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Treponema pallidum

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computational models. The treponemal outer membrane proteins are key factors for the bacterium's pathogenesis, persistence, and immune evasion strategies. The relatively low protein content prevents antigen recognition by the immune system and the proteins that do exist protrude out of the OM, enabling its interaction with the host.
625:. The latent stage that results can last a lifetime in many cases. In a few cases, the disease exits latency and enters a tertiary phase, in which destructive lesions of skin, bone, and cartilage ensue. Unlike yaws and bejels, syphilis in its tertiary stage often affects the heart, eyes, and nervous system, as well. 450:. TP0965 causes endothelial barrier dysfunction, a hallmark of late-stage pathogenesis of syphilis. It does this by reducing the expression of tight junction proteins, which in turn increases the expression of adhesion molecules and endothelial cell permeability, which eventually leads to disruption of the 658:
allows it to move in a corkscrew motion through mucous membranes or enter minuscule breaks in the skin. In women, the initial lesion is usually on the labia, the walls of the vagina, or the cervix; in men, it is on the shaft or glans of the penis. It gains access to the host's blood and lymph systems
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The clinical features of syphilis, yaws, and bejel occur in multiple stages that affect the skin. The skin lesions observed in the early stage last for weeks or months. The skin lesions are highly infectious, and the spirochetes in the lesions are transmitted by direct contact. The lesions regress as
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in vitro was reported in 2017. This was achieved using Sf1Ep epithelial cells from rabbits, which were a necessary condition for the continued multiplication and survival of the system. The medium TpCM-2 was used, an alteration of more simple media which previously only yielded a few weeks of culture
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outer membrane proteins have not been unambiguously identified. In contrast, some of the known antigens are intracellular, and antibodies are ineffective against them to clear the infection. In the last century, several prototypes have been developed, and while none of them provided protection from
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repeat family of proteins (Tpr) are proteins expressed during the infection process. Tprs are formed by a conserved N-terminal domain, an amino-terminal stretch of about 50 amino acids, a central variable region, and a conserved C-terminal domain. The many different types of Tpr include TprA, TprB,
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Outer Membrane Protein A (OmpA) domain-containing proteins are necessary for maintaining structural integrity in Gram-negative bacteria. These domains contain peptidoglycan binding sites which creates a "structural bridge between the peptidoglycan layer and the outer memebrane." The protein TP0624
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is characterized by its helical, corkscrew-like shape. To avoid antibodies attacking it, the cell has few proteins exposed on the outer membrane sheath. Its chromosome is about 1000 kilobase pairs and is circular with a 52.8% G + C average. Sequencing has revealed a bundle of 12 proteins and some
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group and its reduced genome is likely the result of various adaptations, such that it no longer contains the ability to synthesize fatty acids, nucleic acids, and amino acids, instead relying on its mammalian hosts for these materials. The recent sequencing of the genomes of several spirochetes
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has several features that have made it historically difficult to research. These include details such as its low protein content, its fragility, and that it contains fewer gene sequences related to other gram negative outer membranes. Progress has been made using genomic sequencing and advanced
518:. This process mainly happens in TprF, TprI, TprG, TprJ, and TprL, and it consists of a reversible expansion or contraction of polymeric repeats. These size variations can help the bacterium to quickly adapt to its microenvironment, dodge immune response, or even increase affinity to its host. 502:
Antigen variation in TprK is regulated by gene conversion. In this way,  fragments of the seven variable regions (V1–V7) present in TprK and the 53 donor sites of TprD can be combined to produce new structured sequences. TprK antigen variation can help
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was found to rely on its host for many molecules typically provided by biosynthetic pathways, and it is missing genes responsible for encoding key enzymes in oxidative phosphorylation and the tricarboxylic acid cycle. The
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In the past century since its initial discovery, culturing the bacteria in vitro has been difficult. Without the ability to grow and maintain the bacteria in a laboratory setting, discoveries regarding its metabolism and
397:, while its cytoplasmic membrane is covered in lipoproteins. The outer membrane's treponemal ligands' main function is attachment to host cells, with functional and antigenic relatedness between ligands. The genus 2774: 834:
to be effective. Efforts to develop a safe and effective syphilis vaccine have been hindered by uncertainty about the relative importance of humoral and cellular mechanisms to protective immunity, and because
562:. The challenge likely stems from the organism's strong adaptation to residing in mammalian tissue, resulting in a reduced genome and significant impairments in metabolic and biosynthetic functions. 550:'s requirements for growth and gene expression may occur and in turn, yield research beneficial for the treatment and prevention of syphilis, outside of a host. However, continuous efforts to grow 466:
structure that is characteristic of other membrane proteins, and does not traverse the outer membrane. This protein's function has been hypothesized to be involved with control of nutrient uptake.
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to evade a strong host immune reaction and can also allow the reinfection of individuals. This is possible because the newly structured proteins can avoid antibody-specific recognition.
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effects. Penicillin overall is the most recommended antibiotic by the Centers for Disease Control, as it shows the best results with prolonged use. It can inhibit and may even kill
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has one of the smallest bacterial genomes and has limited metabolic capabilities, reflecting its adaptation through genome reduction to the rich environment of mammalian tissue.
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indistinguishable. These bacteria were originally classified as members of separate species, but DNA hybridization analysis indicates they are members of the same species.
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Harper KN, Fyumagwa RD, Hoare R, Wambura PN, Coppenhaver DH, Sapolsky RM, Alberts SC, Tung J, Rogers J, Kilewo M, Batamuzi EK, Leendertz FH, Armelagos GJ, Knauf S (2012).
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through tissue and mucous membranes. In more severe cases, it may gain access to the host by infecting the skeletal bones and central nervous system of the body.
