ANALYSIS OF SOVIET ARTICLE ON SYMBIOSIS OF VIRUSES AND MICROBES

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• Publication Date: July 20, 1954
• Content Type: REPORT

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CONFIDENTIAL
CENTRAL INTELLIGENCE AGENCY
INFORMATION REPORT
COUNTRY USSR
SUBJECT Analysis of Soviet Article on Symbiosis of
Viruses and Microbes
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comments on the Soviet article entitled, e ym os s o ruses an
Microbes," by L.'?A.'Ziltber Uspekhi Sovremennoi Biologii, 22: ill-100, 1952.
A copy of the translation of this article is available on loan at the CIA
Library.
I; This article begins with the usual references to historical figures in Soviet
science and their contemporary successors, together with the customary
allusion to the dialectic method. There follow several statements totally
without foundation and a number of notable non-sequiturs. For example, on
page one, it is stated that the viruses in the opinion of most investigators
are, "deprived of the possibility to exist in an outer natural medium." In
actual fact, no one denies that viruses may exist extracellularly, they
merely cannot by definition multiply in theifouter medium" which is quite a
different thing. Furthermore, the fact that viruses may occasionally come
in contact with microorganisms is no indication, taken alone, that "special
attention must be paid to their interaction with the microbes" (page 2,
paragraph 3). On cross infection, viruses also come in contact with argon
in the atmosphere, but no one assumes this to have any particular significance.
It appears that the author regards adaorption of viruses by bacteria as
important because he believes in the Lamarkian view that the viral heredity
can be modified by "contact" (page 3, paragraph 2). There are some
exceptional instances of an effect on bacteriophages dependent on the genetic
properties of their hosts, but I know of no analogy if bacteriophages are
excluded. Presumably Dr. Ziltber is not writing about phages, but about
viruses-that infect animal tissues.
The experiments mentioned on pages 4 and 5 discuss the adsorption of viruses
by yeast cells. While it may be true that the cells were able to remove
virus particles from their environment, a more convincing experiment would
ave shown that virus particles were concentrated on the yeast surface,
rather than merely present. Furthermore, a useful control mould be
repetitioncf the experiment with inert particles in place of the yeast cells,
to see if loss of virus in the supernatant was the consequence of adsorption,
or of inactivation. These experiments of Turevich and Ianushevich were
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evidently not not performed as carefully as those of Zeitlenk described on page
6.
3. In reference to the work of Kling and collaborators (page 9) the author
appears willing to accept the possibility, that poliomyelitis virus multiplies
in sewage by growth within protozoa of the genus Bodo. Although science
advances by an open minded attitude, the role of protozoa is here only a
hypothetical device intended to explain a high titer of polio virus in the
sewage. It is well known that most people infected with polio will not show
clinical symptoms, and hence the assumption of a protozoan host capable of
supporting viral multiplication may not be required.
4. On page 10 the author draws a distinction between physico-chemical processes
as opposed to biological processes. Such a distinction is regarded as invalid
by most Western scientists. Up to this point there has been no evidence to
support the authors thesis, that adsorption contributes to adaptation of the
virus.
5. Most of the experiments described on pages 11-16 are concerned with the
attempts to cultivate the virus of variola on yeast cultures. Very few of
the experiments are quantitative in nature, but depend at best on the some-
what imprecise test of activity of specific dilutions. Quantitative methods
for the study of numbers of infective particles, long used by bacteriophage
workers, have only recently been developed for the titation of animal viruses.
The observation of TuIrchinskaia (page 14) that the virus in yeast cultures
was detected up to the 44th generation, but control sowings from bouillon
cultures became negative in the 3rd-4th sowing, does not prove virus
propagation. It is entirely possible that the conditions when yeast is
present are more favorable for the survival of virus, a possibility the
author does not seem to consider important. A virus preparation of high
titer could be diluted many times on transfer and still produce an infection.
6. Numerous additional work is cited to support the authors theory of
sYmbiosis. It would be easier to evaluate these pages if fewer references
were cited and more critical attention was directed to their evaluation.
The author has made an effort to include SOMB references that contradict
his beliefs.
7. There are certain fairly simple conclusions. that emerge from A study:of.
'."General Results" on pages 28734 of the ZDAber communication, and that are.
apparent to any: microbiologist:
The material disaassed by the author is notable for the fact
that about 85% of, the references appeared in the literature
before 1940, notwithstanding the keen interest shown by
microbiologists of all countries in the possible extracellular
or non-specific cellular propagation of viruses. For? example,
Krueger, in this country, has recently claimed that bacteriophage
titers can be raised extracellularly, but hiSwork probably
involves activation rather than dwRlication. There is no doubt
that uniuspected symbiosis tetween 'Virus and cell zaY well exist,
since it is usual for a virus-oell relation*410 to be detected
only if the adjustment between virds and host is comparatively
poor, i.e., there is a definite pathology.. On the other hand,
the remarkable thing about the relationships between viruses
and their hosts is the specificity involved. A human strain
of virus:might grow in the tissues of monkeys, but is hardly
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likely
tb be maintained in the genetically unrelated cells of
yeast Or bacteria. As previously stated, it appears far more.
,probable that yeast cells provide an extracellular environment
-104hIy favorable for the survival of some kinds of viruses.,
The .virtues of YOast extract ag a contributor to the growth
Or survival of factitious microorganisms are well known to
bacteriologists.'
In the last analysis, modern knowledge.of viral growth maintains that the virus
draws' upon intracellular amino acids, iourine s and pyrimidines for the
synthesis of its replicating units, and that the host range is so specific
that even closely related species may tot serve as host. ,.The authors complete
lack of knowledge of modern biochemistry and his heatr, emphasis on out-
dated literature indicate that the paper is not worth serious attention.
His idea is neither original nor esoteric, and would have been long since
confirmed by many microbiologists, if capable of quantitative verification.
The numerous qualitative experiments described are for the most part without
means for critical evaluation. Since some Russian scientists are undoubtedly
very capable, it would he unwise to take this as a typical example of Soviet
science.
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fro: Uspekhi Sovremennoi Diologii
Vol. 33, pp.81-1400, 1952
On the Symbiosis of Viruses and
Ficrobm!
By
L. A. Ziltber (roscow)
Soviet biological science has ascertained the decisive role
of the outer redium in the development and variability of organisms.
The great Soviet scholars, T. P. ravlov and T. V. Ifichurin in dife
ferent ways, while studying the verious phenomena of nature, came
to one and the rare position on the unity of the organism and mediut.
Numerous scholars of our country with T. D. Lysenko nt the head are
successfully solving the questions of development heredity, varia-
bility, and formation of species in centinuous hareem, with condi-
tions of life.
Productive use of this principle, which has enriched Soviet
science with the greatest theoretical and practical achievements,
requires a detailed study of orgnnisms in close cooperation with
the medium during various and changing conditions of this inter-
action. Offering special difficulties is a sillier study of the
different parasitizing organisms for which the inner medium of
other organisms is, in certain periods of their existence, the
outer medium and which do not breed in the outer natural medium.
The viruses too belong to such parasitizing organisms. The
viruses, being obligatery cellular parasites, in the opinion of
the majority of investigators are deprived of the possibility to
exist in an outer natural medium. it is quite natural that con-
temporary virology has paid much attention to a study of the
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interaction of the viruses with the cells of macroorganisms. The
question even of the interaction of viruses with the outer natural
medium has been studied almost exclusively on the plane of the
investigation of the effect on viruses of temperature, humidity,
and other such factors It is easy to see that such a study
cannot give a proper ilea of the interaction of viruses with the
outer natural medium. This medium is never sterile; it is always
in one or another deuee populated with microbes, and the constant
presence of microbes in this medium cannot be ignored during the
study of the interaction of viruses with the natural medium,
In characterising the dialectic method, I? V. Stalin wrote:
"...any phenomenon in any sphere of nature can be converted into
an absurdity if ie is examined out of connection with the surround-
ing conditions, en breaking away from them; and, vice versa, any
phenomenon can le> understood and substantiated if it is considered
in its indissoluele content with surrounding phenomena and its de-
pendency on the phenomena surrounding it" ("History of Vie (a),,
Short Course", p. 101; 19,50,). This position of I. V. Stalin should
be a guiding fele in a study of the relationships and connections
between the Greeniam and the medium,. Only in this case, if we
shall study flee interaction of viruses with the outer natural
medium in Oese connection with those conditions in which the virus
is found in ehis medium, we shall be able to understand the basic
rules of We reciprocity and take into consideration its importance
for the exitence and propagation of the viruses,
One o: the constantly existing conditions or the outer
natural medium is the presence in it of microbes. An overwhelming
majority o: the viruses is inevitably4*ound with them, not only when
they are e) be found in the outer medium but also during their
direct peesing over from one Organism to another, since the tipsues
of the oveanism through which the viruses infiltrate are not
sterile, Hence, tt follows that during the study of the inter-
action ef the viruses with the outer Medium it is necessary that
specie', attention be paid to their interaction with the microbes,,
he capacity of the viruses for intracellular parasitism
arose as a result of their many centuries of evolution, in the
courg of which the outer medium was altered, and the macroorganieme
and venises were altered as well as the microbes that populated and
were n contact with the outer medium of the tiesue of the maero-
orgaresmle,
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It is difficult to suppose that many centuries of close cone
tact between different living beings have proved inseneible to them
and have not led to any Corms of their coexistence. It is natural,
therefore, to suppose that among the viruses and microbes there may
exist both the phenomena of eymbiosis and the phenotena of antagonism
so prevalent in nature.
