JPRS ID: 9809 USSR REPORT METEORLOGY AND HYDROLOGY NO.1, JANUAY 1981

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- JPR,S L/ 10071
26 October 1981
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~ SS~ Re or~
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LIFE SCIENCES
BIOMEDICAL AND B~HAVIORAL SCIENCES
CFOUO 14/81)
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JPRS L/10071
26 Gctober 1981
~ USSR REPORT
_ LIFE $CIENCES
BIOMEDICAL AND BEHAVIORAL SCIENCES
(FOUO 14/81)
CONTENTS
_ BIOCHENISTRY .
Factor Promoting ComplPte Translation of Encephalomyocarditis
Viral Genome 1
IntegratiQn of Genomes of ttussian Tick-Borne Encephalitis Virus
(TEV) and Cell During Chronic Infection Induced by TEV and
CV40 4
MEDICINE
_ Interferons in Medical Theory and Practice 8
PHYSIOLOGY
- How We See What We See i4
Methads of Mathematical Bio~ogy, Book 2: Methods for Synthesis
of Algebraic and Probabilistic Models of Bioiogical Systems..... 17
Methods af Mathematical Biology, Book 3: Methods of Synthesizing
Dynamic Models of Bialogical Systems (VITZ Textbook) 22
Changes in Endocrine Gland Function During Exercise: End~crine
Mechanisms of Regulation of. the Body's Ada�~tation to Muscular
. Activity 28
RADIATION BIOLOGY
Biochemical Bases of Action of Radioprotective h.gents 30
- a- [III - USSR - 21.a S&T FOUO]
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HUMAN FACTORS
Mathematical Model of Human Operator Used in Flight
Simulators.�~�~�����~�~��~~.�r��.���~.~~��~~��~~~�~��~~~~~��~��� 34
Reduction in Adaptiation Time in Ergatic Control Systems........... 40
Automatic Speaker ldentification by Voi..e........��������.�������� 45
Specific Laws of ~iological Regulation............�.��������~��.�� 69
Visual Situation Simulators in Aircraft Trainers 73
PSYCHIATRY
Clinical Aspects of psychopathic Dynamics 76
PSYCHOLOGY
Problems of Group Unity 85
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BIOCHEMISTRY
UDC 57E.858.23
FACTOR PROMOTING COMPLETE TRANSLATION OF ENCEPHALOMYOCARDITIS VIRAL GENOME
- Moscow DOKLADY AKADEMII NAUK SSSR in Russian Vo1 258, No 4, 1981
_ (manuscript received 13 Feb 81) pp 1013-1015
[Article by Yu. V. Svitkin and V. I. Agol, presented by Academician A. S. Spirin
on 1 February 1981~ Institute of Poliumyeliris and Viral Encephalitides of the USSR
- Academy of Medical Sciences~ Moscow~ M., V. Lomonosov Moscow State University]
[Text] The RNA of the encephalomyocarditib (EMC) virus with cell-free protein-
sqnthesizing systems, depending on the conditions is considered either completely,
or the translation is limited primarily by the 5'-et.~ region of the viral genome
[1-5]. Limited translation is governed ~y the functioning of a relatively weak
"terminating signal" which is located in the viral RNA somewhere on the boundary
between the 5'-end region that codes the precursor of the structural protein (preA),
and the central region, in which the nonstructural polypeptide F is coded.
Under optimal conditions, termination does not occ~~r at this site, however, ~~he
rate of translation sti11 is noticeably reduced [6]. Similar, although weaker,
"terminating" or"inhibiting" signals are apparently found on other sections of the
RNA of the EMC virus. Slowing down of translation on definite sections of the
matrices has been described in nonviral systems.
We recently advanced a hypothesis on the existence of a special factor of trans-
lation which promotes overcoming of similar weak "terminating signals," and called
it the elongation factor~X (EF=X)' [4,5]. This report presents the first data which
confirm this hypothesis. Our ~:cperiment~? wgFreach consisted oF the following. We
initi~l ly c~blained ~i system by ~~rtificial meana in which tr~nslati~n on the bc,uncl~iry
1,~~~wc~rn ~I~~ R~A sec~t ic~n5 oC the f~:~1C virus which code the pr.eA ~nd 1~ components
was drastically slowed ~:own. We then attempted to overcome this translation
barrier by addition of different protein fractions isolated from the uninfected
ceLls.
t:lycerine to a final concentration of 1~% was added to an extract made of Krebs-2
cells (fraction S30) [1,2] treated with micrococcal nuclease. Nine millimeters of
this extract were superposed on 4.5 ml of 1.25 M of sucrose prepared on buffer
solution A(65 mM KC1; 3.3 mM '.`:gC12; 30 mM tris-HCl; pH 7.6; 0.1 mM EGTA; 2 mM
dithiotreite, DTT) and centrifuged (12 h, 44,000 rpm, 2�C) in a Ti-50 rotor (geckman).
1
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- The test tube contents were divided into ~ portions: upper 6 ml (fraction S120)'
lower 4 ml ("sucrose layer") and precipitate (ribosomes). whereas the prot~ins
contained in the sucrose layer were preci.pitated out by ammonium sulfate, the pro-
teii~ precipitates were dissolved in bufrer B(100 mM KC1; 20 mM tris-HC1; pH 7.6;
0.1 mM EDTA; 1 mM DTT; 5% glycerine), dialyzed against the same buffer and stored
at --70�C. Protein concentration was defined according to [7].
The reconstructed cell-free system (25 ul) contained 12 ul (0.05-4.07 units of A26Q)
of f'raction S (preliminarily passed through a column with sephadex G-25 balanced
by t~uffer A wi~~ 3.3 mM DTT), 0.5 U1 (0.4 units of AZ 0)of ribosomal suspension
in t~uffer A, 2 ug c~f RNA of EMC virus, 2.5 of ~~e stu~ied factor preparation, KC1
(fir..al concentration 130 mM), MgC?2 (3.3 mM), S-methionine (10 uCi, 2 uM), 19
unma.rked amino acids, nucleotides and ATP-regenerating system [1,2J. Incubation
was doiie at 30�C under SO ul of Bayol F. The reaction was stopped by adding
SO }~1 of dissociating mixture and the products were analyzed by electrophoresis
in a 15-% polyacrylamide gel [1].
~.ccnrding t~ the previously obtained results [1,2J, when an unfractionated
~ extract (S30) i~ used as the cell-free system, complete translation of viral RNA
occurs (figure lA, track a, see insert). This is indicated by the presence
among the products of translatfon, besides the preA polypeptide and products of its
splitting (A, B, D1, a, G, p 14, p i2) coded in the 5`-end section of the genome,
_ nJ_so of polypeptides F(centrul region), C, D. and E(3'-end region). In contrast
to this, thz ~ystem which is reconstructed from the ribosomes and S120, having
higl-: total ~protein-synthesizing activity, does not guarantee effective formation
of F~olypeptides coded in the central a.nd 3'-end regions of the genome ~figure lAs
track c). Since in the high-molecular region of the ge1, certain nonstructural
pol}'peptides.are covered by incomplete chains of preA, it is convenient to judge
the presence of a barrier ~ihich prevents translation of the central region of t.:he
~enome irom the presence or ab~ence of polypeptide F. Incomplete rranslation in
the re~onstructed system is apparently not associated with a deficiency of tRNA,
- sinc�e ~ddition of a surplus of tRNA practically does not affect the nature of the
forcied products (figure lA, tr:;ck d) .
The capacity of the reconstructed system to implement complete translation of
RNA o~ the EMC virus is restored as a result of addition of protein fractions
whi~~h are isolated from the "sucrose layer" by precipitation with ammonium sul-
fate (fig lA, tracks e-i). We assume that the fractions contain EF-X. For further
pt~rification, material was taken which was precipitated from the "sucrose layer"
ti~ith saturation of (NH4)2SO4 betwe~n 50 ~nd 70%, since in this material, there
are relati~:iely few ballast proteins. As is apparent on figure 1B, the activ~
beginning (EF-X) is adsorbed on DEAE-sephatsel and can be eluted at 225 (but not
at 100) mM KCL.
The obtained result can be explained by the presence in the "sucrose layer" of a
factor which promotes the overcoming of weak terminating signals (i.e � EF-X).
Hourever, the possibility of another interpre.tation also existed. It c4uld be
as~~umed that our reconstructed system is defective not for elongation, but proteo-
lyt.ic activity requir.ed for the formation of viral proteins (including F) from the
corresponding precursors. From this viewpoint, the appearance of F in the
pr~~sence of active fractions of the "sucrose layer" would be explained by the
pr~~~ence in them of the necessary proteases. In order to clarify the correctness
of. tt~is hypothesis, the reconstructed system was incubated with viral RNA for 3 h.
2
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Elongation was stopped by adding cylcoheximide, and incubation was continued for
another 15 h either in the presence or in the absence of material con~aining EF-X.
It was found that incubation of polypeptides synthesized in the reconstructed system
_ with EF-X preparation did not result in the appearance of polypeptide F(.fig 1C).
Th�s, the hypothesis on protease nature of the factor we studied was not confirmed.
_ We note that during lengthy incubztion (18 h), a certain quantity of polypeptide
F is formed in the reconstructed system sven without the additic~n of ma.terial from ~
the sucrose layer. This agrees with the already mentioned concept that the block
preventing synthesis of polypeptide F can be manifest not in complete stopping,
but only in a considerable delay in translation. It still remains unclear
whether this slow overcoming of the translation block is associated with residual
quantities of EF-X, probably present in the reconstruct~ed system, or with the fact
that slow movement of the ribosomes in the correspon~ing regian is also possible
with complete absence of EF-X.
An important question is whether EF-X corresponds to the known components of the
protein-synthesizing apparatus. The answer to this question requires further
studies. in particular, purification of EF-X. However, even no~a one can note that
EF-X is apparently not the RI~;A-bonding grotein, since it is r.ot adsorbed on the poly-
(V)-~ellulose (fig lI'). In this respect, it is distinguished from the previously
described eukaryote factors of elongation (EF-1 and :F-2) and the majority of
factors of initiation of eukaryote translztion [8,9].
BIBLIOGRAPHY
1. Svitkin, Yu. V. and Ahol, V. Z. DAN, VoZ 238, 1973, p 744.
2. Svitkin, Yu. V. and Agol, V. I. FEBS LETTS., Vol 87, No 7, 1978,
3. Pelham, HRB, EUROP. J. BIOCHEM. Vol 85, 1978, 457.
4. Agal, V. I., Chumakov, K. M. et al, in SOV.SCI. REV (BIOLOGY), Vol 1, 1980, 319.
- Agol, V. I. in PROGR. MED. VIROL. Vol 26, 1980, p 119.
~ 6. Shih,D. S., Shih, C. T.. et al. J. VIROL., Vol 30, 1979, p 481.
