JPRS ID: 9501 USSR REPORT LIFE SCIENCES BIOMEDICAL AND BEHAVIORAL SCIENCES

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JPRS L/9501
23 January 1981
USSR Report
LIFE SCIENCES
BIOMEDICAL AND BEHAVIORAL SCIENCES
(FOUO 1 /81)
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.4
JPRS L/9501
23 January 1981
USS R REPORT
LIFE $CIENCES
BIOMEDICAL AND BEHAVIORAL SCIENCES
- (FOUO 1/ 81)
CONTENTS
ADVANCED BIOTECfINOLOGY
The Development of Molecular-Biological Research and Multilateral
Cooperation
1
Some Questions of the Theory of Swimming of Fishes and Dolphins
8
Book on Biological Bases of Behavior Control of Fish in ilowing
Water ..................................o...................
13
Applied Mathematics .1n Biology
24
' ADVANCED MEDICAL TECHNOLOGY
Mathematical Descriptions of Cardiac Bioelectrical Activity
27
- Applications of Focused Ultrasound
30
Monograph Analyzes Cryogeni'c Preservatives
35
Lasers in Traumatology and Orthopedics
38
ENVIRONMENTAL HAZARDS
Radiation Exposure, Reduction of and Methods for Chemical Protection
41
~ Radiation Injury Diagnosis Based on Analysis of Blood Nucleic Acids
50
Biophysics of Canplex Systems and Radiation Injuries
55
; - a- [ III - tTSSR - 213 S&T FOUO]
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Radioactlve Agents and Wounds 7$
Prediction of Ionizing Radiation Transfer by Photometric Methods 82
- Radiation Safety and Shielding of Nuclea.r Power Plants 85
Biological Aspects of Radioactive Contanaination of the Seas 90
-b-
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ADVANCED BIOTECHNOLOGY
UDC 577.23
THE DEVELOPMENT OF MOLECULAR-BIOLOGICAL RESEARCH AND MULTILATERAL COOPERATION
Moscow VESTN.IK AKADIIKII NAUK SSSR in Russian No 6, 1980 pp 82-88
[Paper presented by Academician J. Kiman of the Czechoslovak Academy of Sciences
at the XI Meeting of Representatives of the Academies of Sciences of the Socialist
Countries in Tallinn]
[Text] The topic of this presentation is closely related to one of the most.
important tasks of bilateral and multilateral cooperation among socialist countries
i.n the field of natural sciences, which is to ensure the foremost development of
the most promising scientific investigations. Specifically, the discussion will
- focus on the necessary, in our opiniori, measures that must be taken to ensure the
developcnent of molecular-bioiogical research currently being coordinated by the
New Program of,Multilateral CoopE.ratton.
As part of the topic which interests us, we would first like to briefly charac-
terize the goals and methods of those studies which are basically molecular-genetic
in nature. Tn general the goal of this research is to explain the molecular basis
of the various functions of biological systems--from the simplest forms, i.e.,
functionally active biological macromolecules (nucleic acids and proteins) and
molecular complexes (nucleoproteides and pha.ges, viruses, procaryotic and eucaryotic
cells to multicellular plant and animal organisms.
Biological phenomena, which serve as the subject of molecular biology, are simul-
taneously studied as phenomena by other biorogical disciplines. This involves
; the fundamental behavior of living systems, such as nourisrnent, growth, reproduc-
tion, mobility, irritability, secretability and specific characteristics such as
infectability, resistance, differentiation, abnorrnal cell transformations, immunity
and tissue histocompatability as well.
= The spectrum of experimental methods used in molecular-biological research is very
wide and attests to the complexity of the problems being investigated.
Molecular-biological analysis, based on the biochemistry of nucleic acids and
proteins, requires, in addition to complex physical-chemical and physical methods,
x-ray diffraction techniques, nestron scattering and nuclear magnetic resonance,
and now, mathematical models, not only the necessary equipment but also a wide
assortlnent of special reactants, enzymes and radioactive isotopes. The application
, of this arsenal of techniques can be effective only if there is a well characterized
experimental biological model, whose value increases the more its genetic structure
is defined.
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It is necessary to emphasize the qualitative specifics of the experimental model
in molecular-biological research in comparison to that used in other areas of
~ biological research. If the model is biologically and genetically simple enough,
- then iC becomes not only the subject of the research, but its instrument as well,
since it becomes necessary at all stzges of research - from the statement Qf the
vroblem to the interpretation of the results. This approach allowed us to postulate
the existence of a nucleotide triplet as the basic unit of the genetic code some
20 years ago, while direct biochemical proof of this was obtained only in 1961.
In the last decade, the application of this relatively simple experimental bio-
logical model was partially resgonsible for the transformatior~ of oncogenic
viruses and their host cells into valuable tools in the study of not only malignant
cell changes but of several nonnal cell functions and the structure of cell genoma
as well. Therefore we now have numerous data on the molecular basis of virus and
microbe functions.
~ Today this approach, together with the wide appl.ication of recombinant DNA tech-
niques as we:Ll as the whole c anplex of immunological, i.mmunogenetic, cytological,
physico-chemical analysis and mathematiaal models, offers wide possibilities for
the study of the molecular basis of fundamental and specialized functions of
eurocaryotic cells, amon.g these human cells. This is supported by the wealth of
new data obtaized in the past few years which have substantially changed our ideas.
- This in part includes the data on the dual role of biological molecularly functioning
elements, i.e., those transmitted in replication and those replicating in trans-
mission, and data on nucleases in r.eplication (after the discovery of functioning
;nacromolecule complexes).