The 'probability of such a hypothesis is confirmed further by
the following considerations. It has long been known that microbes
can adsorb the most diverse substances: dyes, colloid metals, etc.
On the other hand, the capacity of viruses to be adsorbed by differ-
ent substrates is known and used for development of methods for
their purification. Finalle, the capacity has long been known of
killed microbes to adsorb viruees. All theSeltacts make valid too
the hypothesis of the adsorAion of vireseStvy live microbes.
Similar adsorption, in whiee closest contacttocoure between them
could in itself be the factor that has stipulated in the range of
evolution the appearance cd any forts of adaptation of viruses to
micrebee,
Thus, the study of the interaction of viruses from the outer
natural medium requires a study of their interaction with the
microbes.
Part of this great problem the phenomenon of the syebiosis
of microbes and viruses, eus studied in our laboratory from 1932 to
1937 in collaboration eith the late F. I. Vostrukhova to the
memory of whom this week is dedicated. Our investigations attracted
the attention of other authors, and in subsequent years rather con-
siderable literature has been compiled on this problem. Now when
the position of the harmopy of organism and medium has become a
guiding principle of Soviet biology it is opportune to supply certain
results of the study of this problem posed and studied by Soviet
scientists.
We shall not set forth here in chronological order the data
to be had in literature to the present time, It is more expedient
for us to give an account of these materials in their logical de.
velopment. Sincl the question of adsorption of viruses by microbes
is quite essential to the study of the symbiosis between them ands
together with it, has separate theoretical and practical.signifi-
cancel, we shall give an account of it before the others.
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Adsorption of Vieuses byjel.ve Aerobes
The adsorption of viruses by different sahatrates has re,.
peatedly served as the subject of investigation at different authors,
however, the question of the adsorption of viruses by living miarobes
has alm:st not teen studied at all.
In our investigations (Miter and dOliaeva, 2935) the ad.
Sorption was studied of the virus of variolar vaccine by the Torula
jr yeast. The choice of the yeast for eXpertmente on the
of the syebioais of viruses and microbes was stipulated by
the fact that yeasts are *widely distributed microbe* often found
in the air And on the skin amen. Furthermore, they are large,
easy to culture, and well. studied ricroorganisms. The very same
strain TertAle kephir, was the first strain with ehich experiments
On symigiairiellnieup, and since these experiments gave positive
results we continued working with this strain further.
Experiments on the study of the adsorption were set up in
the following way. To a suspension of yeast in physiological
solution of sodium chloride or in bouillon the virus of variolar
vaccine. was added in the form of a centrifugate or filtrate of
moist ?soskoban (scraping?) from calf or rabbit or of testicular
rabbit virus. The mixture was left standing in a thermostat at
370 for 2 hours, and in part of the experiments items let stand
for 24 hours in a refrigerator at 5-6?. For control, the same
amount of sirup was added to bouillon and preserved with the same
conditions. At the expiration of the timelndioated the mixture
was centrifuged, the deposit was washed twice with large amounts
of physiological solution, and the content of virus was determined
than in the super.charged fluids the wash waters, and the doposit.
To the supercharged fluid obtained fresh yeast was again added,.
and the experiment was repeated again. Determination of the virus
was made by titration on the skin of rabbits or the cornea of pigs
(Translator's note: probably guinea pigs).
In the overwhelming majority of experiments the washed'
deposit of yeasts contained the virus of variolar vaccine, and
positive results were obtained at 1: 10 and 1: 100 dilution of the
deposit. The first wash waters contained the virus, sonatinas in
considerable amount Cup to a is 1000 dilution). The second wash
waters did not disclose the virus in a single experiment. The
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super.oluirged fluid after first contact with the yeast did not
always give an apparent reduction of titer, *specially if it was
high, but after a second contact the titer was reduced from ten
to a thousand timee.
In experiments in which the yeasts were added to the virus.,
the filtrate of which had a low titer (1400), the virus could
not be detected in the super-charged fluid after first contact.
These experiments revealed to us that living yeast can adsorb
the virus from variolar vaccine.
Turevich and Ianushevieh (1937) studied this question in
more detail in our laboratory. They took into account in their
exPeriments the pH of the medium in which the adsorption of the
virus by the yeast was studied and the yeast was precipitated and
washed (three to five times) during 3000 revolutions of the con-
trifuge in the course of 5 minntes.
Centrifugation was carried out a short enough time to avoid
even an insignificant deposit, together with the yeast of the
variolar virus too. Suitable experiments showed. that yeast,
the use
of moist scrapings containing a, vast amount of elementary bodies
the latter are deposited, even if in an insignificant amount, during
centrifugation for a period of 10 or more minutes. Experiments set
up during the maintenance of all these precautions with filtrates
of moist scraping revealed that yeast, after contact with the
variolar virus in the refrigerator for a period of 2h40 bows and
after washing three times contain this virus and provoke in
animals typical variolar lisease.
Adsorption of the virus in the most considerable' *mount was
no in the acidsone at pH el 5.4 - 6.2. At pH is 7,1 8.2 it vas
loss sharply expressed and was not observed in all oases. Attempts
to study the adsorption of the virus by microscopic examination
of smears did not give Turevich convincing result.. At the
present time by means of electron optics (microscope?) this wee..
tion can be investigated in great detail.
The question of the adsorption of the viruses by live mi-
crdbes was studied also by Zeitlenk (1950) on types of virus of
infectious ectromelia. Filtrates of this virus were blended
through a Berkefe/d V or N candle filtomrwithsuspensions of
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various microbes, the absence of the pathogenicity of which for
mice was verified in preliminary experiments. The minture was let
stand in a thermostat 1.5 to 3 hours, then in the refrigerator for
20 hours, after which the fluid with microbes not yet precipitated
was removed, and the microbe deposit MIS washed with repeated con.
trifugation. For control, the same *meant of virus was added to
the buffer mixture not containing microbes/ and with this mixtura
the same handling was carried out as in the main experiment. The
content of virus was determined by inoculation of mine 1) by
filtrates 2) by a mixture of filtrate and buffers 3) by the last
wash fluid of microbes, Why the last wash fluid of the contras
5) by the deposit of microbes washed frcm ths filtrate, and 6)
by the control precipitate. The following cultures of microbes
were tested by the author: the yellow Swains., the white and the
yellow Ltubylogggeti., prcoides, ands species of yeast
(kephir, kouniass the rose.colored, the thermophilic, and others)a
Aasorption was studied in relation to the pH of the mediums the
temperature, the consistency of the microbe suspension, the time
of contact of the virus with microbes, and so forth.
All these tests disclosed a definite capacity of the virus
or ectromelia to be adsorbed by microbes and revealed, together
with this, that the different microbes potties* an entirely different
capacity to adsorb the virus, Thus, for examples the staphylococci
and sarcina in all 35 experiment., disclosed this capacity 0 *areas
the rose.colored yeast adsorbed the virus only in one out of 14
experiments, but in the knumies yeast this capacity was not
generally observed. In those experiments in shich the phsnomenon
of adsorption, was observed the washed microbes contained virul,
but the last wash water and the controls did not contain virus.
It should be noted that in certain oases the virus was adsorbed
by microbes almost wholly, as the experiments revealed this in
which the deposit was titrated for content of virus.
In the experiments of Zeitlenks the same es in the experiments
of Turevich and Ianushevichs the importance or the pH of the median
for adsorption was noted. As a rules the adsorption proceeded con-
siderably more fully during a slightly acid reaction, with the pH
equal to 6.0-618. In conditions of a slightly alkaline modioa
(pH * 7.4-8.4) the number of positive results was considerably
reduced. The substitution of a buffer solution of horse serum
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did not inhibit adsorption. The virus in the adsorbed state proved
more durable both to preservation in the conditions of the themea
stet and refrigerator and to heating at 55? C.