7. Warburg, 0. and Christian W., BIOCHEM. ZS� Vol 310, 1941, p 384.
8. Vlasiky S. P.; Dom~gatsky, S. P~ et al. FEBS LETT. Vol 116, 1980, p 8.
9. Ovchinnikov, L. P., Spirin, A. S. et al., FEBS LETT., Vol 88, 1978, p 21.
~ COPYRIGHT: Izdatel'stvo "Nauka", "Doklady Akademii nauk SSSR", 198i
9035
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i'nR (1N ~1('1 ~1 i !C}' (1N1
UDC 576858.25
INTEGRATION OF GENOMES OF RUSSIAN TICK-BORNE ENCEPHALITIS VIRUS (TEV) AND CELL
_ DURING CARONIC INFECTI~N INDUCED BY TEV AND OV40
Mascow DOKLADY AKADEMII NAUK SSSR in Russian Vol 258~ No 4, 1981
(manuscript received 16 Jan 8I) pp 1000-1002
[Article by I. D. Drynov, L. V. Uryvayev, V. V. Nosikov and Active Member of the
USSR Academy nf Medical Sciences V. M. Zhdanov, D. I~ Ivanovskiy Institute of
Virology of the USSR Academy of Medical Sciences, Institute of Molecular Biology
of the USSR Academy of Sciences, Moscow] ~
i
[TextJ The development of a chronic viral inf ecti~n, according to one of the i
theories, may be associated with integration of genomes of the virus and the ~
chr~nically infected ccll [1]. Experiments on transfection of sensitive cells with
- DNA preparations made of chronically infected cells in a number of cases confirm
the possibility of including the DNA-copy of the genome from an RNA-containing
infectious nononcogrnic virus in the cPl~ular genome [2]. The presence of a lir.iited
' number of virus-sp~-cific sequences in nuclear DNA of chronically infected ce11s [3]
has been shown by the met~od of molecular hybridization.
I-~ order to study precisely the integrat~ion variant of. viral persistence develop-
ment, we use;i combined infection of cells with Ru~sian tick-borne encephalitis virus
(TEV) and CV40. In combined infection, it is hypothetically possible to conjugate
rhe processes of reproduction of the genomes from both viruses, and to incorporate
the TEV genome into the cellular DNA similarDirect detectiontofnvirushs eci~ficenome
P
into the DNA of the transformed cells [5].
- sequences in the cellular DNA is posaible by the method especially designed to
establish localization of individual genes in the DNA fragments which develop under
the influence of differ.ent rzstrictases [4]. This method was successfully used to
' study the nature of OV40 genome incorporation into the DNA of the transformed
cell~ [5].
L1 10 cells were infected with TEV, strain "MF" which was isolated from chronically
in~ected cells grown foY several years (11]. The original TEV strain "Sof'in"
durin5 infection of the cells jointly with OV40 did not induce the appearance
of transfixing activity of the DNA preparations made of infected cells. After
incubation of the culture cells with TEV, reinfection with OV40 was done. The
cells were then repassivated several_ times. DNA was isolated from the nuclei of
the infected cells by the method of phenol deproteinization with prThemobtained
treatment with pronase and subsequent treatment with RNA-ase [6].
DNA preparations were used for transfection and for work with the restrDNAasre-
- In the experiments to transfect kidney cells of pia embryos with SPEV, p
parations were used for the infection (20-40 ug/10~ cells). The control was DNA
preparations treated with DNA-ase an3 RNA-ase (50 ug/ml).
4
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RNA of TEV was isolated from a purified virus by the standard method of phenol
deproteinization. The genome RNA of TEV is typical messenger RNA with molecular
weight on the order of 3.0 MD (40 S).
Re~trictases were isolated by ths method of Green (7] ar.d Bickle [8J. DNA pre-
parations from infected, uninfected cells and thymus of a calf were ex~osed to
splitting by restric~ases. DNA from uninfected cells L121~ and the DNA of calf
thymus were used as the control. The latter was selecfed as the control because
during synthesis ~f 32p-kDNA by TEV RNA, the seed crystal was thymus DNA that was
fragmented into short chains under the influence of DNA-ase I. The material was
_ further applied to lamellar gel of 1% agarose. Fragments of DNA of the a-phage
formed under the influence of restrictase were used during fractionating of the
material by electrophoresis as the markers to determine the dimensions of the DNA
fragments.
DNA fragments from the agarose gel were transferred to nitrocellulose filters by
- the method of Southern [4]. After drying, the filters were incubated for 24 h
at 65�C in Denhardt's solution [9]. After addition of labelled preparations, kDNA
hybridization was done in the same solution at 65�C for 48 h.
The drie~ filters were exposed after hybridization and washed of the free labelled
material with RT-1 films at room temperature for 3-12 days.
32P-kDNA was synthesized with the help of revertase using scattered or statistical
inoculation, short DNA fragments formed under the influence of DNA-ase I on DNA of
calf thymus. The reaction was done according to th~ method of Taylor [10] with
concentration of the seed crystal 0.25 ug/ml.
After a preliminary series of experiments to select suitable restrictases, the
virus-specif~c sequences were detected in the DNA fragments of the cells infected
with TEV and OV40. Figure 1~see insert between pp 916-917) presents the results
of radio autographic finding of virus-specific seqaence~ in the restriction frag-
- ments of nuclear DNA. The following are clearly revealed: DNA fragment after
resctriction of Eco RI with molecul.ar weight of 6.0 M'il, DNA fragment after res-
_ triction ofLgl II with molecular weight of 11.0 MD, and DNA fragment after res-
triction of Pst I. In the case of DNA t-reated with Hind III, no obvious bands of
hybridized material were noted. The latter can be govern~d by the fact that small
- DNA fragments developed, while the rate and level of hybridization were propor-
= tional to the molecular weight of the interacting fragments. With equal experimen-
tal conditions, part of the DNA fragments with high molecular. weight first becomes
a hybrid. In the control DNA of the L1210 cells and the DNA uf the calf thymus,
no hybridization band was observed.
The sizes of the restricticn fragments of DNA that hybridize with kDNA by TEV RNA,
according to the molecular weight ~f one chain coincide with the dimensions of the
~;enome RNA of TEV in the ~ase of Eco RI, or exceE~d the dimensions of TcNA in the
case of fragments wt~ich develop under the influei~~e of Bgl II and Pst I restrictases.
Tl~e latter can be governed by the presence in the fragments of DNA sequences that
adjoin the continuous sequence of the DNA-copy of the viral genome from two sides.
Th~s may ~e entirely associated with the fact that under the influence of various
restrictases, one DNA fragment develops which bears the virus-specific sequences.
5

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~
The findings first of all confirm the possibility of incorporating the DNA-copy
of the TEV genome as a single block. For more complete discussion of the results
it is expedient to present data on the transfixing activity of cellular DNA [11],
reducing them to Table 1 which is convenient for examination. The data presented
in the table indicate the absence of infectious properties of the DNA from tlie
cells infected with TEV only. The infectious properties of cellular DNA appear
after combined infection with TEV and QV40 (TCD50/m1=6.0). Infection of the cells
with TEV jointly with the virus of the Rous sarcoma (RSV) results in the er~ergence
of infectious properties of DNA (TCD ~/m1=3.5). The greatest ~nf ectiousness
is noted in DNA from cells jointly in~ected with TEV + OV40 + RSV (TCD ~/m1=6.25).
It is noted that DNA infectiousness is completely removed by the eff ec~ ~f DNA-ase,
but not RNA-ase. Data on titration of full-value infectious virus TE which
develops after transfixing of sensitive cells with DNA preparations made of
chronically infected cells not only confirm the incorporation of the full DNA-
copy of the TEV genome into cellular DNA, but , mainly, indicate the complete
' expression of the DNA-provirus.
Table 1. Transfection of SF'EV Cells with DNA Preparations Made of Cells Infected
with TEV Jointly with Oncogenic Viruses
Viruses Original quantity Final dilution Titer of transfixed virus, ,
of DNA, ug per 105 of DNA TCD50/ml
cells without antiserum ~oith antiserum
TGV 22.0 ~ ~ ~
TEV+RSV ?9.0 1:8~ 3'S 0
TEV+OV4n 24.4 1:120 6�~ ~
TEV+RSV~{)V40 37.0 1;160 6�25 ~
T'~e presencc~ of onco-RNA-viruses in the TEV-infecCed cells piays a significant
role in the formation of the DNA copy o~ the TEV genome, the D~3A from cells which
arejointly infected with TEV and RSV during transfection induce the appearance of
an infection virus. The absence of infectiousness of the ce~lular DNA infected
only with TEV forces us to assume a stimulating effect of OV40 on the activity
oF the onco-RNA-virus usually present in the cells. Stimulation of activity of
oncc-RNA-viruses on ~he part of OV40 is quite probable. The OV40 stimulates the
output of adenovirus with the combined infection of cells [12],and OV40 in its
action in this respect is analogous to the effect of iododesoxyuridine [13]. In
terms of the studied question on incorporation of the TEV genome, all.'that has been
listed in relation to OV40 is additionally strengthened by the known data on
stimulation of cellular DNA synthesis during inf ection with OV40 [14].
Based on the available data, one can hypothasize a possible path for formation
of a DNA-copy of the genome of an RNA-containing nononcogenic virus and its
incorporation into ctiromosomal DNA of cells jointly infected with TEV and OV40.
The initial stage, synthesis of the DNA-copy can be associated with manifestation
of the activity of onco-Rl~A-viruses usually present in the cells. Their inverse
transcriptase does not have specificity in relation to the matrix. The subsequent
stage, incorporation of the DNA-copy can be associated with the possible conjugation
nf the processes of preproduction of the TEV genome and the processes of repro-
duction of viruses which reproduce in parallel and are capable of incorporation.
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BIBLIOGRAI'HY
1. 'Lh~3anov, V. M. NATURE, Vol 256, 1975, p 471.
2. Zhc3ai~ov, V. M. MOI~. CEI.L. BIOCHF.~1. , Vol 15, 1977, p 45.
3. Andzhaparidze, 0. G. and Drynov, I. D. DAN, ~ol 239, 1978, 970.
4. Soutilern, E. M. J. MOL. BIOL., Vol 98, 1975, p 503.
5. Botchan, N., Topp, W. and Sambrook, J. CELL, Vol 9, 1976, p 26q.
6. Temin, H. CELL, Vol 18, 1978, p 387.
7. Grc~en, F'. J. and Heyneker, H. L. T~UCL. ACID RES. Vo] S, 1978, pp 2373.
_ 8. fiickle, 'T'. A. and Pirrotta, L. IBID, Vol 4, 1977, p 2561.
- 9. Denhardt, D. BIOCHEM. BIOPHYS. RES. COMMUN.,Vol 23, 1966, p 641.
10. Taylor, J. M. ar.d Il~~n~~nsee, P. BIOCHIM. ET BIOPHYS. ACTA, Vol 4, 42, 1976,
p 324.
11. Uryvayev, V., Parasyuk, N. A.; et al. in "Virusy raka i leykoza" [Cancer and
Leukosis Viruses], Moscow, 19�~0, p 41.
12. Jerkofsky, M. and Rapp, F. VIROLOGY, Vol S1, 1973, p 466.
13. Jerkofsky, M. and Rapp, F. IBI~ Vol 15, 1975, p'253.
~I
, l4. Eckhart, W. COMPREHENS. VIROL., Vol 9, 1978, p 1.
~ COPYRIGHT: Izdatel'stvo "Nauka", "Doklady Akademii nauk SSSR", 1981
i
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MEDICINE
UDC 615.3~9:578.245
INTERF'EROhS IN MEDICAL THEORY AND PRACTICE
Moscow INTERFERONY V TEORII I PRAKTIKE MEDITSINY in Russian 1981
(signed to press 12 kt3r 81) pp 2-4, 398-400
[Summary, introduction and table of contents from "Interferons in Medical Theory
and Practice" by Valentin Dmitriyevich Solov'yev and Tagir Abdullayevich
Bektemirov, USSR Academy of Medical Sciences, Izdatel'stvo "Meditsina", 2021
copies. 400 pages]
[TextJ Eleven years have elapsed since the publication of the first edition of
this book. During this interval of time a very great amount of data~ both
- experimental and clinical, has been amassed; these data dictated the need for t~e
present edition. The monograph has been almost completely rewritten. It gives a
description of viral interference, considering in detail interference not caused
by interferons.