These are also data obtained by virtue of the discovery of reversible transcrip-
tase (1970) and the restriction enzymes, and later by the development of techniques
for determining DNA and RNA sequences, by obtaining recombinant DNA, in other
words, data obtained by virtue of the application of the technology of genetic
engineering (after 1972).
In addition, these are new data on the "maturity" of the information transcript
and pri.marily on the previously unknown technique of genetic recombinaCion by
combining information carrying RNA fragments after a preliminary breakdown of the
molecular predecessors (splicing). This relatively simple technique of genetic
recombination in eucaryotic ce11s, which was also probably the earliest evolutionary
phenomenon, allows us to postulate that specifically RNA and not nNA serves as
the priunary information material. It is probable that a simple ribonucleotide
(for example, nicotinamidadenine-dinucleotide) could have become in the procPSs
of evolution the pri.mary information unit, the genetic monomer which served as
the basis of cell metabolism as the precursor of reactions which are catalyzed
by enzymes which evolved later. This new idea about the prisnary information function
of RNA is not even contradicted by data on protein which recognize necleotides.
To these proteins, in addition to the already known lac-repressor, belong the
enzymes of RNA-RNAse II maturation which is a nucleotide and probably also the
_ proteins of inem.branes which are responsible for the transport of specific nucleic
acids within the cell.
- 2
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This, in the final analysis, is a very striking result obtained in the process
of studying the structure of eucaryotic genes; the discovery of their mosaic
- structure in which coding and non-coding sequences alternate, the so-called exones
and intrones. The recEntly obtained p.roofs of the fact that individual exones
within the gsne determine individual functional regions in the structure of
specialized proteins (on the example of the heavy chain in immunoglobulin) are
- directly related to this discovery.
In Jeneral, it can be said that the application of new recombinant DNA techniques
is approaching the possibility of molecular analysis of eucaryotic cells on a
level which is now reached only by studies of viruses and microbe cells. This
is supported in part by the preli.minary results of experiments on determining the
functional properties of catalytic regions in the information molecules of eucariotic
cells.
_ The detailed analysis of the structure af some'animal genes, including human
genes, became possible thanks to the newly discovered capability of producing
- significantly greater (several milligrams) quantities of individual genes in
microbe cells which received these genes through genetic engineering. This is
the method used in the production of, for example, globin, immunoglobulin, oval-
bumin, histone genes and the genes of some viruses. The recently discovered prin-
ciple about the increased selective transcription of weak quantities of infomnation
RNA will enable us in the near future to obtain larger quantities of genes such
_ as the gastrin gene, which was recently isolated in precisely this way, ar,d it
will facilitate detailed study of these previously inaccessible genes.
Comparative studies of dEfective and non-defective oncogenic viruses led to the
discovery and retransmission of the gene sarc. Finally, by means of direct
biological evidence, the region in the molecule of plasmidic DNA (one of the plasmids
- of the bacteria Escherichia coli) which is responsible for its virulance was
recently identified.
The use of the genetically determined biological model as an experiteental tool,
together with current techniques of isolation, cleaning, amplification, immuno-
logical and molecular analysis of specific components of living systems, allows
us to study in detail the mechanisms of vdrious functional phenomena of the
eucaryotic cell on all of its structural levels even to the level of cell organelles.
This approach has led to important results in the area of cell nucleus research
and in studies of cltromatin and cytoplasmatic cell organelles.
Studies of cell membranes - their morphogenesis and various functions: transport,
signal, recognition, receptor, and among these the function of the nerve cell
membrane during the transmission of a nerve irnpulse proceed in similar ways.
This method is likewise used in the study of cell mobility carried out by its
- actomyosin apparatus which directly participates in the formation of the cell
cytoskeleton, and especially by the geodome which reacts to external signals.
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v
In addition, enzyme mechanicros of the so-czlled biologiczl clock which determines
the daily biorhythm of sane birds and mammala were discovered. Thtse are usually
conditioned by the level of synthesis of the hormone epiphase-melatonin. A hypo-
thesis was also set forth about the molecular mechanism of genetic clocks, where
- the primary role is assigned to the activity of the reatriction-modification
enzymes.
_ Final.ly, there have been successful results in the field of bioenergetics, especially
- photosynthesis in plants as well as photosynthetic bacteria.
In this far from complete survey of the successes of contemporzry molecular biology
we have not yet touched on its applied aspect. The przctical zpplication of
genetic engineering is well known, for example, the production of importani: phy-
siologically active natural substzncer in significant quantities, primari?y the
various substances produced in the human organism such as insulin, the growth
hormone, interferon and others, and the production of a series of bacterial en-
zymes important for industry. Moreover, data have recently been published on the
- successful identification of the infectious nature of the DNA zs well as in the
, DNA of the lambdoidic phage and reproducing in the bacteria Escherichia coli.
Of extremely practical significance is the perfecting of substitute enzyme therapy
of genetic diseases of man, in which the chemical recognition signals of the en-
zymes are used to heighten the selectivity of cell activity. ':nother accomplish-
ment is the recent synthesis of the first pigunent-protein complex (v(-pyrochloro-
phyll-pheophytin), which imitates the function of the photosynthetic reaction
center of green bacteria (Rhodopseudomonas sphaeroides), and which represents the
first level of transfoxming solar energy into electrical energy.
This brief characterization of goals,. methods and some of the most recent results
in the field of molecular biology represents not so much a survey of the state
of current molecular-biological rpsearch or an example of its great scientific and
practical significance but serves as a basis for the recommendations made below
for ensuring the development of this type of research within the context of mul-
tilateral scientific cooperation among the socialist countries.
The development of molecular-biologzcal research depends on two interrelated
- factors.
