Historical Observations of Spirochetal Cysts
and L-Forms

Summary of historical observations of spirochetal cyst (and L-forms) prepared by an anonymous volunteer of actionlyme, to whom we all are VERY grateful. I have been given permission to distribute.

Date: Sun, 09 Apr 2000 19:03:08 -0400
From: Kathleen Dickson
Organization: ActionLyme

Any errors or typos are my own.

The data is quite stunning. I have seen some of the Scanning Electron Micrographs. Guaranteed to be thrilling.

I will be eventually getting some of these docs scanned into scilyme, but I'm not giving myself a deadline ;)

It is inarguable that spirochetes are extremely adaptable to environmental extremes. This was known to genuine scientists before 1975 when Polly Murray discovered Lyme disease. I don't know what to make of the fact that rheumatologists at the time, in ~1982 (when Willy Burgdorfer discovered the Borrelia), did not research this historical scientific evidence of spirochetal pathogenesis and virulence. (Kidding)

(P)= interesting photos, (R)= Highly recommended reading, (F)=foreign language

Borrelia - burgdorferi

Microbiology, 2000 Jan;146 (Pt 1):119-27

Serum-starvation-induced changes in protein synthesis and morphology of Borrelia burgdorferi.

Alban PS, Johnson PW, Nelson DR

B. burgdorferi spirochetes converted to a cystic form when placed in a deficient culture media. These cysts reverted to motile spirochetes when transferred to growth media.
[From the abstract:] "Cyst opening and recovery of spiral-shaped non-motile organisms was induced within 1 min by the addition of either BSKrs+ or rabbit serum (6%v/v, final concentration). Cells regained motility with additional incubation in BSKrs+."

APMIS, 1999;106(12):1131-1141

A rapid method for generating cystic forms of Borrelia burgdorferi, and their reversal to mobile spirochetes.

Brorson, O., & Brorson, S.

Motile B. burgdorferi spirochetes converted to cysts within 1 minute when placed in distilled water. The cysts reverted to spirochetes after transfer to a growth medium.

APMIS, 1999;107(6):566-576.

(R) Brorson, O., & Brorson

An in vitro study of the susceptibility of mobile and cystic forms of Borrelia burgdorferi to metronidazole.

B. burgdorferi cysts were degraded upon incubation with metronidazole (in vitro).

12th International Conference on Lyme Disease and Other Spirochetal and Tick-Borne Disorders., April 8-9, 1999

Keynote Address - The Complexity of Vector-borne Spirochetes.

Burgdorfer, W.

"This relatively large Borrelia [Borrelia burgdorferi] is not readily detectable in blood smears or thick drops of Lyme disease patients and susceptible host animals, yet engorgement on infected hosts results in up to 100% infected ticks.... RML [NIH's Rocky Mountain Lab] scientists Dave Dorward and Claude Garon using silver staining, transmission and scanning electron microscopy investigated the nature of naturally elaborated membrane blebs on the surface of cultured B. burgdorferi or free in the medium, and found both linear and circular DNA (Fig.13)... These most recent findings [of RML researchers and others] do confirm the development of membrane-derived cysts, blebs, spherules, vesicles and the potential transformation to motile, helical spirochetes...as a "survival mechanism" of spirochetes to overcome or escape unfavorable conditions." [Willy Burgdorfer, Ph.D., of the National Institutes of Health, is the discoverer of Borrelia burgdorferi.]

Infection, 1998;26(3):144-50

(R) In vitro conversion of Borrelia burgdorferi to cystic forms in spinal fluid, and transformation to mobile spirochetes by incubation in BSK-H medium

Brorson, O., & Brorson S.

. B. burgdorferi transformed into cysts (spheroplast L-forms) within 1-24h of inoculation into spinal fluid. When transferred to a growth medium, the cysts converted back to normal spirochetes after 9-17 days of incubation. The authors suggest that encystment of Bb may explain why cultivation of spinal fluid may be negative in patients with neuroborreliosis. The cysts may be recognized by microscopy.

American Journal of Dermatopathology,1996;18(6):571-9.

(R) Heterogeneity of Borrelia burgdorferi in the skin.