The book examines the properties of interferon, the interferonogenic activity of
different families and groups of viruses and their sensitivity to interferons.
- C~nsiderable space is given to data on interferon production by leucocytes and
macrophages, as well as to methods of obtaining and purifying ~eucocyte,
lyenphoblastoid and fibroblast interferon. The role of interferons in the
pathogenesis of viral infections and their significance as an index ~f reactivity
under normal conditions and in pathology are presented in detail. The book
correlates materials on experimental and clinical application of Exogenic
interfer4n and stimulators of endogenic interferan of viral and nonviral nature.
Great attention is given to nonantigenic inducers of interferon. Data are
presented on the non-antiviral action of interferons and their inducers, including
the effect on antibody production and cellular i~muni~y. A special chapter is
devoted to oncogenic viruses and interferon. In it the mechanisms of the
f~~mation and action of interferons are described. In conclusion, an evaluation
is given of the theoretical s~gnificance of interferons and the prospects of their
use in practice.
The monograph is intended for virologists~ immunologists, biologists and
physicians.
The book contains 33 tables and 37 figures; the bibliography, 585 titles.
For summary see page 396.
Reviewer: Yu. Z. Gendon, doctor of inedical sciences, professor~ head of the
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Department of General Virology of the NII [Scientific Research Institute] of Viral
Prep~rations of the USSR MZ [Ministry of Health].
INTRODUCTION
The first edition of our book~ "Interferon in Medical Theory and Practice" was
published in 1970. In that edition the existing studies in the literature were
correlated; the results of our own research were presented, and the outlook for
further research was given. The book recPived a high appraisal from readers and
was awarded the D. I. Ivanovskiy Prize. In 1973 the book was translated into
- Czech and published in Prague by the Avicenum PubTishing House. In the same year
a somewhat sugplemented edition was translated into English and published by
Plenum Press in New Yark.
Since then many experimental findings and results of clinical observations have
been amassed. This fact accounts for the need for the present edition. Into it
went completely revised materials and the results of studies by Soviet and foreign
, scientists up to 1980. We are unable to give a complete bibliograph;~ of studies
on interferons since it would exceed the scope of the present monograph.
Accordingly the bibliography gives only studies published since 1970.
The range of research has also expanded incommensurately with the past. Evidence
of the existence of several types of interf'erons and their action has been
_ obtained not only in viral infections but also in normal cells and cells
undergoing malignant changes. It has also been demonstrated that interferon '
- obtained from human blood leucocytes is capab].e of being highly concentrated and
purified of inactive proteins. Such a concentrated preparation, containing up to
1 million or more activity units per 1 ml of volume, may be, and is being,
employed in diverse pathological conditions, not solely those of viral etiology.
- The newest trend is the use of interferons in clinical oncoiogy. It is more
advisable to use concentrated preparations for therapeutic use in this pathology.
The difficulties associated with obtaining human donor blood~ which serves as a
source for obtaining leucocyte interferon, necessitated a search for other
producers. Foll~wing up the species specificity of interferon, we unexpectedly
discovered that swine leucocytes are capable of producing interferon which acts on
~ human diploid cells in the same way as human interferon.
In the USA, Englanr; and other countries research is being conducted in the
direct�ion of obtaining interferon-forming cells outside the organism. Here two

~7 methods have been established: one consists of using cultured lymphoblastoid
cells; the other, of using diploid producer cells. Both these methods are also
being used in our country's laboratories. Up to now~ however, they have been at
the stage of experimental study and have not entered medical practice.
Ten years ago we formulated the so-called interferon reaction of leucocytes. It
consists of the fact that the general immunoreactivity of an organism can be
evaZuated according to the activaness of interferon product~:ion by lymphoid tissue.
_ After namerous tests we recommended this reaction for evalu3ting the condition of
the reticuloendothelial systen~ under normal conditions and in pathology.
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Interferons are now viewed as an indispensible component of iromunological
reactivity which to a significant extent ensures the stability of the organism's
internal environment. The significance of interferons as factors of antivirai
imrnunity is being increasingly evident. It is most likely that interferons are
the regulators of many immunity reactions. They are known to influence cell
- division, the activity of normai killer cells, DNA synthesis in lymphocytes, the
delayed type of hypersensitivity reaction and transplantaton immunity.
Recent years have been marked by numerous studies in immunobiological and
c~togenetic directions, which have revealed new facts characterizing the
mechanisms of the formation and action of interferons.
- In conclusion~ a word must be said about the stages of development of the study of
~ interferons. For many years the scientists of the USA and England did not
consider studies on a;:ogenic interferon of current interest and sought substances
and means inducing the formation of endogenic interferon, working on different
synthetic preparations of the type of polynucleotides. But beginning in 1976,
foreign scientists' interest in exogenic interferon--i.e., in the direction which
Soviet investigators had been developing since 1964--sharply increased.
At the recent Fourth International Virology Congress the reports of foreign
scientists were devoted to presentation of experimental and clinical experiments
on successful use of exogenic interferon obtained fram leucocytes of donor blood.
The monograph offered to the reader retains the previous title in only slightly
changed form. The title successfully reflects the contents~ which have been
almost completely rewritten.
- The authors sought not only to present the current data as accurately as possible
but also~ principally to reflect the resulcs of studies by Soviet scientists
engaged in the study of interferons. Realizing the complexity of the task in
hand, we will gratefully accept critical remarks and suggestions.
CONTENTS ~~e
Foreword. .......................................................................3
Chapter 1. Viral Interference ..................................................5
General considerati.ons ......................................................5
Interference between infectious and inactivated viruses .....................6
Canditions fo~ producing inteference .......................................12
Practical significance of interference .....................................19
Methods of inducing interference ...........................................22
The mechanism of interference ............................................24
Autointerference by viruses ................................................33
Chapter 2. The Concept of Interferons, Their Formation and Antiviral Action in
Vitro ......................................................~...................42
Basic properties of interferons...��������~�����������~�~��~~~������������~~'2
Physicochemical properties .................................................43
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~~ciR ~F~~~t.�. ~~tiN' (1N11'
Species specificity .........................................................53
- Antigenic properties .......................................................56
Chapter 3. The Interferogenic Activity of Viruses .......,,....................~.60
Smal~pox viruse~ ............................................................60
Herpes viruses .............................................................61
Adenoviruses.............e ..................................................63
Myxoviruses .................................................................64
Rhabdoviruses ...............................................................66
Arboviruses .................................................................67
- Enteroviruses ..............................................................69
Chapter 4. Factors Influencing the Intensity and Dynamics of Interferon
Production ......................................................................70
Cell type ...................................................................70
Viral properties ............................................................73
Other ~'actors ...............................................................75
Dynamics of interferon formation in cell cultures ...........................81
Chapter 5. The Action of Interferons in Cell Cultures .........................�85
The sensitivity of viruses and cells to interferons .........................85
Methods of inducing and titering interferons ................................90
Blocking of the formation and actxon of interferons by viruses .............�95
Chapter 6. Interferon Formation by Human Leucocytes in Vitro ..................100
Production of virus induced interferon by animal leucocytes ................100
The formation of interferon by human leucocytes..����.�����~~���������~��~�104
The role of different elements of white blood in interferon production����~~�1~
Nonviral induction of interferon in leucocytes Iromune interferon..�.�~����117
Chapter 7. Lymphoblastoid and Fibroblast Interferon ...........................123
Lymphoblastoid interferon and its purification .............................123
Fibroblast interferon and its purification .................................126
Chapter 8. Interferon Formation and Vira~ Infections...����.~�~~~���~����~���~13~
Interferon formation in experimental viral infections�...���������~������~~13~
Interferon formaticn in viral and nonviral diseases of man��������~~�~~�~~�134
Chapter 9. The Role of Interferon in the Pathogenesis of Viral Infection�.��..142
The action of interferon in infection atria and the pathways of tr.e
spread of viruses in the organism....�������~~����~~��~���~�����~���~����142
Virulence of viruses and interferon..��~������~��������~�~����~~~��������'�1~'7
Age and individual properties of interferon formation in animals...........152
The influence of stress and other factors on the formation of
interferon in viva .......................................................156
The correlation between interferon formation and the susceptibility
- of animals to certain viral infections.~����������~~�~~~�~~��������~�����159
Chapter 10. The Interferon Reaction of Leucocytes--An Index of the Reactivity
- of the Organism Under Normal Conditions and in Patholagy.���~������������~~���.166
General aspects .........................................................156
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FnR (1F'F1('1:11. l~ti~' (1N1.1'
...............1~~
Age ar~d nutritional properties
..............................17U
The factor of individuality
The influence of cooling, irradiation and immunodepressive preparations�172
....i73
The formation of leucocyte interferon in viral diseases...�~�~�~��~~~��
The formation of leucocyte interferon in patients with endocrine
disorders ................................................................175
The formation of leucocyte interferon in patients receiving therapy��~�����i76
Chapter 11. The Effect of Exogenic Interferon and Stimulators of ~ndogenic
Interferan on Viral Infectior~s ...................................������~��~����1~9
The effect of exogenic interferon on experimental viral infections........179
Clinical testing of exogenic interferon.........�.�����.��~��~~��~~��~�~'�187
Pharmacokinetics of interferons ...................................���~�~~�199
Stimulation of serum interferons by viral inducers ........................204
Serum interferon producer cells 208
T h e e f f e c t o f v i r a l i nducers on ex perimental viral infections.���~������~�214
Chapter 12. Stimulation of Productian of Endogenic Interferon by Double Strand
RNA and Its Effect on Viral Infection .........................................�218
General considerations.. " " " " " " " " ~~~~~~~~~~~218
Interferon induction in cell cultures and in animal organisms�������~�����219
The effect on experimental viral infections....��.�.��~~�~~�~�����'�" 221
Correlation between interferonogec;ic activity and the structure of
polyribonucleotides ..........................................o..........225
- Interaction between polyribonucleotides and cells .................�����.��230
Correlation betwe~n interferogenic zctivity and induction of antiviral
resistance............~ .................................................234
Toxicity of polyribonucleotides for humans and animals~�����~~���~�~'��~��235
Modification of polyribonucleotides active in primate~j in vivo........�..�240
Modificatian of polyribonUCleotides of different molecular masses.........250
Chapter 13. Stimulation of Endogenic Interferon By Other Nonviral Inducers and
................z55
Their Effect on Experimental Viral Infections..�~�������������'
General data ..............................................................258
Synthetic inducers
.......261
Tiloron and its anologues " " " '
- Stains~ pyrimidi.nes and other inducers ...........................~..��~���263
................266
Bacteria, yeast and products isolated from them....������~
~ Chapter 14. The Influence of Different Factors on the Action of Nonviral Inducers
of Interferon. Superinduction of Interferon����~��~���������~�������~��~������276
Effect on double strand RNA .............................................�.276
Effect on other nonviral inducers .........................................279
Superinduction of interferon " " " " " " '280
Chapter 15. Refractoriness in Interferonogenesis 286
- General considerations ....................................................286
Correlation between refractoriness and antiviral resistance ����~��~~��~�~�2 89
Mechanisms of refractoriness " " " " " '�'�'�����~~������290
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Chapter 16 Clinical Testing of Viral and Nonviral Stimulators of the
Formation of Endogenic Interferon .........................................295
Viral inducers ............................................................295
Double strand polyribonucleotides ........................................~302
Tiloron and other low-molecular inducers ..................................305
Chapter 17. Non-Antiviral Action of Interferon and its Inducers ...............307
Action against nonviral infectious agents and their decomposition
products ................................................................307
Effect on antibody formation~ cellular immunity and cell growth���~~������310
Chapter 18. Oncogenic Viruses and Interferon�..����~�����������~~�����~������~321
Induction of interferon by oncogenic viruses ..............................321
- Action of interferon and its inducers on the development and course of
neoplasms and leucoses ..................................................324
Mechanism of the effect of interferon on oncogenesis����~��~���~���������~328
Interferon therapy in oncological clinical practice .......................341
Chapter 19. Mechanisms of Formation and Action of Interferon ..................345
Structural genes of interferon format.ion ..................................34-5
Priming mechanisms of interferon formation ................................347
Matrix RNA for interferons ................................................352
Regulation of interferon synthesis............ ...........................�354
Interaction of interfer~n with cells .................................~...�356
The mechanism of the suppression of virus reproduction by interferon.....�358
;
Conclusions ....................................................................36~
' Bibliography ...................................................................367
COPYRIGHT: Izdatel'stvo "Heditsina" Moskva 1981
9380 .