It is interesting to note that the strain at hephir yeast
which adsorbed well the virus of variolar vaccine in.?W-experi.
rents and the experimente of Turevich adsorbed the virus of ec.
tromelia poorly in the experiments or Zeitlenk. Thai, in the
phenomemme of adsorption both the properties at the microbe and
the properties of the virus play a role.
All these data have left no doubt of the fact that at least
certain viruses are adsorbed by microbes in the conditions of the
experiment, Does this process take place in natural conditions?
Falikovich and Ianushevich (1937) have devoted special investigation
to this question.
They sowed fresh variolar calf 2r4Oh in is 3.0 dilution on
blood agar poured intoa Petri dish. After 24 hours of standing
in a thermostat all species of the colonies cultured were reamed
from the dish to bouillon. The bouillon cultures were introduced
intracutaneously into rabbits, which were killed with e blow an
the 4th.5th day with the typical reactions present. The papules
on the skin that were separated off from these rabbits were readily
cut into from the inner side, and a spell amount of exudate was
collected, diluted with sterile physiological *elution, and in-
vestigated for the presence of the virus of the vaccine after
verification of the sterility. The presence of the virus was es.
tablished by inoculation into the cornea of rabbit or into the
dhorio-allantoid membrane of chick eibryo? eith subsequent di
closure of the typical reaction and of the elementary bodies.
During the investigation of 90 strains of different microbes
isolated from 30 series of alit lymph it was determined that out
of 39 strains of white staphylococcus 34 were bearers of the virus
of the variola vaccines out of 24 strains of yellow sarcine two
strains were bearers of the virus, out of 15 strains of yellow
staphylococcue six strains were bearers at the virus, and out of.
22 strains of diphtheroid bacillus not one contained virus..
Thus, these experiments showed that the virus of variolar
vaccine can be adsorbed by certain microbes and, in natural csindi
tams, adsorption here too bearing a selective character.
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All these investigations of Soviet anthers dinonstrets with
certainty the possibility at adsorption at islet of certain virtue*
by microbes both in everimental and in natural Asonditions. Itoredga
autbers until recently paid no attention to this quseitiOn.
Thddric de Is Riviera (1930) observed that staphyl000ceue isolated
from variolar calf lymph provokes at inoculation into the skin of
rabbit variolar pustule, and at inoculation keratitis on the (soma
of (guinea!) pig. But the role of the StePhifiaebeeue in the re's
&dim provoked by variolar vaccine .interested his mainly.
In connection with our works, Amiss (1936) studied the ad.
sorption of the virus of the vaccine by thebij yeast
and by white staphylococcus sad did not iiid?iption of
the virus by these microbes. In the overwhelming majority of eases
Andes used in his experiments a washed sluspension of elementary
bodies, in the preparation of which their adsorption capacitor was
possibly changed. Moreover, he did not take into account the pH
of the :medium in whieh the adsorption occurs. It is surprising
that Amies did not observe the precipitation of the elementary bodies
during their centrifugation for a period of 30 minutes at 3500 rem-
lutions per minute, In the above-mentioned experiments at Turevich
the precipitation, as indicated, occurred at a far earlier period,
and Amiss should have been able to detect the virus in the yeast
and staphyl000ccuse-precipitates if not in an adsorbed state, then
as a result of the settling during centrifugation. However, Amiss
reports no such data, It is entirely likely that Assimit negative
results were stipulated by the use of washed elementasy bodies
and the unfavorable hydrogen figure of the reedit* in which the
experiments of adsorption were conducted.
In connection with our data cited same, as also in connect.'
tion with observations that established the frequent occurrence in
the outer medium of the virus of poliomyelitis, foreign investigator*
too have occupied thenrelvee in recent years with the question of
the possibility of adsorption and the carrier ability in general
at viruses by microbes.
In our works it has been pointed out that the survival at
Times in the outer medium is possibly etipulated by their spa
-
biota* with micro:bee. This opinion too is placed at the base of
the torts of foreign investiigators. Kling and his collaborators,
Olin, rahriews and Norlins (1942) studied the presence Of the virus
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of poliomyelitis in sewer waters of one of the Markham district..
The number of virus detected was so considerable that it was Eleossi.
saw to ammo either the possibility of the marrying of the 11X011
by all the residents of this district (shalt 100,000 persons) or
ite seatiplicetion in the sewer waters. The author" =Pr...Fed the
hypothesis that this multiplication occurs is igabissie with protosca
of the genus Bodo.
Evans ani Ostend (1946) tried to esoartain whether the vine
of policegelitis propagatee in cultures with protases (the _Bodo
gems, Mouas? and several others) by *Xing to cultures grown?Tia
the warraorable conditions a au*Pension of spinal cord contain.
int the virus. After incubation, for a period of 544 dales mow-
wars nada. The fluid of different pnerationst treated with
ether and centrifuged for riddance of bacteria, was introduced into
the brain of monkeys. The experiments disclosed no propagation of
virus in the protosoa cultures. The authors did not study the Phew,
nomenon of the adsorption of the virus by the miarcerganisms.
The experiments of other authors have disclosed, however, a
certain possibility of adsorption and the ability of protosoa and
bacteria to carry the virus of policeeralitis.
Toomey and coworkers Taksics and Schaeffer, WO) studied
the possibility of the abilitt of Amoeba proteus to carry the virus
of poliomyelitis, The authors cultured the bu in water to which
bacteria, killed by boiling, bed been added. After contact vitth the
virus of poliomyelitis the amebae were repeitedly meshed (up to
times) with water and were then introduced into the brain of riLso4
Experiments revealed that amebas can, trap the virus of poliomyelitis
or adsorb it securely on their surface. Washed uLth 50 cc. of
sterile water from five to 11 times, they still infected mice, pro-
ducing the typical disease. The authors not* that the virus was not
detected in the fifth wash water, but was detected in the litho
and they explain this by the fact that on3,y prolOnged washing
destroyed the neabrane of the ameba and admitted the virus to the
surrounding fluid, Microscopic investigation actually disclosed
coniiderable demolition of the amebae after the llth wash(ng. The
*true was not discovered in the amebae 3, lt, and 6 days after con.
tact.
Recently Ioung, Felsenfeld, and Byrd (1949) (here printed
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? 10 ?
as Young, in the bibliography as Ibung) diselosed the poesibiliAy
of the virus of poliomyelitis (the Laming strain) to be adsorbed
byte eta ea as well as by the intsetinal becalm* erid by a
? bacillus. It is interesting to mote that the
authors observed the great iMMUnit0 of the virus adsorbed btr tbe
amabss and bacteria to the temperature at the thermostat. The
mambos contained the adsorbed virus after I hour, of incUSation
of the cultures in the thermostetm No promotional` tbe Virus
mas abservedby the authors.
The aggregate of all of the tboveomentioned facts leaven no
doubt that the viruses ars adsorbed by microbes and protosoa both
in experimental and in natural conditions.
This adsorption is not the ueual physico?chemicel procera
although it proceeds during definite physico?chemical conditions.
The adsorption of the viruses by the micrase bears a seleetive
Character stipulated by the biological and not the physico?chemical
peculiarities that have a hand in the reaction of th0 microorganisms,
One and the same virus is adsorbed to an entirely different degree,
or is not adsorbed at all, by the various microbes in Ore end the
same conditions. On the other hand, different viruses can be ad.
embed by one and the same microorganism. All this permits think.
ing that the phenomenon of adsorption of viruses by microbes la a
biological phenomenon connected, mith the adaptation of the virus to
the outer mediate'
Blubiotic Cultures
The above?mentioned consideratiorm have given occasion to
suppose the possibility not only of the adsorption err/raise by
microbes but also their maltiplication in the microbe cells,
Suitable experiments were set up by us and later on by other investi-
gators mith viruses of variolar vaccine, fowl variolsvmeasles,
herpes, rabies, encephalomyelitis of horses, Seine plague, and hoof
ana mouth disease, The circumstance that the ridoettsies are not
dietinguished from the viruses by the conditions of their culture
has given oacasion to study in syMbiotio matures the virus of
exanthematous typhus. The symbiotic cultures of the virus of variola
with different species of yeast wore studied in greatest detail.