Aberer, E., Kersten, et al

Encysted B. burgdorferi were found in a skin biopsy. The authors conclude that these are in vivo variant forms of Bb and not staining artifacts. Includes remarkable color videomicroscopy photographs as supporting evidence.

[Diagnosis:] "Borrelia may escape immune surveillance by colony formation and masking within collagen, resulting in seronegativity."

Infection, 1996;24(3):218-26.

Formation and cultivation of Borrelia burgdorferi spheroplast L-form variants.

Mursic, V.P., Wanner, et al

[Cysts:] In vitro investiation of morphological variants of B. burdorferi, in an effort to explain the clinical persistence of active Lyme borreliosis despite antibiotic therapy. The authors suggest that these atypical forms may allow Borrelia to survive antibiotic treatment. [Persistence:] "...clinical persistence of Borrelia burgdorferi in patients with active Lyme borreliosis occurs despite obviously adequate

Antimicrobial Agents & Chemotherapy, 1995;39(5):1127-33.

(R) Effects of penicillin, ceftriaxone, and doxycycline on the morphology of Borrelia

Kersten, A, Poitschek C, Rauch S, Aberer, E

B. burgdorferi cultures gradually developed cysts and blebs [granules] when incubated in antibiotics. The degree of alteration was strongly correlated with dose and duration.The alterations occured most rapidly with ceftriaxone, then with penicillin; doxycycline caused a smaller percentage of organisms to undergo morphologic changes. Those organisms that remained in typical spirochetal form in the doxycycline cultures showed decreased motility. The authors suggest that these morphologic changes may shed light on the ability of B. burgdorferi to survive

Infection, 1994 ;22(6):401-406.

Ultrastructure of Borrelia burgdorferi after exposure to benzylpenicillin

Schaller, M. et al&

B. burgdorferi were observed to form cysts and blebs when treated with penicillin G. "These structures were not found under optimal culture conditions. One may speculate that the borreliae could escape the action of the antibiotic by developing such spherical bodies." (p. 404)

Abstracts of the V International Conference of Lyme Borreliosis, Arlington, VA.1992

Borrelia burgdorferi in the skin: a morphological and immunohistochemical study of the heterogenous appearance of this microorganism

Aberer, E., Kersten, et al

B. burgdorferi granules were detected in skin biopsies of erythema migrans and acrodermatitis chronica atrophicans lesions.

Annals New York Academy of Sciences, 1988;539:468-470.

Concurrent neocortical borreliosis and Alzheimer's disease: Demonstration of a spirochetal cyst form

MacDonald, A.B.

In vivo finding of Borrelia burgdoreri cysts in an autopsy of a human brain. "An unexpected observation was the identification of cystic forms of the Borrelia spirochete in dark-field preparations of cultured hippocampus, and in imprints of hippocampus... A cystic form of the Borrelia spirochete would explain the ability of the microbe to persist in the host during a prolonged period of asymptomatic clinical latency, which spans the period between primary infection and the expression of tertiary manifestations of neuroborreliosis."

Borrelia - duttoni

Lancet, 1920 ;2:1237-1244.

(R) The Horace Dobell lecture on an experimental investigation of Spirochaeta duttoni, the parasite of tick fever

Leishman, W.B.

The author argues that S. duttoni, when inside a tick, reproduce by a process of budding and extrusion of granules; the granules grow into young spirochetes. The granules are also themselves capable of multiplication. Their development into spirochetal form within a vertibrate host is an exceptional occurence, brought about by certain environmental circumstances. This theory rests on an accumulation of observations by the author and by other researchers cited, including a correlation of the temporary disappearance of spirochetes from the tick's stomach with the appearance of large numbers of granules. Temperature was found to influence the development of classic-shaped spirochetes.

Annales de l'Institut Pasteur, 1918;32:49-59.

A note on the "granule clumps" found in Ornithodorus moubata and their relation to the spirochaetes of African relapsing fever (tick fever).

Leishman, W.B.

Granules developed into spirochetes. Innoculation of tissues containing only granules produced spirochaetosis in mice. Observed periods of several days where few, if any, spirochetes could be found inside a tick, followed by sudden re-invasion of tissues with mostly young and vigorously motile spirochetes; then this sequence would repeat in a regular pattern. Concludes that this has to do with the reproductive habits

Parasitology, 1912;Vol. IV, p.133.