The first factor is scientific personnel who, in the course of the experiment
_ and depending on the specific problem, are able to effectively use ideas and
methods from a variety of related biological and non-biological areas and dis-
- ciplines.
The second factor is the material base of molecular biology, including the fol-
lowing: special equipment, that is, preparatory, analytical and cultivation
technology and among these, special laboratory equipment for microwork as well
as for the purpose of increasing the effectiveness and safety of experimental work;
ultrapure reactants, special compounds and biologically active material including =
= a growing assortment of specific antibadies, serums and serum components for tissue
= cultures, and biologically active materials and radioisotopes with great specificity; =
4 -
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a sufficient quantity and the necessary assortnent of genetic biomodels, e.g., ~
mutant strains of plasmids, phages, viruses, microbe cells, plant and animal cells ~
as well as pure strains of laboratc+ry animals.
Both of these factors should correspond to the level of experiments being Gonducted.
We shciuld take into account the fact that in the past twenty years in the countries
of the West the development of molecular biology, i:l spite of the deepening
= economic crisiss is ensured by large investments of capital, by a rapid development ~
of special industries and by an increase in the production. of products which are
- required by all types of research. At the present time this tendency can be
noted not onZy in the United States, Great Britain, France, Italy, but in average
and smaller countries as well, for example, Sweden, The Netherlands, Switzerland
and even Luxembourg and Liechtenstein.
Until 1970 the material support for molecular biology in tie West consisted of,
_ in general, the production of ursique instruments, equipment and the required
assortsnent of reactants and special biologically active substances. The commercial
production of biomodels was practically limited to pure strains of experimental -
animals. The discovery of reverse transcriptase in 1970 led to a mass commercial
production of the AMV virus in the U.S. as a source of reverse transcriptase, -
and soon after, to the mass production of a whole series of various oncoviral
strains which literally xesulted in an explosion of information in that area of
molecular biology.
In 1971, that is, a year before the official announcement about work in genetic =
engineering, thP commercial production of basic technological elements for these
experiments was already organized in the U.S. In the U.S. and Western Europe
specialized centers for the mass production of not only compounds such as insulin, -
the growth hormone, interferon, industrial enzymes but also of separate genes for
basic research were set up. .
During the las"L two years, for example, there has been a sharp increase in the
production of biomodels, among them cell hybrids used in the production of certain
antibodies. The method of cultivating the B-lymphocyte antibody producer by' -
joining it to a tumor cell enables scientists to select a specific ceYl clone =
for the mass production of any given antibody. This method allows the production
_ of a practically unlimited range of experimental antibodies for immunology,
endocrinology and toxicology. According to several Western companies, the capital
investment required for the production of certain biomodel types pays for itself
five times faster than in the area of electronics. In the mass production of
biomodels Iike viruses, cell hybrids and individual genes, small specialized
companies are more effective and profitable since larger ones do not adjust readi?y
_ to the constantly changing requirements.
Molecular-biological research is supported in all remaining areas, including 'the
production of equipment designed to make research more ef�ective.
5
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' 0�lr primary goal is to increase the effectiveness of molecular-biologica.l research
- and we should explore and realize alI of our potential in reaching this gozl.
Let us now examine the current state and developciental potential of molecular
biology in the socialist countries.
- Today we can with all assurance say that in the past decade molecular-biological
research developed at a fast rate both quantitively znd qualitively. Molecular
biology has reached-a high level in the USSR based on its own traditions as we21
as on the most promising international accomplishments and enjoys the support
of the CPSU Central Committee and the USSR Council of Ministerc. Molecular-
biological research in other brother countries, Bulgaria, Sungary, German Democratic
Reputlic, Poland and Czechnslovakia, has also had great successes and progress.
The results obtained in the fields of microbiology and genetics of microorganisms,
virology and oncovirology, ismnunology and immunogeneties, in membranology, in the
field of protein 3nd n;zcleic acid biosyr.thesis and reeently in recombinant DNA
~ techniques in3icate a good preparation on the part of our zcientific personnel,
an ability to rpsolve any problems which the development of molecular biology
poses in successful competition with scientific developments in capitalist cauntries.
But in addition to personnel, the successful development of molecular-biological
research requires a material base of its own production. Therefore we must, to
the extent that we are able and by mutual cooperations find and develop everything
that can contribute to the independent development of our molecular biology and
ensure our experimental and technological competitiveness in this area which will
facilitate a mutually agreeable exchange of technology and data with the capitalist ~
countries.
Molecular-biological experiments are of a very fundamental nature. Therefore
there is no doubt that in time we wi11 have our own well organized and growing
- material base in whose creation would have participated all of our countries.
We should immediately begin building this base, quickly and with initiative,
everyone to the best of his ability. Certain meaaures can be taken imnediately,
and some already have beer., for example, the mobilization of the internal resources
of our Academies. With relatively small capital expenditures it is possible to
make a decision about a bank for registering and safe-keeping special biological
materials.
_ We have already acquired some experience in creating a coumnon material base for
- molecular-biological research in our mutual support of the project on "Reverse
transcription (within the program of cooperation among the Academies of Sciences)
by the AMV virus, by the enzyme revertase, by lesoxyribonucleotide triphosphates,
by oligo-inoculation (dT) and by special information molecules and in our mutual
support of the topic "Oncovirology and its molecular-biological aspects" (within
the framework of the CEMA program 1�HaYwful new formations) by providing fetal
calf serum, several pieces of equipment and cell and virus models.
