JPRS ID: 10527 USSR REPORT BIOMEDICAL AND BEHAVIORAL SCIENCES

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JPRS L/10527
18 May 1982
- ~JSS~ Re ort
p
- BIOMEDICAL AND BEHAVIORAL SCIENCES
CFOUO 3/82)
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JPRS L/10527
18 May 1982
USSR REPQRT
LIFE $CIENCES .
. LIOMEDICAL AND BEHAVIORAL SCIENCES
(~ouo 3/ 8 2
CONTENTS
BIONICS
Bioacoustics of Insects 1
Method for Telemetric Recording of Blood Pressure and
Respiration of Freelp Swimming Dolphins 12
~BIOTECHNOLOGY
- Kovats Indexes Used To Identify Sex Pheramones...........o...... 17
Seismosen~orY System and Classificat3on of Cottidae ~ish
_ ~Myoxocephalinae, Artediellinae)�������������~��������~~������ 23
ENVIRONIlrIENT
Determination and Quantitative Evaluation of Mutagenic Factors .
of Anthropogenicallp Varying Habitat Using Bacterial Test
Systems 31
90Sr and 137Cs Accumulation by Some Lower Plants in Vicinity
~ of Beloyarskaya.Atomic Power Plant in rhe Urals 43
Collaboration of U.S. and Soviet Scientists in Legal
Protection of the En~viro~ent 47
- MEDICINE
Low~-Frequency Acoustic Vibrations in Industry 56
_ a_ [III - USSR - 21a S&T FOUO]
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HUMAN FACTORS
Analysis and Optimization of Contrnl Systems for Manned
Aircraft 60
PSYCHIATRY
Joint Symposium on Biological Psychiatry of USSR Academy of
Medical Sc~.ences and Swedish Academy of Sciences 66
Conference of Neuropathologists and Psychiatrists in Perm'...... 69
b
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BIONICS
UDC: 591.582.591.185.5:595.7
BIOACOUSTICS OF INSECTS
Moscow BIOAKUST~KA NASEKOMYKH in Russian 1981 (signed to press 14 May 81) pp 2-8,
217-226, 256
[Introduction, article, conclusion and table of contents from book "Bioacoustics
of Insects by Rustem Devletovich Zhantiyev, Department of Entomology, Moscow
State University, Izdate.l'stvo Moskovskogo universiteta, 1800 copies, 256 pages]
[Text] This monograph submits the results of studying acoustic communicatian
systems of insects. It describes the structure and functions of�acoustical organs,
physical characteristics and adaptive properties of acoustical signals. The
latest data are submitted on the structure and functions of hearing organs and
mechanisms of processing acoustical information in the central nervous system of
insects. Much attention is given to the problem of identifying acoustical signals
and localizing the source of sound, There is discussion of the possibility of
u sing ~:ounds to control insect behayior. It is i~ntended for workers, graduate
and unde.rgraduate students specializing in entomology, physio~logy, biophysics and
bionics. References 445; figures 88, tables 12, appendix 6 tables.
Introduction
Bioacoustics is one of the new directions of research developing on the boundary
of biological and physical sciences. It is usually defined as the science of
acoustical signaling and orientation of animals. Both of these phenometi3 play
an extremely important and diverse role in vital functions of most animals; for
this reason, a wide spectrum of acoustical, mathematical and biological methods
is used to study them. The results of bioacoustical studies are of first and
foremost importance to the solution of many problems confronting such biological
disciplines as systematics, ecology, ethologys neurophysiology and genetics.
Some of the advances in bioacoustics are used for bionic studies and solving
applied problems related primarily to the control of behavior of animals that
are of economic importance.
The conditions for inception of bioacoustics existed long ago, since studies in
- this direction were pursued for many decades; however,.the First Bioacoustics
Congress, which convened in the United States in 1956, is generally considered
the official beginning of its existence. ~
Of course, one can view insect bioacoustics as a part or section of general bio- .
acoustics. The goals and tasks of this direction are largely determined by the
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specifics of the insect class. In assessing it as the object of bioacoustic
studies, one must take into consideration three circumstances. In the first
. place, insects do not use sound for active ranging (echoranging), while hetero-
specif ic communication did not obtain significant development in them, so that
researchers can concentrate mainly on the study of intraspecific communication
systems. In the second place, these systems appear numerous times in insects
and independently in different taxons; they are notable for remarkable diver-
sity and, in spite of the conceptions that prevailed until recently, they are
extremely widespread. In the third place, insects are the only group of
- terrestrial invertebrates in whom sonic signaling underwent rather significant
development, and since its inception is apparently referable te the Permian
period (Sharov, 1968), it can be assumed that acoustical communicatibn systems
reached considerable refinement in the course of prolonged evolution.
On the whole, this form of communication plays a very large part in vital func-
- tions of many insec*_s; it is instrumental in encounters between specimens of
the opposite sex, maintaining reproductive isolation of close species, regula-
tion of intrapopulation and intrafamily relations. In addition, some insects
- use sounds as protection against enemies and for detection of prey.
All of this circumstances determine the importance of bioacoustic studies and
their intensive development in the last few decades. In order to gain an
idea about the theoretical and practical implications of such work, it is
enough to list the entomological and general biological problems, in work on
which bioacoustical data play a considerable and, sometimes, deciding role.
In this regard, it must be noted, first of all, that in the course of acoustical
communication and orientation ins~cts solve problems that are as difficult as
those of vertebrate animals, but they do this on a diff erent morphofunctional
basis. For this reason, the results of studying their acoustical systems con-
stitute necessary and interesting material for comparative studies in the
fields of morphology, physiology, ethology and ecology of animals.
The importance and place of bioacoustics in entomology are attributable chiefly
to the fact that acoustic signaling is directly related to insect reproduction
and intrapopulation relations. This explains the heightened interest of ecolo-
gists and ethologists in acoustic studies. At the present time, it is growing
increasingly obvious that the most important patterns of distribution of
insects in biotopes, regulation of population density, dynamics of population
. size, migration, etc., could not be studied without consideration of acoustical
communicatinn systems. Knowledge of the "language," by means of which many
insects communicate during contact or at considerable distances from one
another, makes it possiblr_ to adopt a different approach to the study of their
behavior during reproduction, competitive relations, establishment of rank
(ranking), protection against predators, spatial orientation and other etholo-
gical problems.
Animal bioacoustics is equally important to the study of inechanisms of species
formation, particularly in those for whom sonic signals serve as the main
isolating factor. Species-specificity and stability of sonic signals makes it
possible to use them as reliable taxonomic characters, which are needed to solve
such complex problems as differentiation between twin-species, determination of
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,
species statss, degrEe of similarity of close taxons, etc. For this reason,
some characteristics of sounds have found a firm foothold in systematics
concerned with the study of insects wirh well-developed communication systems
(Orthoptera, Cicadidae and others).
The vast area of insect bioacoustics related to the study of processes of
receiving and processing acoustical ir~Pormation is, in essence, a part of phy-
siology of sensory systems. For this reason, the study of auditory analyzers of
insects helps disclose mechanisms that permit identification of conspecific
signals and orientation in relation to the s~und source, and it contributes
many new elements to conceptions of rhythmic processes in the central nervous
system, integrative functions of some parts of the brain, interactions of
different analyzers, etc.
Investigation of acoustical communication systems of insects is of both theo-
retical and practical importance. Knowledge of the principles of organization
of these systems enables us, on the one hand, to use the discovered patterns
for development and refinement of technical systems (primarily acousticaY, diraction
finding, radio measuring) and, on the other h2nd, to gain information about
vital functions of harmful insects or actively influence their behavior. As a
result of these studies, devices and equipmer.t could be developed for the de-
tection, estimation of number, diagnosis and monitoring the physiological state
of pests, as well as acoustic repellents and attractants.
There are very many articles and several surveys dealing with descriptions of
_ various aspects of organization and function of differerit elements of acoustical
communication systems of insects (Busnel, 1955,~ed.; Haskell, 1961; Busnel,
1963, ed.; Alexander, ?967; Bennet-Clark, 1975a; Elsner, Huber, 1973; Michelsen,
- 1974; Michelsen, Nocke, 1974; Elsner, Popov, 1978, and othe~s). No comprehensive
description of these data could be offered in this book, nor was it included in
our objective. At the same time, analysis of literature sources shows that most
works dealing with bioacoustics converge on two~main problems--recognition and
localization. The reason for the increased interest of many researchers in
these problems is tha~ insects solve essentially two main problems in the
course of acoustical aommunication and orientation: they recognize conspecif ic
communication signals and determine the spatial position of the sound source.
For this reason, we have tried to discuss in this book primarily material tnat
enables us to come as close as possible to understanding the mechanisms involved
in recognition and localization of sonic signals. The second and third secL
tions are concerned with these problems. Section 1 describes the main components
of acoustical communication systems. Data on some of them are submitted in a
generalized form, but many aspects of their organization are discussed in greater
detail* in subsequent chapters.
This book was based on the results of 15 years of studies pursued by this author
in the department of entomology, Biology Faculty of Moscow State University.
*We virtually fail to touch upon audio signaling of social insects in this book,
since a summary of the literature on this subject is giv~en in the monograph of
Ye. K. Yes'kov (1979).
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This author expresses his profound gratitude for valuable advice and assistance
to Prof G. A. Mazokhir~ Porshnyakov, Prof N. P. Naumov, Prof S. N. Rzhevkin, N. N.
