ml at day 15=20 (Fig. 5).

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MATERIALS AND=20 METHODS

RESULTS

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DISCUSSION

F. tularensis is a highly infectious and facultative=20 intracellular bacterium that causes various types of tularemia in = mammals=20 (18).=20 It has been considered a potential biological weapon and possible=20 bioterrorism agent (7),=20 so its survival in nature is of interest from an epidemiological=20 perspective. The main reservoir of the bacterium is not known, but = it has=20 been shown that other intracellular bacteria, such as = Legionella,=20 Mycobacterium, and Chlamydia, utilize protozoa as = natural=20 reservoirs (2,=20 26,=20 38).=20 In this context, it has been shown that growth of F. = tularensis is=20 enhanced in media preconditioned by A. castellanii (15).=20 It has also been speculated that F. tularensis may persist = in A.=20 castellanii (4).

In the present study, we therefore aimed to study the = interaction=20 between F. tularensis and A. castellanii more = thoroughly.=20 The results show for the first time that F. tularensis can = survive=20 and grow within A. castellanii. The infection process = begins when=20 trophozoites engulf F. tularensis cells, which replicate = and grow=20 in vacuole structures inside the trophozoites. Thus, the infection = process=20 in amoebae shows a resemblance to Francisella infection in=20 macrophages (3,=20 11).=20 Our ultrastructural examinations showed that growth of F.=20 tularensis in A. castellanii resulted in early changes = in the=20 amoebae, such as disappearance of the nucleus and condensation of = the=20 cytoplasm (Fig. 4).=20 Moreover, cell organelles such as mitochondria and endoplasmic = reticulum=20 were recruited to the vacuoles containing bacteria (Fig. 4C=20 and D). This phenomenon has also been observed in A. = castellanii=20 infected with Legionella (1).=20 Infected trophozoites were found both intact, filled with vacuoles = containing F. tularensis, and in the process of cytolysis,=20 excreting vesicles containing F. tularensis. Some infected=20 trophozoites were also seen undergoing encystation, with two = observed=20 sequences of events: in one F. tularensis was contained = inside the=20 double wall, while in the other F. tularensis cells lined = up in the=20 space between the layers of the double wall (Fig. 3=20 and 4),=20 as previously described for Mycobacterium avium and = Simkania=20 negevensis (22,=20 38).=20 Thus, taking the observations together, the infection cycle of = F.=20 tularensis in A. castellanii seems to display many = features in=20 common with Legionella infection in A. castellanii = (17).

Viable counts of cocultured F. tularensis showed that = the=20 presence of A. castellanii enhanced its growth. This = finding is in=20 accordance with previous results (15)=20 and is probably due to the F. tularensis using = CO₂=20 produced from live A. castellanii cells and nutrients = derived from=20 dead ones. In addition, intracellular F. tularensis may = escape into=20 the culture medium after lysis of the A. castellanii = cells.

Counts of A. castellanii cells showed that there were = 25% fewer=20 cells when they were cocultured with F. tularensis than = when they=20 were grown alone, apparently because F. tularensis killed a = substantial number of them. Furthermore, the increase of = fluorescence=20 inside A. castellanii cells with time showed that the = number of=20 F. tularensis per amoeba increased over time. Accordingly, = the=20 viable counts of F. tularensis released from A. = castellanii=20 by deoxycholate treatment also increased over time.

In this study the attenuated vaccine strain F. = tularensis LVS=20 was used. Compared to virulent strains of F. tularensis, = this=20 strain obviously is not highly virulent for humans. However, it is = still=20 highly virulent for mice and guinea pigs (34).=20 The genetic change(s) that results in attenuation is unknown. It = is=20 possible that the LVS strain could have a toxic effect on the = protozoan=20 host that is different from that of fully virulent F. = tularensis=20 strains.

Many types of bacteria have been found in free-living amoebae,=20 including Legionella spp., Burkholderia pickettii, = Vibrio=20 cholerae, Mycobacterium avium, and Listeria=20 monocytogenes (17,=20 39).=20 These bacteria are able to survive inside the protozoa as = endosymbionts=20 and to take advantage of the amoebae as vectors for their = dissemination.=20 However, Francisella shows no close relationship to these = genera,=20 being most closely related to arthropod parasites and symbionts, = including=20 Wolbachia persica, the Dermatocor andersoni = symbiont, and=20 Ornithodorous moubata symbiont B (10,=20 27,=20 28).=20 Other related strains, outside the Francisellaceae, include = members=20 of the Pisciricketsiaceae, in particular the intracellular=20 bacterium Piscirickettsia salmonis, which is a fish = pathogen (37).

Francisella bacteria are commonly transmitted to = vertebrates via=20 bites by ticks or mosquitoes (18,=20 19).=20 However, ticks may not comprise a natural reservoir for=20 Francisella, since infected ticks show a high mortality = rate and=20 transovarial transmission remains dubious (12,=20 19,=20 20).=20 Moreover, the bacterium has been isolated from about 250 wildlife = species=20 (18,=20 37),=20 clearly demonstrating that F. tularensis should be = considered a=20 highly successful intracellular parasite with a very broad host = range and=20 ability to survive and spread in widely differing natural=20 environments.