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The Virus of Variolar Vaccine
In 1932 we reported (Zillber and Vostrukhova - translator's
notes the latter is spelled Vostroukhova in the bibliography .10
1932, 1933) on the possibility of growing the virus of variolar
vaccine for a prolonged period in a bouillon culture of yeast,
The filtrate of damp nsoakoban (?scraping) was added to a 24-hour
culture of Torula Itephie yeast, and this culture was let stand in
the thermoiara 370-iT; Reamings were made every three days in
a test Ube with ordinary meat-peptone bouillon, In control test
tubes the virus was added to the bouillon without yeast, and sowe
info: were made at the same time. The content of virus in the test
tubes was studied by intracorneal inoculation of pigs, an well as
by intracutaneous and intratesticular inoculation of rabbits. In
these experiments we used the Torula h21.51. strain
Symbiotic cultures of the variola virrs? which can be trans-
ferred ten times with preservation of virulency, were obtained by
the aboveementioned method. One of the cultures was transferred
over more than one and a half years and still kept its activity
after the I2Oth reaming, A study of these cultures had revealed
that they have a nueber of peculiarities that distinguish the virus
in these cultures from the ordinary virus of the vaccine. One of
these peculiarities consisted of the fact that the cultures did not
take at cutaneous inoculation:, but took well at intracutaneous
inteednetion? as well as at intratesticular and intracerebral
inoculation of rabbits and pigs, provoking typical alterations and
the ammulation of elementary bodies (translator's note: same as
inclusion bodies) in the tiepues, The cultures took well also at
inoculation into the cornea of animals. The leek of the capacity
in syMbiotic cultures to take at cutaneous inoculsedenItus
stipulated possibly by the close connection of the virus with
the yeast cell, Experiments in which such inoculation was sue
-
easeful if the cultures were earefully pulverised beforehand in a
mortar with sand served as proof of this hypothesis, The viruleney
of eyebiotic cultures wts inthe first ten generations rather high)
Thus, for example, strain 662 in the 39th generation took at dilw.
tion of 1: 100,000, strain 305 took :in the 29th generation at dills-
tion of 1: 50,000, etc.
The greatest dilution at which inmulaticemas observed to
take was that of ls 5000000. However, far from all strains had
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similar virulency, and it OA not remain constant. Atter Several
edscadess (tens) of generations the virulency usually dropped
rather considerably. A special investisationmie together with
AG Timakov shamed that not every yeast ceIl is s, vine bearer
A person is easily convinced of this at sowing a eyebiotie culture
on agar and at studying the virus content at esCh colony. At
selecting colonies that contain virus and at reviewing thee on
bouillon it vas possible to increase considerably the virulency of
the syebiotic culture and to maintain it by this eatised at a high
levele
In all cases at successful inoculation the culture always
succeeded in disclosing elementary (Inclusion) bodies in the
tissues. But all attempts to disclose elementary bodies in the
cultures produced no results. A study of the antigenio properties
of syMbiotic cultures disclosed their capacity to Cause immunity
and the formation of antibodies to the variolar virus a certain
tine after inoculation, However, the intensity of this process
was not very sharply expressed, Thus, for example, the sere of
rabbits ismunized by symbiotic cultures completely neutralized
3.0 infectious doses of variolar calf lymph neutralisation of 20 or
100 doses was not constant, figs and rabbits isesuStsed by cultures
proved immune to subsequent inoculation by 10 infectiousdosesof
variolar calf lymph? but this immunity WS relatively brief and
weal; not observed in all cases. Thus, the stedy of the antigenic
properties of symbiotic cultures disclosed the presence in them of
the variolar virus, but with somewhat altered properties,
A histological atudy of the skin of a rabbit inoculated with
symbiotic cultures disclosed typical alterations and Ousrnieri
bodies (Khurgina? 1935).
It should be pointed out that the symbiotic suites.* ob.
timed by us of variolar virus were turned over to three institutes .
to the L. A. Tarasevich Control (Institute), to the TsIeM? and to
the Kharkov Mechnikav (Metchnikoff) Institute. Investigation of
these cultures, both in the generations obtained by us and in sub.
sequent ones, discloacd the presence in them of the variolar virus
(Gil' gut,1935; Korolikova? 1936; Khastovich and Shur, 19$5)0
The aggregate of all these data have permitted us to make
the conclusion that the virus of variolar vaccine combo propagated
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in syebiotic cultures with certain yeasts and Web cultures mutes
reaown for along time in the laboratory. It is entirelr natural
that adaptation to yeast cells causes an alteration of the properties
of a. virus. This alteration is not, however, stable, and the virus
from a yeast culture passed through animal theme quickly return
to its initial properties.
Our investigations on istbiotic cultures of the Tires of
variola were verified and confirmed by a number of authors.
Oilogut (1935) atudied the synbiotic cultism of 16 strains
of variolar vaccine withIreeteir yeast. In eVerimosnita from cix of
these strains symbiotic cIis showed quick wakening in the first
generations and in all the rest the virus was detected in cultures
of the litho 9th, 17th, 19th, 20th, and 234 generations. Also
successful were certain of the author's experiments at cultivation
of the virus in syMbiotic cultures with staphylococcus and sarcira.
Determination of the presence of the virus in the cultures was dons
by inoculation into the cornea, elementaey bodies sleep being die..
covered in positive cases. Cutaneous inoculation of rabbits vitt
oyebiotic oultures produeed no results as was also the case in cur
experiments. At intracutaneous inoculation of yeast cultures into
rabbits the authors observed only slight reactions and not once
observed generalisation. One of our yeast cultures of the virus
of variola, brought by us in the laboratory to the 74th generstitee
was sent to Dr. Giligute and she still detected the virus in this
culture after 13 further generations (up to the 87th inclusivaly:e
The immunity test in seven rabbits inoculatedwith eyMbiotic
cultures by smearing on their skins undiluted calf lymph gave neve.
tive results. The euthort, however, indicates that these results
were stipulatedi by negligible reaction or by its absent* at
inoculation of the cultures.
Worollkova (1935) studied three strains of eyebiotic cultures
of the virus of the varicaar vaccine, two of which were obtained
tran our laboratory (leth and 8th generations) and one was obtained
by sowing in ITggr yeast the virus of the neurovaccins. The first
two proved stIIItive in the 34th and 20th generations, the latter
in the 26th generation. The activity was controlled by inoculatleen
into pig cornea and into the skin of rabbit, and the specificity
in a considerable portion of the experiments was oonfiemed by the
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discovery of elementary bodies, The author notes that the intra
cuticular inoculatiote of rabbits "were attended by the formation
of pockmarks over the whole surface of the integument, reaction
ustally having been lacking at the place of the introduction of
the needle and the pockmarks having appeared at a'distance of
2.3 cm, from the place of injection", Inoculations of cultures
into heifers were uneueceseful.
Positive results of cultivating the virus of variola in syme
bleeds with the yeast were also obtained by Isabolinskii, Levtsov?
and Cherniak (1935), The activity of the cultures was maintained
in their experiments up to the 17the2Oth generations. Attempts to
cultivate the virus on sareinas produced no results, least cultures
the virus kept their activity at reuovieg on agar, as well as at
preservation for a prolonged period at rook temperature,
Timakov (1936) cultivated the virus of variolar vaccine in
symbiosis with kephir yeast, staphylococci, and sarcinse Cultures
vith yeast kept?ariirus active to the 24th generation, the same
as certain cultures with staphylococci, Cultures with gamines
ware nonevirulent after the 13th to the 21st generations.
The author not only studied the presence of virus in the
cultures, but also determined its titer in many resowings, The
titer of the virus proved eufficiently high, and certain cultures
of distant resowinge (15 . 17) were active in a dilution 'up to
1: 25,0000 The cultures took for the rabbit only at intracutanoous
introduction and the smearing of cultures onto the ekin gave no
positive results. In these experimenta there was also noted the
a pacity of yeast cultures to produce in rabbits a generalized
process. By selection the author succeeded in considerably bconte
ing the virulency of the cultures obtained by him. The ke hir
yeast both in his works and in the worte of Gil' gut and Karol kova
was entirely &Pathogenic,
Tullchinskaia (1936) studied in detail the question of the
culture of the variola of fowls in symbiosis vithlAireast.