' CSO: 1840/297
i
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PHYSIOLOGY
HOW WE SEE WHAT WE SEE
rioscow KAK MY VIDIM T0, CHTO VIDLK in Russian 1979 (signed to press 29 Mar 79)
PP 2-5, 207
[Annotation, foreword by Academician 0. G. Gazenko and table of contents from book
"How 41e See What We See", by Vyacheslav Yevgen'yevich Demidov, "Science and
Progress" series, Izdatel'stvo "Znaniye", 100,000 copies, 208 pages]
[Text] The problem of perception of the outside world with the organ of sight is
one of the interesting natural scientific problems. We find the names of many out-
standing people of different times and nations am~ng those who made their contribu- ~
tion to it. At present, major scientific teams, armed with the mos~ modern ,
. investigative methods, are working on this problem. Among them is the Laboratory
of Physiology of Sight, Institute of Physiology imeni I. P. Pavlov, USSR Academy
_ of Sciences, in Koltushi near Leningrad, which used the principles of holography
to explain the "mechanism" of processing visual information of our brain. In
discussing the work at this laboratory, the author makes broad use of material
obtained by other Soviet and foreign researchers. This scientific and literary
book is intended for a broad circle o.f readers.
Foreword
The author of this book is not only a reporter, but an engineer. The book itself
is the result~f 5 years af fruitful collaboration with scientists from the Labora-
tory of Physiology of Sight, Institute of Physiology imeni I. P. Pavlov, USSR
Academy of Sciences. For these entire 5 years, the author attentively watched
the progress of its staff and constantly published articles about their work.
- The problems discussed in this book are presented by V. Demidov on the basis of the
- holographic hypothesis of brain function, which is being actively developed in t�ecent
times by scientists of the entire world (in all fairness, let us mention that Lhis
conception is not shared by all researchers). Among these studies, the work of
Soviet scientists occupies a prominent place, particularly those working at the
Ir.stitute of Physiology imeni I. P. Pavlov. And, since holography is the child
of engineers, it becomes understandable why one of them was able to make sense,
easily and without constraint, out of the sea of diverse information that appears
scattered outwardly but actually demonstrates a profound unity of material
essences of the world revealed to us, using a holographic compass.
The solitary approach the author used to the most varied problems brought him
deserved success. Demidov finds good explanations tor "mysterious" phenomena
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and advances plausible hypotheses by examining from the same positions the ques-
tion of identifying visual images and causes of the "whims" of fashion, perception
of color and structure of diff erent systems of the optical tract, visual illusions
_ and formation of the internal model of the world. Such discoveries include, For
example, the hypothesis of reasons for changes in fashions, the explanation of
the "secret" of Penrose's triangle and "impossible pictures"; the author uses a
unique and curious approach to the problem of essence of abstractions and concept
of beauty. There is convincing disclosure of a seemingly paradoxical thesis that
' visual illusions are a reflection of the automatic accuracy of function of the
visual system, a reflection of accuracy of the model of the world formed as a
result of man's prior experience.
The lucidity, understandability and, at the same time, scientific strictness of
presentation of material are definite qualities of the book that you have in your
hands. The bibliography used by the author is referable to the works of the
greatest scientists dealing with problems of vision, both Soviet and foreign.
Demidov knows many af his heroes personally; he has visited scientific laboratories,
witnessed experiments and, for this reason, he has conveyed the atmosphere of
scientific search in an entertaining and convincing manner.
A good, graphic Zanguage is used to present extremely complex problems of neurophysio-
logy and psychology, cybernetics and medicine; the author never falls into vulgari-
zation. He ouerates freely with the concepts of many sciences, cites good and
vivid examples, so that it will be unquestionably interesting to the reader to
- follow the logic of development of the subject. The problem of holography is a
mathematical problem in many respects, and it is all the more pleasing that he
succeeded in explaining it without formulas, on a level that is quite understandable
to the broadest reader. It is very important that Demidov refers to the most recent
works, the results of which were literally just published, in 1977, and one feels
the pulse of modern times in the book, it is timely and fresh.
The history of learning about the mechanisms of function of the visual system is
the history of the struggle of science against idealism. The results of current
- research confirm, again and again, the materialistic thesis that it is possible to
gain knowledge about nature in all of its manifestations, including such extremely
complex ones as sight and thinking. Science puts electrochemical processes in
neuronal networks of the brain, which are amazing in their refinement, in the place
~ of the "soul." With each year, experimental techniques grow more and more sophisti-
- cated, and we are delving deeper and deeper into the essence of things. Man is
learning more and more about himself and penetrates into mysteries that make
those of the oceans and space paler. At the same time, the language of science is
growing increasingly complex; sc~ence is breaking down into more and more narrow
disciplines, so that scientists working in one laboratory have difficulty in
discerning problems of their neigtibors in the next laboratory. The volume of in-
formation is growing like a snowball, and for this reason the role of scientific
popularization, particularly when it generalizes the achievements in related and
allied disciplines, is ever growing in our times. Not infrequently, a scientist
will glean some useful information from such works. Demidov's book is one of them--
a successful attempt at summarizing results obtained by specialists working in the
most varied areas of knowledge. He not only was able to summarize, but relate
these results to vital problems that are close to literally everyone, blending the
seriousness of the scientist's approach to the vitality of a writer's style.
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Contents Page
Cr~apter 1. Range of the Preconscious 9
Ctiapter 2. The Prediction of Galen 23
Cliapter 3. The Traps of Image Details 33
Cl.iapter 4. The Tree of Knowledge of Good and Evil 49
_ C~napter 5. When We Cannot Say "a" We Say "b" 57
Chapter 6. C,ycles That Have a Reason 6~
Chapter 7. The World is Built of Parts [Details]
Chapter 8. All Roads Lead to Rome 93 I
(:hapter 9. The Flat Three-Dimensional World 109
Chapter 10. The Immediate Consequences of What Was Overturned 125 ~
~~hapter 11. The Palette 133
Chapter 12. A Sieve for Images 149
Chapter 13. Deceptions Caused by the Desire for Truth 165
Chapter 14, A New Key for Old Mysteries? 181
Bibliography 206
COPYRIGHT: Izdatel'stvo "Znaniye", 1979
10,657 j
CSO: 1840/999 (
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J
UDC: (577.4+577.1)001.57
~
METHODS OF MATIiEMATICAL BIOLOGY, BOOK 2: M~THODS FOR SYNTHESIS OF ALGEBRAIC AND
PROBABILISTIC MODELS OF BIOLOGICAL SYS'PEMS
Kiev METODY MATEMATICHESKOY BIOLOGII, KNIGA 2: METODY SINTEZA ALGEBRAICHESKIKH I
VEROYATNOSTNYKH MODELEY BIOLOGICHESKIKH SISTEM in Russian 19~31 (signed to press
9 Jan 81) pp 4-8, 10
[Annotation, introduction to Part I and taL~le of contents from book "Methods of
Mathematical Biology. Book 2: Methods for Synthesis of Algebraic and Probabilistic
Models of Biological Systems", by Viktor Mikhaylovich Glushkov, Yuriy Gur'yevich
Antomonov, Vasil Kabulovich Kabulov, Khusan Kadyrovich Kadyrov, Svetlana zvanovna
, Kiforenko, Andrey Mikhaylovich Klochkov, Alina Borisovna Kotova, Todor Angelov
' Popov and Anatoliy Aleksandrovich Stogniy, edited by Academician V. M. Glushkov
' (chief editor), approved by the IIkrainian Ministry of Higher and Secondary
I~ Specialized Education as a textbook f~r stud2nts in biological specialties at
~ VUZ's, Izdatel'skoye ob"yedine.r.iye "Vyshcha shkola". 2000 co~ies, 312 pagesJ
; [TextJ There is discussion of inethods of choice of experimental material in the
i
, structure of an algebraic model and determination of the coefficients of the mode
~ for describing various static functional modes of biological systems. The use of
functions and regre~sioi: equations is demonstrated for synthesis of mathematical
models of biological sy:;tems.
Methods are described for primary proc~ssing of experimental data, for equalizing
statistical series, determining the reliability of differen~es, determination of
' laws of distribution in correlation and spectral analysis of random biological
processes and use of Markovian random processes.
This book is a textbook for courses on "Biological Cybernetics," "Bionics,"
"Medical Cybernetics," "Mathematical Modeling of Biological Systems" and "Biophysics"
for srudents at universities, medical and pedagogic institutes and institutes of
physical culture; it may be useful to scientists concerned with ;:roblems of mathe-
matical biology. Figures 76, tables 37, references 69.
Introduction [to Part I]
Methods of modern algebra are used extensively in various branches of science--
mathematics, mechanics, physics, cybernetics and mathematical biology. They include
linear algehra, logical algebr~, matrix albegra, set algebra and algorithm algebra.
In constructing biosystem theory and solving various problems, modern biology and
medicine make extensive use of inethods of quantitative analysis of the link between
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s
output coordinates and input factors. Use is made of the conception of a link in the
- form of tables and graphs, from which one turns to mathematical. descriptions. The
simplest mathematical description of the link between output coordinates of biologi-
_ cal systems and input factors can be obtaj.ned by means of various algebraic functions.
In this case, the mathematical description is a functional model of the biosystem.
The diversity of interactions between biological systems and the environment makes it
necessary to use both elementary functions and superpositions thereof varying in
complexity in order to describe them. There is discussion here of algebra methods
that permit solving problems of synthesis of various functional models of biological
systems; however, listing all of the methods of functional relations and equations
used in modern mathematics is not an end in itselF. Attention is devoted chiefly
' to the methods and specific types of functions that have now become quite popular
in mathematical biology.
This book, which deals with applied algebra as a science but theory of biology,
is concerned with methods of searching for types of functions according to experi-
mental data, investigation of applicability of algebraic models to solving biological
and medical problems.