Dubrovin, I. N. Kalinkina, 0. S. Korsunovskaya, A. G. Chernyy and V. S. Chukanov,
and expresses his appreciation to A. V. Popov, Prof F. Huber, Prof K. Kalmring,
Dr. J. Rheinlaender, Dr. H.-G. Rehbein and Dr M. J. Samways who kindly furnished
_ drawings from their works.
Applied Problems of Insect Bioacoustics
At the present time, it is difficult to de:.~ermine when the idea of using acoustic
vibrations to control harmful insects appeared. The first attempts to use
acoustical methods to solve applied sntomological problems yielded encouraging
results. Thus, as far back as 1935, an instrument was designed in Ge~cmany to
determine latent infestation of wood by the house longhorn beetle (Schwarz et al.,
_ 1935); ir~ Canada, a device was developed and successfully tested for chasing
the beet webworm away from crops (Belton, Kempster, 1962); acoustic mosquito
traps were tested in Cuba (Kahn, Offenhauser, 1949), the "apidictor," an instru-
ment for controlling bee swarming, was developed in England (Woods, 1959), etc.
- However, in the years that followed there was no further development of acoustical
methods and there were virtually no practical applications thereo�.
As we analyze the causes of diminished attention to such studies, we must men-
tion three circumstances: in the first place, this period was characterized by
excessive enthusiasm for chemical methods of control; in the second place, acous-
tical methods seemed to be unprofitable at that time, since they required the
use of cumbersome and expensive electronic equipment and, in the third place,
applied bioaccustics did not yet have the proper theoretical base.
By the end of th~ 1960's, the situation in this area of research began to changa
rapidly. The search for new methods of controlling pests that would not pollute
the environment and appearance of relatively cheap and compact instruments
again raised the question of developing acoustical methods and introducing them
into agricultural practice. At the same time, the results of unceasing experi-
mental studies revealed that insecrs have more complicated and refined acoustical
communication systems t~at was previously assumed. For this reason, it became
apparent to most rese.archers that applied problems can only be solved on the
basis of comprehensive analysis of the principles of organization of these sys-
tems. A new approach to solving this problem has yielded the first practical
results in the last few years, but apparently there is still a large amount of
work to be done before acoustical methods will take a worthy place in the .
arsenal of modern means of regulating the number of economically important
insect species.
What are the mc:st promising directions in this area of research? Experience in
studyitig acoustical communication systems shows, first of all, that acoustic
signals are used by insects to solvP many vital problems. For this reason,
having learned to record and analyze these sounds, we can gain some very 3.mport-
ant information about the species of interest to us. At the very least, it is
possible even now to devalop devices for detection of pests, determYnation of
~ their species c~mpositio:i, number, level of activity and distribution in agro-
cenoses or reservoirs. It is desirable to gather all of this information by
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remote control, using diverse sensors, to process it on a computer and use it
to forecast or control pests.
~ Knowledge of the principles of organization of acoustic communication system's
of insects arms us with yet another effective tcol, the possibility of interven-
~ ing in the process of transmission of acoustic sig~als and thereby exert the
desired influen:e on insects or, in other words, controlling their behavior.
Devices can be created for this purpose that disrupt in some way or other the
normal communication and orientation of insects: those that attract them to
traps, scare them away from protected objects, create interference in the
communication channel, etc. Nor should one overlook the fact that sound and
high-power ultrasound have a depressing effect on ~nsects or lead to their
death. Thus, the acouGtical resources that we have at the present time enable
us to perform three main types of tasks: 1) to obtain and analyze information
about pests; 2) to control their behavior; 3) to exert a direct physical
effect on them.
Let us consider these questions in somewhat greater detail.
Obtaining Acoustical Information
Acoustical instruments could become irreplaceable tools for determining the
latent infestation by insects of all sorts of materials, stock and even plants.
Devices that are usually recommended for.this purpose include microphones or
vibration detectors, amplifiers and recording elements, as well as filters that
permit isolatian of the useful signal from noise. With such instruments one
can pick up both the communicating signals of insects and noise associated with
various manifestations of their vital functions.
A device of this type was first developed in Germany to determine infestation of
wood by the Xylotrupes bajulus longhorn beetle (Schwarz et al., 1935). Sub-
sequently similar instruments were developed in England~ the United States and
USSR. In 1953, an acoustical device was proposed for determining latent in-
festation of grain (Adams et al., 1953). This method was found to be quite
effective and gained practical use in several countries (Zakladnoy et al., 1970;
Zakladnoy, Snyurina, 1977).
Acoustical methods of detecting soil pests appear to be less promising. However,
the results of studies in this direction warrant the belief that such methods
cvuld be usec~, at least in experimental research (mainly under laboratory
- conditions).
If the sounds emitted by insects are loud enough, one dan determine from their
intensity not only the location of pests, but their number. True, to obtain
satisfactory results, one must adhere to certain conditions, in particular,
one should place the pickups at a specific distance from the object, take into
consideration the noise level, etc., but theoretically they do not i.mpose
appreciable limitations on use of the method. The use of acoustical methods
is desirable, not only to keep a record of number of invisible insects, but
in cases where other recording methods are ineffective for some reason or other.
Since the sounds of most insects can be well-distinguished by their time and/or
frequency parameters, it is theoretically possible to develop methods of
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. l'Vt~ Vt'l'I~IAL vvi. V1~L1
identifying the species of pests. One can use for this purpose both communica-
tion signals and sounds that are made when insects move or feed. In the former
case, tables or atlases and acoustic instruments can be develope3 even now that
permit identificaticn of Orthoptera and Cicadidae in the field. It is harder to
develop equipment to record the sounds of small and concealed species. The
efforts made in this direction do not always yield positive results. For example,
Wojcik (1968, 1969), who used very complicated equipment, was unable to record
the communication ~ignals of several pests of stored products, including
Cryptolestes pusillus, which has stridulation organs, and two species of Pyra-
loidea, Ephestia kuehniella and Cadra cautella. However, this doe s not mean
that these insects do not emit sounds, since precopulation signals have been
recorded in another borer species, Corcyra cephalonica (F. coro, verbal
communication). Experience in such studies indicates that, in addition to a
meticulous choice of acoustical equipment, one must have comprehensive knowledge
about the biology of the species under stu~iy.
~11 of these difficulties grow significantly in the study of noncommunicative
sounds, which have less marked species-specif ic features, but progress in the
field analysis of diverse industrial noises may facilitate substantially
the solution of this problem.
As noted nreviously, special acoustical equipment permits not only detection
and identification of insects, but obtaining information about various aspects
of their vital functions and, first of all, the nature of their activity. The
most elementary devices of this type, which include a microphone, amplif ier and
automatic recorders, are used with success to study the circadian rhythms of
insects (see, for example, Medioni, 1964). Analogous devices can be used in
- the field to gain information about changes in level of pest activity in the
course of a day or season. At the present time, there is an opportunity to
introduce acoustical methods to monitor insect cultures and, in part~cular, for
- automatic regulation of their upkeep conditions. The interest of beekeepers
in such control methods is not waning either. Many types of bee activity are
associated with certain sounds; for this reason, analysis of the acoustical
background in the hive yields important information about the condition of
the bee family or changes occurring in it. The data accumulated to date mak~
it possible to design and refine devices similar to the "apidictor" we mentioned
before (Woods, 1959; Schauff, 1975; Yes'kov, 1~79).
Control of Insect Behavior
Studies in this branch of insect bioacoustics are being pursued in three main
- directions: developmen*_ of acoustic attractants, repellents and interference in
communication systems (H. Frings, M. Frings, 1965).
A t t r a c t an t s: To attract insects to sound sources, it is expedient to use
calling, assembling signals or flight sounds, i.e., signals that have an
attractive Pffect on insects in their natural habitat. By reproducing such
sounds or models thereof one can virtually always attract or detect a certain
number of specimens of the desired insect species that are situated in the
vicinity. For example, Spooner (1968b) proposed the use of simulated response
signals of females to tLip male Phaneropterinae crickets, and in the above-
mentioned experiments with mosquitoes, it was possible to attract some males
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to female sounds (Kahn, Offenhauser, 1949). However, these methods must h4ve
immeasurable greater efficacy for them to gain practical use. Efforts made at
insect attraction on a mass scale to sound have usually failed. In our opinion,
the causes of these failures are attributable to the following factors.
In the f irst place, it is not always possible to use adequate stimuli to attract
insects. For example, the sound of female mosquitoes in flight recorded on tape
do not attract males if two conditions are not met: the intensity of the sound
exceeds 60-70 dB and its source (usually a loudspeaker) is too large (preventing
formation of spherical waves). In the second place, many insects have a settled
life style and do not migrate beyond a certain range. In the third place, insects
are distracted from acoustic traps by natural sound sources, i.e., sounds
emitted by specimens of the same species.
To eliminate the f irst of these obstacles, it is necessary to make a comprehen-
sive study of signals and acoustical environment. And, as we have seen, it is
sometimes not enough to know the frequency and time parameters of sounds, and
one has to take into consideration their intensity, emitter radiation pattern
and other parameters. At the same time, having information about the informative
elements of the signal, we should not necessarily strive to reproduc? it exactly.
As noted previously, time and, occasionally, frequency parameters of signals
are markedly dependent on temperature. For this reason, use of tape recordi~gs
to attract insects does not appear to be too promising. In our opinion, one
must use for this purpose special generators of model signals, which automatically
adjust their characteristics with change in temperature. The complexity of such
generators (and their cost) depend, of course, on the complexity of the si.mulated
signal. For this reason, in developing tlie scheme of an instrument, we should
provide for reproduction of only the si~nal elements that determine its attrac-
tion to insects.