It is also possible now to take some measures to develop a common supply of
special enzymes to ccopexating countries. Many of our institutes independently
and often with no exchange of information are isolating, for example, restrictase
6
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of various types and even of identical types. In particular, our institute has
isolated nine restrictases, seven of these independently and two in cooperation
with the Hungarian Academy of Sc iences. An inventory of all restrictases isolated
in the course of our work will probably show that we already have most of the
80 currently known different types of restrictases. This holds true for the isola-
tion of other enzymea of DNA recosnbination and in general for special enzyme&,
the production of which could be increased.
Analogous measures can also be taken in the area of cell models, including the
already existing original models of lymphocyte hybrids.
= In our opinion, the most purposeful and realistic way to ensure a reliable material
base for cooperative molecular-b iological studies is to gradually proceed from
the level of separate themes which appear to be most promising to the development
of a common project on their basis. Such theu:es, as part of our program of multi-
lateral cooperation, should be identified and analyzed with a view to establishing
concrete tasks for the pa:ticipa.ting countries in the development of these themes.
Some effective coordination of the research on these themes should also be found.
The coordinating programs will have to change their function from simply registering
work to a level of actual coordination. The commission "Molecular Biology" should
know the technological requirements of individual themes as well as the potential
of the pari-icipating countries, in other words, the xeserves in the area which can
be mobilized for the mutual supplying of project requirements. A supply service
_ could be organized at each base laboratory. This commission should also have a
general idea of all molecular-biological research and in particular a rating system
as to its relative importance.
Molecular biological research is by its nature already suited to integration.
Qn the other hand, integration in this area is for us the most effective way
of ensuring the successful continued development of molecular-binlogy. The
guarantee of i ts continued prograss is a real task of the Academy of Sciences of
_ the socialist countries. COFYRIGHT: Izdatel'stvo "Nauka", "Vestnik Akademii nauk SSSR", 1980
[23-9233]
9233
- CSO: 1840
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UDC 591.524,1:591.177
SOME QUESTIONS OF THE THEORY OF SWIMMING OF FISHES AND DOLPHINS
- t4oscow DOKLADY AKADEMII NAUK SSSR in Russian Vol 253, No S, 1980 pp 1082-1085
[Article by Ye. V. Romanenko, Institute of Evolutionary Morphology and
Ecology of Animals imeni A. K. Severtsov, USSR Academy of Sciences]
[Text] In recent years there have been several attempts to creaCe a mathe-
matical model of the swimming of fishes and dolphins. However, mainly
two dimensional problems have been examined [1,2], and this has permitted
only qualitatively examining the mechanisms of swimming and effects con-
nected with that process.
One of the most successflzl solutions of the spatial problem of the mechanism
of swimm:ing of fishes was given by Acadesni.cian of the Ukrainian Academy of
Sciences G. V. I,ogvinovich [3]. He examined the motion of a slender body in
the inertial coordinate system x,y,z, which moves in an unlimi.ted liquid
medium in the direction of the Ox axis. Figure 1 presents a diagram which
clarifies the formulation of the problem. The diagram has been taken from
the work of G. V. Logvinrnrich, as have aLl the symbols: the abscissas of the
ends of the body xl and x2 and, consequently, the length of tbe body LP =
xl - x2, R(x) is the semi-major axis of an elliptical cross sect-Lon of the
body, the value of dR/dx is small along the entire length of the body, and S
--the longitudina,l curvi.linear axis of the body--deviates little from the
- axis of abscissas. Put as the basis of the theory is the concept of the
"permeable layer," accord.ing to which the body, i.n passing through a certain -
"permeable iayer," which is iffinobile in relation to a quiescent liquid, gen-
erates in it an alinost plane cross.evrrent, similar to the flow of an ideal
liquid. A flux of pulseG m~Vv flaws in that case from the tail fin of the
slender body under consider2ti$n along the tangent to it. Here ~ is the
virtual mass, eque1 to -r~PR (~cl), P is the density of the liquid,~ V is the
velocity of the body, and v is the layer velocity normal to the curvilinear
axis of the body, determineR with the expression
_ an an .
- a - v aX
s
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~ In such a formulation oP the problem for the case of periodic change of
= swimming movements of a body G. V. Logvinovich obtained the following gen-
eral expressions for the thrust caused by encounters of pulses, suction
and kinetic energy remaining in the wake per uni.t of path of the tail fin: 77 (1) 1=m'(x)V(al - V ar)8x at x=xl,
(Z) P=-2 xI1. . ~d_ dx Xat - vax 'dx,
l- ~
2
-
(3) E_'n 2 x) (Li? a_ v aX ) at X= X, .
Expression (1) is valid on1y for fishes sw-im*+iz1g by the "eel" method, when
the amplitude of the wavy oscillations of the body is constant at all its
points. A very general expression for the t2lrust, one without the indicated
limitations, also was obtained by Academician G. V. Logvinovich [4] and has
the form:
x' ~ � [M*W(at V ~-dx.
(a) r = I (aX ~ar ax
xt
S.
z Figure 1. Diagram explaining
Tne formulation of the problem.
i
~
N
.xt
The forimalas obtained by G. V. Logvinovich permit estimating the hydro-
- dynamic characteristics of fishes on the basis of experimentaly measured;
kinematic characteristics. Academician G. V. I,ogvinovich applied that ,
theory to describe onl,y one particular case of the swimm.ing of fishes by
the "eel" method. In reality not one fish species swims tn that way, a1-
thoughin some species, eels i.n particular, the anrplitude of oscillations
of the head in the process of swimming amounts to about 30-40 percent of
the amplitude of tail oscillations. In other fish species, however, and
also in dolphins, it is even less than that.