Contents Page
Part I. Methods for Synthesis of Algebraic Models of Biological Systems
- Introduction 10
Chapter 1. Functions "11
1.1. Elementary functions 11
Definition of function 11
Linear function 14
In~ersely proportional function 14
Exponential function 15
Step �unction 17
Logarithmic function 18
Trigonometric functions 19
Circular (inverse trigonometric) functions 20
1.2. Determination of parameters of functions from experimental data 21
Plotting graph of functions through typical points 21
Least squares method 24
Integral criterion 26
1.3. Elementary algebraic models of biological systems 27
Model of threshold properties of nervous tissue 2~
Empirical functions 33
Chapter 2. Polynomial (Regression) Models 34
2.1. Linear regressions 34
Types of regressions 34
Linear single-factor regression 3~
Linear multiple-factor regression 41
2.2. Nonlinear regressions 46
Nonlinear single-factor regression 46
Nonlinear multiple-factor regression 4~
Regression models based on the method of group consideration
of arguments (MGUA) of A. G. Ivakhnenko 48
Method for insertion of time or parameter into regression
equation 53
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2.3. Regression models of ~iologieal processes 58
- Modeling an epidemic process. Base data 58
Model of an epidemic process 68
Method of retrieving adequate model of an epidemic according
to mathematical criteria 71
Insertion of time in regression model of an epidemic 73
Example of synthesis of regression model of the ecological system
of the Rybinskiy Reservoir by the MGUA method 78
Chapter 3. Methods for S,ynthesis of Algebraic Models and Biological
Identif ication 84
3.1. Schematization of modeling 84
General approach 84
Example 85
3.2. Choice and comparison or algebraic models 89
Biological identification--criterion for choice of structure of
algebraic model 89
Example of choice of m~,del structure ~ 91
Chapter 4. Methods for Solving Equations 96
4.1. Matrix algebra 96
Main definitions 96
Properties of determinants 99
Operations with matrices 100
4.2. Solving systems of linear equations 106
Systems of linear equations 106
Kramer method of solving linear equations 108
Gauss method 109
General case of solving systems of linear equations 110
Systems of homogeneous linear equations 112
4.3. Algebraic equations 114
Lowest degree algebraic equations 114
Determination of real roots of algebraic equations 117
Bibliography
Part II. Methods for Synthesis of Probabilistic Models of Biological Systems
Introduction 124
Chapter 1. Methods of Primary Processing of Results of Biological
Experiments 125
1.1. Specificity of biological experiments 125
Correlation between factors 125
Investigation of dynamics 128
1.2. Distributior~ series of random quantity, its parameters and
estimation 131
Main definitions 131
Parameters of series 133
Set of sequences of random quantities and their characteristics
according to experimental results 137
Analysis of time of animal death 137
Analysis of number of animal deaths 142
1.3. Choice and calculation of parameters of variable 3istribution
series 144
Risk of death due to tumor 145
General risk of death 146
Dynamics of relative number of animal deaths due to cancer 153
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~,58
1.4. Systems of random quantities and their c?iaracteristics ~58
Dependence of random quantities
Numerical characteristics of a system of two random quantities 159
Numerical characteristics of a system of several randon quantities 163
Evaluation of numerical characteristics 163
Coefficient or multiple correlation
Coefficient of correlation as a criterion for choi~e of linearly 164
independen'~ param~ters
- 1.5. Estimation of reliability of differences between mean values of 166
system~ of random quantities 167
Criterion of Student 168
Simplified criterion 1~2
- 1.6. Formula of Bayes 173
Bayes formula and its use in differential diagnostics 176
Chapter 2. Laws of ~Jistribution of Random Quantities 176
2.1. Function and density of distribution 1~8
Statistical function of distribution ~79
- Histogram 181
2.2. Equalization of statistical series 181
Choice of theoretical distribution curve 188
Criterion of agreemen* 191
2.3. ThPOretical distributions of continuous random q~antities 191
Normal law of distribution 1~~
Distributions related to normal ones 198
Law of uniform density 201
2.~. Theoretical distributions of discrete rar.3om quantities 201
Binomial distribution ' 2p2
Polynomial distribution 203
Hypergeometric distribution 204
Geometric distribution 204
Pascal distribution 204
Poisson distribution 205
Chapte.r 3. Correlation and Spectral Analysis of Random Processes 205
- 3.1. Main properti.es of random processes 206
Definition 206
Classification of random processes 209
Evaluation of stability 213
Ev aluation ofEEG stability 215
3.2. Characteristics of random processes 216
Mathematical expectation and dispersion of a random function 219
Laws of distribution 22p
Evaluation of law of distribution 222
The XZ square criterion 22g
The ~ criterion of Kolmogorov-Smirnov
Use of characteristics of random process~s in electroencephalo- 224
graphy 234
3.3. One-dimensional correlation-spectral analysis 234
Coefficient oi correlation 235
Autocorrelation function 236
Evaluation of autocorrelation function 238
Use of autocorrelation function for analysis of EEG 241
Spectral density 243
Evaluation of spectral density 248
Use of spectral analysis in electroencephalography
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- 3.4. Two-dimens ional correlation-spectral analysis 26Q
_ Reciprocal correlation function (VKF) 261
Use of reciprocal correlation function in encephalography 263
Reciprocal spectrum 265
Amplitude spectrum 266
Reciprocal spectrum of phases 266
Spectrum of coherence 26~
Evaluation of r~ciprocal spectra 268
Dispersion and confidence intervals of reciprocal spectrum
evaluations 270
Planning the required accuracy of evaluations 2~2
Example of the reciprocal spectrum method in
electroencephalography 2~3
Chapter 4. Methods of Describing the Dynamics of Markovian Random
Processes 289
4.1. Main theses of Queueing Theory 28~
Subject of theory 289
Flow characteristics 290
Markov random process 292
4.2. Distinctions of information flow in medical institutions 295
Distinctions of health care organization 295
Biomedical information 2y~
4.3. Analysis of inedical care systems 298
System of service [care] with loss of patients 298
System of service with expectation 301
System of inedical service with limited waiting line 305
Bibliography 308
COPYRIGHT: Izdatel'skoye ob"yedineniye "Vyshcha shkola", 1981
10,657
CSO: ].840I999
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UDC: (577.4-~577.1~001.5(07)
METHODS OF MATHEMATICAL BIOLOGY~ BOOK 3: METfiODS OF SYNTHESIZING DYNAMZC
MODELS OF BIOLOGICAL SYSTIIy1S (WZ TEXTBOOK) ~
Kiev METODY MATEMATICHESKOY BIOLOGII, KNIGA 3: METODY SINTEZA DYNAMICHESRiKH
MODELEY BIOLOGICHESKIKH SISTEM (UCHEB. POSOBIYE DLYA WZOV) in Russian 1981
_ pp 4-10
[Annotation, introduction and table of contents from book "Methods of Mathematical
Biology. Book 3: Methods of Synthesizing Dynamic Models of Eiological
Systems (Textbook for WZ's)" by A. M. Molchanov, Yu. G. Antomonov, A. B. Kotova
and M. A. Khanin, Izdatel'skoye ob"yedineniye "Vyshcha shkola", 328 pages]
[Text] Methods are described of theory of differential equation~, which permit
synthesis of mathematical models of the dynamics of biological systems. There is
discussion of linear equations with constant and variable coefficients, typical
nonlinear equations, first and second order partial [differential?] equations. Much
- attention is given to proper investigation of equations for phase and structure
proriles. Methods are described for writing differential equations for biological
- systems with gradual complication of physiological conditions, which permit
expression in the model of different properties of biological systems.
This book is a text for course of "Biological Cybernetics," "Bionics," "Medical
~y~ernetics," "Mathematical Modeling of Biological Systems" and "Biophysics" for
students at universities, medical and pedagogic institutes and institutes of physical
culture; it may be useful to scientists concerned with problems of mathematical
biology. Fi~ures 136, tables 2; references 46.
Introduction
In matt?ematical biology, wide use is made of inethods of differeatial equation theory
for the synthesis of mathematical models of dynamic modes of biological systems.
Thus, to describe the function of biological systems, use is made of theory of
linear differential equations; first or second order differential equations are used
to describe the function of biological systems with reference to one parameter; sys-
t~ms of linear differential equations are used to synthesize mathematical models
describing the dynamics of many para~neters; the entire spectrum of formal mathema-
tical methods for solving differential equations, including methods of computation
["computing mathematics"?], is used for concrete modeling.
- It is known that the dynamic behavior of biological systems is quite complex and
so multifaceted that mathematical models based on linear differential equations can
only be considered modeling in the first approxi~mation.
22
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In this textbook, the biological exampl.es are incl,uded virtually eyerywhere in the
descriptions of inethods of writing and solving the appropriate di~ferential equa,
~ tions; they are often an adequate description of the dynamics of variables and
- conform with the objectives and tasks of modeling.
The distinction of the presentation also lies in the fact that the sections of
differential equation tY?eory that are the easiest to comprehend and most used to
solve biological and medical problems are very concise. The mo.re difficult sec-
tions dealing with nonlinear equations, equations with v2riable coefficients are
more comprehensive.
. Unity of structure and function is inherent in the function of '~iological systems.
Whenever this must be reflected in mathematical models, it is often (and will be
more and more often in the futur.e) necessary to use methods of de^cribing the form
of different structures (organ, vessels) and distribute among strurtures the
_ variables (concentration, potential, pressure) that determine functional states.
For this reason, we devoted much attention to description of tneory of partial
equations with the use of examples.
It is known that partial differential equations have found wide applications i.n
physics as an effective tool that permits d escribing differ.ent physical phenomena
, which include, in particular, heat conduction diffusion, electric and magnetic
fields, propagation of diverse waves and quantum fields. The intensive development �
of mathematical biology in the last decades ~as led to expansion of the area of
application of partial differential equations of the second order. We can arbi-
- trarily single out two main directions. The first is related to development of
mathematical theory of biological phenomena, which are based on physical processes
(diffusion, head conduction, etc.), described by second order partial equations.
~ Use of partial second order equations to describe the propagation of pulse waves
~ in bloo~ vessels, diffusion of oxygen and other substances in ~ells, in t~ie micro-
circulatory system and other phenomena are illustrations of this direction.
~i
-j The second direction of using second order partial equations is related to quantita-
~ tive description of biological processes which are in essence biological, even
though they present some analogy to certain physical processes. This direction can
~ be illustrated with examples of theory of inf lammatory processes.
Discussion of biological applications of second order partial equations is
limited to a small section, which briefly describes the classification of second
order partial equations and the most frequently encountered boundary value problems.
The methods of solving partial equations are described in the sections dealing with
specific biological problems. Of course, the material presented in the foregoing
does not cover all of the branches of mathematical biology making use oi second
" order partial equati_ons developed to date. The examples are given here in such
a manner as to illustrate as broadly as possible the different methods of solving
problems in the field of biol~gy to which these problems refer.
In the presentation of specific sections, considerable attention is devoted to
biological validation of the mathematical system used.