As for the second of the above-mentioned difficulties, to eliminate it acoustical
attractants must be used during periods (which are sometimes quite lengthy)
when insects migrate. Mole crickets and many grasshopper species are a good
example in this respect. One can increase the number .of *_raps per unit area and
the power of emitters to attenuate the distracting influence of natural sound
sources. However, it is not always desirable to amplify the sound, since such
signals have a repellent effect on some insects. All of these statements can
be illustrated on the example of the acoustic traps we developed for mole
crickets.
As noted previously, the first experiments on attracting mole crickets of the
genus Scapteriscus to sounds were conducted in the United States (Ulagaraj,
Walker, 1973, 1975). In these experiments, tapes of model signals were used.
On the basis of the data we obtained from studies of the acoustical communica-
tion systems of mole crickets (Zhantiyev, Korsunovskaya, 1973), we developed
and constructed, together with V. S. Chukanov and D. A. Yusipov, an acoustical
system to attract and collect two mole cricket s~ec3.e~~ Gryllotalpa unispina and
G. gryllotalpa. The main element in this system is an electronic unit that
generates pulses corresponding to the calling signals of the male mole cricket
in time and frequency parameters. After amplification, these pulses are fed to
10-W dynamic loudspea.kers installed in cone-shaped plastic traps (1 m in diameter).
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A container with fixing liquid is attached to the tapered end of the trap. The
trap is buried in the gr~und to its top edge. As mentioned previously, the
pulse repetition frequency in insect sounds is usually a linear function of
ambient temperature. For this reason, a thermistor is connected in the generator
circuit that automatically regulates pulse repeti tion'frequency in accordance
with the results ~f prior acoustic readings.
The acoustic traps were tested for two seasons in Turkmenia.(Baba-Durmaz station).
Mole crickets (G. unispina) start to fly with onset of darkness and stop at 0200
hours. The ~nsects flew to the trap mainly from the leeward side and fell into
the funnel or desce nded atsome distance from the trap, traveling the remaining
distance on the 6round. Even under adverse weather conditions (relatively low
temperature, wind), up to SO mole crickets were caught in the trap in 1 h, and
95% were females.
The results of these tests revealed that raising sonic pressure beyond the
levels inherent in the natural calling signal had no repellent effect; on the
contrary, it increased the attraction of the sigr~al for mole crickets. For
this reason, by increasing the power of the emitter one can broaden appreciably
the area covered and improve the eff icacy of the traps. Analogous data were
obtained previously from experiments with two Scapteriscus species (Ulagara},
Walker, 1973, 1975). Moreover, we established in preliminary experiments that
the shape of the pulse envelope does not affect insect reactions. This enabled
us to subsequently use square-wave acoustic signals and simplify appreciably
the circuitry of the generator. ~
Re p e 11 e n t s: Development of acoustic repellents is one of the most promising
but little-elaborated directions of r.esearch. Theoretically, such repellents
could be used against all insects that ha~~e receptors for soinic vibrations or
oscillations. In order to elicit negative phototaxis, one can apparently use .
two types of sounds: biologically important signals (or an imitation thereof),
which have a repelling effect under natural conditions, and oscillations at
frequencies that elicit discomfort.
. The most vivid manifestation of negative phonotaxis is observed in nocturnal
Lepidoptera subject to attacks by bats. For this reason, it is not surprising
that in the f irst tests related to development of acoustic repellents ultra-
sounds simulating the echoranging signals of these animals were used (Belton,
Kempster, 1962). The results of field trials revealed that exposure of corn
fields to ultrasound (SO ms duration, 50 kHz frequency) lowered by 50% infesta-
tion thereof by the beet webworm. Analogous results were later obtained on
- cabbage and lettuce fields. Use of 20 kHz ultrasounds lowered plant infesta-
tion by Trichoplusia ni moths by 66% (Payne, Shorey, 1968). However, trials
of this method to control the Heliothis zea moth on corn and cotton did not
yield positive results (Agee, Webb, 1969a; Shorey et al., 1972). The authors
consider the cause to be that insects perhaps become accustomed to continuously
pulsating ultrasound, while large plants prevent propagation thereof. These
explanations do not appear quite eonvincing, since they are in contradiction
with the results of the above-~mentioned tests.
- In general,'experience with ultrasound to scare off moths and Pyraloidea has
~ shown that acoustic repellents could become an effective means of control of
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-
many pests of the order Lepidoptera. But to develop such methods, it is neces-
sary to conduct special studies of the life style and behavior of each species.
There are grounds to believe that it is expedient to use analogous methods for
protection of both crops and stored products.
In time, acoustic repellents may find applications not only in agriculture, but
in the control of blood-sucking Diptera. This question has been repeatedly dis-
~ussed in the scientific literature. Some foreign firms produce devices for
personal protection against mosquitoes, of the "mosquito repeller" type, which
consist of portable sound generators. However, tests conducted in our country
(Rasnitsyn et al., 1974) and abroad (Kutz, 1974) showed them to be entirely in-
effective. A search for sonic oscillations that elicit negative phonotaxis in
mosquitoes is being pursued in many laboratories. And, althaugh the desired
results have not yet been obtained, this does not mean that there are some
basic difficulties that prevent development of such repellents.
Eradication of Insects
Eradication of insects with sound is more a technical than a biological problem,
so that we shall not deal with it here. Let us merely mention that, according
to data obtained to date, ultrasound and sound of suff iciently high power have
a depressing eff ect on insects. For example, there are reports that sounds at
frequencies of 0.2-2.kHz inhibit development of Plodia interpunctella moths
(Kirkpatrick, Harein, 1965), while powerful ultrasound (0.2-4 W/cm2) has a
devastating eff ect on Drosophila pupae (Pay et al., 1978).
Apparently sources of sonic and ultrasonic oscillations will be used to destroy
insects in stocked or stored materials; however, it is unlikely that they will
be used in the field (H. Frings, M. Frings, 1965).
Conclusion
In assessing our knowledge on sonic communication and orientation of insects, it
must be noted that the results of intensive experimental research cenducted for
the last two decades had a deciding influence on forming current conceptions of
the principles of organization of their acoustical communication systems.
The greatest progress was made in the study of.. mechanisms of sound emission,
the auditory system and some forms of acoustic behavior. For this reason, we now
have a rather complete idea about the n~uromuscular system of acoustic organs,
acoustical properties of sound-emitting and sound-receiving structures, ultra-
structure of auditory receptors, morphofunctional organization of auditory organs
and physical characteristics of acoustic signals. Use of combined methods of
research made it possible to obtain important data on the functional distinctions
of ttie auditory system in the course of identification and localization of
communicative acoustic signals. However, in spite of the effor~s of many re-
searchers, we have rather limited information as yet concerning processing of
acoustic information in the central parts of the auditory analyzers, and we know
virtually r~othing about the conneetions between auditory interneurons and other
parts of the central nervous system. As for the neurophysiological mechanisms
involved in recognition of conspecific signals, the data available to us enable
us to develop only pre liminary hypotheses or conceptual models.
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e vn va ~ war~.. v~~i. i
ln view of these circumstances, we can be sure that most researchers will con-
centrate in the near future, as they did before, on tne problem of recognition
and certain other problems of basic importance. But we can also expect further
widening of the circle of objects studied and increase in number of comparative
bioacoustic studies that will help demonstrate the adaptive distinctions of
acoustical communication systems and the patterns of their inception and evolution.
~ The following main tasks will have to be done in the course of future neuro-
physiological studies: study of morphofunctional organization of acoustic centers,
i.e., neuronal nets implementing the function of diverse acoustic organs of in-
_ sects; completion of investigation of peripheral and central par~s of the audi-
tory analyzers; trace the functional connections between auditory interneurons
and associative centers and effector systems. The main results of these studies,
which will have been obtained in acute experiments, should then be checked in
chronic experiments on mobile insects. Perhaps only such combined studies will
enable us to form a definitive idea about the mechanisms of recognition of
- communicative signals and localization of sound sources.
In the near future, we should expect further development of Pthological studies,
including not only laboratory experiments to test functional hypotheses and
models, but observations in the field, which permit analysis of complex forms
of acoustic behavior of insects.
There is still quite a lot to be done with regard to investigation of acoustic
signals of insects, particularly their variability, transformation in the course
of species formation and adaptive distinctions.
~ Genetic studies aimed at determination of patterns of inheritance of species-
specific distinctions of various elements of acoustical communication systems
should yield much interesting and important information.
The results of all these investigations will not only help form a general con-
ception about the principles of organization of insects' acoustical communication
systems, but will serve as the basis for developing new and effective methods
of controlling the behavior of economically important species.