The Virus in the yeast cultures was detected 0 to th
generation, whereas in control resowings in bouillon without the
yeast it was possible to discover the virus no further than in the
34th reeowing, In skin affections observed at introduction of
cultures elementary bodies were found. In certain cases at intro-
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duction into the skin of a yeast culture (10th generation) a
generalised process and the appearance of pockmarks on the mucosa
of the mouth, eyelids, etc were observed, The author observed
weakening of the virulency from the 16th to the 18th generations
and the disappearance of it from the 35th to the 44th goner Mons,
Yeast cultures of chicken pox in the 12th generation. and of pigeon
pox in the 16th generation provoked, moreover, distinct reaction
and eebaequent immunity to the natural virus. Immunity was verified
in chickens by the smearing of natural chicken virus on the
scarified skin of the comb and wattle and by implantation from sick
chickens to immunieed? and in pigeons by smearing pigeon view onto
the scarified follicles of the breast, and in both cases 45 aye
and 5.6 months (? respectively) after the initial affection, 11 of
the chickens so tested and 19 of the pigeons did not fall sick from
the variola, while there was sickness of one and all of the controls,
An analogous experiment with yeast cultures of 20th to 21st
generations gave negative results., and in the birds made ill again
after infection with yeast cultures no immunity was discovered at
aubsequent introduction of natural TirkW, Tulachinskaia noted a
further interesting cireumstanoe, She added healthy fowl to fowl
that had been diseased by yeast cultures, provoking in them di
tinct disease, These healthy birds that had been in close contact
eith those that were inoculated did not fall ill1 however, which
permite supposing the modification of the virus in the symbiotic
cultures and loss by it of contagiousness.
Delanov and Riakhovskii (1936) tried to culture the virus of
variola? the diphtheria of fowls in symbiosis with brewers a yeast
and sarcinas, The authors tested only the second generation of
their cultures, inoculating it into two chickens by smearing on
acarified club and wattle, At obtaining negative results, the
authors did not continue their eipeeimentu. Negative results were
also obtained by Aries (1935), who in his main experiments used
washed elementary bodiceof variolar vaccine, as well as Voet
(1935) in some experiments with nsurovaceine and Arbeit (1935)
in experiments with vaccine and exanthematous typhua.
All the aboveementioned data leave no doubt of the fact that
the virus of variolar vaccine can keep its activity for a long time
in symbiotic cultures with yeast and certain bacteria? All
investigator? who followed our method and did not abandon their
work after unsuccessful exteriments with the first cultures of
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16 -
microbes and strains of virus that fell into their hands received
positive results*
iether. Virusste
Besides the symbiotic cultures of the virus of variola:
there were studied symbiotic cultures of other viruses also,
Smirnov and coworkers (1935) cultivated the virus of measles in
eyebiosis with kephir0 bread, and brewers' yeast. The blood Of
measle Patientsets Rn at onset: was diluted with Tyrode 's
oolution in the proportion it 7 . 1: 100 and 3$ drops of this
blood were sown in a test tube with freshly sown yeast., Relsowe
ings were made every 6.7 dare, and detection of the virua in tho
cultures was made by inoculation of rabbits and in other eXperi-
ments. The control rabbits were inoculated with the blood of
patients: taken at onset, as well as by the initial yeast cultu.ee,
Parallel with these experiments there were set up experiments of
the inoculation of rebbits with measle virus cultures on Riverle
endium. Judgment NW made of the specific reaction of the rabbite
by the rise of temperature 5-14 days after infection: by the
Appearance of the catarrhal phenomena of the conjunctivee and of
the mucosa of the nose: by leukocytoeis in the first 2148 hours
after infection: and subsequent leukemia and alteration of the
henegremo The specific pathogenicity was disclosed in other
experimehts too. A comparison of all the data obtained permitted
the authors to conclude that the measles virus is preserved in
yeast cultures for a period of 35 or more generations*
Unfortunately the authors did not harn of our instruction
e411 the neceasity to use for cutaneous inoculation pulverized
eulturee. It is entirely likely that had they so done their
eeoulte would hair been oven more demonstrative* Afanasleva and
hapiro (1936) also studied the possibility of cultivation of
oeeslec virus in sytbiosie with, IhsEtle! yeast, They came to the
eonclnsion that such Cultivaeion does not turn out well in all
we:mei; and symbiotic cultures possess weak pathogenicity for rebbtee,
degvia (1527) observed prolonged survival of measles virus in
eultures with different coccie
Izabolinskii, ievtsov, and Cherniak (1935) reported on the*
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obtaining symbiotic cultures of rabies virus with kephir yeast.
The data cited by them, however, are unconvincing.--Tiii-loss
(or killing?) of rabbits on which testing of the cultures was
carried out was observed on the second day when there were
phenomena of paralysis. No bacteriological controls were set
up. Insufficient passages were made. No histological investi-
gations were made of the brain.
Rosengolets and Karnaukhova (1934) cultivated the virus of
rabies on Micro lyp.odeicticus. The cultures were tested after
5-12 days f growtE7WinoCiaiiion of rabbits and pigs subdurally
and of mice into the muscles of the. femur.
One of the cultures (the 10th generation) was inoculated
into 27 mice, 13 rabbits, and 4 pigs. Dying from rabies with the
confirmation of further passages were /1 mice, 3 rabbits, and two
pigs, Incubation in the majority of cases was from 25-60 days,
although the three-day was also observed. The authors reported
likewise on certain successful experiments at cultivating the
virus of rabies with kephir yeast, In cultures of micrococcus
the authors also grew tiii-Virus of street rabies. A dog was in-
oculated with the 6th generation; it died with the phenomena of
paralysis 7 days after inoculation. Its brain was passed through
eabbits, in 18 out of the 26 animals paralysis having been observed.
In two rabbits inoculated intramuscularly with the brain of the
dog, inclusions were observed similar to Negri bodies, and in sub-
oequent passages into animals these formations were not observed,
Akker and Florinskii (1935) obtained no results in attempts
to cultivate in _symbiotic cultures the virus of rabies. In their
experiments with kephir and brewers e yeast there were observed not
only propagation Bit -also survival of the virus, Moreover, the
yeasts at intracerebral inoeulation proved pathogenic. Experiments
eith sarcina were successful*
Palavandovs Sertbrianaia, and Pugach (1936) cultivated the
virus of rabies in mixed culture with yeast and Micr. lysodeicticus,
The culture contained the virus only to the 3d ienerairodp-iihieli
the authors explain by the survival of the virus. Since in control
sowings in bouillon the virus was not detected in these generations,
the authors think that in conditions of mixed .culture the virus
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finds conditions better for its preservation than in bouillon.
Entirely negative results were likewise Obtained in the
carefully set up experiments of Vinogradova (1936), Waldhecksr
(1936)a and Portniagina (1939),
All these materials permit our thinking that there were
obtained no syebiotic cultures of the virus of rabies with the
yeast, The positive results reported by certain authors are une
convincing and act not contain all the proofs needed of the spe-
cificity of the Pathological processes caused by the cultures
being investigated.
Offering far more interest are the works of Minervints
laboratory. Minervin studied the possibility of the carrying of
the virus of rabies by microbes in natural conditions,
Minervin and Rapoport (1936) studied the microorganisms of
the mouth cavity of rabid dogs in the character of bearers of the
rabies virus.
The authors sowed the mucus of the oral cavity of dogs
destroyed by rabies on dishes of blood agar and then after 2 days'
growth transferred the isolated colonies to bouillon. The bouillon
generations (up to the 7th) that had developed in the course of
3.5 days were tested by inoculation of rabbits, with subsequent
pa$sages of their brain when disease was present. The cultures
ware introduced into rabbits subdurally in very small dosage, from
five to ten million microbe bodies. With the use of this method
there wan observed in the rabbits after 9.60 days of incubation a
disease with the clinic characteristic to experimental rabies.
The disease proved to be possible to pass several times from one
rabbit to another, the organs and blood of the rabbits having
been bacteriologically sterile. The authors did not observe in the
brain of rabbits inoculated with cultures the typical Negri bodies,
but saw formations similar in certain respects to these bodies,
The clearest results were obtained with one gram-positive bacillus
and with streptococcus.
These observations indisputably have great interest. Un-
fortunately they were not continued and set up sufficiently erten
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sively, Meanwhile, namely in this direction, it is expedient also
to study the problem of the symbiosis of the rabies virus with the
nicrdbes. At escaping from the central nervous system into the
mouth cavity the visus of rabies is encountered in the saliva with
many microorganism, and it is necessary to seek among them the
objects of possible syMbiosis. It is necessary to take into con.
sideration that tais symbiosis is certainly not attended also by
complete preservstion of all the biological properties of the
virus - it, mayte expressed simply in the fact that the virus in
the microbe cu! ire is preserved in those conditions of the medium
in which by itself it dies. Unfortunately this question, a
principal one from our point of view, has remained outside the
field of vis/on of many investigators, namely from the need there
would be to jegin the study of the eyMbioais of the rabies virus
with the mixdbes.