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is not obtainable in a pure culture, meaning that this sequencing played an important role in filling gaps of understanding regarding the microbes' functions.
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strain 14 is believed to derive from a single-point mutation that increased the organism's livability. Many of the syphilis treatment therapies only lead to
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allowed for a long-term drug treatment. These experiments established the groundwork that modern scientists use for syphilis therapy. Penicillin can inhibit
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Tang Y, Zhou Y, He B, Cao T, Zhou X, Ning L, Chen E, Li Y, Xie X, Peng B, Hu Y, Liu S (19 October 2022). "Investigation of the immune escape mechanism of
191:. It is known to be transmitted only among humans and baboons. It is a helically coiled microorganism usually 6–15 μm long and 0.1–0.2 μm wide. 3280: 1711:
Hazlett KR, Cox DL, Decaffmeyer M, Bennett MP, Desrosiers DC, La Vake CJ, La Vake ME, Bourell KW, Robinson EJ, Brasseur R, Radolf JD (September 2005).
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Luthra A, Zhu G, Desrosiers DC, Eggers CH, Mulay V, Anand A, McArthur FA, Romano FB, Caimano MJ, Heuck AP, Malkowski MG, Radolf JD (2 December 2011).
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has been proposed to facilitate this structural link, as well as interactions between outer membrane proteins and corresponding domains on the thin
3824: 2653: 381:, which is found in the outer membrane of other Gram-negative bacteria. It has an endoflagellum (periplasmic flagellum) consisting of four main 3863: 3315: 462:
TP0453 is a 287 amino acid protein associated with the inner membrane of the microbe's outer membrane. This protein lacks the extensive
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in 6–8 hours, though the cells still remain in lymph nodes and regenerate. Penicillin is not the only drug that can be used to inhibit
739: 385:, a core structure, and a sheath. The flagellum is located within the periplasmic space and wraps around the protoplasmic cylinder. 3811: 3105:: progress toward understanding their structural, functional, and immunologic roles. Treponema Pallidum Polypeptide Research Group" 1494:
Hawley KL, Montezuma-Rusca JM, Delgado KN, Singh N, Uversky VN, Caimano MJ, Radolf JD, Luthra A (8 July 2021). Galperin MY (ed.).
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Outer Membrane Protein Repertoire: a Road Map for Deconvolution of Syphilis Pathogenesis and Development of a Syphilis Vaccine"
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species are small, about 1.14 Mbp. Their DNA sequences are more than 99.7% identical. About 92.9% of DNA was determined to be
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has ribbons of cytoskeletal cytoplasmic filaments that run the length of the cell just underneath the cytoplasmic membrane.
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These virulence factors are thought to contribute to the bacterium's ability to evade the immune system and cause disease.
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reputation as a "stealth pathogen" is primarily due to this unique OM structure, which serves to evade immune detection.
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Fraser CM, Norris SJ, Weinstock GM, White O, Sutton GG, Dodson R, et al. (July 1998). "Complete genome sequence of
2706:"Treponema pallidum subsp. pertenue Displays Pathogenic Properties Different from Those of T. pallidum subsp. pallidum" 499:
TprC, TprD, and TprE, but variability of TprK is the most relevant due to the immune escape characteristics it allows.
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permits a thorough analysis of the similarities and differences within this bacterial phylum and within the species.
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Zobaníková M, Mikolka P, Cejková D, Pospíšilová P, Chen L, Strouhal M, Qin X, Weinstock GM, Smajs D (October 2012).
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Infection in the Wild Baboons of East Africa: Distribution and Genetic Characterization of the Strains Responsible"
301:, remains a separate species because no isolate is available for DNA analysis. Disease transmittance in subspecies 2770:"Vorläufiger Bericht über das Vorkommen von Spirochaeten in syphilitischen Krankheitsprodukten und bei Papillomen" 1770:
outer membrane-associated lipoprotein TP0453 involves membrane sensing and integration by two amphipathic helices"
2907:"Humoral immunity in experimental syphilis. I. The demonstration of resistance conferred by passive immunization" 3868: 1896:
Centurion-Lara A, Castro C, Barrett L, Cameron C, Mostowfi M, Van Voorhis WC, Lukehart SA (15 February 1999).
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the infection, some prevented bacteria from disseminating to distal organs and promoted accelerated healing.
3471: 2220:Šmajs D, Strouhal M, Knauf S (July 2018). "Genetics of human and animal uncultivable treponemal pathogens". 1595:
McKevitt M, Brinkman MB, McLoughlin M, Perez C, Howell JK, Weinstock GM, Norris SJ, Palzkill T (July 2005).
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passage during the later stages of pregnancy, giving rise to congenital syphilis. The helical structure of
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major sheath protein homologue Tpr K is a target of opsonic antibody and the protective immune response"
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Protein Tp0965 Activates Endothelial Cells and Increases the Permeability of Endothelial Cell Monolayer"
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Tp0624 Reveals a Modular Assembly of Divergently Functionalized and Previously Uncharacterized Domains"
2772:[Preliminary report on the occurrence of Spirochaetes in syphilitic chancres and papillomas]. 45: 3679: 3660: 3626: 203: 2769: 3633: 3582: 684:, which is transmissible by direct physical contact between people. Unlike syphilis, one strain of 211: 3672: 2761: 2472:"Comparison of the genome of the oral pathogen Treponema denticola with other spirochete genomes" 219: 2470:
Seshadri R, Myers GS, Tettelin H, Eisen JA, Heidelberg JF, Dodson RJ, et al. (April 2004).