CONTENTS Page
Introduction 5
Part I. Acoustic Communication Systems in Insects 9
- Chapter 1. Sound-Producing Mechanisms 19
Chapter 2. Physical Characteristics of Acoustic Signals
Chapter 3. Auditory Organs 30
Structure of the auditory organs 31
Fine structure and functional character~istics of the auditory 36
receptors (chordotonal sensilla)
Functional organization of the auditory organs 41
Tympanal organs 41
Johnston's organs 61
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Chapter 4. Central Parts of the Auditory System 63
Auditory interneurons 63
Connectians between the aLditory system and e~fector systems 87
Descending neurons in the cervical connectives of crickets 88
Descending neurons in the abdominal nerve cord of orthopteran
insects 92
Chapter 5. Acoustic Behavior 94
Types of acoustic signals 94
Reactions of insects to acoustic signals 99
- Part II. Recognition of Acoustic Signals
Chapter 6. Comparative Analysis of Acoustic Signals of Sympatric
Species 106
Chapter 7. Reactions of Insects to Acoustic Signals and Their Models 121
- Informative parameters of acoustic signals 122
Noiszproof factor of acoustic communication systems 135
Chapter 8. Features of Functions of the Auditory System during
Recognition of Acoustic Signals 138
- Recognition of the intraspecific signals 139
Functions of the: auditory organs 139
Functions of th~~ central auditory neurons 142
Recognition of the conspecific calling songs 148
Fu~~ctions of the auditory organs 149
Functions of the central auditory neurons 151
~ Chapter 9. Mechanisms of Recognition of Acoustic Signals in Insects 156
Part III. Sound Source Localization
- Chapter 10. Adaptive Features of Acoustic Signals and Conditions
for Acoustic Orientation ~ . 166
Comparative analysis of acoustic signals 167
Experimental data 168
Chapter 11. Acoustic ~rientation 176
Acoustic orientation during flight 176
, Reactions of flying insects to terrestrial sound source 178
Acoustic orientation in terrestrial environments 180
Chapter 12. Directional Hearing 186
Functions of the auditory organs 186
Functions of the central auditory neurons 202
Applied Aspects of Insect Bioacoustics 217
References 22~
COPYRIGHT: Izdatel~stvo Moskovskogo universiteta, 1981
10,657 ~
CSO: 1840/121
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~
UDC: 612.14.08
METHUD FOR TELEMETRIC RECORDING OF BLOOD PRESSURE AND RESPIRATION OF FREELY
SWIMMING DOLPHINS
Leningrad FIZIOLOGICHESKIY ZHURNAL SSSR IMENI I. M. SECHENOVA in Russian Vol 67,
No 11, Nov 81 (manuscript received l~ Dec 78) pp 1744-1748
[Article by V. G. Dargol'ts, Ye. V. Romanenko,Ye. A. Yumatov and V. G. Yanov,
Laboratory of Morphology and Ecology of Higher Vertebrates (headed by V. Ye.
Sokolov); Laboratory of Bioacoustics (headed by Ye. V. Romanenko), Institute
of Evolutionary Morphology and Ecology of Aniunals imeni A. N. Severtsov, USSR
Academy of Sciences; Department of Normal Physiology (headed by K. V. Sudakov),
First Moscow Medical Institute imer.i I. M. Sechenov, Moscow]
(Text] At the present ti.me, various techniques have been developed to study
cardiovascular and respiratory functions of terre8trial animals, which permit
recording arterial pressure, the electrocardiogram, volumetric and linear blood
flow, pneumogram, etc., in different states of an organism, including un-
restrained animals [1, 2].
At the same time, there is also a need for such studies on aquatic animals,
for example, dolphins, when their behavior is unrestricted. In particular,
it is interesting to make a comparative analysis of cardiovascular and respi-
ratory functions during emotional reactions of animals situated in different
environments. A need a1:so arose to examine regulation of arterial pressure
and respiration in delphinids in connection with problems of hydrodynamics and
bioacoustics.
- However, there are a number of technical diff iculties involved in recording
these functions on freely swimming animals. It is necessary to develop a
sealed telemetry device for this purpose, with its own power pack that would
pick up useful information and transmit it from the aquatic environment to
receiving equipment. We describe here an instrument that meets these require-
men[s.
The functional flowchart of a radiotelemetry instrument, which we designed
and constructed to measure blood pressure, as well as air pressure in the res-
piratory system of the bottlenosed dolphin, swi.mming freely in a restricted
body of water (tank, marine pen), is illustrated in Figure 1.
An Elema-Schonander (EMT-34) type. ensor 1 served as the device that picked
up pressure directly, and its wor~ing chamber was connected to a blood vessel
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- or tested part of the dolphin's respiratory system by means of a catheter 1.5-2 ~n
in diameter. To r~easure blood pressure, the working chamber of the sensor and
catheter were first filled with saline, which transmitted blood pressure to the
sensitive element of the sensor. Output voltage which changes slowly is the
information signa~, whose amplitude is proportional to excess pressure, in
relati~n to atmospheric, from the symmetrical output of the sensor, and it was
fed to converter 2. In the converter, the ~Clock pulses coming from the clock
pulse generator with a repetition frequency Fn = 2 kHz 3, are submitted to pulse-
- amplitude modulation by the information signal. The same pulses divided into
four frequencies, 4, 5, served as the calibration signal. We determined the
necessary correlation between level of calibration signal and pulse-amplitude
modulated information signal by means of two dividers, R1-R3 and R2-Rs.
The overall signal then passed from adder 7 to the generator with frequency modu-
lator 8, where it modulated the carrier frequency which h~.d a frequency of
31 kHz in the "silent" mode. After the amplifier 9, the signal was emitted into
salt wa.ter. We used narrow-band f requency modulation with a modulation index of
m = 0.4.
Transmission of information about pressure coded in this manner was effected
via an underwater radio channel.
The generated power of the transmitter constitutied 50 mW on a resistance of 7 SZ,
whicti provided for a range of certain reception of up to 8-10 m. This range
makes it possible to receive a signal distinctly over the entire volume of the
pen or clased tank. The construction of the transmitting and receiving antennas
is described in [3].
And now a few words about the choice of coding method. In view of the fact that
a signal with slowly changing frequency has a rather small amplitude with actual
values of blood pressure and air pressure in the dolphin's respiratory system,
it was found expedient to use a subcarrier frequency (in the form of clock
pulses), amplitude modulated by the useful signal, and subsequently work with
this subcarrier frequency signal. The concurrent presence of a clock signal
makes it possible to form a calibration signal required for absolute readings
that is a multiple thereof in frequency.
Reception of radiotransmitter signals is effected by means of a specially developed
receiver of frequency-modulated signals, which consisted of two functional units
[block units]: the unit of selective amplifiers 10 (with central tuning
frequency f= 31 kHz and band of 2 L~f = 8 kHz) and demodulation unit 11. The
carrier. frequency was demodulated and overall signal, which consisted of the
pulse-amplitude modulated information signal at a frequency of 2 kHz containing
information about pressure and calibration signal of SOU Hz was isolated in the
receiver. This overall signal af ter the necessary div ision 12 was recorded on
one o� the channels of an Elektronika-100 type tape recorder 13, while the
explanatory text was recorded on another channel from microphone 14.
In the course of processing the recorded results, the overa].1 signal was repro-
duced on the same tape recorder and fed to tfao filters: band filter with central
frequency of 2 kHz and band of 2 ~f = 650 Hz 15 and a f ilter for lower frequencies
with cut-off frequency of 1020 Hz 16. The band filter served to isolate the
intermediate frequency of 2 kHz with pulse-amplitude modulated information sig-
nal, while the lower frequency f ilter served to isolate tl.~e calibration signal.
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u
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The coefficients of transmission of the band filter at a frequency of 2 kHz and
lower frequency filter at a frequency of 500 Hz equaled 1. Then, the information
' and calibration signals divided by frequency were recorded alternately on
paper of a Bruel and Kjxder 2305A xecorder. The scales of recorder levels were
calibrated at the beginning and end of each tracing with the signal from an
external type GZ-102 generator 18.
BO a
00
x ~o
~
so
~ ~
90 �s
b
00
x
~ ~
~
a
2 .
~ ~ ~
?.0 s
_ Figure 2. Samples of pressure tracings ~
a) blood pressure in vessels of tail fin
b) air pressure in main nasal passage during emission of echoranging series
1) pressure curve
2) echoranging time
If one knows the pressure to which corresponded the constant amplitude of the
calibration signal and graduation characteristics of the pressure sensor, one
could estimate absolute pressure in blood vessels or airway system of the
dolphin at each point in time.
The transmitting equipment 1 together with pressure sensor 2 were attached to
the dorsal fin of the dolphin. A flexible catheter, 1.5-2 mm in diameter,
filled with saline and heparin (to prevent blood clotting) was introduced into
one of the arteries of the dorsal or tail fin and connected to the pressure
sensor. Before taking each reading the catheter was flushed with the same
solution. Access to the artery was provided by cutting off a 2-3 cm end of
the fin. To measure air pressure in the respiratory tract of the dolphin, the
air-f.illed catheter was introduce d into the choanae or blowhole above. them and
attached with a suction cup near the entrance into the blowhole.
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Figure 2a is a sample of a tracing of blood pressure in vessels of the tail.
Blood pressure i~ vessels of the tail f in of a free-swimming dolphin is of
~ the order of 60-80 mm Hg. It should be noted that the dolphin did not submerge
to any depth during this time. Blood pressure in vessels of the dorsal fin is
1.5-2 ti.mes higher, and this is apparently related to the closeness of dorsal
fin vessels to great arteries, thei~ larger diameters, as compared to vessels
of the tail. In addition, it was possible to introduce the catheter deeper
into vessels of the dorsal fin than into the smaller vessels of the tail.
Air pressure was recorded in the region of the main nasal meatus (blowhole),
between the external sphincter and muscle plu g shutting entry into the choanae(P~ig-
ure 2b). Yressure curve 1 is`correlated to echoranging time 2. Typically,
the echoranging process is always associated by approximately 30-50 mm Hg in-
increase in pressure in the blowhole, whereas pressure in the choana does not
change.
- It should be noted that we set the pressure sensor to zero at the start of
each experiment. For this purpose, we equalized pressure in the working chamber
of the sensor with atmospheric pressure. The sensor was then adjusted fo~ a
minimal output signal in the absence of excess pressure in the catheter. The
remaining dysbalance signal at the output of the sensor was recorded on tape,
and its value served as the zero reference point.
