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
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
Breinl
[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.
Leptospira
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.
Non-Spirochetal
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)
Other/Mixed/Unknown
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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