3829: 3715: 3665: 3506: 3307: 543:(MEM) with CMRL 1066, a complex tissue culture medium. With development, new discoveries about 374: 2797:. Lippincott's Illustrated Reviews Series. Hagerstown, MD: Lippincott Williams & Wilkins. 1319:: novel flagellum, periplasmic cone, and cell envelope as revealed by cryoelectron tomography" 779: 3640: 3571: 390: 130: 3785: 3619: 3612: 3562: 3442: 3410: 2483: 2424: 2287: 2229: 1847: 1665: 930: 803: 559: 8: 3388: 3303: 2407:
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in Berlin in 1905. This bacterium can be detected with special stains, such as the
362: 215: 3855: 3022: 1388: 3697: 3381: 3333: 2973: 2437: 2299: 2241: 1860: 1678: 1020: 943: 876:(4th ed.). Galveston (TX): University of Texas Medical Branch at Galveston. 712: 531:
sensitivity were greatly impaired.{ However, successful long-term cultivation of
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at low to high doses, with each increase in concentration being more effective.
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Weinstock GM, Hardham JM, McLeod MP, Sodergren EJ, Norris SJ (1 October 1998).
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Proceedings of the National Academy of Sciences of the United States of America
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3747: 3555: 3530: 2930: 2873: 2375: 1224: 1145: 1044: 807: 451: 382: 173: 77: 67: 1515: 389:'s outer membrane has the most contact with host cells and contains few 3816: 3496: 3268:(CDC Fact Sheet). Centers for Disease Control and Prevention. May 2004. 1441: 767: 666:
infection is usually around 21 days, but can range from 10 to 90 days.
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Centurion-Lara A, Molini BJ, Godornes C, Sun E, Hevner K, Voorhis WC,
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subsp. pallidum strain Amoy: An Asian isolate highly similar to SS14"
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culture have been unsuccessful, indicating that it does not satisfy
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During the early 1940s, rabbit models in combination with the drug
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researched was not vertically transmissible in a guinea pig model.
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1493: 987:: Correlation of Functional Activities With Genome Sequence Data" 823: 806:
results, unless larger concentrations of penicillin are used for
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Edmondson DG, DeLay BD, Kowis LE, Norris SJ (23 February 2021).
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strain 14 has built-in resistance to some macrolides, including
3790: 3154:"Syphilis vaccine: challenges, controversies and opportunities" 2558: 1594: 1018: 711:
was first microscopically identified in syphilitic chancres by
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Tomson FL, Conley PG, Norgard MV, Hagman KE (September 2007).
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for syphilis is available as of 2024. The outer membrane of
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TP0965 is a protein that is critical for membrane fusion in
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is a motile spirochete that is generally acquired by close
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results in minimal metabolic activity. The treponemes have
188: 2528: 2469: 2273: 912: 2951: 2603: 2170: 2061: 1763: 646:, entering the host via breaches in squamous or columnar 2606:"Microbial Vertical Transmission during Human Pregnancy" 3214: 2006:"Long-Term in Vitro Culture of the Syphilis Spirochete 605:
putative hemolysins are potential virulence factors of
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outer membrane proteins: current status and prospects"
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growth. This success was the result of switching out
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1766:"The transition from closed to open conformation of 1206: 704:
cultured on epithelial cells of cotton-tail rabbits.