As fa- other viruses that infect the central nervous system,
then, inasmuch? as the majority of them penetrate into the nervous
system, escaping into the nonsterile cavities and tissues of the
animal erganism, it is possible to judge their symbiosis with the
micrdbc,: only in respect to those microbes with which these viruses
are enrountered in the organism of the conveyor. From this point
of V1%711 it would be interesting to study the "ire of tick en-
cephalitis, inasmuch as it is transmitted by ticks from generation
to gAeration and circulates in their organism for a long time. Of
course, it is first. necessary to clarify whether the virus cirou-
latia in the organism of the tick in. nonsterile tissues.
In order to complete the account of data obtained during the
e,udy of the symbiosis of viruses that affect the nervous system
lith microbes, it is necessary to dwell briefly on data obtained
sith the virus of poliomyelitis and herpes.
Brutsaert and coworkers (1946) tried to cultivate the virus
of mouse poliomyelitis in cultures of tick bacillus, a mixture of
intestinal flora, of Leptospira, and of various protozoa. In cul-
tures of Trichomonas haninis the virus may be found in the lith
generstionTEr crai-iisults were obtained with a killed
culture of the sane Trichomonas. It should be noted that the
authors used media marriRiiI)e to the growth of the protozoa
being studied, departing from our instructions on this problem.
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In natural conditions obviously such media are lecking. Moreover,
in a nueber of experiments they introduced into the animals fil-
trates of symbiotic cultures, evidently assuming that the virus
is propagated in the medium and not in the microorganisms.
Certainly these experiments could not give positive results. It
is possible that they would not be positive too with observance
of the necessary conditions of the experiment. Inasmuch as the
herpes virus can be found with microbes of the integuments it
would be possible to doubt in it the capacity to adapt itself to
them.
Grundfest (1934 - translator's note: spelled Gruntfest in
the bibliography) cultivated the virus of herpes with Ilp_ole
yeast. The virus was observed in six subsequent generation,
provoking typical affection and creating specific immunity in
reinfected animals. In further generations the virus was not
detected. Histological alterations in the brain of rabbits that
died after inoculation with symbiotic cultures into the cornea
were fully typical to the herpetic. Symbiotic cultures kept their
virulence for A period of 3 months when preserved at room tempera-
ture.
Interesting data were Obtained during the study. of symbiosis
with microbes of the virus of hoof and mouth diseaie. As early as
19330 together with Eel. Vostrukhova, we tried to obtain sym-
bietic cultures of the hoof and mouth diseaee virus. Experiments
of obtaining these cultures by means of our strain of Torula "Tpl.tr
gave no positive results. More successful Were investTianniTIE--
which we tried to detect the carrier of the vireo of hoof and
mouth disease in natural conditions. At isolating from guinee
pigs infected with hoof and mouth disease various cultures from
their diseased seems E. I, Vostrukhova separated out a sluggishly
growing streptococcuss which indisputably was the carrier of the
hoof and mouth disease virus and provoked in the pigs the typical
disease, This property was preserved in it in eight resowingss
whereas in control resOwings:of the virus on the same* however with..
out a microbe medium, it was no longer detected in the third gener-
ation, These investigations were not continued and were not pub-
lished.
Frenkel (1934) obtained negative results at endeavoring to
obtain symbiotic cultures of the virus of hoof and mouth disease
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with Torula kephir yeast. Poppe and Busch (1936) likewise detained
negative results in experiments with this yeast. However, the
latter authors received indisputably positive results at cultivation
of the virus of hoof and mouth disease in symbiotic cultures with
Torula =bra and with certain other species of yeast. Cultures of
orra mouth disease virus with Torula rubra in the 60th
generation provoked in guinea pigs not only iI? but also a
generalized process. The breeding of the hoof and mouth disease
virus initialAy sown in the yeast culture reached the gerstration
figure of 103'. With yeast of certain other species, Torula Kiel
and Z. (sic!), the experiments were also positive and the cultures
contained virus in the 10th and 28th generations. With other
species of yeast (Torula pulcherrima, the wine yeasts, and others)
and with certain biZteiIa75U-AMEoccue? sardine, and others) no
symbiotic cultures were obtained.
The authors explained the importance of the presence of a
neutral reaction of the medium (bouillon),, in which the syMbiotic
culture is grown, as well as of the medium in which the virus is
suspended at extraction of it from the tiesuee For the latter pure
pose a phosphate buffer proved completely inadequate, whereas with
Tyrode 'a solution good results were obtained. Thus, Poppe and
Buschis experiments established with certainty the possibility of
obtaining symbiotic cultures of the virus of hoof and mouth disease
mith certain specieseof yeast.
In systiotic cultures the virus of the plague of swine use
likewise studied. Likhachev (1937), proceeding from the fact that
enterococcus and sarcinas are most often isolated from the blood
of swine affected with plague, tried to cUitittete the virus of
this disease in symbiotic cultures with these microbes. The 5the
10th? and 21st generations of these cultures.preduced at intro...
duction into young pigs a pig plague typical in clinic and path?.
logicaleaeatomical picture. The blood or the sick animal, tested
in one case produced the same disease *t introduction into A
healthy animal. For control there was an Absence of virus in the
installation in which the experiments were carried out and there
leas a lack of contagiousnese for the. disease?preehiced by the
eymbiotic cultures; in adjoining pens healthy young pigs were
kept. These young pigs fell ill with plague on the 17th and 28th
day after the start of the experiments and succumbed on the 24th
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and 41st days which indicates their infection from the experi-
mental animals.
Cultures of enterococcus and sarcina without the virus
produced no pathological phenomena in the young pigs. The
animals that survived the introduction of the 21st generation
of symbiotic culture were tested for immunity and did not din-
close ouch. Earlier generations were not subjected to such
testing. The experiments with two other strains of virus gave
no positive results, but a total of three generations, the Mho
5ths and 9th, were tested.
All the materials stated show that not only the virus of
variolar vaccine can survive and multiply in conditions of
nymbiosiseaith certain microbes, but that this capacity is inherent
to other viruses too.
Experiments with Exanthematous Typhus
The provoker of exanthematous typhus is not, as known, an
ultravirus (filtrable virus), but is a rickettsia. However, the
rickettsias in the conditions of culture are quite similar to the
viruses. They, as the latter, are obligatory cellular parasites
and are incapable of multiplying on artificial nutritive media-,
These properties of theirs can make possible the adaptation of the
rickettsias to other microbes, the more to that the provoker of
exanthematous typhus is constantly found with microbes in the in-
testines of lice. The possibility of the alteration of Certain
properties of microbes under the effect of the Virus of
exanthematous typhus was demonstrated by us an early as 1922 when,
at cultivation of Proteus vulgerie in the abdomen of a young pig
with exanthematouirtt?iira--iilfiairewaa Obtained of the
exanthematous typhus proteins, distinct from any whatsoever isolated
from the exanthematous typhus organism. Minervin (1935), at
introducing his exanthematous typhus into pig testicle, observed
an analogous alteration of the Proteus vulgaris. It should be
noted that tila virus of exantheaTais this possess no pronounced
eytotropicity to cells of a definite species and type. Thus, it
infects mans guinea pig, rabbits and mice, and it multiplies in
not only their brain but also inthe tissue of the lungs, spleens
and other tissues, All these data have also given us basis to
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try to obtain symbiotic cultures of the exanthematous typhus virus.
Experiments set up by approximately the same method as with the
virus of the variolar vaccine (Ziltber and Dosser, 1934) disclosed
the presence of the exanthematous typhus virus in cultures with
kephir yeast and sarcinas? in which the blood of exanthematous
iiP%-i (victims) was sown in a dosage of 0,2-0.3 ml. or 1/200 of
the brain of exanthematous typhus pigs, The presence of the
virus in these cultures was demonstrated first of all by experi-
ments of inoculating pigs 'with them, the pigs having disclosed a
temperature reaction typical to exanthematous typhus, the disease
hewing been passed bacteriologically into sterile brain up to five
times. In the diseased pigs rickettsias were discovered in the
tunica vagiao, as too the histopathological alterations character-
istic to exanthematous typhus. In lice affected by symbiotic
cultures by the rectal method rickettsias were discovered: and
they died 15-20 days after inoculation. Inoculation of mice by
the nasal route was not carried out since this method was not
known at that time. Rabbits inoculated with symbiotic cultures
produced an accumuletion of agglutinins to Proteus X19 in a titer
of 12 80, however not in all easel).
This work of ours attracted the attention of many investi-
gators. 0elltser and Hemshilov (1934) cultured the virus exanthematous
typhus (the blood of patients) with Saccharomyees cereviseae. The
15th, 16th, and 24th generations of these cultures were introduced
into pigs, in which on the 12th37th day after inoculation there
was observed a rise of temperature within the limits of 0.8.20 and
a duration of from 4 to 6 days. When passages were made with the
blood and brain of these diseased pigs, the rise in temperature
occurred on the llth.18th day and the fever lasted 7.15 days (most
often a days), In all the affected pigs there was noted a drop
in weight. At introduction of cultures, as well as of brain of
the passage pigs into rabbits there was observed in them the
appearance of the WeileFelix reaction with a titer of 11 40 to 1t 60.