The transmission of Spirochaeta duttoni.

Hindle, E.

[According to W. Burgdorferi, 1999:] Hindle reported that in infected ticks kept at 21SC, the spirochetes disappeared from the midgut by the 10th day and could no longer be detected either in the gut or in the tissues. However, triturates of such ticks were able to infect mice, and an increase in temperature to 35SC led to the reappearance of typical spirochetes.

Annals of Tropical Medicine and Parasitology, 1907;No. 3

The morphology and life-history of Spirochaeta Duttoni


[According to Dutton, 1907:] Observed encysted forms of S. duttoni in the spleen. The cysts broke into granular bodies from which new generations of spirochetes emerged.

Lancet, 1907;ii:1523.

(R) A note on the morphology of Spirochaeta Duttoni.

Dutton, J.S. & Todd,

Spirochetes within sporocyst-like bodies were found in the blood even when other forms had disappeared. The authors suggest that reproduction may occur by direct division, but possibly also by a process involving extrusion of granules which subsequently develop into new spirochetes.

Borrelia - recurrentis

Compt. rend. acad. sci., clix, 1914;pp. 119-122.

(F) De la periode de latence du Spirille chez le Pou infecte de fievre recurrent.

Sergent, E. & Foley,

[As described by Leishman, 1920): After ingestion into ticks, the Leptospira studied disappeared after 24 hours. After the 6th day, new, actively mobile spirochetes reappeared suddenly in great numbers. Infectivity was highest on the 6th day prior to this reappearance, despite of the absence of demonstrable spirochetes. Transverse fission of the spirochetes in the louse was only rarely observed. When classic-shapedspirochetes were present, their numbers did not increase.


Journal of Bacteriology 1954;67:619-627.

(R) Formation of granular structures by Leptospirae as revealed by the electron microscope.

Czekalowki, J.W., et al

Leptospira began to show granulation after 2 weeks in a culture. The granules were spaced regularly within the bodies of the spirochetes. After four weeks a larger type of granule appeared which was broader than the body of the spirochetes. These were later "shed free." By the 5th to 7th month, there were no spirochetes observed; the culture contained only granules. The granules consisted of "what appears to be short segments of leptospiral body embedded in homogeneous substance." The authors conclude that the "formation of granules represents a rhythmic and constant process and hence these granules must play a role in the life-cycle of leptospirae."

J. Hygiene, 1949;47:390-392.

The morphology of the genus Leptospira as shown by the electron microscopy.

Babudieri, B.

Acta Biologica Belge, 1943;3-4:245.

(F)Coloration des granules leptospiriens.

Herreweghe, E.

Bull. ass. diploms microbiol. fac. pharm.,1942;61:72-80.

(F) Le micro-manipulateur et les granules d'une souche de Leptospire aquicole non-pathogene.

Bessemans, A.,

[According to Czekalowski, 1954]: Granules from the culture of a leptospira were isolated using a micromanipulator and grown from single cell

Compt. rend. soc. biol., 1942;136:141-144.

(F) Sur l'evolution du L. ieterohaemorrhagiae, granule leptospirogene.

Gastinel, P. et al

Quoted by Van Thiel, P.H., 1948.

The leptospiroses.

Timmerman, H. 1927

Universitaire Pers, Leiden.

Granules develop in response to physical and chemical changes.

Journal of Experimental Medicine, 1916;23:377-402.

The etiology and mode of infection and specific therapy of Weil's disease.

Inada, R., Ido, Y., et al,

Interprets atypical spirochetal forms as "degenerative." Spirochetes were seen inside phagocytic cells and epithelial cells. Classic-shaped spirochetes were found in the blood in small numbers, and only in the early stages of the disease.


Journal of Bacteriology, 1942;44:37-70.

The significant of the large bodies and the development of L-type colonies in bacterial cultures.

Dienes, L.

"If they [L type colonies] represent a variant form [of bacterial morphology], the observations indicate that the variability of bacteria extends much further than is commonly supposed. The bacteria apparently are able to grow in the form of soft granules and to multiply by the development and germination of large bodies." (p.60)


CRC Press, 1993

Cell wall deficient forms: Stealth Pathogens

Mattman, L.H.

Eur. J. Clin. Microbiol. Infect. Dis., 1989.