1886: 1550:Chen J, Huang J, Liu Z, Xie Y (27 September 2022). 3053: 2792: 2219: 980: 313:is the most invasive pathogenic subspecies, while 2760: 2120:"In Vitro Cultivation of the Syphilis Spirochete 2057: 2055: 2053: 2003: 1075: 234:each of which has a distinct associated disease. 3883: 1999: 1997: 1995: 1648:Zhang RL, Zhang JP, Wang QQ (16 December 2014). 3215:Althouse BM, Hébert-Dufresne L (October 2014). 3004: 2117: 1713:"TP0453, a concealed outer membrane protein of 1549: 1365: 1127: 2795:Lippincott's Illustrated Reviews: Microbiology 2554: 2552: 2406: 2050: 1825: 1823: 1647: 1489: 1487: 1423: 1421: 1419: 1417: 1076:Marks M, Solomon AW, Mabey DC (October 2014). 3288: 3054:Penn CW, Bailey MJ, Cockayne A (April 1985). 2847: 2845: 2843: 2775:Arbeiten aus dem Kaiserlichen Gesundheitsamte 2754: 2113: 2111: 1992: 1951: 1123: 1121: 976: 974: 972: 2998: 2945: 2904: 2786: 1947: 1945: 691: 2827:Infectious Disease and Antimicrobial Agents 2549: 2269: 2267: 2004:Edmondson DG, Hu B, Norris SJ (June 2018). 1820: 1484: 1427: 1414: 3295: 3281: 3217:"Epidemic cycles driven by host behaviour" 2854:"Global Challenge of Antibiotic-Resistant 2840: 2108: 1200: 1118: 969: 906: 353:is a helically shaped bacterium with high 31: 3494: 3240: 3187: 3169: 3128: 3075: 3030: 2981: 2881: 2820: 2816: 2814: 2737: 2629: 2580: 2505: 2495: 2446: 2436: 2383: 2213: 2196: 2147: 2118:Edmondson DG, Norris SJ (February 2021). 2091: 2033: 1942: 1925: 1869: 1859: 1803: 1785: 1740: 1687: 1677: 1624: 1571: 1545: 1543: 1523: 1467: 1397: 1387: 1366:Alderete JF, Baseman JB (December 1980). 1342: 1273: 1232: 1153: 1101: 1052: 952: 942: 865: 863: 861: 859: 857: 855: 853: 3538: 2264: 1434:Spirochete Biology: The Post Genomic Era 1069: 981:Norris SJ, Cox DL, Weinstock GM (2001). 695: 615: 510:To introduce more phenotypic diversity, 337: 263:, which causes bejel or endemic syphilis 179:with subspecies that cause the diseases 3221:Journal of the Royal Society, Interface 2793:Fisher B, Harvey RP, Champe PC (2007). 1306: 1128:Giacani L, Lukehart SA (January 2014). 680:The causative agent of yaws disease is 218:, treponemes are visible only by using 3884: 3098: 3005:Cameron CE, Lukehart SA (March 2014). 2811: 2543: 2328: 1540: 1368:"Surface Characterization of Virulent 869: 850: 317:is the least invasive of the species. 187:(also known as endemic syphilis), and 3714: 3713: 3276: 2862:Antimicrobial Agents and Chemotherapy 2851: 2565:: new light on the agent of syphilis" 2533:. 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"The 1033:Journal of Clinical Microbiology 228:T. p. pallidum, T. p. endemicum, 44: 3145: 3092: 3056:"The axial filament antigen of 3047: 2898: 2823:"Treponema Pallidum (Syphilis)" 2722:10.1128/IAI.68.6.3219-3225.2000 2697: 2672: 2646: 2597: 2537: 2531:Brock Biology of Microorganisms 2522: 2463: 2400: 2347: 2322: 2189:10.1016/j.antiviral.2016.11.002 2164: 1757: 1704: 1641: 1617:10.1128/iai.73.7.4445-4450.2005 1588: 1359: 1213:Nature Reviews. Disease Primers 377:, but its outer membrane lacks 328: 3121:10.1128/MMBR.57.3.750-779.1993 2409:"Whole genome sequence of the 1249: 1170: 1025:"Molecular Differentiation of 1012: 798:. Resistance to macrolides in 226:consists of three subspecies, 1: 3023:10.1016/j.vaccine.2013.09.053 2364:Standards in Genomic Sciences 2356:"Complete genome sequence of 1389:10.1128/iai.30.3.814-823.1980 1134:Clinical Microbiology Reviews 1078:"Endemic treponemal diseases" 843: 754:immobilization reaction, and 746:, treponemal antibody tests ( 700:Electron micrograph image of 582:was first sequenced in 1998. 342:Electron micrograph image of 333: 325:are intermediately invasive. 273:The three subspecies causing 237: 3689:Streptobacillus moniliformis 3099:Norris SJ (September 1993). 2974:10.1016/j.micinf.2007.05.018 2852:Stamm LV (1 February 2010). 2438:10.1371/journal.pone.0182768 2300:10.1126/science.281.5375.375 2278:, the syphilis spirochete". 2242:10.1016/j.meegid.2018.03.015 2078:(1): 10.1128/mbio.03536–20. 1861:10.1371/journal.pone.0166274 1679:10.1371/journal.pone.0115134 1496:"Structural Modeling of the 1323:Journal of Molecular Biology 1186:Immigrant and Refugee Health 1130:"The endemic treponematoses" 944:10.1371/journal.pone.0050882 761: 662:The incubation period for a 309:is considered non-venereal. 