~ 9
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There was an attempt to apply G. V. Logvinovich's theory to describe the
swimmi.ng of fishes by the "scambroid" method on the assumption of a linear
law of increase of amplitude of a locomotur wave duri_ng its propagation
from head to tail along the fish body [5]
x1 - x rCt x1- x 1
(S) n-n0 LD s`"VL L f' where C= constant.
It can readily be shown, hawever, that representation of the shape of os-
cillations in such a form is incorrect, as it leads to a negative value of
the thrust under that cont:ition V= C. Here C is the phase wave velocity.
To prove that statement, let us represent the shape of oscillations of a
- fish bod,y in the form
- (6) ri _ 17a(x)sin [cjt - k(xz - x)] , where k = constant.
If we use expressions (2), (4) and (6) we obtain the sum of oscillations
of the thrust and suction averaged during the oscillation period for V= C
z
(7) I I}+ {P} m'(xi) ano s
_ /
` az) ,
x=xi
We will consider in such case that at x= x2, m* = 0.
From the noted fact follows the inevitable conclusion that in formula (6)
the value of k must be a function of the coordinate, and since k= c,/C,
- then the loconotor wave phase velocity C must be a flinction of the coordin-
ate. Then the law of fish body deformation in the process of swinmi.ng must
have the following form
8) n-1lo(x)sin wt - w (C~xz X) - z)
I (
The form of the wave phase veloci ty- coordinate relation is difficult to
- predict, and so we will assume that it is approximately linear. As for the
shape of Lhe function jo(x), if one starts from obvious experimental data
- which are reduced to the fact that the locomotor wave amplitude of a fish
or dolphin increases monotonica]1y from head to tail, it can be approxima.ted
by an exponent with a minimum number of undetermined parameters. Then the
lakT of fish or dolphin boci,y deformation in the process of Syrimming can re-
presented definitiv�ly in the form
(9) n= n i(Kr - 1+ exp ~OAx z- x)/LP)"] sin t- xz - x
Cr[1 +b x= -x)]
Here -I1 is the amplitude of tail oscillations, Kr is the ratio o.f i;he ampli-
tudes of head and tail oscillations, G1 is the circular frequency, 't. is the
time, C is the value of the locomotor wave pYiase velocity in the region of
- the heaa. The values of ~1, Kr, ~ g b, w and Cr are unlazown and must be
- determined experimentally. The coefficient 3( can read.ily be expressed
th-rough K from the boundary condition. Actually, the amplitude of tail
oscillations at x- x must be equal to-11, that is, Kr - 1+ ed = l, whence
kTe have of = 1n ( 2-r).
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By using expression (9) and the evident condition
(10) jlJ+ jp} =0 at V=C,
' it would be possible to estimate the values of the most interesting para-
meters e arid b. However, the expression for the sum of the thrust and
suction proves to be exceptionally unwieldy and little suitable for numer-
ical calculations in that case. Still, such an estimate can be made if it
is recalled that the thrust is determined in accordance with expression (4)
by projecting on the Ox axis the pulse flawing from the tail fin in a unit
of time, and the pulse projection is determined in turn by projection on
the same axis of the liquid layer velocity normal to the sa.me axis
_ (a,1 anlan
- t i) Unx - ~ar - V arlax The equa_tity to zero of the thrust at V= C is equivalent to equality to
zero o'L expression (11). Therefore to estimate the values of Y and b we
will mak.e use of expression (11), equating it to zero under the conditions
= V= C and x= xl. Here we will make two additional assumptions: firstly,
_ we will make the estimate at a point corresponding to the ed.ge of the Ju-ail
(x = x1) That assumption evidently is tiTalid, since the thrust is accom-
pllshed with the tail. Secondly, we will neglect suction, which according
to the data of [3,5] constitutes a very small portion of the total thrust.
To sum up, if we use expressions (9) and (11), we obtain
e2 s_ t~= 1 f 1 l
(12) L: ry {r [1 +LPb ll 1+Lpb J'
- P
In the case of the swimming of dolphins, there is a clear connection between
the values of !v and V/L
~ P
~ (13) w= 27., (1,1 V/Lp +0,15).
~ In addition, observations of dolphins shaw that the value of Kr is about
0�25. If we take this and expression (13) into consideration,we obtain a
, correlating corli-iecting the values of Y and b with respect to bottlenose
dolphins ~ 6,41 0,15\
(14) 7= i{ LPb(l,t + Y~p ) LPb.
\ /
- The results of calculations according to formula u4) for the most typical
' values of the relative felocity of a dolphin are given in Table l.
' What sort of correlation between ~ and b exists in the case oP real swim-
ming of fishes and dolphins can be shovm only by careful experiments on
animals.
I sincerely thank Academician G. V. Logvinovich, Doctor of Physical and
Mathematical Sciences K. A. Naugol'nykh and S. G. Pushkin for their interest
in the work and valuable comments.
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Table 1
p, M'
ry at Y/LP-�1
~&t V/Lp-2
b,
T 8t '/Lp�1
I y&'t YjLp- 2
0.05
2,30
2,17
0,3
3,88
3.65
011�
2,99
2,81
0,4
3,98
. 3.74
0,2
3,62
3,40
1,0
5,64
5,31
.
- BIBLIOGRAPHY
1. Stecknann, J. ENG. ARCH, No 31, p 214 (1962).
2. Wu, T. Y.-T. FLUID MECHAATICS, Vol 45, Par t 21 p 337 (1971).
~ 3. Logvinovich, G. V. BIOiiIKA, Vol 4, p 5(1970
4. Logvinovich, G. V. Gidroclinamika techeniy so svobodnymi granitsami
(Hydrodynamics of Flows With Free Boundaries). Kiev, 1969.