Unusual in vitro formation of cyst-like structures associated with human intestinal spirochaetosis

Gebbers, J.O. et al

In vitro findings suggest that spirochetes may develop in cysts, contrary to the traditional view that transverse fission is their main mode of reproduction. As cysts were found in centrifugates of cultures but not in biopsy speciments, the authors speculate that this mode of reproduction may occur only when in sub-optimal environments outside the host. Electron micrographs of maturation of spirochetes within the

Microbiol. Immunology, 1980;24:321-334.

Electron microscopy of the spherical bodies of oral spirochetes in vitro.

Umemoto, T., et al

J. Gifu Dent. Soc. 2:1-15.

Spherical body formation of oral spirochetes following addition of sucrose

Umemoto, T. et al 1974

Am J. MS, 1958;236:677-91.

Preliminary observations of the cultivation and morphology of a microorganism from the cerebrospinal fluid of patients with MS.

Myerson, R.M.

Journal of Neuropathology, 1954;13:221-29.

Morphology of Spirochaeta myelophthora in multiple sclerosis

Steiner, G.

Journal of Neuropathology, 11:343-72.

(R) Acute plaques in multiple sclerosis, their pathogenic significance and the role of spirochaetes as etiological factor.

Steiner, G. 1952

Journal of Bacteriology, 1951;62:347-349.

(P) Further studies on the significance of spirochetal granules.

Hampp, E.G.

American Journal of Syphilis, 1950;34:122-125.

Studies on the life cycles of spirochaetes: I. The use of phase contrast microscopy.

Delamater, E.D., et al

Includes several small photos of spirochetes emerging from "gemma," which the authors interpret as reproductive forms.

J Am Dental Assoc, 1950;40:1-11.

Morphologic characteristics of the smaller oral treponemes and Borrelia vincentii as revealed by stained smear, darkfield and electron microscopic technics.

Hampp, E.G. et al

Archives of Neurology and Psychiatry , 1939.

(R) "Silver cells" and "spirochete-like formations" in MS and other diseases of the central nervous system

Hassin, G.B. et al

Review confirms the findings of G. Steiner and other researchers who found "silver cells" [spirochetal granules that take a silver stain] in brain autopsies of MS cases. (G. Steiner contended that MS is an infectious disease caused by a spirochete that changes from a classic spirochetal form to a granular form, and which destroys myelin.) While the authors found granules in the CNS of all 8 MS patients they studied, they dispute Steiner's belief that they are the causative agent of the disease.

Deutsche Ztschr. f. Nervenheilkunde, 1928;107:112.

(F) Demonstration von Spirochäten im menschlichen Gehirn bei Multipler Sklerose.

Steiner, G.

(Demonstration of spirochetes in the human brain in multiple sclerosis)

Das Handbuch der pathogenen Protozoen, Leipzig, Johann Ambrosius Barth, Band 3, 1925

(F) Die Spirochäten

Zuelzer, M.

Münchner med. Wochenschrift, Med. Klin., 1919;66:1245

(F) (R) Forschungen über die Ätiologie der multiplen Sklerose.

Kuhn, P. & Steiner,

(Research on the etiology of multiple sclerosis)

Münchner med. Wochenschrift, Med Klin, 1917;13:1007

(F) (R) Über die Ursache der multiple Sklerose.
[The cause of multiple sclerosis.]

Kuhn, P., & Steiner,

British Medical Journal, 1916;i:409-411.

(R) Spirochaetes and their granular phase.

Fantham, H.B.

Observed the process of spirochetal formation of granules, and the emergence of small spirochetes from these granules. "It must also be borne in mind that coccoid bodies may be present when spirochaetes as such cannot be detected." (p.410)

Julius Springer, Berlin, 1914.

(F) Studien über die Fortpflanzung von Bakterien, Spirillen and Spirochaeten.

Meirowsky, S.E.

British Medical Journal, 1911;1:752.

(R) The infective granule in certain protozoal infections, as illustrated by the spirochaetosis of Sudanese fowl.

Balfour, A.