7: 3448:Louse borne relapsing fever 682:Treponema pallidum pertenue 640:Treponema pallidum pallidum 628: 409:The outer membrane (OM) of 373:. It is often described as 255:Treponema pallidum pallidum 10: 3923: 3902:Bacteria described in 1905 3472:Tick borne relapsing fever 3171:10.3389/fimmu.2023.1126170 2923:10.4049/jimmunol.117.1.191 2622:10.1016/j.chom.2017.04.007 1970:10.1007/s15010-022-01939-z 1315:"Cellular architecture of 817: 673: 632: 521: 3722: 3680:Fusobacterium polymorphum 3661:Fusobacterium necrophorum 3650: 3627:Capnocytophaga canimorsus 3602: 3529: 3485: 3398: 3341: 3332: 3323: 2569:FEMS Microbiology Reviews 1335:10.1016/j.jmb.2010.09.020 1275:10.1016/j.tim.2022.09.010 692:Laboratory identification 565: 469: 457: 437: 421: 204:oxidative phosphorylation 136: 129: 41:Scientific classification 39: 30: 23: 3634:Porphyromonas gingivalis 3583:Lymphogranuloma venereum 541:minimal essential medium 446:, and is located in the 200:tricarboxylic acid cycle 3673:Fusobacterium nucleatum 3158:Frontiers in Immunology 3109:Microbiological Reviews 2821:Fantry LE, Tramont EC. 2497:10.1073/pnas.0307639101 2331:Prescott's Microbiology 1787:10.1074/jbc.M111.305284 1721:Journal of Bacteriology 1504:Journal of Bacteriology 669: 570:The chromosomes of the 257:, which causes syphilis 220:dark-field illumination 3507:Leptospira interrogans 3233:10.1098/rsif.2014.0575 2962:Microbes and Infection 2710:Infection and Immunity 1605:Infection and Immunity 1573:10.1093/femspd/ftac023 1376:Infection and Immunity 1262:Trends in Microbiology 705: 391:transmembrane proteins 347: 346:, highlighted in gold. 3641:Prevotella intermedia 3572:Chlamydia trachomatis 2911:Journal of Immunology 2084:10.1128/mbio.03536-20 2026:10.1128/mBio.01153-18 1918:10.1084/jem.189.4.647 1560:Pathogens and Disease 1094:10.1093/trstmh/tru128 699: 616:Clinical significance 341: 3620:Tannerella forsythia 3613:Bacteroides fragilis 3563:Chlamydia pneumoniae 3443:Borrelia recurrentis 3411:Borrelia burgdorferi 2874:10.1128/AAC.01095-09 2658:www.pennmedicine.org 2376:10.4056/sigs.2615838 1225:10.1038/nrdp.2017.73 1146:10.1128/CMR.00070-13 1045:10.1128/JCM.00784-06 1023:(1 September 2006). 731:is also detected by 165:, formerly known as 16:Species of bacterium 3907:Pathogenic bacteria 3666:Lemierre's syndrome 3389:Treponema denticola 2660:. 10 September 2022 2488:2004PNAS..101.5646S 2429:2017PLoSO..1282768T 2292:1998Sci...281..375F 2234:2018InfGE..61...92S 1852:2016PLoSO..1166274P 1670:2014PLoSO...9k5134Z 1516:10.1128/JB.00082-21 935:2012PLoSO...750882H 744:rapid plasma reagin 576:open reading frames 484:peptidoglycan layer 359:peptidoglycan layer 269:, which causes yaws 167:Spirochaeta pallida 3856:treponema-pallidum 3754:Treponema pallidum 3724:Treponema pallidum 3551:Chlamydia psittaci 3377:Treponema carateum 3354:Treponema pallidum 3304:Bacterial diseases 3103:Treponema pallidum 3058:Treponema pallidum 2956:Treponema pallidum 2856:Treponema pallidum 2563:Treponema pallidum 2411:Treponema pallidum 2358:Treponema pallidum 2329:Willey JM (2020). 2276:Treponema pallidum 2177:Antiviral Research 2122:Treponema pallidum 2066:Treponema pallidum 2008:Treponema pallidum 1954:Treponema pallidum 1900:Treponema pallidum 1834:Treponema pallidum 1832:"The Structure of 1768:Treponema pallidum 1715:Treponema pallidum 1652:Treponema pallidum 1599:Treponema pallidum 1554:Treponema pallidum 1498:Treponema pallidum 1442:10.1007/82_2017_44 1430:Treponema pallidum 1370:Treponema pallidum 1317:Treponema pallidum 1180:Treponema pallidum 1027:Treponema pallidum 985:Treponema pallidum 917:Treponema pallidum 756:syphilis TPHA test 709:Treponema pallidum 706: 379:lipopolysaccharide 351:Treponema pallidum 348: 295:Treponema carateum 202:and processes for 162:Treponema pallidum 140:Treponema pallidum 25:Treponema pallidum 3879: 3878: 3716:Taxon identifiers 3707: 3706: 3598: 3597: 3525: 3524: 3521: 3520: 3481: 3480: 3101:"Polypeptides of 2804:978-0-7817-8215-9 2610:Cell Host Microbe 2340:978-1-260-21188-7 2128:Current Protocols 1727:(18): 6499–6508. 1451:978-3-319-89637-3 1432:Outer Membrane". 