This technique permits satisfactory reading of both blood pressure in blood
vessels and air pressure in the dolphin's respiratory tract when the animal
swims on the surface without submerging. Artefacts were found in the course
of recording blood pressure, which were related to suhmersion.of the animals,
and with measurement of pressure in the caudal artery with movement of the
flukes while swimming.
With submersion, the distance between the sensor and reading site has an
effect. This occurs because of the flexibility of the walls of the long cathe-
ter (1.0-1.5 m), which transmit water pressure and distort the true pressure
- of both blood and air in the respiratory tract. To rule out such artefacts,
we placed the sensor as close as possible to the measurement site on the dorsal
fin, thus also reducing the length of the catheter to a minimum.
BIBLIOGRAPHY
1. Tkachenko, B. I. (ed.), "Methods for Testing Circulation," Izd. "Nauka",
Leningrad, 1976, pp 1-278.
2. Skotselyas, Yu. G., Ivanova, L. I. and Yumatov, Ye. A., "Dynamic Recording
of Rat Arterial Pressure in Chronic Experiments and Witr? Unrestrained
Behavior," FIZIOL. ZH. SSSR, 63, 1, 1977, pp 154-155.
3. Yanov, V. G. and Romanenko, Ye. V., "Radiotelemetry System for Measuring
Hydrodynamic Parameters of Dolphins in a Confined Water Area," Izd.
"Bionika", Kiev, 6, 1972, pp 92-99.
COPYRIGHT: Izdatel'stvo "Nauka" Fiziologicheskiy zhurnal im. I. M. Sechenova,
1981
10,657
CSO: 1840/117
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BIOTECHNOLOGY
- UDC: 543.544:547.996.02
KOVATS INDEXES USED TO IDENTIFY SEX PHEROMONES
Tashkent KHIMIYA PRIRODNYKH SOYEDINENIY in Russian No 4, Jul-Aug 81 (manuscript
received 19 Jan 81) pp 501-505
[Article by S. F. Nedopekina, B. G. Kovalev and A. N. Kost'h, All-Union Scientific
Research Institute of Biological Methods of Plant Protection, Kishinev; Moscow
"Order of Lenin," "Order of the October Revolution" and "Order of Red Banner of
Labor" State University imeni M. V. Lomonosov]
[Text] In view of the development of new methods of protecting plants, which
are safe to the environment, there is special interest in substances of natural
origin--sex pheromones of insects--use of which is considered quite promising [1]. ~
Isolation and identification of the structure of natural pheromones involve a
number of difficulties, due chiefly to the small amounts of this substance con-
tained in insect glands (nanogram range). Hence the need to use highly sensitive
physical and physicochemical methods, as well as combinations thereof with chemical
~ identification.
The pheromones of Lepidoptera identified to date consist mainly of unsaturated
aliphatic alcohols, their acetates, aldehydes, esters and epoxides containing 10-18
atoms of carbon [2].
Identification of a natural pheromone involves determination of the length of
the molecule's carbon chain, quantity, position and configuration of dual bonds,
as well as the nature of other functional groups it contains.
Relative gaso-chromatographic retention level is often used to describe f unctional
~ groups of analyzed compounds [3J. Kovats [4] proposed the use of the difference
between indexes of compo~r.nd retention in two phases of different polarity ~I12 as
a characteristic parameter:
~l,~ _ /~-1.,, (1)
- where I1 and I2 are the retention index on stationary liquid phases 1 and 2, res-
pectively. ~I12 is the characteristic of the functional group; it depends on the
nature of the stationary phase and depends little on column temperature. Experi-
mental determination of the value of ~I12 makes it possible t~ relate the ana-
lyzed compound to a specific class, provided the values of ~I12 have been tabulated
for a large number of known substances.
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The retention index of a compound is the sum of the retention index of the carbon
shell and increment$ corresponding to functional groups. This makes it possible
to determine the retention index of the carbon shell, if tne contribution of the
functional group and length of the carbon chain are known, bearing in mind that
the contribution of the methylene group is 100 units.
Our objective here was to construct tables of retention indexes for unsaturated
aliphatic alcoho~s, their a.cetates and some epoxides. The practical usefulness
of the tabulated data is demonstrated on the example of identification of natural
pheromones of female cabbage (Mamestra brassicae) and tomato (Mamestra oleracia)
moths.
We used n-alkanes (C nH2n+2, where n= 12-17, 19, 20, 23, 24) as standards for
determination of I. We calculated I using the following equation [5]:
t.= l 00 �~t ~ n 2
lU0 (n ~ -~g 1"~
~ ' i lo i,~,-lg ' . ~ )
where nl and n2 are the number of atoms of carbon in the corresponding rralkanes,
Znl, Zy~Y and Zx are distances on the chromatogram from introduction of sample to
height of corresponding peak (in millimeters), and
l~~ ~ lX ~ ~n~.
The increments of functional groups for each type of phase were calculated as
follows:
. ~J,= J.~r~ -100�n,
~l~ _ (.re-6o-1 OO � !Z, .
where l~a~~, 1c`- is the retention index of the compound at the (3)
corresponding phase;
~t is the legnth of the carbon chain.
The difference between retention indexes of the compound on polar and nonpolar
phases was determined using the formula:
~I,' = I''~`-~o-InpL . . . ~4~
All of the obtained values are listed in Tables 1-4.
The tables show that use of the difference between retention indexes on polar
and nonpolar phases (~I12) makes it possible to identify rather accurately
ethylene alcohols (~I12 = 371-429 units), their acetates (DI12 = 286-345 units)
- and saturated epoxides (DI12 = 264-276 units). However, we see that there is
some overlapping of values of ~I12 (Table 4) for ethylene acetates (286-345 units)
and saturated alcohols (337-364 units), as well as saturated acetates (260-299
units) and epoxides (264-276 units). In this case, for exact identification one
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should use the increments of functional groups for each type of phase (QI1, pI2,
Table 4), which differ substantially.
Table 1. Retention indexes for ethylene acetates with the general formula
cis
R-CH = CH(CH2)n OAc and some saturated acetates
Compound ( T, oC I ~APL I I /a~t-60 I~1, I a/u
{
~ C~1i~sCH=Cf-I-CH.QAc 1338 336 1624 6_4 286
CeH13CH=CH (CH.)_OAc 1330 3,'i0 1616 616 216
C;,H�C=1=CH (CH,)30Ac 1314 314 1626 626 312 �
C3H,CH=CH (CHz)y~ ~Ac 'r 1337 337 . 1644 644 307
C2H~CH=CH (CH,)gOAc 1346 :i46 I654 654 308
C,~H�OAc 1359 359 1626 626 267
C7H~bCH-CH (CH1)3OAc 1520 320 1831 ' S31 311
C3H�CH-CH (CH~)sOAc o 1527 327 1844 ~ 64~t 317
C~H~CH-CH(CH~IeOac ~ 1~33 333 184U 64) , 3~7
C.H;,CH=CH (CH.)st)Ac 1540 340 186G ~ 6G6 ' 326
C,~ ~;OAc '~:i~;2 362 I 1822 F22 I 26'1
C~H,~CH=CH (CH~)30Ac ' ~ 1719 319 2045 6~~G I 327
C,HyCH-Cfi (CH:)80Ac ' 1734 334 2073 673 33~
C~H7';11=CH (CI-i~)flOAc ~ 1734 33~ 20~~1 674 337
C~HSCH==CH (i H;)~nOAC ' ' 1743 343 20SS 688 34~i
C, , H~~0;\ c I 7~7 357 2052 G 52 29:i
CeH13CH=CH (CH,)s'~Ac 192~~ 325 2267 667 I 342
CSH�CH=CH (CHz)~OAc o 192~ 329 2272 6"2 343
C,H9CH-Gf-1(CH.),,,nAc ~ 1436 326 1278 678 342
C~H7CH-CH (CH2)�OAc 1934 33~ 226y F6~~ 335
C, BH3~0 A c ~ 19G0 36U 225 ~ 6F;~.i 29~
Table 2. Retention indexes of ethylene alcohols with general formula ,
cis
R-CH = CH(CH2)n OH and some saturated alcohols
Compound . I T�, C /.~PL ~.rt-GO p!, ~/,a
I I
GH,;,CH -C 1-CFI..OH 1221 221 ~'608 608 387
C~ ~�CH=CH (CHz)tOH o 1225 225 1596 5~.i96 371
C-,' i�CH=CH (C:-I ~)30H ~ 1213 213 i614 G14 401
C,o~-1.,ON 12~3 2~3 1i90 590 337
C6H,~;CH=Cf 1(CH._),OH 1419 219 I 1821 621 402
C;H�CH=CH (CH~);Ot! 0 1~119 219 182u 620 4)1
CH,CH=CH (CH.~.,UH ^ 145U 250 1x61 6:1 411
C,_Hz5 )H 1.4~3 2~i3 18U4 604 351
C;,H�CH--CH (Cf(.,);C) ~ 16?6 226 I 2035 63~ 409
C.,H-CH-C i(CH~),Ufi o 1629 229 2048 ' 648 419
Cfi:;Cfl=C1f (CH_)�OH 165/ 257 ~ 2U86 686 429
C�Hl,U 1 I 1 Go8 258 I 2U21 622 364
C,H.,Cfi-C~i ~CH_),,,OH I I 1833 I 233 I 22~3 I 6ti3 I 420
C~,;H;~OH Ib62 262 2126 626 364 ~
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rvn vrr~~,ana. ~..sa: v.~a.a
Table 3. Retention indexes of epoxides with the geaeral formula
R~ \/(,ti^-R_
O
re-c., I I
Com ound I~�uL ~i~~~ ~~2 e112
P
C i3\CH (CH._), CH-CH-C,H,; 1795 95 2060 36Q I 26i
3/ ~~o~ I
t
Ch3/C'H(CH_):,"H-CH-CeH1e 1998 ~8 2272 373 ?74
\ ~
- O
CH3%CH (C",),GH-CtI-C,,,H� 1896 96 2160 360 264
\ /
O
CH3~CH (CH,),CH-CH-CI�H~, I 1:~90 90 2266 366 276
3 ~O ~
Table 4. Increments of retention indexes for functional groups
Compound o~ ei ~i
I , I ~
R-CH=CH (CH.:~n CH 213-257 596-G86 371-429
R-CH~CH (CH_)n O.'1c 314-346 616-668 `?86-345 '
. C~ H,~�t10H 253-262 590-626 337-3E4
' C~ H2nT~0Ac 357-362 622-659 260-299
R~-~'H-CH-R, 90-5'3 36')=372 264-276
~ / .