Spirochetes were observed to discharge large numbers of granules. "...the spirochaetes undergo an astonishing change. They discharge from their periplastic sheaths spherical granules, and it is apparently these granules which enter the red cells, develop in them and complete a cycle of schizogony...In process of time the spirochaete loses its activity, becomes difficult to see, and eventually all that is left of it is the limp and lifeless... [that the granules] do not appear to take on the Romanowsky stain may explain why they have not previously been noticed... I have found these granules to be resistant forms and their presence in countless numbers in the tissues might explain part of the mechanism of relapse and the difficulty of curing completely some of the more chronic spirochaetal infections, as, for example, syphilis and

Journal of Infectious Diseases, 1906;3:291-293.

Studies on the Spirillum obermeieri and related organisms.

Novy, F.G., & Knapp,

Found that the classic spiral form is not the only form that spirochetes may assume.

Treponema ...

Zbl. Bakt., 1994;280:297-303.

Formation of multiple treponemes.

Wolf, V. & Wecke, J.

[From the abstract:] "It was calculated that the formation of spherical bodies may reduce their surface by up to 75% as compared to the single form. Thus, the reaction surface for antibodies or other compounds produced by the host is considerably diminished. Therefore, such spherical structures being at resting states may represent a survival strategy of spirochetes. ...the spherical bodies may be the starting point of the new inflammatory episode. This wavelike process is typical of many spirochetal diseases."

Vestn Dermatol Venerol, 199?;4:32-6.

(F)The cytoarchitectonics of hard chancre in rabbits with experimental syphilis exposed to oliusulfon and cefamezine

[In Russian; English abstract available]

Delektorskii VV.,

Describes T. pallidum ultrastructure, and the process of formation of a granule. Cefamezin did not effect spirochetal cysts in the treatment of (snip)

Microbiol. Immunolo.,;1984;28:11-22.

Colonial morphology of treponemes observed by electron microscopy.

Umemoto, T., et al,

[From the abstract:] "Scanning and transmission electron microscopy revealed that the colonies of Reiter treponemes contained spherical forms almost up to 5 am in diameter, each consisting of an outer membrane and a treponemal main body."

Journal of Applied Bacteriology, 1983;55:417-428.

(R)A proposed life cycle for the Reiter treponeme.

Al-Qudah, A.A.

Demonstrates the viability of cysts and the existence of a complex manner of reproduction. "Although transverse fission may be the main mode of reproduction of Reiter treponemes in optimal growth conditions, the spontaneous formation of cysts increases in aging cultures to the extent that it is rare to find a typical treponeme in old cultures. We conclude that such cysts... [serve to] by-pass adverse environmental conditions and to ensure the propagation of the organism. ...the existence of the causative agent of syphilis in a nonspirochetal form has long been hypothesized to explain the latency of syphilis and the infectivity of tissues devoid of demonstrable treponemes...This agrees with what usually happens in protozoa in nature; ...the majority of cysts in protozoa are a means of protecting their contents against unfavorable conditions... Later, depending on conditions when the harmful exposure is past, protective cysts may become multiplication cysts. They are not

Microbiol. Immunolo. ,1982;26(3):191-198.

An internal view of the spherical body of Treponema macrodentium as revealed by scanning electron microscopy.

Umemoto, T., et al,

"External observation of a spherical body by scanning electron microscopy clearly revealed the main bodies [spirochetes] running beneath the inner surface of the spherical body membrane [cyst]. " Includes a freeze fracture photograph of a cross-section of a multispirochetal cyst,

Acta Pathol Microbiol Scand [A],1977;pertenue (sic KMD) Electron microscopy of lymph nodes of hamsters experimentally infected with Treponema

Blom J.

Treponemes were found intracellularly in macrophages. These treponemes did not show their typically helical shape, but were present as spherical forms or cysts.

J. Am. Vener. Dis. Assoc. , 1976;3(2):109-127.

Biopharmacology of syphilotherapy.

Rein, M.F.

British Journal of Venereal Diseases, 1971

(P) Current concepts of the morphology and biology of Treponema pallidum (syphilis) based on electron microscopy

Ovcinnikov, N.M., et al

[Granules:] "Another mode of reproduction resorted to in adverse circumstances consists in the formation of spores which subsequently develop into new treponemes. The breakdown into granules is especially pronounced under the action of penicillin and immune sera." [Cysts:] "Under stressful conditions, the treponeme 'packs' itself into a compact roll (Fig. 8) and becomes covered with a transparent mucoid capsule, which resists the pentration of drugs and antibodies." "Encystment as a mechanism of survival and mode of reproduction is widespread in nature, especially among protozoa." [Intracellular:] T. pallidum were found inside a cell taken from the site of a chancre; and L-forms were found

New England Journal of Medicine, 1971; 284: 642-653.