5. Kozlov, L. F., and Oleynik, R. A. BIONIKA, Vol 12, p 3(1978).
. 6. Kinn, V. P., and Pyatetskiy, V. Ye. BI0IVIKA, Vol 11, p 36 (1977).
COPYRIGHT: Izdatel'stvo "iVauka", "Doklady Akademii nauk SSSR", 1980
[439-2174]
2174
cso: 1840
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3iOOK ON BIOLOGICAL BASES OF BEHAVIOR CONTROL OF FISH Iv FLOWING WATER
~ ;toscow BIOLOGICHESKIYi OSNOVY UPRAVLENIYA POVEDENIYEM RYB V POTOKE VODY
(Biological Bases of Behavioral Control of Fish in Plowing Water) in Russian 1980 signed to press ^3 Apr 79 pp 2, 3-10, 317-319
[Annotation, Introduction and Table of Contenta from book by Dmitriy
Sergeyevich Pavlov, 1,250 copies, 318 pages]
[Text] Patterns of fish behavior are examined on the basis of original ma-
terial: orientation, swimaning speed, behavior in a rheogradient, daumrivpr
, and spawning migrations, behavior in some hydroengineering atructures. Much
attention is devoted to the ecological study of rheoreaction. Basic stereo-
; types are revealed of behavior of pelagic and bottom fish in flaaing water.
~ The book is of interest for zoologists, ichthyologists, specialists engaged
, in research on animal behavior and also for hydraulic engineers and person-
; nel engaged in fishery.
Ir.troduction
The development of the theory and methods of control of biological processes
constitutes one of the chief directions of biological research. The problem
~ of behavior control of animals is an important part of it. It concerns one
of the most ancient problems of concem to maakind. Nanetheless thE scien-
tific bases of behavioral control are beg_`nning to be established only naw.
~ Their further development is of major importaace to the rational utiliza-
tion, reconstruction and conservation of the anirnal wrorld.
i
Behavior is one of the most complex forms of manifestation of animals' vital
~ activity. As A.N. Severtsov has pointed out (1922), it is of extreme
; biological importance, ensuring plasticity of animals in regard to rapid
environmental changes. For this reasaa~lit is quite logical for research on
animal behavior to be included on an increasingly large scale at the present
time in the general complex of ecological-evolutionary problems.
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Mobility of the physical medium of habitation (air and water, to a lesser
degree ground) is a factor acting on the duratian of all evolution of ani-
mal and vegetable organisms. In the animal world, this factor is of espe-
cially great importance for aqueous inhabitants, many of them spend their
entire life in flowing water. In regard to the water environment, animals
have developed an adaptatio?: pertaining to different aspects of their vital
activity. These adaptations are manifested both in a_nfmals' strlicture and
in their fuactions, including behavior. The special features of fish in
flowing water constitute an important side of their biology.
- The purpose of the given work lies in the development of the biological bases of behavioral control of fish in flawing water. Such a formulation
of the questions provides first of all for the study of the basic laws of
this behavior and then determinatian of the possibilities and methods of
control.
The pertinence of the sub ject of research is connected principally witlz one
of the most important problems of contemporaneity--the problem of "man and
tiie biosphere. Water bodies as one of the elemEnts of the biosphere are
- increasingly becoming subject to the influence of man's activity. As a re-
sult of the construction of dams and withdrawal of water for irrigation and
odler purposes, conditions of currents have sharply changed in many internal
water bodies. Conditions have correspocidingly changed for the manifestation
. of behavioral adap*_ations of fish connected with life in flowing water, par-
ticularly spawnitiQ and dawnriver mi.g.rations. Freservation of natural repro-
duction of many species of_ fish in these water bodies would be impossible
without the development of scientific bases -of behavioral control of fish
- in f lawing water and the practical implementation of the proper measures.
_ At the same time, the importance of investigation of the basic laws of be-
havior of fish in flawins water is determined by the necessity of under-
s tanding the role of behavior in the system of their adaptation first of
all to dwelling in a mobile environment. This promotes a deeper examina-
tion of the different aspects of the ecology of fish, especially their�
migrations.
The complexity and many-sidedness of the phenomenon of behavior is respon-
sible for the fact that work connected with its study is conducted by repre-
sentatives of different specialties (physiologists, zoologists, psycholo-
gists, mathematicians, bionicists (bioniki] and others) and along different
lines. The basic theses of the theory of behavior of animals were analyzed
and described by a nunber of authors (Promptov, 1940, 1956; Anokhin, 1945;
I. Pavlov, 1951; Voronin, 1957; Biryukov, 1958; Krushinskiy, 1960, 1972;
Slonim, 1965, 1967; Tinbergen, 1969; Shoven, 1972; Fabri, 1975; Khaynd,
1975; Lorenz, 1939; Tinbergen, 1955; and others). The objectives of our
work, cannected with benavioral study at the level of organisms and popula-
tions of a number of species, bring it close to the work of zoologists con-
ducted in regard to the ecological aspect. A comgarable direction of re-
search appeared in the second half of the 19th century, undergoing
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significant development in the first half of the 20th centuxy in the works
of zoologists and ecologists (Panov, 1975; Novikov, 1975).
At the present time, it is becoming practically iiupossible to examine such
traditional problems of ecology aF feeding, reproduction, defense and ecol-
og} of populations without an analysis of the beh,,vior of animals (Nikol'-
~ skiy, 1961; Naumov, 1963; rtakfed'ye*~, 1965; Odum, 1975; and others). A
number of researchsrs conside: the study of the behavior of animals in the
ecological aspect as one tasks of ethology (Fabri, 1975; Novi.kov, 1975;
Panov, 1975; and others). B.P. Manteyfel' (1970a, 1974) takes this divi-
sion _:1L science of behavior gnd proposes to call it "the ecology of animal
behavior" (p 13).