O
Within a specific class of compounds, the value of the retention index (I) at
a given phase permits determination of the length of the carbon chain provided
we know the contribution of the functional group at this phase (~I,~,DI2):
I~- ~1, 1.-e1.: ~5~
n=-1Q0 --l0U '
For all of the tested alcohols and acetates, the value of I is an overall one,
including the contribution of the alcohol or acetate group and unsaturated bond.
If the retention indexes of saturated compounds are known, one can assess the
contribution of the multiple bond.
The value of ~I12 is determined primarily by dipole-dipole interaction between the
dissolved substance and stationary phase. With increase in polarity of the mole-
cule, there is increase in value of ~I12, and this is evident on the example of
the compounds studied:
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- el1z ~C,~H',n-~IOH~;~ ~I12 ~CnH2n1~0AC~~ ~1~2 (R,-CH-CH-Rz), as weli as
\ /
~O
~jt~ ~R-CH = CH (CH~)~ OH~ ~ ~I12 ~CnHsn+~ OH)" and
~112 (R-CH = CH (CHy~~ 0AC) ~ ~112 ~C~H2n+1 OAC~.
The existing scatter of values o~ T and ~I for each class of compounds is attri-
butable to the influence of the length of the molecular chain and position of
multiple bond, a change in which alters the mutual infl.uence of functional groups,
which affects general polarity of the molecule.
The obtained retention indexes were used to identify the sex pheromones of female
cabbage and tomato moths.
We determined the retention index of the chromatographic peaks on two columns
of diff erent polarity, corresponding to the length of the chain and functionality
of pheromones to determine these parameters.
Female extract consists of a mixture of substances. In order to refer a specif ic
peak on the chromatogram to a pheromone, preparative selection of 1-minute extract
fractions was made, followed by electroantennographic te$ting, as described in
[6]. We then identified peaks contained in fractions, on which male antennae
showed maximum responses. It should be noted that, in view of the small amount of
active substance contained in insect glands, the pheromone peak is not traced on
- the chromatogram, whereas the antennae respond to the corresponding fraction. In
this case, it is quite helpful to use Kovats indexes that can be calculated
from the time of appearance of the fraction that yields the maximum response of
male antennae.
The pheromone of female cabbage moths consists of one main component II with
retention indexes Ii = 1936, I2 = 2278 and pI12 = 342. The pheromone of the
tomato moth has two components (III, IIII}, and the retention indexes are
Ii= = 1833, I2I = 2253, ~I12 = 420, IiII = 1936, I2II = 2278, ~I12I = 342.
If we compare the values of ~I12 of natural pheromones to the tabulated data
(Tables 1, 2, 4) it is easy to see that the I?, II~= (~I12 = 342 units) peaks
fall into the range of ethylene acetates, while the III peak is in the range
of ethylene alcohols (~I12 = 420 units). The coincidence of retention indexes
on nonpolar and polar phases of natural pheromones with hexadecenol and hexa-
decenyl actetate models leads us to the conclusion that the length of the carbon
chain of the compounds to be identif ied is 16. In the general case, one should
use formula (5) to determine chain length.
Experimental Section ~
The retention indexes were determined on a Khrom-42 chromatograph with flame-
ionization detector. The glass columns are 2.Sx3 mm in size. The carrier gas is
~ extra pure nitrogen. We used N-AW-HIrIDS 60/80-mesh chromaton as the solid carrier.
Apiezon L(8%) served as the nonpolar phase and XE-60 nitrile silicone rubber (15%)
as the polar phase.
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� V~~ V~ ~ ~t~/~L, v.../V V~\L�
Conclusions
Determination was made of retention indexes of a series of unsaturated alcohols,
acetates and saturated epoxides on two columns of different polarity. The
obtained tabulated data were used to determine chain length and functionality of
pheromones of two moth species.
BIBLIOGRAPHY
1. Jacobson, M., "Sex Pheromones of Insects," Moscow, 1976.
2. Minyaylo, V. A. and Kovalev, B. G., "Advances in Science and Technology," Ento-
mology Series, Moscow, 2, 1973.
3. Reynolds, W. 0., "Gas Chromatographic Retention Data," Evanston, 1966.
- 4. Kovats, E., HELV. CHIM. ACTA, 41, 1915, 1958.
5. Idem, ADVANCES CHROMATOGR., 1, 229, 1966.
6. Kovalev, B. G., Nedopekina, S. F., Lebedeva, K. V. and Kost, A. N., HCOORGAN.
KtIIMIYA, 5, 912, 1979.
COPYRIGHT: Izdatel~stvo "FAN" UzSSR, 1981
10,657
= CSO: 1840/103
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UDC: 597:585:591:488.9:471.3+591.9(261,265,266)
- SEISMOSENSORY SYSTEM AND CLASSIFICATION OF COTTIDAE FISH (MYOXOCEPHALINAE,
ARTEDIELLINAE)
Leningrad SEYSMOSENSORNAYA SISTEMA I KLASSIFIKATSIYA KERCHAKOVYKH RYB (COTTIDAE:
MYOXOCEPHALINAE, ARTEDIELLINAE) in Russian 1979 (signed to press 28 Aug 79)
pP 2-7, 207-208
[Annotation, introduction and table of contents from book "Seismosensory System
and Classification of Cottidae Fish (Myoxocephalinae, Artediellinae)" by Aleksey
Vadimovich Neyelov, approved by the Zoological Institute of the USSR Academy of
Sciences, Izdatel'stvo "Nauka", 1100 copies, 208 pages] ,
[Text] This monograph deals with a comparative morphology of the seismosensory
system of 54 species and subspecies of Cottidae. Original studies are preceded
by analysis of the worldwide literature on morphology of the seismosensory system
of Anamnia vertebrates, different aspects of function, nature of variability
and use in systematics and phylogeiiesis. A classification of both subfamilies
studied is validated on the basis of the structure of various systems of organs.
Characteristics and identification of various taxons through subspecies levels
are submitted. There is discussion of the nature and patterns of evolutionary
. transformations of diff erent elements of the seismosensory system and phylogenetic
relations and evolution of Cottidae. References 642, illustrations 85.
Introduction
The Cottidae family is the largest group of fish, with regard to number of species,
in the ichthyofiauna of the USSR and.adjacent waters. Expressly the Cottidae con-
stitute the most substantial element of the fauna of our northern and far eastern
seas. Thus, Andriyashev (1939b) reports 297 f ish species in the Bering Sea,
65 of which are Cottidae, i.e., over one-fif th (21.9%) of its entire ichthyofauna.
Cot`idae are represented by 10 species in the Kara and Chukotsk seas of the 56 and
44 ~pecies of fish, respectively, known in these seas (Andriyashev, 1954). Schmidt
(1950) cites about 50 species of Cottidae for the Sea of Okhotsk, or almost one-
fifth of the entire fish fauna. Wili.movsky (1964) stresses in particular that
45 out of the 140 species of fish in coastal regions of'the Aleutian Archipelago,
or almost one-third, consist of bullheads of the Cottidae family, and no other
tamilies can compete with them in this regard. Cottidae play an excepti~nally
large part in the fauna of Lake Baykal; all 26 bullh~ad speciesl are endemic
to this lake (Taliyev, 1955).
As yet, very little has been done with regard to working out the morphological
bases of the system of the Cottidae family and particularly with respect to
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� Vl\ Vl'~ 1~.~~1a+ v~Li Vi ~L l
evulutionary relations between the taxons it includes. In this respect, the
liaykal bullheads were the "luckiest," since comprehensive studies thereof resulted
in the excellent monograph by Taliyev (1955). In this work, the author devoted
several pages to evolution of bullheads of the Cottidae family in connection with
the origin and formation of endemic bullhead fauna of Lake Baykal. Current con-
ceptions of the scope and relations within Cottidae and families close to it
are based on the rather concise, but mor~hologically validated work of Taranets
(1941), "Classification and Origin of Bullheads of the Cottidae Family."
A large group of father-lasher bullheads, which Taranets (1941) placed in the
Myoxocephalinae subfamily numbering 14 genera with 48 species, according to Taranets,
occupies the central position in this family. Schmidt (1950), Taliyev (1955) and
Bolin (1947b) stressed the leading role of Myoxocephalinae among the Cottidae.
Not only are the species of this subfamily numerous, but they are extremely
diverse. Father-lasher bullheads are among the most common elements of the
benthic ichthyofauna of all our northern and far eastern seas. Moreover, they
are quite numerous within the range of their habitat, inhabiting seas from the
littoral zone to depths of about 2000 m, i.e., they are also among the deepest
water fish in the Cottidae f amily. Some species form concentrations of commercial
magnitude. Thus, in 1970, 2653 tons of bullheads were caught in the Northeast Atlantic
(Statistical Information on USSR Fishing in the Northeast Atlantic ...,1971b),
and 1283 tons were caught in the Baltic Sea (Statistical Information 1971a).