Diagnosis and treatment of syphilis.

Sparling, P.F.

Includes a review of recent [as of 1971] evidence indicating that penicillin treatment is not always curative in patients with late syphilis. "Penicillin therapy of neurosyphilis has not been as effective [as in early syphilis]. Several studies have reported relapses... Clinical progression of symptomatic neurosyphilis is relatively common despite antibiotics." (p.650) [Diagnostic issues:] Some infected patients also

British Journal of Venereal Diseases, 1968;44:1-34.

(P) Further study of ultrathin sections of Treponema pallidum under the electron microscrope.

Ovcinnikov, N.M., et al,

Observations of T.pallidum cystic and granular formations under the electron microscope. "...under unfavourable conditions of existence, treponemes form real cysts as a method of persistent survival and multiplication, as occurs not infrequently among protozoa." "As the treponeme moves, the thickness changes. This indicates that the body possesses a capacity for contraction... The sharply-marked structural elements of the treponeme and its complex and characteristic structure indicate that cysts are not a product of degeneration. In addition, in cultures where there are many cysts, they are very mobile, which is another argument against degeneration... When transfers are made from cultures containing cysts and almost no ordinary spiral forms, growth of ordinary spiral forms occurs." Includes photo of a treponeme packed into a cyst

British Journal of Venereal Diseases, 1968

Further observations on the persistence of Treponema pallidum after treatment in rabbits and humans.

Yobs, A.R.

Results of a 4-year study of rabbits treated with penicillin for late latent syphilis. Confirmed persistence of syphilis in numerous subjects after antibiotic treatment. Cortisone treatment can reactivate clinical disease. Offers various theories to explain the persistence of T. pallidum, including morphologic changes in the organism. However, the author believes that the existence of a complex life cycle with differing

Vestn Akad Med Nauk SSSR. 1965;20(8):46-50.

[L-forms of Treponema pallidum].

Ustimenko LM

in Russian. No abstract available.
PMID: 5328461 UI: 66154896

British Journal of Venereal Diseases , 1964

Significance of spiral organisms found after treatment in late human and experimental syphilis.

Collart, P., Borel, et al

Persistence of T. Pallidum after treatment. Organisms are still present but have lost their virulence. Cortisone reactivates clinical disease.

Journal of Bacteriology, 1963;85:932-939.

Morphology of Treponema microdentium as revealed by electron microscopy of ultrathin sections

Listgarten, M.A., et al,

[Findings:] Spirochetal granules were found in cultures of T. microdentium. They were more numerous in older cultures. [Observations pertaining to the classification of spirochetes:] "The [cell] envelope had an irregular contour, was easily disrupted during processing, and did not appear esstial in maintaining the shape of the protoplasmic cylinder. It is therefore probable that this envelope is quite distinct from bacterial cells walls, which in ultrathin sections appear as regular, well-defined, electron-dense structures." (p.938)

Journal of Bacteriology, 1961;82:967-978.

Influence of osmotic pressure on the morphology of the Reiter treponeme.

Hardy, P.H. & Nell

"[Reiter] Treponemes in saline solution were observed while distilled water was pulled into the preparation by capillary action, and it was found that although all treponemes in a field were not changed to spheres simultaneously, the conversion of any single one took place

American Journal of Syphilis, 1953;37:29-36.

Treponema pallidum buds, granules and cysts as found in human syphilitic chancres and seen in fixed unstained smears under darkground illumination.

Coutts, W.E. et al

"Spirochetogenic granules are by far more numerous than the cysts."

American Journal of Syphilis, 1951;35:164-179.

Studies on the life cycles of spirochaetes: V. The life cycle of the Nichols non-pathogenic Treponema in culture.

Delamater, E.D., et al

Formation of reproductive cysts.

American Journal of Syphilis, 1951;35:216-224.

Studies on the life cycles of spirochaetes: VII. The life cycle of the Kazan non-pathogenic Treponema pallidum in culture.