_ 23
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Conten.ts ~a~~
9
Introduction 11
Chapte~ 1. Linear Equations 11
1.1. Concept of derivative and differential 12
. Definition of derivative function 13
Def inition of differential 14
Second order derivative 14
Partial derivatives and par*ial differentials 1~
Concept of differential equation
1.2. Linear f irst order diff erential equations with canstant coefficients 18
General appearance ~A
Equations with P(x) = 0 22
Equations with P(x) = const
- 1.3. Linear second order differential equations with constant coefficients 26
General appearance 2~
Homogeneous equation , 30
Partially homogeneous equation 31
Hete:cgeneous second order equation 39
1.4. Higher order equations. Differential equation of nth order 40
System of f irst order linear equations 4L
Chapter 2. Linear Equations With Variable Coefficients 42
2.1. General concepts 42
Normal systems. Coordinate and vector methods of writing 43
Derivative along solution 45
Self-contained systems. Concept of first integral 46
Cauchy's problem. Equivalent integral equation
2.2. Theorem of existence and uniqueness of solution to Cauchy's problem. 49
Continuous dependence of solutions on base data 49
Picard's method 51
Theorem of existence and uniqueness 53
Nature of convergence 54
Euler's method of broken [lines?] 55
Concept of nonuniqueness
Continuous dependence of solution on base data 5g
Local continuity 61
A priori evaluation 63
Local theory of continuous dependence of solution on base ~lata 64
Local theorem of uniqueness 65
Theorem of uniqueness as a whole 6~
- 2.3. Neighborhood of regular solution 6~
Systems containing the parameter 68
Variation of parameter 69
Equations in variations ~1
Variation of base data
2.4. Linear systems with variable coefficients. Definitions. Theorem 72
of existence and uniqueness 73
Homogeneous systems. Elementary properties 74
Wronskian determinant 76
Properties of fundamental matrix 78
Liouville's formula 79
Definition of fundamental system of solutions 80
Structure of general solution of linear homogeneous system 81
Heterogeneou;; systems. Method of variation of arbitrary constants
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Chapter 3. Nonlinear Equations 83
3.1. Hamiltonian systems 83
� Hamiltonian form of system 84
First integral of Hacailtonian system 85
Investigation of geometry o~ Newtonian movement with set potential U(x) 86
Kinetics of system $8
Integration of Newtonian movement with U(x) = ax2 89
Planar Hamiltonian systPm. Necessary and sufficient condition of
Hamiltonian system 91
Periodic movement (general case) 93
General appearance of s~Gtem with specified first integral 96
3.2. Special integration procedures 99
Lowering order of system ~ 99
Equation in complete differentiaZs 100
Equation with separating variables i01
_ Verhulst-Pearl equation 103
Distinctions of straight parts of system (3.60) 105
Limit cycles 106
Homogeneous equations 108
Linear equations 109
3.3. Equations that are unsolved with reference to derivative. Formulation
~f the question 111
Addition of parameter 112
Integration of Lagrange equation 115
Clai.raut's equation 116
Geometric aspect of t.he problem 118
3.4. Resistance [stability] 118
Formulation of the question 118
Neighborhood of stationary solution (stationary points) 120
Definitions of stability 124
Technique for testing stability. Lyapunov's function 125
Lyapunov's function in linear system 128
Differential inequality. A priori evaluation 129
Nonlinear systems 131
Chetayev's function 133
Chetayev's function for linear system 135
Chetayev's function for nonlinear system 137
Algorithm for testing stability 138
Chapter 4. ~lements of Qualitative [or goodJ Differential Equation Theory 142
- 4.1. Phase prcfile of system 142
- Phase space 142
Phase profile of Hamiltonian planar system 144
Algebraic Hamiltonian 146
Phase profile of linear system. Saddle, point, focus 147
Case of real roots 149
Case of complex roots. Focus 151
Correlation between geometry and kinetics 152
Algorithm for constructing phase profile of l~near system 153
Phase profile of Volterra system 154
'Cold flames' lEzl
Simplified model of glycolysis 162
Stability of stationary mode lb4
Analysis of spe.cial points 17Q
Local characteristics of pt:ase profile 17b
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Critical value of parameter 176
Generation of limit jr~aximum?] cycle l~g
Destruction of limit cycle 179
~+.2. Structural profile 18~
Structural profile of model of immune system ~ 186
- Local phase profile 190
Phase profile as a whole 192
Structure profile. Immune profile of population 193
Single-barrier immunity (.'step') 195
Structure prvfile of single-step i~ununity 200
Neighborhood of triple point 2~2
Multi:barrier immunity. Mechanical analogy 205
Chapter 5. Partial Equations 209
5.1. Quasilinear partial equations 209
Linear equation. Cauchyts problem 210
_ Quasilinear equation. Riemann's method 218
Lagrange's coordinate system
Euler's equation 22~
'Flow [or current] tube. Cauchy's problem Z23
Dynamic systems 224
Curvilinear coordinates 22~
Partial linear equations. Theoretical analysis
Practical methods. Lowering order 23Q
General quasilinear equation. Method of immersion 233
5.2. Second order paxtial equations 236
Classification of equations 236
Canonical appearanc.e of equations 238
Boundary problems 239
- 5.3. Use of parabolic equations to solve problems of mathematical 243
biology 243
Mathematical model of early stage of inflammatory process 249
Model of oxygen transport
Mathematical model of sluggish inflammatory process in the presence 252
of necrotic area
Mathematical model of inflammatory process in heterogeneous area 258
with consideration of spread of necrotic area
5.4. Asymptotic solution of nonlinear and quasilinear equation of tlie 264
parabolic type
Mathematical model of cavernous tubercular inflammation with
consideration of the effect of partial oxygen tension as a 264
limiting factor
Incidence of dominant gene with selective advantage (with consideration 2~7
of migration)
5.5. Asymptotic and periodic solutions of system of parabolic nonlinear 281
equations
Dynamics of two populations (predator-prey) with consideration of
281
migration 286
Dynamics of two populations without migration of prey 289
5.6. Solution of elliptic type of equations
Stationary distribution of biologically active substances in the cell 292
Oxygen transport in the microcirculatory system
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Chapter 6. Methods of Composing Equations for Dynamics of Biological Systems 298
6.1. Modeling the dynamics of nervous processes on an elementary level 298
Capabilities of differential equations as a modeling tool 298
Description of form of action potential 299
' Model of dynamics of axonal excitation 301
Model of dynamics of conduction 304
Model of dynamics of synaptic processes 307
6.2. Modeling the dynamics of biocenoses 311
Model of elementary correlations ~1~-
Modeling the dynamics of populations and biocenoses with consideration
of structural and functional parameters of individuals and ecological
conditions 313
3ibliography 325
COPYRIGHT: Izdatel'skoye ob"yedineniye "Vyshcha shkola", 1981
10,657
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CHANGES IN ENDOCRINE GLAND FUNCTION DURING EXERCISE: ENDOCRINE MECHANISMS OF
REGULATION OF THE BODY'S ADAPTATION TO MUSCULAR ACTIVITY
- TartuIZMENENIYA FUNKTSIY ENDOKRINNYKH ZHELEZ PRI FIZICHESKIKH NAGRUZKAKH: '
ENDOKRINNYYE MEKHANIZMY REGULYATSII PRISPOSOBLENIYA ORGANIZMA K MYSHECHNOY
DEYATEL'NOSTI in Russian 1980 pp 166-167
[Table of contents from book "Changes in Endocrine Gland Functions With Physical
Loads: Endocrine Mechanisms of Regulation of the Organism's Adaptation to
- Muscular Activity", by N. N. Yakovlev et al., 167 pages] ~
~
Contents Page
Somatotrophic Pituitary Hormone and Adaptation to Muscular Activity
(N. N. Yakovlev) 3
Evaluation of Condition and Capabiliti.es of Athletes in the Aspect of .
Humoral-Hormonal Parameters (G. N. Kassil') 19
Changes in Concentration of Tropic Pituitary Hormones During Lengthy
Ski Trip (A. A. Viru, A. P. Kallikorm, K. E. Tomson, T. A. Smirnova,
�c. A. Masso, T. A. Matsin, Ya. P. Pyarnat, T. K. Savi, A. K. Eller) 29
Dynamics of Levels of Growth Hormone, Insulin~ Metabolites ~f Carbohydrate
and Fat Metaboli~m in Blood of ~`.thletes During Bicycle Ergometer
Test Varying in Power (T. D. Bol'shakova, V. A. Siluyanova, Ye. P. Gitel',
A. B. Burkashov, E. V. Sokova, A. Nasonov) 34
Feedback Mechanism of Hypothalamus--Hypophysis-~-Adrenal Cortex System
and Stressor Reaction During Athletic Activity (G. L. Shreyberg,
N. N. Sharov) 43
Correlation Between Glucocorticoid Reaction and Physical Work Capacity 57
(T. A. Smirnova)
Stereological Analysis of Changes in Myocardial Resistance During Exercise 66
_ (R. Masso)
Actomyosin ATPase Activity of Myocardium and Skeletal Muscles of Thyroid- .
ectomied and Adrenalectomied Rats During Physical Conditioning 84
(T. P. Seene, K. E. Tomson, A. K. Eller, K. P. Alev)
Effect of Muscular Activity on Thyroid Homeostasis of the Body (K. E. Tomson) 95
Evaluation of Some Elements of the System of Energy Supply at Rest and
During Exercise When There Are Excessive Thyroid Hormone Levels
(Ye. S. Rom-Bugoslavskaya, M. R. Ozerova, G. I. Kharag) 117
Nonspecific Resistance in Ontogenesis During Muscular Conditioning in 131
the Presence of Hyperthyroidism (V. Ya. Rusin, I. V. Bykov)
Effect of Exercise on Blood Serum Testosterone Level in Conditioned 138
[Trained] White Rats (T. P. Kotsegub, B. I. Fel'dkoren)
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Involve~ent of Calcitonin in Man's Reaction to Muscular Loads
(I. A. Drzhevetskaya, N. N. Limanskiy) � 147
Hormonal Regulation of Calcium and Phosphorus Content of Osseous Tissue
During Dynamic Physical Exercise (G. G. Tsybizov) 152
COPYRIGHT: Unknown
10,057 .
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RADIATION BIOLOGY
UDC: 535.23:577.1:591.443
BIOCHEMICAL BASES OF ACTION OF R?,DIOPR(1TECTIVE AGENTS
I~oscow nIOKHIPtICHESK.IYE ~SNOVY DEYSTVI"YA RAI)IOPROTEKTOROV in Russian 1980 (signed
to press 3 Jul 80) pp 2-5, 167-168
[Annotation, list of abbreviations used, foreword and table of contents from book
i ~~Biochemical Bases ~f Action.of Radioprotective Ag~nts", by Yevgeniy Fedorovich
Romantsev, Vera Dmitriyevna Blokhina, Zoya Ivanovna Zhulanova, Nikolay Nikolayevich
_ Koshcheyenko and igor' Vladimirovich Filiup~vich, Atomizdat, 119~ copies, 168 pagesJ
[Text] This book analyzes ~~he mechanism of action of agents that modify radiation I
lesions on the molecular level. Much attention is devoted to molecular interactions ~
between radioprotective agents, radiosensitizers and biologically important endo- !
genous macromolecules. An original conception is developed of a complex biochemical i
- mechanism of action of agents that modify radiation lesions; special attenti~n is
given to processes of temporary inhibition of replicative processes and stimulation
of DNA repair processes. Analysis is made of data on the significance of temporary
formation of mixed disulfide bonds between radioprotective agents--amino thiols--
- ~nd protein-enzymes having a sulfhydryl group. The existing hypotheses conc~xning
the mechanism of action of radioprotective agents are discussed critically. This
book is intended for radiobioiogists, biochemists, physicians, se nior year students
at biological VUZ's and medical institutes. Tables 1, figures 14; references 570.