The Cottidae species studied are very important to work on zoogeography and origin
of ichthyofauna of different bodies of water.
The foregoing explains, to some extent, the reason for the increased interest of
~ ichthyologists all over the world in bullheads of this group and Cottidae in
general. Hence it is understandable that this book, which has some prior history,
was the logical development of *he interest of Russian ichthyologists in this
group of fish.
Originally, the work was planned as a study on the "Fauna of the USSR," where
- attention would have been focused on osteological and external morphological
distinctions of structure, which are generally used extensively and constantly in
studies of fish. However, in the course of this work, I consistently encountered
instances where the status of the species was quite vague and the range diffuse,
i.e., it was impossible to off er a clearcut description of the species and, quite
often, of the genus as well. It was necessary to f ind some morphological criteria,
which had not been discovered before (if they existed at all) that would permit
- demonstration of rather distinct species-specific differences in the group under
study.
As a result of comprehensive studies of the morphology of different systems, in-
cluding osteology and external morphology of many species of the Myoxocephalinae
s. l. subspecies, it was found that the species and entire groups of species
are quite distinguishable, primarily according to quantity, structure and,
first of all, topography of integumental canals and pores of the seismosensory
system of the head and body. Expressly the structural distinctions of the seismo-
sensory system or, as~ it is usually inaccurately, but commonly called the"lateral
line organs," enabled us to comprehend, to a certain degree of course, the sys-
tematic correlations in the Myoxocephalinae s.L subfamily. Having some degree
_ of species specificity, the general structural plan of the seismosensory system
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turned out to be profoundly different in the two tribes with the most species--
Myoxocephalini and. Artediellini, subfamily Myoxocephalinae(in the scope adopted
by Taranets, 1941). Structural differences in the seismosensory system of
hooked-horn bullheads (Artediellini of Taranets) and all other representatives of
thie subfamily made it possible to validate the independence of the hook-horns
in a separate subfamily (Artediellinae), along with the use of other morphological
structural distinctions. Its species-specificity made it possible to clearly
distinguish within these two subfamilies lower taxons through species. This,
in turn, made it possible to make broader use of the distinctions of external mor-
phology, which it had been sometimes impossible to detect before exact differ-
entiation between species. .
In the course of our work, we ecountered a number of inethodological difficulties.
The main one was the absence of developed terminology to designate the numerous
canals and pores of the seismosensory system of the fish head. Usually, no atten-
tion was paid to such structural distinctions of the peripheral branches of the
seismosensory system when it was studied, since it was essentially the route and
nature of the canals that were studied. Apparently, this is why insufficient de-
tails were provided about the peripheral branches of the seismosensory system and
it is the reason for absence of conventional terminology for them (Makushok, 1958,
1961c; Jakubowski, 1965; Cowan, 1970, 1971). It was necessary to not only find a
- convenient method to study the system proper and stock material, but primarily
to systematize the large quantity of pores and cutaneous canals which appeared,
at first glance, to be quite disorderly in arrangement. After this, it was
necessary to develop a system of designations, with which it would be convenient,
to some extent, of course, to work in describing the structural distinctions of
the fish sense organs under study. Since this is, to some extent, the first
attempt at a comprehensive description of the morphology of the peripheral
branches of the seismosensory system, this author realizes that it is far from
perfect, and would be very appreciative of all critical comments and additions
that would improve it.
Comprehensive studies of the topography of the seismosensory system as applied
to systematics was largely instrumental in developing the conceptions submitted
in this book concerning the validity.of different taxons and their systematic
rank, which led to development of conceptions concerning the evolutionary relations
of the fish species studied.
The large number of species and their considerable morphological variety made it
possible to trace on vast (in the sense of species) and moreover related material
ttie following: in the first place, the diversity of structure, primarily the
topo~;rapfiy of peripheral branches (canals and pores) of the seismosensory system
uf closely related forms with the same general structural plan thereof; in the
second place, the significant differences in structure of central parts of the
seismosensory system (morphology of canals) and communication with the environment
and, in the third place, which is the main point, this diversity enabled us to
demonstrate and trace the direction and sequence of evolutionary changes in struc-
ture of the seismosensory system of species of the same genus, different genera
and similar genera. As a result, we found instances of morphological parallelism,
wliere species formed homologous series in the meaning of Vavilov (1967) and
ecological parallels in the meaning of Berg (1935). As a result, all this made
- it possible to propose, of course with the use of the entire set of studied
morph~l.ogical structural distinctions, a system of subfamilies, their position
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among Cottidae and to submit a scheme of probable phylogenetic ~elations between .
- taxons in this group. It is expressly on the basis of comprehensive morphologi-
cal analysis of structural distinctions of the seismosensory system of all repre-
sentatives of the Myoxocephalinae and Artediellinae subfamilies that I demonstrated
(Neyelov, 1967a, 1967b, 1971a, 1971b, 1973a, 1976, 1977) that simplification of
structure of peripheral branches--oligomerization according to Dogel'--should be
considered one of the main directions of evolutionary transformations of this
system of sense organs in bullheads and sculpins.
In working out the system of subfamilies, we also used some of the osteological
distinctions of structure of rhe skull and axial skelet~~n (mainly for substantia-
tion and characterizati~n of ;~ubF~.milies). For characteristics of various taxons,
extensive use was made of ext~~Y�.t~.= morphological structural features, which are
generally used in systematizin~; aud identifying Cottidae (along with new or pre-
viously insufficiently used morphological structures). This enables us to make
use of the abundant factual material already accumulated from all previous studies
and to include in the identifications and determination tables external morpholo-
gical structures that can be readily seen with the mc~st cursory examination of
fish at:'~ could always be compared to already exis*_inE; descriptions.
Such synthesis.of morphological structures whieh'~were used for the f irst time and
external morphological distirictions used previously was instrumental in more
accurate utilization of these fea,tures and made it possible to,describp more
fully a given taxon within ttie foxm of a brief diagne~sis. It i~s expressly because
of these considerations that I resisted the temptation.of constructing idEnti-
fication tables and offering taxon diagnoses solely on the basis of the dis-
tinctions of the seismosensory system, a~though~this could have been done (as,
for example, was done by Illick, 1956 and Nelson, 1972).
In view of the foregoing, it was necessary to alter the initial direction of
research, devoting more attention to morphology than fauna proper.
This book consists of an introduction and five chapters. The first chapter deals
with the history of research on the seismosensory system, general conception of
its structure, function and use of its morphological distinctions in systematics
and pliylogenesis.2 The second chapter is concerned with morphology of the seismo-
sensory system of cottoid bullheads and sculpins and contains the following:
information about the volume of material studied, terminology for peripheral ,
branches (canals and pores) of the seismosensory system which was developed and
is proposed for Cottidae, general morphological description of this system in
both subfamilies, description of genera and species with reference to seismo-
sensory system, and it is illustrated with original drawings. The third chapter
- contains a morphological validation of the ind~pendence of Myoxocephalinae and
Artediellinae subfamilies and lower ranking taxons. Chapter 4 has information
about the scope of the subfamilies under study and changes in the system of these
fish, as compared to the classification of Taranets. This chapter includes identi-
fication tables for all taxons through the subspecies, characteristics of sub-
families, tribes and genera, and diagnoses for all lower taxons. The diagnoses
are listed in the identification tables in the form of an expanded thesis or
antittiesis. There too is brief information about the distribution and habitat
depth. After the description of each genus, its scope is indicated--list of
species and the main synonyms. The synonyms are provided in full for taxons
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from the genus level up. Since the complete validated name, with its author, is
given for all of the species studied, the author's name is omitted in the rest of
the book where only their Latin names are given. The Latin natne of all other
mentioned species is given along with its author.
In view of the fact that the International Commission for Zoological Nomenclature
adopted a decision to consider 1814 the year of publication of "Zoographia
Rosso-Asiatica" by Pallas (Vol 3, Fishes) (Svetovidov, 1976), unlike the previously
used year of 1811, this change is taken into consideration in indicating the year
of description of the relevant Pallasian species.
The book concludes with Chapter 5, in which there is discussion of direction of
specialization of the seismosensory system, its evolution as a system of sense
organs that implement direct contact between the organism and habitat, questions
of evolution of the systematic groups studied and offers a scheme of phylogenetic
relations within the Myoxocephalinae and Artediellinae subfamilies.
This work constitutes an attempt at systematization of morphological details of
structure of the seismosensory system and to use this system of sense organs for
the purpose of classification and identification of taxons on the subspecies-
subfamily rank.
This work was done in the laboratory of ichthyology of the Zoological Institute,
USSR Academy of Sciences. I wish to express my profound appreciation to the
management of this institute and laboratory for g~anting me the opportunity to work
in that laboratory and make use of its rich fish collection. I wish to express my
sincere appreciation and profound gratitude to A. P. Andriyashev, corresponding
member of the USSR Academy of Sciences, for his considerate and attentive attitude
throughout my work on the monograph and for taking on the job of editing the
manuscript. The advice and constant aid of Anatoliy Petrovich were largely
instrumental in following the adopted direction of research. I am also profoundly
grateful to the never to be forgotten G. U. Lindberg for his constant interest .
in my work, valuable advice and availability for consultation whenever necessary,
as well as the opportunity to use the abundant bullhead catches of the Kuril-
Sakhalin expedition. I am sincerely grateful to V. M. Makushok, who unintentionally
was the inspiration for the chosen direction of investigation for his advice in
making the morphological drawings. To all of the laboratory staff, my friends
and coworkers and particularly senior laboratory technicians A. A. Korovkina,
M. S. Morozova and V. P. Prirodina I am profoundly grateful, and I thank all of
tt~em for their constant assistance in everyday work and support, without which
the book would hardly have been completed. I offer my sincere gratitude to
the staff of the Zoological Institute library, and particularly to the bright me-
mory of Ivan Georgiyevich Musatov, for their constant aid in bibliographic work.