In the brain of the affected pigs there were observed histopathological
alterations characteristic to exanthematous typhus. In a subse-
quent work Oelttser and Hemshilov (1935) tested a further series
of generations of symbiotic cultures, using for this purpose also
kephir yeast, and obtained by this principally the same results.
In later generations of certain strains (18th-36th) the virus
was not discovered, the weakening of the virus having proceeded with
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different rapidity in different strains (strain I in the 34th
generation, and strain III in the 18th generation). The experi-
ments of the authors with the obtaining of immune aera by symbiotic
cultures are quite substantial. These sera were obtained by
immunization of horse, calf, and sheep and disclosed a considerable
titer of the Weil-Felix reaction, Thus, the sera of horse and
heifer raised the titer from 1: 100 to is 1200, and the sera of the
sheep from 1: 50 to 1: 1200. These sera, tested after 42 days of
preservation, in all experiments neutralized the passage virus of
exanthematous typhus. Pigs into which a mixture of immune serum
and virus was introduced gave no signs of exanthematous typhus and
disclosed no immunity at their subsequent inoculation with passage
virus, An attempt to vaccinate guinea pigs with symbiotic cultures
killed in various ewe produced no results, which is fully under=
atandable in the light of contemporary data, since the number of
virus in the symbiotic cultures was not sufficient for this purpose.
Iakovlev (1934) observed in cultures of the exanthematous
typhus virus with yeast and sarcinas formations of various structure,
which he considered to be different stages of the metamorphosis of
the rickettsias, There have been offered no direct proofs of this.
Kenna and Danishevskaia (1933) in a preliminary report
stated the positive results obtained at inoculation of the 24th and
49th generations of a yeast culture of the exanthematous typhus virus
that had been transferred daily or every other day. However, the
material cited was little conclusive since the typical temperature
curve was observed in a total of one of the three pigs.
Symbiotic cultures of the exanthematous typhus virus up to
the 6th and 8th generationa vere studied in detail by Tokarevich
and Kliachko (1935). The disease typical to exanthematous typhus
wan observed in 13 out of 24 of the pigs inoculated with these
cultures, the main indices characterizing the temperature curves
? having disclosed considerably greater diversity than is observed
at inoculation of the pigs with passage virus. Histological altera-
tions were discovered in four out of 12 of the pigs investigated,
they having been discovered in three out of five cases with typical
fever and in one out of seven cases with atypical fever. The cul-
tures in a number of cases provoked the appearance of the Weil-Felix
reaction in rabbits in a titer of 1: 40 and 1: 160. The immunity
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was tested in a total of five pigs, one of them not having had a
general fever at introduction of 001 of brain of passage pig and
the other having given a brief (2-3 day) rise in temperature with
major remissions. Levkovich (1934) also Obtained principally the
paw) results.
Afanasleva and Sterkhova (1934) cultivated the European
virus of exanthematous typhus in symbiosis with kephir and brewers'
yeast. They came to the conclusion that euch callgElon
"succeeds in rare cases and in a limited number of generations",
the exanthematous typhus virus proving slightly pathogenic and
lessening its immunizing properties. Beet results were obtained
with yeast cultures of rat exanthematous typhus, which "disclose
all the characteristic reactions natural to this virus and die- .
tinguish it from the epidemic, such as: the fever reaction in whits
micas the fever and scrota/ reaction in pigs, and the slight affec-
tion in them by granulomatosis of the cerebrum", However, all these
indications in the cultures were expressed considerably more weakly
than these were observed at affection by passage virus. The
disease in pigs, provoked by yeast cultures of the virus, communi-
cated immunity to the rat passage virus, but did not transmit
immunity to the European. One of the cultures lost pathogenicity
only in the 34th generation.
Mins Frenkman (given as Freknan in the bibliography), and
Urban (1935) tried to cultivate the virus of exanthematous typhus
in symbiosis with yeast (with which or the yeasts is not indicated)*
At affection by soma yeast cultures not containing the virus they
observed in the pigs the temperature and histological alterations
typical to exanthematous typhus. They also observed a positive
Weil-Felix reaction in rabbits into which only the yeast was
introduced.
The authors promised to report on further investigations of
this paradoxical phenomenons but this report did not follow,
Gnutenko and Friauf (1935) cultured the virus of exanthematous
typhus in symbiosis with yellow sarcinas on a solid nutritive medium.
Out of the 12 experiments positive results were obtained in four
cams,
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ompaneete aid coworkers (1936) studied cultures of the
virus er exantheme,ous typnas in symbiosis with yeasts (kephir,
bread, and brewer1?). The cultures were tested by inoculation of
guine4 pigs and b/ observation for the Weil-Fekix reaction in
rabbfras and goati that had received these cultures..
The authees note the drop in virulency of the virus as be-
giyeing alread: in the second generation, However, rabbits which
was immunizec4 by the cultures, beginning with the 4th generation,
veva in a cowiderable percentage of the cases a transition from
negative Weil-Felix reaction to the positive with a titer up
ea 1: 40, ari in two the transition of the positive reaction was
eevelt a titee of ls 60 to a titer of 1: 320. No virucidal properties
are discovired in the goat serum.
Ottt (1934) and Voet (1935) obtained negative results in a
few attewts to cultivate the virus of exanthematous typhus in
symbiotiv cultures,
Heaervin, Ziltberman, and Getbillskii (according to the
bibliography, Gebrillekii) (1936) isolated from the intestines of
lice takmn from exanthematous typhus patients bacterial cultures
which proved capable of producing in pigs at subcutaneous intro-
duction typical exanthematous typhus fever and histopathological
alterations entirely characteristic to exanthematoue typhus,
In certain cases the disease with a typical picture was passed by
the brain of affected pigs bacteriologically sterile,
In a subsequent work Ziliberman and 0erbillskii (1938)
reported that the 4th and 6th generations of the virus cultures
from bacteria isolated from lice did not disclose the presence of
the virus at inoculation of pigs, Likewice? attempts to cultivate
the exanthematous typhus virus in cultures of Proteus vul aria,
yeast, and narcinas proved unsuccessful. Geriarrir ne o
Proteus emlgeeiej formerly in contact with the blood of
exanthematous typhus patients: acquired the ability to by aggluti-
nated by the serum of rabbits immunized. 'with Xle? Cultures of
prOtevas yeast, and earcina that were taken into the experiment
by the authors, by themselves without the virus.: provoked in the
pigs a nueber of pathological phenomena in a series of cases.
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In judging the above-mentioned data it is necessary to take
into consideration that all authorS who have studied the properties
of symbiotic cultures have compared them with a passage virus well
adapted to the guinea pig organism. Of course, syMbiotic cultures
could not disclose in full measure those properties which are
observed in a passage virus, but the presence of the exanthematous
typhus virus in these cultures in many experiments was sufficiently
demonstrated in a persuasive way. This was recognized by all these
authors who carefully noted the absence of a complete conformity
of the properties of the cultures and of the passage virus
(rokarevich and Kliachko, Golltser and Nemshilov, Levkovich and
others),
In certain cases the cultures of yeast that were taken for
the experiments were insufficiently carefully selected and pro-
duced in the pigs certain pathological symptoms. However, in the
majority of authors who studied this problem the cultures of the
microbesynbionts were completely apathogenic to guinea pigs and
produced no reaction in them, whereas the same cultures after
contact with the exanthematous typhus virus produced in the pigs
a number of symptoms typical to exanthematous typhus. The virus
certainly weakens and modifiwi in symbiotic cultures, but,
together with this, it is preserved in them in those conditions in
which it dies when it is without microbes (for instance, during
multi-month keeping in a thermostat).
Both in our experiments and in the experiments of other
investigators rickettsias were not discovered with certainty in
the symbiotic cultures, as the elementary bodies of the variola
virus were not discovered. Apparently the rickettaias of
exanthematous typhus can be represented either as formations of
considerably lesser dimensions or as formations with a different
than usual capacity for staining. By this apparently is explained
too the difficulty of discovering the rickettsias in the brain of
affected pigs. The problem of the symbiosis of the virus of
exanthematous typhus with the microbes could be productively
studied at the present time owing to the reproduction of pulmonary
exanthematous typhus pneumonia in mice and the possibility of
studying symbiotic cultures by the electron microscope.