List of Used Abbreviations
tsAMF-- adenosine 3':5'-cyclic phosphate
APAETF--aninopropylaminoethyl thiophosphate (gammaphos)
ATP--adenosine-S'-triphosphoric acid
AET--2-aminoethylisothiuronium
BSA--bovine serum albumin ~
GTF--guanosine-5'-triphosphoric acid
GED--guanidoethyl disulf ide
dATF~-deoxyadenosine triphosphate
dGTF--deoxyguanosine triphosphate
Diamide-bis--(N,N-dimethylamide)-diazene of dicarboxylic acid
DNK--deoxyribonucleic acid
- DTT--dithioerythrite
dTsMF--deoxycytidine manophosphate
3TsTF--deoxycytidine triphosphate
:iRNK--messenger RNA
rIPA--S-mercaptoethylamine
N'EG--2-mercaptoethylguanidine
30
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NADN--nicotinamide adenine dinucleotic~e
NADFNZ--nicotinamide a3~nine dinucleotide phosphate
OMF--orotidine monophosphate
PGPS--prostaglandin-like compounds
PPS--peroxide-like compounds
PKhMB--n-chloromercuribenzoate
RNK--ribonucleic acid
rRNA--ribosomal RNA
yaRNA--nucleic RNA
TMF--thymidine monophosphate
TTF--thymidine triphosphate
' UMF--uridine-5'-monophosphoric acid
Foreword
The problem of change in radiosensitivity of the organism by means of various chemi-
cal compounds--modifiers--is still one of the most pressing ones that are being
worked on intensively in modern radiobiology.
In the Soviet Union, much attention is given to development of this direction. The
first monograph analyzing the state of the ques~ion of chemical protection of or-
ganisms against ionizing radiation was published by Ye. F. Romantsev and A. V.
Savich in 1958. Since that time, many books have been published shedding light
on different aspects of modification of radiation damage by means of chemicai
compounds (Yu. B. Kudryashov, P. G. Zherebchenko, A. G. Sverdlov, N. N. Suvorov,
V. S. Shashkov, S. P. Yarmonenko, L. Kh. Eydus, A. S. Mozzhukhin, F. Yu. Rachinskiy
and others).
_ However, radiobiology is developing so rapidly, that there is an urgent need to
periodically report the achievements in this branch of natural science in the
form of summarizing works. This book describes the concept of complex biochemical
mechanism of radiation lesion modifiers, which is developed by the authors, and
analyzes the status of the problem as a whole.
Ir is of basic: importance to comprehension of the triggering effects of radiation
to conduct studies of the molecular mechani~ms of action of radiation modifiers.
At the sar~e time, identification of the molecular mechanisms of the effects of
radiation on cells open~ up the prospect of new approaches to the search for
effective radioprotective and radiosensitizing agents. For this reason, we
deemed it expedient to shed light on a number of basic sections of radiation bio-
chemistry, with emphasis on biochemical mechanisms of interphase death of irradi-
ated cells. Modifiers intervene actively in virtually all biochemical processes
that determine individual and species-specific radiosensitivity. For this reason,
analysis is made in relevant chapters of data on the effects of radioprotective
and radiosensitizing agents on DNA metabolism, RNA biosynthesis, processes of DNA
replication and repair, protein synthesis in animals and energetic processes in
the cell. It was judged necessary to discuss briefly the state of the question
of possible role of cyclic adenosine monophosphate and prostaglandins in the
mechanism of radioprotective agents. Much attention is devoted to analysis of mole-
cular. mechanisms of manifestation of activity of radiation damage modifiers,
questions of temporary inhibition of replicative processes and optimization of
c~nditions for DNA repair. At the end of the book, there is discussion of some
~eneral patterns in the molecular mechanism of radiation damage modifders, and a
scheme is offered for different stages of their biological activity.
31
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It so happened, historically, that radioprotective agents were studied as compounds
intended to protect eukaryotes against absolute minimal lethal doses of ionizing
radiation ("hemopoietic form" of radiation sickness). At the sa*_ne time, it is
already ap~arent that it is possible, in principle, to provide protection against
considerably larger doses of penetrating radiation. This increases ?nterest in
the problem of modifying radiation effects with chemical compounds. .
Modern molecular radiobiology and radiation biochemistry are d~hel~~egthistbook
- rapidly. For this reason, we are aware of the fact that, by
is published, new data will appear concerning the mechanism of radiation damag,e
modifiers.
The authors will be grateful to all readers for any critical comments they may have.
Contents Page
4
Foreword 6
Chapter 1. Radiobiochemical Processes in Mammalian Tissues
Possibility, in principle, of modifying radiosensitivity of mammalian cells 8
Direct and indirect effects of ionizing radiation on living cells 9
Target molecules ~
Effect of ionizing radiation on biochemical processes in mammalian cell 11
_ culture 12
Biochemical mechanisms of interphase cell death
Changes in structure and function of nucleic acids and nucleoprotein 12
complexes 22
- Impairment of bioenergetics 22
Ionizing radiation and membrane permeability 24
Biochemical mechanisms of reproductive cell death 25
Bibliography
Chapter 2. The Role of Endogenous Thiols in Action of Radioprotective Agents 39
Bibliography
Chapter 3. T:~e Role of Functional Protein-Enzyme Groups in Binding Amino 43
Thiols and Disulfides
General conceptions of the role of SH and SS groups in biological 43
activity of enzymes 44
Reaction of thiol-disulfide exchange
Formation and properties of miYed disulfides between proteins and 46
low molecular thiols and disulfides
Formation of mixed disulfides between radioprotective amino thiols and 50
disulfides 52
Enzymology of thiol-disulfic~e exchange [metabolism?]
Bibliography 63
Chapter 4. Effect of Amino Thiol Protective Agents on DNA Synthesis
- General conceptions of correlation between intensity of inetabolism 63
and radiosensitivity of the organism 64
Effect of amino thiols on some aspects of DNA metabolism 66
riechanism of inhibitory effect of amino thiols on DNA synthesis 70
Bibliography
Chapter 5. Molecular Bases of the Biochemical Mechanism of Radioprotective 73
Action of Amino Th~ols
Evaluation of the role of formation of mixed disulfides of amino thiols 73
and proteins in the radioprotective action
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Amino thiols as a means of preventing formation and expression of primary
damage in DNA molecules 77
Bibliography 85
Chapter 6. Molecular Mechanisms of Radioprotective Action of Agents on
- Biosynthesis of Ribonucleic Acids 90
Introduction 90
Possible mechanisms of action of amino thiols and disulfides 90
Some information on the effects of indolyl alkylamines 99
Radioprotective eff icacy of ATP and j.ts derivatives 100
Effects of exogenous nucleic acids on postradiation metabolism of RNA 103
Conception of the mechanism of action of antibodies to sin.gle-helix DNA
on the postradiation transcription process 105
Bibliography
Chapter 7. Effects of Radioprotective Agents on Protein Synthesis in Animals 113
Isolated protein-synthesizing systems and effect of radioprotective
agents on them (in vitro experiments) 113
Effect of protective agents on protein synthesis in nuclei of intact
rat spleen and thymus 115
Effect ~f protective agents on protein ~ynthesis in splenic nuclei of
- rats exposed to radiation at different dose rates 126
Bibliography 12$
Chapter 8 Effect of Chemical Protective Agents on Energy Processes in Cells 135
Bibliogr iphy 143
Chapter 9. Role of Adenosine 3':5'-Cyclic Phosphate (cAMP) in the
Radioprotective Action of Amino Thiols 145
Bibliography
Chapter 10. Prostaglandins and Mechanism of Action of Radioprotective Agents 152
Bibliography
Chapter 11. General Patterns of Molecular Mechanisms of Action of Radiation
Damage Modifiers 157
Bibliography
COPYRIGHT: Atomizdat, 1980
10,657
csu; 1840/999
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HUMAN FACTORS
_ UDC 6297.06(082)
MATHEMATICAL MODEL OF HUMAN OPERATOR USED IN FLIGHT SIMUI~ATORS
Moscow AVIATSIONNYYE TRENAZHERY in Russian 1978 (signed to press 5 Jul 78) pp 14-20
[Section 1.3 from book "Flight Simulators", by ~,'asiliy Afanas'yevich Bodner, Rafael'
Abdullovich Zakirov and Inna Ivanovna Smirnova, Izdatel'stvo "1K3shinostroyeniye",
3600 copies, 192 pages]
[Text] 1.3. Mathematical Model of the Human Operator
We will ~:xamine some psychophysiol~~~ical ~haracteristics of the onerator which should
be considered i.n formulating the information similarity criteria. Of these charac-
t:eristics, the most important are the transmission functions and transmission
information capacity on visual, acceleration and auditory channels.
The operator's trans~:~ssion runctions are determined by his psychophysiological
features and properties of the control loop in which the operator is working. If
the operator controls a stationary loop of control, for example, FV [flight vehicleJ,
then his transmission function in the perception of visual information and trans-
m:ssion of signals to the control units will look like
ke `s(atis + 1) (1.19)
WO~S~ (tls + 1)(tss + 1) '
. where k--amplification coefficient; T--time for formation of response reaction to
inp�t signal; T1 and T2--time constants charactErizing the transmission of signals
on nerve fibers; a--coefficient which takes into consideration the degree of train-
ing, experience, simulation, fatigue and type assignment~ The average values for
the transmission function coefficients (1.19) are presented in table 1.
It is apparent that for different frequencies of the perceived signal, the para-
meters of the transmission function adopt different values. This indicates the
adaptive properties of the operator. The operator, working in the control ].oop,
~~dapts by changing the parameters T]~, T2 and k, striving, as studies have shown,
to minimize the standard errur and Eo maintain the phase margin on the order of 40�.
In other words, the operator is a natural system of optimization.
- The operator's transmission function, presented in the form ~f expression (1.19)
does not take into consideration all the features of the operator (nonlinearity,
adaptivity). Expressions were therefore suggested which take more complete consi-
deration of the psychophysiological characteristics of the operator. One of these
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Table 1.1.
Signal z1 sz ~a a k
frequencies I I I
0,16 25 0,67 0,15 0,08 100
0,32 9,1 0,22 A,20 0,055 40
0,48 5 0,09 0,25 0~067 15
mathematical models which describes the operator's transmission properties is pre-
sented in figure 1.4. This model takes into account the zone of insensitivity in
signal perception, displacement equivalent to the noise generator~ (remnant),
dynamics of formation of the response reactions in the central nervous system, and
dynamics of signal transmission to the arm muscles.
K~ M r .
a 1J+1
e~t -r: sx +~+1 + ) ~ i' ~~t~
a, N~ (i.~,s, ~ir m1=+nt.f
Figure 1. Structural Plan of Model of Perception of Human Operator
The examined model of transmission properties of the operator can be presented in
the form of an analytical expression
Wo (s) = kTke-~: ( s+ 2E S~- 11 W~ ~S) kA, (1. 20)
t Wrt ~R 1
where Wk(s)--transmission function of the arm, and
do~ + 1 ; (1. 21)
WR ~S) = atss azss + a3s + 1!
i QTIz
a 2 raTl e~`�T~ ;(1. 22)
kr=1-erf~=~+~ n \arl
at
2 aA (1. 23)
k~ - V ~ ,
�a
k--amplification coefficient.
' In these expressions, aA and aT are mathematical expectations, and Q~ and aT are
the standard deviations of the element characteristic~ on the assumption of
normal distribution. The numerical values of the coefficients which are included
in expression (1.20) are given in table 1.2.
Acceleration information plays a large role in flight control, therefore we *.~ill
examine the features of its perception. The main biolagical sensor which per-
ceivesaccel.Erationsand supplies man with information on the change in spatial position
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is the vestibular analyzer. Its purpose is static and dynamic orientation of
the head and orientation of the eyes in any position of the head.
Table 1.2.
w
c, c I ao, c I al, c3 ~ az~ ~2 ( a3~ ~ I rad/s E I b~
0,2 0,05 0,098 0,007 0,002 4,17 0,523 2
- The vestibular analyzer contains otoliths and semicircular canals.