Researches on behavior in its ecological aspect are beinR developed increas-
ingly in contemporary ichthyology. These works are conducted inseparablv
from the study of reception of fish. A general characterization
of works on this plane has been given by B.P. Mantayfel' (1967, 1970b).
Detailed information on individual reactions and receptor systmes of fish
_ (school reactions, lip,ht reaction, hydrostatic reactians, optomotor reac- -
tion, swimming speeds, learning, migrations, vision, hearing, organs of the
~ lateral line [bokovaya liniya], . chemical reception, electrical reception _
and others) can be found in suffiarizing works of Russian and foreign authors
(Baburina, 1972; Barannikova, 1975; Girsa, 1970; Disler, 1960; Disler, Smir-
nov, 1974; Zusser, 1971; I.eshcheva, 1974; Malyukina, 1955; Malyukina et al.,
1969, 1974; Pavlov, 1970a; Pavlov, Saburenkov, 1974; Podlubnyy, 1971; Prazd-
- nikova, 1970; Protasov, 1965, 1968a, b, 1972; Pushkov, 1954; Radakov, 1972; Flerov, 1962; Tsvetkov, 1969, 1974; Blaxter, 1969, Braan, 19',7; Dijkgraaf, _
1933, 1962; Harden Jones, 1968; Hasler, 1966; Hoar, 1951, 1953; Fage, Fon-
taine, 1958; Teichmann, 1962; arid others). It may be said with confidence
- that 'oehavioral research occupies an increasingly important place in fish
ecology. But, despite the long history of such research, the behavior of _
fish in flowing water up to the present time has hardly been subjected to
special study. At the same time, it is impossible to understand many of -
the features of fish ecology in flowing water bodies: migrations, distrib-
ution, feeding, reproduction, defense against enemies, daily and seasonal
rhythms, dynamics of populations and so an. We therefore hope that the
study of the basic laws of fish behavior in flowing water, representirg a
new direction in the sCudy of fish ecology, will contribute to a deeper
understanding of different aspects of their life.
- As shawn schematically (Figure 1), the problem of fish behavior in flowing
water is cannected not only with ecology but also with other scientific
disciplines: physiology, especially with the study of orientatior and
locomotion in flowing water; morphology (for example, with the study of the
role of the bodily form of fish in maintenance in flowing water) ; zoo-
" geography in connectian with the dispersion of fish and varying composition
of ichthyocenoses depending on the speed of the current in the water body;
- with hydraulics and hydroeng,ineering. This problem finds a broad outlet
in fishery practice.
Turning to the concrete content of the work, it should be emphasized that
a basic behavioral reaction of fish connected with dwelling in flowing,
~ 15
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water, as in the case of many other aquatic animals, is rheoreactions. Tt is of an inborn character and consists of the fact that animals in flaaing -
water move as a rule against the current. Such movement keens them from be-
ing swept daan by the current. All oCher features of behavior in flawir.g -
- water develop against the background of this specific reactions. For this
reason rheoreactirnY does not just have a specific meaning but actually is
- the chief reaction of the organism in flawing water, reflecting the essence
of behavioral adaptations to flawing water. It should be noted in this
connection that the basic theoretical thesis of this work are principally
~
based an actual material characterizing this reaction. We examine the rheo-
reaction systematically beginning with composite elements and elementary
forms of the manifestatian of its significance in such complex forms of be- ~
havior as migrations.
The rheoreactian is displayed in representatives of different classes o':
aquatic animals, beginning with the infusoria and ending with amphibians
- (Stahl, 1884; Dewitz, 1899; VeYworn, 1899; Lyon, 1904; Prosser, Bratm, 1967;
� Pavlov, 1972; Arnold, 1974; and others). The innate character of the rheo-
- reaction in fish, as in other aquatic a:r,imals, was emphnsized by E. Lyon
(1904), who called it rheotropism. Subsequently, a number of authors have
more frequently designated this reactian as rheotaxis (Fraenkel, Gunn, 1940,
1961; Puchkov, 1954; Harden Jones, 1968; Shoven, 1972; and others). But
the occurrence of this reaction of fish to a current as pointed out by
Arnold (1974) exceeds the meaninR of such narraa terms as tropism and tax-
is constitutes a significantly broader biological phenomenon. The theory
of tropisms and taxes on the whole possesses a mechanical charactex and
does not disclose the biological significance of the behavioral reactions
of an organism, putting taxes �Lnto the rank of causes of the phenomenon.
Criticism of use of the theory of constrained movements was given in a
monograph by R. Shoven (1972) and, with respect to fish, in the works of
S.G. Zusser (1953, 1971) and B.P. Manteyfel' (1959, 1960, 1961). In the
- belief that the behavior of fish in flawing water should be considered as a
' complex behavioral reaction, we proposed (Pavlov, 1966a, 1970a) to desig--
nate it as a reaction to a current--a rheoreaction.
The number of special works dealing with the rheoreactian and behavior of
fish in flowing water is not great (Lyon, 1904; Schiemenz, 1927; Dijlcgraaf,
1933; Gray, 1937; Martinsen, 1937, 1940; Aslanova, 1952; Harden Jones,
1963a-c, 1968; Pavlov, 1965, 1966a-c; 1970a, b; Arnold, 1969-1974). Many _
auttiors did not adhere to any sort of plan providing for the systematic
study of the problem of f ish behavior in flowing water in general but stud-
ied only its individual features. A part of the information is scattered -
in different works bearing no direct relation to this theme of research.