883:978-0-9631172-1-2 786:can be used. The 560:Koch's postulates 371:periplasmic space 198:s lack of both a 158: 157: 3914: 3872: 3871: 3859: 3858: 3846: 3845: 3833: 3832: 3820: 3819: 3807: 3806: 3794: 3793: 3781: 3780: 3768: 3767: 3758: 3757: 3756: 3743: 3742: 3741: 3711: 3710: 3536: 3535: 3492: 3491: 3467:Borrelia parkeri 3461:Borrelia duttoni 3455:Borrelia hermsii 3433:Neuroborreliosis 3428:Erythema migrans 3417:Borrelia afzelii 3339: 3338: 3330: 3329: 3297: 3290: 3283: 3274: 3273: 3269: 3254: 3244: 3227:(99): 20140575. 3202: 3201: 3191: 3173: 3149: 3143: 3142: 3132: 3096: 3090: 3089: 3079: 3051: 3045: 3044: 3034: 3002: 2996: 2995: 2985: 2949: 2943: 2942: 2902: 2896: 2895: 2885: 2849: 2838: 2837: 2835: 2833: 2818: 2809: 2808: 2790: 2784: 2783: 2758: 2752: 2751: 2741: 2716:(6): 3219–3225. 2701: 2695: 2694: 2692: 2690: 2676: 2670: 2669: 2667: 2665: 2650: 2644: 2643: 2633: 2601: 2595: 2594: 2584: 2556: 2547: 2541: 2535: 2534: 2526: 2520: 2519: 2509: 2499: 2467: 2461: 2460: 2450: 2440: 2404: 2398: 2397: 2387: 2351: 2345: 2344: 2326: 2320: 2319: 2286:(5375): 375–88. 2271: 2262: 2261: 2217: 2211: 2210: 2200: 2168: 2162: 2161: 2151: 2115: 2106: 2105: 2095: 2059: 2048: 2047: 2037: 2010:subsp. pallidum" 2001: 1990: 1989: 1949: 1940: 1939: 1929: 1893: 1884: 1883: 1873: 1863: 1846:(11): e0166274. 1827: 1818: 1817: 1807: 1789: 1780:(48): 41656–68. 1761: 1755: 1754: 1744: 1708: 1702: 1701: 1691: 1681: 1645: 1639: 1638: 1628: 1592: 1586: 1585: 1575: 1547: 1538: 1537: 1527: 1510:(15): e0008221. 1491: 1482: 1481: 1471: 1425: 1412: 1411: 1401: 1391: 1363: 1357: 1356: 1346: 1310: 1304: 1303: 1277: 1253: 1247: 1246: 1236: 1204: 1198: 1197: 1195: 1193: 1174: 1168: 1167: 1157: 1125: 1116: 1115: 1105: 1073: 1067: 1066: 1056: 1016: 1010: 1009: 991: 978: 967: 966: 956: 946: 910: 904: 903: 901: 899: 867: 216:light microscopy 197: 142: 122:T. pallidum 49: 48: 35: 21: 20: 3922: 3921: 3917: 3916: 3915: 3913: 3912: 3911: 3882: 3881: 3880: 3875: 3867: 3862: 3854: 3849: 3841: 3836: 3828: 3823: 3815: 3810: 3802: 3797: 3789: 3784: 3776: 3771: 3763: 3761: 3752: 3751: 3746: 3737: 3736: 3731: 3718: 3708: 3703: 3698:Haverhill fever 3646: 3594: 3517: 3477: 3394: 3334:Spirochaetaceae 3319: 3301: 3264: 3261: 3211: 3209:Further reading 3206: 3205: 3150: 3146: 3097: 3093: 3052: 3048: 3003: 2999: 2968:(11): 1267–75. 2950: 2946: 2903: 2899: 2850: 2841: 2831: 2829: 2819: 2812: 2805: 2791: 2787: 2759: 2755: 2702: 2698: 2688: 2686: 2678: 2677: 2673: 2663: 2661: 2652: 2651: 2647: 2602: 2598: 2561:"The genome of 2557: 2550: 2542: 2538: 2527: 2523: 2482:(15): 5646–51. 2468: 2464: 2423:(8): e0182768. 2405: 2401: 2352: 2348: 2341: 2327: 2323: 2272: 2265: 2218: 2214: 2169: 2165: 2140:10.1002/cpz1.44 2116: 2109: 2060: 2051: 2002: 1993: 1950: 1943: 1894: 1887: 1828: 1821: 1762: 1758: 1709: 1705: 1664:(12): e115134. 1646: 1642: 1593: 1589: 1548: 1541: 1492: 1485: 1452: 1426: 1415: 1364: 1360: 1311: 1307: 1254: 1250: 1205: 1201: 1191: 1189: 1176: 1175: 1171: 1126: 1119: 1074: 1070: 1017: 1013: 989: 979: 970: 911: 907: 897: 895: 884: 868: 851: 846: 820: 782:antibiotics or 764: 713:Fritz Schaudinn 694: 678: 672: 637: 631: 618: 612: 568: 524: 516:phase variation 492: 472: 460: 440: 424: 407: 336: 331: 319:T. p. endemicum 303:T. p. endemicum 297:, the cause of 287:morphologically 261:T. p. endemicum 240: 232:T. p. pertenue, 212:outer membranes 195: 171:microaerophilic 154: 144: 138: 125: 98:Treponemataceae 43: 17: 12: 11: 5: 3920: 3910: 3909: 3904: 3899: 3894: 3877: 3876: 3874: 3873: 3860: 3847: 3834: 3821: 3808: 3795: 3782: 3769: 3759: 3744: 3728: 3726: 3720: 3719: 3705: 3704: 3702: 3701: 3694:Rat-bite fever 3684: 3683: 3676: 3669: 3656: 3654: 3652:Fusobacteriota 3648: 3647: 3645: 3644: 3637: 3630: 3623: 3616: 3608: 3606: 3600: 3599: 3596: 3595: 3593: 3592: 3591: 3590: 3585: 3580: 3567: 3566: 3559: 3546: 3544: 3533: 3527: 3526: 3523: 3522: 3519: 3518: 3516: 3515: 3502: 3500: 3489: 3487:Leptospiraceae 3483: 3482: 3479: 3478: 3476: 3475: 3451: 3438: 3437: 3436: 3435: 3430: 3425: 3406: 3404: 3396: 3395: 3393: 3392: 3385: 3373: 3372: 3371: 3366: 3349: 3347: 3336: 3327: 3321: 3320: 3312:proteobacteria 3300: 3299: 3292: 3285: 3277: 3271: 3270: 3260: 3259:External links 3257: 3256: 3255: 3210: 3207: 3204: 3203: 3144: 3091: 3046: 3017:(14): 1602–9. 