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Goneral Results
Tho data cited above indicate sufficiently conclusively the
existence of symbiosis between the microbes and viruses. The form
of symbiosis described above was in due course designated as an
nallobiophoria" (i.e. as a bearer of another life). Most often
the term "virophoria", i.e. a carrier of viruses, is used. This
last term is more concrete and better expresses the nature of the
phenomenon. Entirely likely too is the existence of other forme
of aymbiosis besides the virophoria. In literature there are
reports (Iakovlev? 1949, and others) in yhi41 it is indicated
that the viruses survive in a medium populated with microbes, not
entering into any closer connection with them. Evidently certain
microbes can produce substances that make possible the survival of
the viruses in the outer medium. This question has been very little
studied.
The great similarity of the phenomena of bacteriophagy an
virophoria attracts attention. Regardless of whichever point of
view is adhered to in respect to the nature of the bacteriophages
it is impossible not to recognize that it possesses the principal
properties that characterize the ultraevirases . the capacity to
propagate only in living cells, the lack of a capacity for growth
on artificial nutritive media, the filtratability, the stability
to certain disinfection substances, etc. Aecently the data of the
electron optics (?microscope) have permitted referring with
certainty the bacteriophages to the ultraviruses. The processes
observed in the presence of bacteriophagy are typically virophore
phenomena. It is known that the phage is readily adsorbed by
suitable bacteri and can in such an adsorbed form be preserved
for a long time in bacterial cultures. If with this it was taken
in comparatively small amount, then there occurs no dissolving
in this culture, and it is a typically virophore culture. Those
cultures of bacteria from which by different manipulations a
bacteriophage is successfully obtained are such virophore cultures.
It is interesting to note that at sowing the bacteriophage culture
on agar the some phenomenon is observed as was noted at selection
of virophore cultures of variola: not every colony is affected
by phage and virus.
However, the bacteriophage is transferred ad infinitum .
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together with the bacterial cells, attaining vast concentrations
and causing the dissolution of the cells, in contrast with the
virus in the virophore culture, which, although it undergoes
certain alterations, yet sooner or later is freed of virus,
especially if it multiplies in favorable conditions, It is possible
to suppose that the phenomenon co:': hacteriophagy in the process of
evolution arose from the phenomenon of virophoria? The virus that
had an affinity also to the animal cells and to the microbe (cells),
and is probably nonpathogenic te the animal, can in the process of
evolution empire a more pronouuced affinity for microbe cells and
become pathogenic for them. We would not have resolved to express
this hypothesis if we hadn't man certain possibilities of its
experimental verification. If us assume that permissibility the
bacteriophage to the provoker cO: typhoid fever rises from the
virophore culture prior to the :eitter microbe that bears the virus
that had an affinity to the celly of the intestines, then it meld
be possible to expect that certain strains of this phage still
preserve remnants of the affinity to the intestinal cells. It is
possible to clarify this extY!rirke'atally by studying the capacity
of these cells to adsorb the phagk and preserve it, Obviously for
control there should be studied also the analogous capacity of the
other cells. Such an investigatioa might also produce material
that would permit using the phage Lore efficiently for purposes
of prophylaxis and the treatment of infectious diseases,
In a further study of the pheLemenon of virophoria it is im-
possible to break away from the thewatical foundation on which it
is based. In virtue of this, first cC all it is necessary:
1, To study the symbiosis of tio viruses with those microbes
with which a given virus in encounterel, in natural conditions,
Hence, it is necessary to study the eyebioside of the virus of
variola and of hoof and mouth disease etth the microbes of the skin
and mucous meMbranes of the cerrespondiea animals, the virus of
rabies with the microbes of the saliva ct does, the virus of
inguinal lymph-ogranulomatosis with tha 'aerobes of the skin, the
virus of grippe with the microbes of the vral cavity and naso-
pharynx, the virus of tick encephalitis will-the microbes of the
ticks, that of Japanese encephalitis with ;..he microbes of
mosquitoes, etc,
2, To study all format of this eyeblosis? and not only the
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propagation of the virus in the symbiotic cultures. None the less,
and more important in practice, is the phenomenon of the carrier
proper, which is not obligatorily attended by multiplication of
the virus in the microbe cell. Namely this virus carrier too
should be studied in greatest detail.
3. To study virophoria necessarily first of all in natural
conditions. By isolating the virophore bacterial cultures from
diseased tissues it is easiest of all to obtain a clear idea both
of the presence of symbiotic phenomena in a given infection and of
their character. Of course, testing practices are an essential
supplement to such experiments.
Ito With this study substantial attention should be directed
to the length of the maintaining of the virus in the virophore cul-
tures and to the conditions of this preservation.
5. The study of the capacity of microbe cultures to become
virophoric likewise should approximate natural conditions. For this
purpose it would be necessary to introduce the culture being in-
vestigated into affected tissue and then to isolate it for several.
days
serious attention during the study of the virophore phenomena
should be paid to those alterations which occur in the virophore
cultures both in the microbe-carrier and in the virile. Meisel'
(1935) observed a number of cytological alterations in yeast in
symbiotic cultures of the virus of variola.
It should be noted that during all these investigations it
is never possible for the determination of the presence of the
virus to be limited only to clinical observations, It is always
necessary to supplement them with the morphological (for instance,
by the discovery of Paschen and Guarnieri bcdies during variola)
and by immunological investigations. This is necessary not only
to determine the presence of the virus, but also to differentiate
the phenomena provoked by it from those provoked by the microbe
carrier, which likewise may be to one or another degree pathogenic.
The simplest method for such differentiation is the inoculation by
virophoric cultures both of healthy animals and of animals that have
been treated, on the one hand, with serum for the vitas and, on the
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other hand, with serum for a given microbe. For instance, in
studying the virophoric culture of the stapnylococcus that bears
the virus of the group, it is necessary to inoculate not only
normal mice but also those which had received (the following
therapy) prior to this introduction: one group - the anti-
grippe serum, the Other . the antiastaphylococcus serum. The
resultsathue obtained permit making reliable conclusions even
in a case in which the pathogenic mircobe is studied in the
character. of carrier.
A study of the phenomena of the symbiosis of the ultramicrdbes
with the microbes ddes not have just theoretiCal interest. It is
indisputeble that la conditions of the symbiceis viruses are
preserved longer than in the absence of microbea. Consequently,
thanks to the viropheria, not only are the conditions of the
dispersienof the viruses altered, but there it also formed a
unique reservoiraefiruses in nature. ataiSaimpossible to under-
value theaimportanCaof this supplementary reservoir for certain
virusesa Earlier: the impossibility of the existence in outer
nature of ultrevirusie pathogenic to manaandaenimals was considered
firmly established, It was thought that they exist only in the
organism of theSeanimaltaand in the organiSMacif the vectors - the
mosquitoes, ticks, 'lice, etc. Now it ila:**Whathat this is not 30
and that viruses, like phages, can existainathe outer medium in
,
symbiosis With microbes. Thi- circumstance should he taken into
consideration in thiaepidemiology of certain ultravirusdiseases,
for which much is still in the dark up toatheapresent time. It is
enough even to recall the epidemiology of peliemyelitis. The
phenomenon of the adsorption of the viruseaby the microbes has
practical importance teo. Sergiev and coworkers (l9!) and others
have revealed that microbes that adsorb the viruses can be used
in certain cases for discovery of antibodies to these viruses, the
reaction of the agglutination of the microbes loaded with the virus
possibly having diagnostic significance.
A study of the phenamena of the syMbiosis of viruses and
microbes gives substantial materials for understanding the in-
accuracies of those investigators who, at isolating during virus
diseases various microbes that are carriers of the viruses, have
mistaken them for the provokers of those diseases. To the same
category of phenomena belong also reports of the conversion of
viruses into microbes.
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It is sufficient to look at the microphotographs presented
in Boshlianis book (1950) to be convinced that, for instance, the
forma described by him as the spherical form of the provoker of
Infectious anemia of horses are yeast. Inasmuch as the yeast can-
not be the provoker of this disease, it is possible to explain
its specific infectiousness only by the fact that it is the
carrier of the virus of infectious anemia,
As has been pointed out above, the phenomena of the symbioais
of viruses and microbes do not deplete by themselves those Inter.
relationswhich exist in them in natural conditions. The presence
in them of antagonistic relationships is definite too. The last
question offers very great interest in connection with the dis-
covery in recent years of the inhibiting effect of bacterial
polyeaccharides on the propagation of certain viruses. The
further study of the problem of the interaction of the viruses
and microbes, first posed nnd studied by Soviet scientists, will
enrich acience with facts of great theoretical and practical
importance,
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