'Phe otolithic apparatus includes the utriculus and sacculus. The structure of these
organs is characterized by a fixed part, the macula, and m~vable parts, like
pendulums, the otoliths. Movement of the heavy otolith above the macula causes a
sensation of i.nclination or movement. It is believed that the utriculus reacts to
linear accelerations, while the sacculus perceives vibrations.
Linear accelerations generated by the gravity force or active forces are the
input quantity of the otolithic appparatus. In this case, the following con-
clusion is correct: the utriculus perceives forces of any direction (vector
quantities) and is stimulated by shearing forces in the plane of the otolith. The
quantity of information obtained by man through the otoliths is determined by
their dynamic characteristics.
Studies have shown that a redamping pendulum which reacts to linear accelerations
of an;~ direction is a mathematical model of the otoliths. The otoliths, like any
pend~.,.lum~ do not distinguish accelerations of active forces and the gravity force.
Man can therefore correctly determine the position of the local vertical only with
u fixed position or with rectilinear uniform movement.
The otolithic transmission function can be presented in the form
~X ~ ) - k' (1. 24)
W S -~T1S + (T~~ + 1~ '
where T1=10...11 and T2=0.66--ti~ne constants, s; k~--ampl.ification co~fficient;
j--inpuf acceleration; x--movement of otolith.
The amplitude-frequency and phase-frequency characteristics of the atolith are
shown in figure 1.5. The characteristics of the otolothic apparatus change de-
pending an the frequency of the perceived acceleration: at low frequencies,
w� 1/Tl, it operates as an acceleration sensor, at medium frequencies 1< W< 1/T2
T1
it operates as a velocity sensor, and at higt? frequPncies w> 1/T2, it operates as
a sensor for human body movement in space. It follows from here that high-fre-
quency, and partially medium frequancy accelerations play the main role in the
human spatial orientation. In addition, because of the property of the body's
adaptation to constant and low-frequency accelerations, information about these
accelerations is not used to determine the spatial position.
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y. -
_ eo -zo(~.coy~)
~ I `
p I I
t~ W�
~
~ ~ '/T~ I '/T T~
-do go~ o,~ fo w

_ Figure 1.5. Amplitude- and Phase-Frequency Characteristics
The otolothic apparatus has a threshold of insensitivity which is not reflected by
transmission function (1.24). A more accurate mathematical model of tae otolithic
apparatus is shown in figure 1.6. Inti~is model, the f irst link depicts the orien-
tation of the otoliths, the second link is the d.ynamics, the third is the threshold
of perception (sensitivity), the fourth is the amplification properties of the
central nervous system, and the f ifth iink with transmission function F(s) is the
boosting properties. Without consideraCion for the zone of sensitivity, the trans-
mission function of the otoliths adopts the appearance of expression (1.24).
r
2 ~ ~3~Om~umei ~ (]~qHC i0~9~t~r~e
.?Beno ~ ~ I ~ d~unrenuA
Cu~a opuenmuquu ~ M ~M ~ f(s) ~
OmonumoB MS~+CS+K I M ~9~
anaMUKa 10 enue
5~ omonumoB ~ 6~vjBcmBum. I ~ Bep
uKan~�
L_--__ _-------`--------.J
- Figure 1.6. Structural Plan of Mathematical Model of Otolithic Apparatus
Key:l. Force 6. Zone of insensitivity
2. Link of otolithic orientation 7. Central nervous system
3. Otoliths 8. Sensation of movement
4. Movement 9. Sensation of vertical
S. Dynamics of otoliths
The mathematical model of the semicircular canals as elements sensitive to angular
accelerations e can be presented in the f orm of a transmission function
' ) k2 (i.2~)
C?~ S =(Tas 1)(Tas.+ 1~ ,
- where T3=9...12 and T4=0.003-0.03--time constants, s; k2--amplification coefficient;
n--di.splacement of endolymph during accelerations.
The amplitude-frequency and phase-frequency characteristics of the semicircular
canal are shown in figure 1.7. At low frequencies, man does not distinguish the
_ constant angular rotatinn velocity and does not have low sensitivity to the Iow-
frequency changes in this velocity. At medium frequencies, 0.07 < w< 300 s-1, the
semicircular canals operate as rotation velocity sensors, and at high frequencies,
w> 300 s-1~ they operate as sensors of human body rotation.
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x . '
+0~ -
3 rc
~r
4 ~ ~ ~ 20
o ~ '
_ o~ o,~ f fo ~00 ;~DD w ~p
~
_4 0
~ E~ deg/s2
~r
- Figure 1.7. Amplitude-Phase-Frequency Figure 1.8. Thresholds of Perception of
Character~stics of Otolithic Apparatus Angular Acceleration
The s~micircular canals have a threshold of perception which depends on the delay
- time, i.e., on the moment of development of the accelerati.on to the moment of its
perception (fig 1.8). It is apparent that the maximum value of threshold accelera-
tion is 0.5 deg/s2, while the threshold of perception with delay time of 0.1 s is
- 5 deg/s2.
F -~-I -
~ 3KCnrpur.r~~,~~ o
0
~ 4 u
~1~ ,oa Qp~1-e~~%~
0
i 1
m
~
~
v
- o zw ye ~1
~3>lYO/1lLV2C/ACO dMFt[
Figure 1.9. Characteristi~~s of Degree of Training
, Key:
1. Degree ef tra~.ning
2. F.xperiment
3. Number of days
During the perception of an~ular velocities, the human operator also has threshold
values which, with a delay of 0.1 s are: 3.2 deg/s for heeling; 2.6 deg/s for
pitching, and 1.1 deg/s for yawing.
The vestibular system and vision are linked so that the image an the retina i.s
stabilized during rotation of the head. Discrepancy between movement of the eyes
~i~c! sensations of the vestibular system does not exceed 0.1 s. Negative illusians
develop with greaCer discrepancy.
The examined transmission functions are mathematical models of pilot activity in
the control loop of a FV or simulator. Acquisition of habits and the knowledge
necessary for correct functioning is attained in the training process, including
on the simulator. During training on the simulator, the pilot acquires knowledge
that increases as the training tinie increases. Experimental studies show [10] that
increase in knowiedge during the training process occurs by the exponential law.
3$
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In examining the transmission function (1.19) the degree of operator training was
characterized. by the c~efficient a. This coefficient increases in the training
process and reaches a certain limiting value at which training can be considered
finished. The transmission function which characterizes the training process
accordin~ to the experimental data may be presented in the form of a lag network
Il'~~a (S) _ ~ ~
r~s 1
where T~--time constant of training which averages 25 days.
The coefficient of training a, according to the transmission function l~.o~(S) can
- be presented in the form
~ ~S~ _ ao
tos T 1 ~
where a~--limiring value of trainin~ coefficient.
Figure 1.9 gives a graph for the process of increase in the degree of training as
a function of time. It also shows the experimental points. It is apparent that
the model of training in the form of the transmission function a(s) agrees well
with the experimental data. The process of training the operator is several orders
longer than the process of control, therefore the coefficient a in formula (1.19)
should be considered constant.
C~PYRIGHT: Izdatel'stvo "Mashinostroyeniye", 1978
9035
cso: si44/1610
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UDC 62-50
REDUCTION IN ADAPTATION TIME IN ERGATIC CONTROL SXSTEMS
Kiev AVTOMATIKA in Russian No 5, Sep-Oct 79 (manuscript received 19 Feb 79) pp 59-62
[Article by V. V. Pavlov and A. M. Meleshev, Institute of Cybernetics, UkSSR Aca-
demy of Sciences]
[TextJ The possibility of rapid compensation for the consequences of failures in
the control system plays an exceptionally important role in ergatic systems in
which the human operator performs so-called manual control of an object and the
control process takes a short time (less than S mi.nutes). Auring failures, the
control object becomes somewhat different for the human operator. Time for "adap-
tation" to the "new object" is needed in order to maintain the previoras control
quality. The adaptation time for the mentioned processes can comprise a signifi-
cant part of the time for the entire control process. Its reduction is therefore
an important task for ergatic control systems. One of the methods for reducing the
"adaptation" time t is visual depiction of the main properties of the control
object (for examplea tables of a ship's maneuver characteristics [3], etc.). This
- may be a graphic relationship, or any other picture illustrations which permit
utiequivocal illustration of the necessary data. Images on cathode-ray tube screens
are the most valuable. They permit rapid illustration in a real time scale of
the characteristics.depending on the technical condition of the control object and
the formed situation. For ergatic systems with manual control, iti is convenient
to depict in axes associated with the cantrol object the quantities of the control-
lin~ factor developed by the control units, depending on the deviation of the
control lever from the neutral position. We will call this graphic picture DCF,
"diagram of controlling factors."
In order to clarify what has been said, we will examine a simple ergatic system
oF compensator tracking (fig 1). The human operator sees in the visual indicator
errors e, e which are proportional to the deviation of the center of mass of the
control obje~t from the zero position in the OX~XO coordinate system (fig 1, b, d),
By deviating the control lever in the necessary direction, the human operator
creates the controlling factor T which is proportional to deviating the control
_ lever from the neutral position Tx=cd, TY=cy (fig l, c d). Thus, the human opera-
tor manually controls the object with the task of keeping, for example, the error
modulus e,e less than the assigned value a(i.e., tracking with the assigned
accuracy ~).y It is easy to see that under the conditions stipulated above, the
orientation of the T vector uf the controlling factor coincides with ~he orienta-
tion of the R vector of the contr.ol lever deviation, i.e., S=a.
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If we now fix the vector modulus R and change the angle of orientatiun a=0-360,
then the end of vector T outlines a curve in the plane of associated OXY coordi-
nates whose visual depiction we will call the diagram of controlling factors. The
DCF prov'des a graphic idea regarding the quantity of the controlling factor in
the direction of control lever deviation. In our case, DCF will have the appear-
ance of a circle with radius proportional .to the deviation of the control lever
from the neutral position.
We now assume that a malfunction develops in the control system (fig 1}. As a
result, the component Ty of the controlling factor is reduced by 1/3: Tx=cd, Ty-
0.66 cy. In this case, the DCF will be considerably altered, i.e., the quantity
of the modulus of the controlling fact.or and the angle of orientation of vector T
will change, i.e., S~a. For this case, figure l,e presents the DCF where the
yr
8' ~ 6' S~
- �,.3 c~~ f e 4,
RN 4�U)~ yY 01i 9~ 3'
yjoa Er !v Y l0~ ~ 6.
L ' S 4 2~
u 1!' l0 9 / ~ 1
I y Ya y 12, n ~ / l
_ ~R _ 1
j`; 4.-;, J I t T 1J / 1
f .~r
E u i ' J' 14 15 Z~ 17 pJ~
\
- j ~ x , ~ !4' i6 ~9 17'
/8
b c � d xo ~s~ 1~'
. 70,
- /8' /9'
Figure 1. Structural Plan of Control System and Basic Correlations
Key:
a. structural plan
b. picture observed by human operator in visual indicator (e , sX--quantities
proportional to deviations of regulated coordinates from ~ssigned);
c. R--vector of control lever deviation from neutral position (a--orientation
angle of vector R in associated axes)
d. T--vector of controlling factor applied in center of mass of control object
(R--angle of orientation of vector T in associated axes; OX~Y~ -system of
reference coordinates)
e, yN, xn--axes of visual indicator for diagram of controlling factors that
coincide in direction with the corresponding associated axes of the con-
- trol object.
- n~imbers with the stroke indicate the corresponding points for the DCF of~the
original system. The points correspond to the 24 fixed values of a. It is
apparent in figure 1, e that, for example, with a=45� (point 3') in the original
system R=45�, and in the system wit'n failure S