2997: 2944: 2897: 2839: 2810: 2803: 2785: 2753: 2696: 2671: 2645: 2596: 2575:(4): 323–332. 2548: 2536: 2521: 2462: 2399: 2346: 2339: 2321: 2263: 2212: 2163: 2107: 2049: 1991: 1964:(2): 305–321. 1941: 1912:(4): 647–656. 1885: 1819: 1756: 1703: 1640: 1611:(7): 4445–50. 1587: 1539: 1483: 1450: 1413: 1382:(3): 814–823. 1358: 1305: 1268:(3): 294–307. 1248: 1199: 1169: 1117: 1068: 1039:(9): 3377–80. 1011: 968: 929:(12): e50882. 905: 882: 848: 847: 845: 842: 819: 816: 804:bacteriostatic 763: 760: 725:Dieterle stain 717:Erich Hoffmann 693: 690: 686:T. p. pertenue 674:Main article: 671: 668: 664:T. p. pallidum 656:T. p. pallidum 652:transplacental 644:sexual contact 633:Main article: 630: 627: 617: 614: 567: 564: 523: 520: 491: 488: 471: 468: 459: 456: 439: 436: 423: 420: 406: 405:Outer membrane 403: 369:cylinder, and 363:inner membrane 335: 332: 330: 327: 323:T. p. pertenue 311:T. p. pallidum 307:T. p. pertenue 271: 270: 267:T. p. pertenue 264: 258: 239: 236: 156: 155: 145: 134: 133: 127: 126: 119: 117: 113: 112: 105: 101: 100: 95: 91: 90: 88:Spirochaetales 85: 81: 80: 75: 71: 70: 65: 61: 60: 55: 51: 50: 37: 36: 28: 27: 15: 9: 6: 4: 3: 2: 3919: 3908: 3905: 3903: 3900: 3898: 3895: 3893: 3890: 3889: 3887: 3870: 3865: 3861: 3857: 3852: 3848: 3844: 3839: 3835: 3831: 3826: 3822: 3818: 3813: 3809: 3805: 3800: 3796: 3792: 3787: 3783: 3779: 3774: 3770: 3766: 3760: 3755: 3749: 3745: 3740: 3734: 3730: 3729: 3727: 3725: 3721: 3717: 3712: 3699: 3695: 3691: 3690: 3686: 3685: 3682: 3681: 3677: 3675: 3674: 3670: 3667: 3663: 3662: 3658: 3657: 3655: 3653: 3649: 3643: 3642: 3638: 3636: 3635: 3631: 3629: 3628: 3624: 3622: 3621: 3617: 3615: 3614: 3610: 3609: 3607: 3605: 3601: 3589: 3586: 3584: 3581: 3579: 3576: 3575: 3574: 3573: 3569: 3568: 3565: 3564: 3560: 3557: 3553: 3552: 3548: 3547: 3545: 3543: 3542: 3537: 3534: 3532: 3528: 3513: 3512:Leptospirosis 3509: 3508: 3504: 3503: 3501: 3499: 3498: 3493: 3490: 3488: 3484: 3473: 3469: 3468: 3463: 3462: 3457: 3456: 3452: 3449: 3445: 3444: 3440: 3439: 3434: 3431: 3429: 3426: 3424: 3421: 3420: 3419: 3418: 3413: 3412: 3408: 3407: 3405: 3403: 3402: 3397: 3391: 3390: 3386: 3383: 3379: 3378: 3374: 3370: 3367: 3365: 3361: 3358: 3357: 3356: 3355: 3351: 3350: 3348: 3346: 3345: 3340: 3337: 3335: 3331: 3328: 3326: 3325:Spirochaetota 3322: 3317: 3313: 3309: 3308:gram negative 3305: 3298: 3293: 3291: 3286: 3284: 3279: 3278: 3275: 3267: 3263: 3262: 3252: 3248: 3243: 3238: 3234: 3230: 3226: 3222: 3218: 3213: 3212: 3199: 3195: 3190: 3185: 3181: 3177: 3172: 3167: 3163: 3159: 3155: 3148: 3140: 3136: 3131: 3126: 3122: 3118: 3115:(3): 750–79. 3114: 3110: 3106: 3104: 3095: 3087: 3083: 3078: 3073: 3070:(4): 635–41. 3069: 3065: 3061: 3059: 3050: 3042: 3038: 3033: 3028: 3024: 3020: 3016: 3012: 3008: 3001: 2993: 2989: 2984: 2979: 2975: 2971: 2967: 2963: 2959: 2957: 2948: 2940: 2936: 2932: 2928: 2924: 2920: 2916: 2912: 2908: 2901: 2893: 2889: 2884: 2879: 2875: 2871: 2867: 2863: 2859: 2857: 2848: 2846: 2844: 2828: 2824: 2817: 2815: 2806: 2800: 2796: 2789: 2781: 2777: 2776: 2771: 2767: 2763: 2757: 2749: 2745: 2740: 2735: 2731: 2727: 2723: 2719: 2715: 2711: 2707: 2700: 2685: 2681: 2675: 2659: 2655: 2649: 2641: 2637: 2632: 2627: 2623: 2619: 2615: 2611: 2607: 2600: 2592: 2588: 2583: 2578: 2574: 2570: 2566: 2564: 2555: 2553: 2546:, p. 499 2545: 2540: 2532: 2525: 2517: 2513: 2508: 2503: 2498: 2493: 2489: 2485: 2481: 2477: 2473: 2466: 2458: 2454: 2449: 2444: 2439: 2434: 2430: 2426: 2422: 2418: 2414: 2412: 2403: 2395: 2391: 2386: 2381: 2377: 2373: 2369: 2365: 2361: 2360:strain DAL-1" 2359: 2350: 2342: 2336: 2332: 2325: 2317: 2313: 2309: 2305: 2301: 2297: 2293: 2289: 2285: 2281: 2277: 2270: 2268: 2259: 2255: 2251: 2247: 2243: 2239: 2235: 2231: 2227: 2223: 2216: 2208: 2204: 2199: 2194: 2190: 2186: 2182: 2178: 2174: 2167: 2159: 2155: 2150: 2145: 2141: 2137: 2133: 2129: 2125: 2123: 2114: 2112: 2103: 2099: 2094: 2089: 2085: 2081: 2077: 2073: 2069: 2067: 2058: 2056: 2054: 2045: 2041: 2036: 2031: 2027: 2023: 2019: 2015: 2011: 2009: 2000: 1998: 1996: 1987: 1983: 1979: 1975: 1971: 1967: 1963: 1959: 1955: 1948: 1946: 1937: 1933: 1928: 1923: 1919: 1915: 1911: 1907: 1903: 1901: 1892: 1890: 1881: 1877: 1872: 1867: 1862: 1857: 1853: 1849: 