The ability of F. tularensis to survive in trophozoites = of A.=20 castellanii and cysts, as demonstrated in this study, may have = implications for the mode of transmission of the microorganism. = However,=20 although we have shown preliminary evidence for the survival of = F.=20 tularensis in amoebal cysts, the potential for its survival = under=20 various types of adverse conditions in these cysts still needs to = be=20 systematically examined. The close connection of tularemia with = water (30)=20 and the isolation of the bacterium from water samples used for = domestic=20 purposes, as well as from natural water systems, as the causal = agent of=20 outbreaks of the disease (14,=20 20,=20 21,=20 32)=20 support the hypothesis that amoebae may have a role in the natural = transmission of F. tularensis.

Acknowledgments

This work has been supported through Swedish Defense Agency = project=20 number A4854.

The opinions expressed in this paper are those of the authors = and do=20 not reflect any official policy.

A. Sj=C3=B6stedt is acknowledged for critical reading of the = manuscript.=20 Electron micrographs were skillfully prepared by V. Baranov, = University of=20 Ume=C3=A5, and monoclonal antibody specific for F. = tularensis was kindly=20 provided by R. Grunow, German Armed Forces Medical Academy, = Munich.

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DISCUSSION

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REFERENCES

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Amann, R., N. Springer, W. Schonhuber, W. = Ludwig, E.=20 N. Schmid, K. D. Muller, and R. Michel. 1997. Obligate = intracellular=20 bacterial parasites of AcanthamoebaAcanthamoeba related to=20 ChlamydiaChlamydia spp. Appl. = Environ.=20 Microbiol. 63:115-121. = [<= IMG=20 alt=3D"Free Full Text in PMC icon."=20 = src=3D"http://www.pubmedcentral.gov/corehtml/pmc/pmcgifs/pmc-freetext.gif= "=20 border=3D0>Free Full text in PMC].

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3a.

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Bhatnagar, N., E. Getachew, S. Straley, J. = Williams,=20 M. Meltzer, and A. Fortier. 1994. Reduced virulence of=20 rifampicin-resistant mutants of Francisella = tularensis.Francisella=20 tularensis. J. Infect. Dis. 170:841-847. [PubMed].

Cormack, B. P., R. H. Valdivia, and S. = Falkow.=20 1996. FACS-optimized mutants of the green fluorescent protein = (GFP). Gene 173:333-338.

Dennis, D. T., T. V. Inglesby, and D. A.=20 Henderson. 2001. Tularemia as a biological weapon: medical and = public=20 health management. JAMA 285:2763-2773. [PubMed]=20 [Full Text].

Eigelsbach, H. T., and C. M. Down. = 1961.=20 Prophylactic effectiveness of live and killed tularemia vaccines. = I.=20 Production of vaccine and evaluation in the white mouse and guinea = pig.=20 J. Immunol. 87:415-425. [PubMed].

Ericsson, M., I. Golovliov, G. Sandstrom, = A.=20 Tarnvik, and A. Sjostedt. 1997. Characterization of the = nucleotide=20 sequence of the groEgroE operon encoding heat shock = proteins=20 chaperone-60 and -10 of Francisella tularensisFrancisella=20 tularensis and determination of the T-cell response to the = proteins in=20 individuals vaccinated with F. tularensisF. tularensis. = Infect. Immun. 65:1824-1829. [<= IMG=20 alt=3D"Free Full Text in PMC icon."=20 = src=3D"http://www.pubmedcentral.gov/corehtml/pmc/pmcgifs/pmc-freetext.gif= "=20 border=3D0>Free Full text in PMC].

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Figures and Tables

FIG.=20 1.

Changes in GFP=20 fluorescence over time in cell populations of A. = castellanii=20 infected with F. tularensis LVS/GFP/ASV (triangles) = and=20 autofluorescence from A. castellanii (squares), = measured by=20 flow cytometry. Mean values from two independent = experiments=20 (more ...)

FIG.=20 2.

Fluorescence = microscopy=20 analysis. (A) A. castellanii trophozoite without=20 intracellular F. tularensis (day 0). (B) Intact A. = castellanii with Francisella-filled = vacuoles=20 (day 10). (C) Disintegrating A. castellanii filled = with F.=20 tularensis LVS/GFP/ASV=20 (more ...)

FIG.=20 3.

Differentiation = between=20 extracellular and intracellular F. tularensis by use = of=20 monoclonal antibodies. Viable intracellular F. = tularensis=20 LVS/GFP/AVS cells expressing GFP appear green, while = extracellular=20 F. tularensis cells appear red after treatment=20 (more ...)

FIG.=20 4.

Electron = microscopy=20 analysis. (A) A. castellanii trophozoite without=20 intracellular F. tularensis (day 0). (B) A.=20 castellanii trophozoite with=20 Francisella-filled vacuoles (day 9). (C = and D)=20 Recruitment of mitochondria (short arrows) and rough = endoplasmic=20 (more ...)

FIG.=20 5.

Viable counts = of F.=20 tularensis cultured alone (squares), F. = tularensis=20 cocultured with A. castellanii (triangles), and = intracellular=20 F. tularensis (circles). Mean values from two = independent=20 experiments are shown (t test, P < 0.05). = Bars=20 show=20 (more ...)

FIG.=20 6.

Counts of A. = castellanii cultured with F. tularensis = (triangles) and=20 without F. tularensis (squares) over time. Mean = values from=20 two independent experiments are shown (t test, = P <=20 0.05). Bars show averages of the absolute deviations of data = points=20 (more = ...)

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