Delamater, E.D., et al

Yale Journal of Biology and Medicine, 1950.

The morphology and staining characteristics of Treponema pallidum. Review of the literature and description of a new technique for staining the organisms in tissues.

Campbell, R.E. et al

Journal of Experimental Medicine, 1950;92:247-250.

(P) Studies on the life cycles of spirochetes: IV. The life-cycle of the Nichols pathogenic Treponema pallidum in the rabbit testis as visualized by means of stained smears.

Delamater, E.D., et al

Experimental Medicine, 1950;92:239-246.

Studies on the life cycles of spirochaetes: III. The life cycle of the Nichols pathogenic Journal of Treponema pallidum in the rabbit testis as seen by phase contrast microscopy.

Delamater, E.D., et al

"...it seems likely from these observations that there are two means of vegetative reproduction, consisting of (1) transverse division (the most important under usual conditions); and (2) the production of gemmae or buds which eventuate into unispirochetal cysts comparable to those described for saprophytic forms, within each of which single spirochetes develop and differentiate, and from which they subsequently emerge."

American Journal of Syphilis, 1949;33:101-113.

Morphology, cultural characteristics and a method for mass cultivation of the Reiter spirochaetes.

Gelperin, A.

Journal of Bacteriology, 1948;56:755-769.

(P) Morphologic characteristics of certain cultured strains of oral spirochetes and Treponema pallidum as revealed by the electron microscope.

Hampp, E.G., Scott

"Typical free granules, the end products of granule "shedding," ... consist for the most part of what appear to be short sections of spirochetes closely packed together...Although it is not possible to determine from these micrographs that the granules are germinative units, their constant rhythmic occurrence in living cultures suggests this possibility. Further support of this hypothesis is provided by the fact that cultures up to 31 months old, showing only refractile granules by dark-field examination, have invariably given normal growths on transfer to fresh medium (Hampp, 1946)." (p.768). Also of interest: the authors did not find cell membranes on the spirochetes they examined.

Bulletin of Hygiene, 1947;23:548.

Study by means of micromanipulation of the virulence of one or several spirochaetes as well as viability of spirochaetes or granular forms of culture of supposed Treponema

Bessemans, A.,

American Journal of Syphilis, 1947;31:109-114.

Transmission of experimental syphilis from mouse to mouse in absence of S. pallida and pathologic changes in presence of successful innoculation

Wile, U.J.

Showed that syphilis can be transmitted by tissues from infected hosts in the absence of spirochetes, suggesting that the infectious agent is present in another form. Note: this study does not specifically mention cysts or granules.

J Am Dental Assoc, 1946;33:201-206.

Morphologic alteration of smaller oral treponemas during aging of cultures; Effect of age on viability of spirochetal cultures.

Hampp, E.G.

Journal of Bacteriology, 1943;46:15-24.

(P) Bacterial morphology as shown by the electron microscope; V. Treponema pallidum, Treponema macrodentium and Treponema microdentium.

Mudd, S., Polevitsky, et al

"Irregularly spheroidal, dense bodies... are often found attached to the spirochetal cell, frequently near the end; such a dense body may be in close apposition to the outside of the spirochetal cell-wall or may be connected to it by a short stalk. The evidence concerning these bodies seems to support the interpretation that they are asexual reproductive bodies." (p.23)

American Journal of Syphilis, 1942;26:565-573.

Some morphologic features of the Nichols strain of Treponema pallidum as revealed by the electron microscope

Morton, H.E. et al,

JAMA, 1942;199:880-881.

(P) The morphology of spirochaeta pallida in the electron microscope.

Wile, U.J., Picard, et al

"...in many specimens a curious knoblike structure was seen at the end of many organisms. Their almost uniform shape and density suggest that these are not extraneous particles of the preparation but a part of the organism itself."

Annales de l'Institut Pasteur, 1940;64:439-455.

(F) Etude morphologique du Spirochaeta pallida. Modes de devision. Spirochetogene syphilitique.

Manouelian, Y.

American Journal of Syphilis, 1938;22:294.

Morphologic variations of the syphilitic germ.

Bessemans, A.

Discusses pleomorphism in T. pallidum.

Annales de l'Institut Pasteur, 1935;55:698-708.