1845: 1841: 1837: 1835: 1826: 1824: 1815: 1811: 1806: 1801: 1797: 1793: 1788: 1783: 1779: 1775: 1771: 1769: 1760: 1752: 1748: 1743: 1738: 1734: 1730: 1726: 1722: 1718: 1716: 1707: 1699: 1695: 1690: 1685: 1680: 1675: 1671: 1667: 1663: 1659: 1655: 1653: 1650:"Recombinant 1644: 1636: 1632: 1627: 1622: 1618: 1614: 1610: 1606: 1602: 1600: 1591: 1583: 1579: 1574: 1569: 1565: 1561: 1557: 1555: 1546: 1544: 1535: 1531: 1526: 1521: 1517: 1513: 1509: 1505: 1501: 1499: 1490: 1488: 1479: 1475: 1470: 1465: 1461: 1457: 1453: 1447: 1443: 1439: 1435: 1431: 1424: 1422: 1420: 1418: 1409: 1405: 1400: 1395: 1390: 1385: 1381: 1377: 1373: 1371: 1362: 1354: 1350: 1345: 1340: 1336: 1332: 1329:(4): 546–61. 1328: 1324: 1320: 1318: 1309: 1301: 1297: 1293: 1289: 1285: 1281: 1276: 1271: 1267: 1263: 1259: 1252: 1244: 1240: 1235: 1230: 1226: 1222: 1218: 1214: 1210: 1203: 1187: 1183: 1181: 1173: 1165: 1161: 1156: 1151: 1147: 1143: 1140:(1): 89–115. 1139: 1135: 1131: 1124: 1122: 1113: 1109: 1104: 1099: 1095: 1091: 1088:(10): 601–7. 1087: 1083: 1079: 1072: 1064: 1060: 1055: 1050: 1046: 1042: 1038: 1034: 1030: 1028: 1022: 1015: 1007: 1003: 999: 995: 988: 986: 977: 975: 973: 964: 960: 955: 950: 945: 940: 936: 932: 928: 924: 920: 918: 909: 893: 889: 885: 879: 875: 874: 866: 864: 862: 860: 858: 856: 854: 849: 841: 838: 833: 829: 825: 815: 813: 809: 805: 801: 797: 793: 789: 785: 781: 777: 773: 769: 759: 757: 753: 749: 745: 741: 738: 737:nontreponemal 734: 730: 726: 722: 718: 714: 710: 703: 698: 689: 687: 683: 677: 667: 665: 660: 657: 653: 649: 645: 641: 636: 626: 624: 613: 610: 608: 603: 599: 594: 589: 585: 581: 577: 573: 563: 561: 557: 553: 549: 546: 542: 537: 534: 530: 529:antimicrobial 519: 517: 513: 508: 506: 500: 497: 487: 485: 481: 478: 467: 465: 455: 453: 449: 445: 435: 433: 429: 426:TP0326 is an 419: 417: 412: 402: 400: 396: 392: 388: 384: 380: 376: 375:Gram negative 372: 368: 364: 360: 356: 352: 345: 340: 326: 324: 320: 316: 312: 308: 304: 300: 296: 292: 291:serologically 288: 284: 280: 276: 268: 265: 262: 259: 256: 253: 252: 251: 249: 245: 235: 233: 229: 225: 221: 217: 213: 209: 205: 201: 194: 190: 186: 182: 178: 175: 172: 168: 164: 163: 152: 148: 143: 141: 135: 132: 131:Binomial name 128: 124: 123: 118: 115: 114: 111: 110: 106: 103: 102: 99: 96: 93: 92: 89: 86: 83: 82: 79: 76: 73: 72: 69: 68:Spirochaetota 66: 63: 62: 59: 56: 53: 52: 47: 42: 38: 34: 29: 26: 22: 19: 3723: 3687: 3678: 3671: 3659: 3639: 3632: 3625: 3618: 3611: 3604:Bacteroidota 3570: 3561: 3549: 3539: 3505: 3495: 3465: 3459: 3453: 3441: 3423:Lyme disease 3415: 3409: 3399: 3387: 3375: 3353: 3352: 3342: 3224: 3220: 3161: 3157: 3147: 3112: 3108: 3102: 3094: 3067: 3063: 3057: 3049: 3014: 3010: 3000: 2965: 2961: 2955: 2947: 2917:(1): 191–6. 2914: 2910: 2900: 2868:(2): 583–9. 2865: 2861: 2855: 2830:. Retrieved 2826: 2794: 2788: 2779: 2773: 2762:Schaudinn FR 2756: 2713: 2709: 2699: 2689:19 September 2687:. Retrieved 2683: 2674: 2664:11 September 2662:. Retrieved 2657: 2648: 2616:(5): 561–7. 2613: 2609: 2599: 2572: 2568: 2562: 2539: 2530: 2524: 2479: 2475: 2465: 2420: 2416: 2410: 2402: 2370:(1): 12–21. 2367: 2363: 2357: 2349: 2330: 2324: 2283: 2279: 2275: 2225: 2221: 2215: 2180: 2176: 2166: 2131: 2127: 2121: 2075: 2071: 2065: 2017: 2013: 2007: 1961: 1957: 1953: 1909: 1905: 1899: 1843: 1839: 1833: 1777: 1773: 1767: 1759: 1724: 1720: 1714: 1706: 1661: 1657: 1651: 1643: 1608: 1604: 1598: 1590: 1563: 1559: 1553: 1507: 1503: 1497: 1433: 1429: 1379: 1375: 1369: 1361: 1326: 1322: 1316: 1308: 1265: 1261: 1251: 1216: 1212: 1202: 1190:. Retrieved 1185: 1179: 1172: 1137: 1133: 1085: 1081: 1071: 1036: 1032: 1026: 1014: 1000:(1): 37–62. 997: 993: 984: 983:"Biology of 926: 922: 916: 908: 896:. 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Index


Scientific classification
Edit this classification
Bacteria
Spirochaetota
Spirochaetia
Spirochaetales
Treponemataceae
Treponema
Binomial name
Schaudinn
Hoffmann
microaerophilic
spirochaete
bacterium
syphilis
bejel
yaws
tricarboxylic acid cycle
oxidative phosphorylation
cytoplasmic
outer membranes
light microscopy
dark-field illumination
subspecies
yaws
bejel
syphilis
morphologically
serologically

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