Gnly three works are of a survey character; in them, a number of aspects of
fish behavior are examined rather fully (Harden Jones, 1968; Pavlov, 1970a;
' Arnold, 1974).
In the manifestation of the rlieoreaction, two aspects of the phenomenon
stiould be differentiated: orientation and movement (locomotion in f lawing
water. These two components correspond to the participation of Cwo systems
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� .
_ �
Morphology E Ecology ~~f-- Physiolopy
. . .
Behavior ~
4---- -
,

= ; - _ . . j
;
Behavior of Fish in Flow-
~
Hydraulics ing Water
.fiydroengineerin9; , Basic Laws and Possibil- Zoogeography ~
ities of Control t i
~ _ .
I
, .
, ,
~ Protection of Fish Stocks , Conmiercial ~
in Connection with Hydro- Fishing
engineering Construction ~
. ,
~ .
� ~
Preservation of :Iatural Artificial ~
Reproduction Reproduction ,
' i
~

OO
f0~+ N 4-4 00 ~l rl v; O}+ r , r C!
r~l 44 !4W 141 + p ~aD 4~-I ~
m G co ~ O Ft+ W m >
co u)' w N cn 0 ,-1 01
0.$4 G~+ O 0 W.' i Gr4 -rl
0 c0 Rt
W 41 iJ 41 W qvi
m i+ + O 11 U ~ bL O O W
w ~ C 3 14 ~ ti-G1 CG7 ~ ~ ~ oG 9 1 ~ ,~i ~ V
1+ ' O u .G U 9 p ~ ~ U r fs+ ; '
W JJ ' ti4 OL' Cn !A tA O M -rl
o p ,4 41 3
u,~ r-r es, ra a d : w
a~ aJ .o cn ~ G~ ' a! > z � m
i0~+ G 4-1 ~ 0. { V ! G O 44 ' (T4 '
~ I I
Figure 1. Principal Connections of the Problem of Fish Behavior in Flawing
Water to Different Branches of Science and Practice.
- 17
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y other
of the organisrLS--receptor and effector--in its realization as in man
behavioral reactions. The researchers when examinina t11e rheoreaction Aaid
the most attention to the investigation of inechanisms of o~~~cetofnE. Lyon's
the same time, it is necessary to emphasize the basic imp _
work (1904) ; ne showed that the orientatidn of fish in flowing caater occurs
on tiie bases of stationary reference points, -basically with the aid of
vision and touch. Ecological analysis of the movement of fish in flawing
water was carried out to a significantly lesser extent; the same was true
_ of analysis of the dynamics of behavior in connection with the influence of
various abiotic and biotic factors. The question of fish behavior under
rheogradient practically anduare
of behavior in flaw g tJ
- reflected in pertinent chapters of the work.
- The rheoreaction, compensati-ag for drifting of the animals caused by the
current, cantributes to preservation of the regiontrcohabitati. nnectednot At
the same time, it is known that the lives of many fish are
only to the retention of a specific region of habitation but also to the ,
use of favorab le conditions throughout the entire area of the species. This
occurs because of fish migrations in a wa~a~i~S~ WiHarden Jonesn(1968) ,
against it (denatant and contranatant mig
summarizing a tremendous b ody of material on fish migrations, showed that, -
as a rule, migration routes are determined by the character of currents in
- a water b ody. Therefore, when speaking of the fundamental laws of fish
behavior in flawing water, it is necessary to pay due attention to such a
complex plienomenon in their life as migration. It would be no exaggeration
to say that the creation of the theory of animal tnigrations is one of the
most important tasks of modern biology. This work examines first and fore-
most the special features of the migrational beh avior of fish connected
with motility of water, particularly the role of the rheoreactionWa e=.re-
action determining the specific character of beh avkar~yye,l~g~ations of _
Special attention has been given to dawn river (poy oung fish whose exd.stence it would appear at first glance is in seeming
On the whole research
contradiction to the existPnce of the rheoreacti~he eriod of their migra-
on the behavior of fish in flowing water during la~e to the adaptation of
tions makes it possible to understand what takes p
fish to their habitat under the concrete conditions of the water body.
We thus consider that the fundamental laaas of fish behavior in flowing
water are first of all the laws of grational
ofhthisr�
These features of fish ecology in our vie unde
rstand phenomenon, the study of possible the mobility oflthef
behavior in the system of adapt ons
water environment and creates a biological b asis for p rediction and control
of fish behavior in flowing water.
- Until recently hydraulic engineers were more interested than biologists in
the problem of fish behavior in flowing water. 'Ihis was connected wi~thinhe over need of creating devices for allowing yish ~SP ~d for protecti g fish g
structures [ryb opropusknyye sooruzheni a]-- )
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- against being caught in structures wittiZrai-ing watcr from a water bodyl
(fish-protecting devices [rybozashchitnyye ustroystva]--RZU). The employ-
ment of these devices has a long history (the RPS--300 years, the RZU-almost
60 years). But their development proceeded mostly empirically, by ttie methcd
of trial and error (Clay, 1961; Bums, 1966; Bol'snov, 1967; Malevanchik,
1973a). A large number of structures was created, but for the great major-
ity of them no bi,ological bases exist for the possibilities of their use
and conditions of work. E1s a result, for example, of 18 RPS built in our
country, only six are operating; at the same timey it has been planned to
erect something li'm-e 20 different structures (Malevanchik 1973b). The
problem of the RZU also demands solution. More youn g are destroyed in
water-gatheri.ng structures than are graan by all of the country's fish-
breeding er.terprises. Such a situation attests to the pressin g need of
developinf; biological bases for the use of these structures; first of all
an elucid