(F) Syphilis tardiva. Forms minuscules du Spirochaeta pallida. Spirochetogene syphilitique.

Manouelian, Y.

American Journal of Syphilis, 1932;16:155-190.

(R) The life history of Treponema pallidum. A Critical review of literature.

Ingraham, N.R., Jr.

Excellent, "must-read" review of the findings and various interpretations of T. pallidum prior to 1932. States that there have been 18 separate experiments in which tissues from infected hosts transmitted infection in the absence of spirochetes, suggesting that the organism is present in another form. Of particular interest to the reader: Roukavischnikilff's belief that the cause of syphilis circulates in the blood of an infected animal in an avisual (submicroscopic) stage; and McDonaugh's theory that Treponema pallidum is the adult male phase of a coccidial

Compt. rend. soc. biol., 1930;104:477-480.

(F) Gommes syphilitiques et formes anormales du treponemes, ultravirus syphilitiques.

Levaditi, C.

Compt. rend. soc. biol., 1930;104:736-740.

(F) Cycle evolutif du Treponema pallidum du Spirochaeta pertenuis et du Spirchaeta cunicola.

Levaditi, C. et al,

Compt. rend. soc. biol., 1930;104:72-75.

(F) Relation entre le cycle evolutif du "Treponema pallidum" et la genese des lesions syphilitiques.

Levaditi, C., et al

American Journal of Syphilis, 1930;14:433-437.

(P) Granular transformation of Spirochaeta pallida in aortic focal lesions.

Warthin, A.S. et al

Findings of T. pallidum spirochetes in atypical forms in aortic focal lesions suggested that the possibility that the spirochete may transform itself into a minute granule by a series of contractions. Atypical forms were found even when typical spirochetes were absent. Includes an interesting drawing of the transitional stages observed as a spirochete transforms itself into a minute granular form. The authors raise the question as to whether this progression represents evolution or involution, but seem to emphasize the possibility of involution.

British Medical Journal of Dermatology & Syphilis, 1913;25:1-14.

(R) The complete life history of the organism of syphilis

McDonagh, J.E.R.

Lancet, 1912;2:1011.

The life cycle of the organism of syphilis.

McDonagh, J.E.R.

Poses several questions: (1) Why is the incubation period of syphilis so long? (2) Why do 1-2 injections of salvarsan [used to treat syphilis prior to the discovery of penicillin] not cure every case? (3) In tertiary syphilis, fewer organisms are found; why then is this stage the hardest to treat? The author argues that the Treponema pallidum is the adult male phase of a coccidial protozoan, and that the spores that result from the conjugation of the two sexual phases are the actual infectious agent of syphilis. The spores were observed to develop inside of

Journal of Experimental Medicine, 1912;16:194-198.

Treponema mucosum (new species) a mucin producing spirochaeta from pyorrhea.

Noguchi, H.

Irregular spirochetal forms and many granules appeared when conditions were unfavorable to the organism. Small spirochaeta were seen attached to round bodies as if they had just sprouted from them. Journal of Experimental Medicine, 1911;XIV:99-112.

A method for the pure cultivation of pathogenic Treponema pallidum.

Noguchi, H.

Observed "spore-like round bodies" connected with young pallida. Also of interest: the author reports that he was unable to cultivate T. pallidum in any medium without the addition of tissue.

Journal of the Royal Army Medical Corps., 1911;Vol. XViI, p.225.

Granule-shedding in Treponema pallidum and associated Spirochaetae.

O'Farrel, W.R. Balfour, A

Münchner med. Wochenschr., 1906;53:310-312.

(F) Weitere Mitteilungen über die Spirochaeta Pallida.

Herxheimer, K.

Found that the classic spiral form is not the only form that spirochetes may assume.

Münchner med. Wochenschr., 1905;53:310-312.

(F) Zur Kenntnis der Spirochaeta Pallida.

Herxheimer, K.

Berlin. Klin. Wochschr., 1905;42:673-675.

(F) Über Spirochaeta pallida bei Syphilis und die Unterschiede dieser Form gegenüber anderen Arten dieser Gattung.

Schaudinn, F., & Hoffman, S.

Note: the American Journal of Syphilis was originally called the American Journal of Syphilis, Gonorrhea, and Venereal Diseases.

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