Without ttiis library and its sharp staff our work would have been simply incon-
ceivable. My sincere gratitude goes to all comrades and friends who were in-
volved in some way or other in this work. I am profoundly grateful to artists
M. M. Zharenkov and N. D. Ogloblina for rendering in India ink the author's
uri~;inal morphological drawings and diagrams sketched in pencil. I take this
opportunity to express my appreciation to V. V.~Fedorov,3 L. S. Kodolov, V. P.
5tiuntov on the staff of TINRO [Pacific Ocean Scientific Research Institute of
Fisheries and Oceanography], 0. G. Kusakin, V. M. Matyushin and V. I. Pinchuk on
the staff of the Soil Biology Institute of DVNTs [Far East Research Center?] of the
Siberian Department of the USSR Academy of Sciences, as well as all other comrades
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for placing at my disposal the Cottidae caught in the bas~in of the Paci�ic Ocean.
I am deeply grateful to doctors D. M. Cohen (U. S. N. M.,j and Nielsen (Univ.
Zool. Museum, Copenhagen) for granting the opportunity of examining bullhead
specimens lacking in our collections, which they were kind enough to send me.
~'inally, I wish to thank with all my heart my Polish colleague, Doctor M. I.
.Takubowski, who was kind enough, during his visit to the Zoological Institute,
to help me learn to use the original techniques he developed for examining
organs of the seismosensory system.
FOOTNOTES
- 1. The Russian name for f ish of the Cottidae family--"kerchakovyye" or
"rogatkovyye" [Father-lashers or horned fish] (Rass, Lindberg, 1971) does
not always apply well to all species, so that in addition to these names,
this author has also used the name, "bullheads," which is old and has been
in wide use f or a long time, and currently u~ed to refer only to the Gobiidae
family; the name, "sculpins," is used for fresh-water species of this family.
2, To hisprofound regret, the author was unable to make use here of the excellent
monograph by Disler, published in 1977, dealing with morphology and develop-
ment of seismosensory system organs of Euselachiae, or for reasons beyond his
control to make use of the interesting works of Cowan (1970, 1971, 1972a,
, 1972b, 1973) and Kartavtsev (1975) dealing with species of the genus Myoxo-
cephalus in the discussion of his own findings (Chapter 5).
, 3. Presently a scientist in the laboratory of ichthyology.of the Zoological
Institute, USSR Academy of Sciences.
CONTENTS Page
Introduction 3
Chapter 1. Seismosensory System of Fish and Methods of Studying It 8
l. Historical survey 8
a. Morphology 8
b. Function 15
2. Use of distinctions of seismosensory system in systematics and
phylogenesis 21
a. History of the question and its present status 21
b. Use of distinctions of seismosensory system in systematics and
phylogenesis of Cottoidei 27
c. Some aspects of variability of morphology of seismosensory
system organs 30
d. Free neuromasts 32
Chapter 2. Morphology of Seismosensory System of Bullheads
(Myoxocephalinae) and horned fish (Artediellinae) 34
1. Material and methods 34
2. Terminology and designations of elements of seismosensory system 35
a. General theses 35
b. Designations for canals and pores 3~
3. Description of Myoxocephalinae and Artediellinae subfamilies with
reference to seismosensory system 42
General information 42
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Subfamily Myoxocephalinae 42
Myoxocephalini tribe 42
Genus Myoxocephalus 43
Genus Megalocottus 57
Genus Triglopsis 58
Microcottini tribe 60
Genus Microcottus 60
Genus Porocottus 62
Genus Argyrocottus 66
- Taurocottini tribe 67
Genus Taurocottus 6~
Genus Trichocottus 69
Enophryini tribe
Genus Enophrys
Genus Aspicottus ~l
Genus Taurulus 73
Genus Micrenophrys 74
Subfamily Artediellinae 75
Genus Artediellus 75
Genus Artedielloides ~ 89
Genus Artediellichthys 90
Genus Artediellina 91
Genus Zesticelus 92
Genus Cottiusculus 93
Chapter 3. Morphological Validation of Subfamilies Myoxocephalinae and
Artediellinae, and Their Place in the System 96
1. History of the question and classification of A. Ya. Taranets 96
2. Seismosensory system 99
- 3. Skull 100
a. Sensory canals of cranial bones and structure of frontal bones Z00
b. Myodome 101
c. Some other distinctions of cranial morphology 103
4. Thoracic cingulum and axial skeleton 104
5. External motphol~gy _ 106
a. Structure of some visceral bones of the head 106
b. Fins 107
c. Nasal and preopercular spines 108
d. Granulation of inembrane bones 110
e. Osseous investment of the head and trunk, and appendages of
the skin ~ 111
6. Coloration 112
7. Biology 112
Chapter 4. System of Myoxocephalinae and Artediellinae 114
Main changes made in the system 114
Table for identifying subfamilies 116
I. Subfamily Myoxocephalinae 117
Table for identifying tribes 118
1. Myoxocephalini tribe 119
Table for identifying genera 119
1. Genus Myoxocephalus 120
Table for identifying species 123
2. Genus Megalocottus 129
3. Genus Triglopsis 131
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2. Microcottini tribe 133
Table for identifying genera 133
1. Genus Microcottus 134
2. Genus Porocottus 135
Table for identifying subgenera, species and subspecies 137
3. Genus Argyrocottus 141
3. Taurocottini tribe 142
Table for identifying genera 142
1. Genus Taurocottus 142
Z. Genus Trichocottus 143
4. Enophryini tribe 143
Table for identifying genera 144
1. Genus Enophrys 145
2. Genus Aspi~ottus 147
Table for identifying species 148
l. Genus Taurulus , 149
2. Genus Micrenophrys 149
II. Subfamily Artediellinae 150
Table for identifying genera 152
1. Genus Artediellus 153
Table for identifying subgenera, species and subspecies 155
2. Genus Artedielloides 159
3. Genus Artediellichthys 159
4. Genus Artediellina 160
� 5. Genus Zesticelus 160
Table for identifying species 161
1. Genus Cottiusculus '161
- Table for identifying species 162
Chapter 5. Main Directions of Evolution of Seismosensory System and
Phylogenetic Relations in Subfamilies Myoxocephalinae and Artediellinae 164
Bibliography 181
Index of Latin Names 201
COPYRIGHT: Izdatel'stvo "Nauka", 1979
10,657
CSO: 1840/101 ~
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ENVIRONMENT
UDC 575+577.4
DETERMINATION AND QUANTITATIVE EVALUATION OF MUTA(~NIC FACTORS OF ATITHROPOGEr1I-
CALLY VARYING HABITAT USING BAC'l~sRIAL TEST SYSTF.MS
Sverdlovsk EKOLOGIYA in Russian No 6, Nov-Dec 81 (manuscript received 21 Apr 81)
- PP 72-81
[Article by R. A. Pshenichnov and Yu. V. Pashin, Institute of Ecology of Plants
and Animals, Ural Scientific Center, USSR Academv of Sciences; and Institute of
General Genetics, USSR Acaderay of Sciences]
[Text] Basi.ng themselves on a review of the literature and
observation results, the authors describe the optimum variants
of bacterial test systems and validate their suitability,
� economy and promise as a means for screening environmental
mutagens. Observation results are presented, and the future
directions of research and possible li.mitations of the method
are noted.
The spectrum of chemicals in the environment having genetic effects changed signi-
ficantly in recent years as a result of swift development of-t~re-~cientific-technical
revolution. Throughout its long evolution the hum~n population has been exposed to
natural mutagens; in recent decades, however, the effects of artificially synthesized
compounds--ones which the human body as wel~. as animals, plants and microorganisms
had not encountered formerly--have assumed dominance. Therefore evaluation of the
effect of mutagenic factors i.n the environment is acquiring important significance,
and the need has arisen for developing and introdu~ing test systems that would per-
mit us to reveal, aiaong many new compounds, those which are genetically active, for
determining their mutagenic potential and for predicting their effects (2).
Among dozens of test objects used to register mutagenicity, microorganisms--bacteria
_ primarily--are the most sensitive and sufficiently representative. The reasons for
this are:
1) Microorganisms are a constant component of all ecosystems, and their diversity,
abundance and contribution to the cycling of matter and energy in nature are ex-
tremely great. Correspondingly a study of their gene pool and heterogenicity, as
formed by the habitat, must become an object of constant attention and supervision.
2) Microorganisms have a minimum reproductive period (measured in minutes and
_ hours), and change in genotypes may be registered within a short time.
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rvn vrriuni. ~o~ vivi,t
3) Bacteria have a haploid set of chromosomes, and disturbance of the genotype is
followed immediately by its phenatypic expression. The methods of determining
change in phenotype are rather diverse. As a rule they are simple and practicable.
4) Many years of experience in testing hundreds of diverse chemi.ca~. mutagens have
revealed a high degree of correlation in their action upon DNA preparations and
- upon genetic structures in cultured cells, plants, ani.mals, man and microorganisms
(7.8~1~)�
It should be noted that the principal achievements of modern molecular genetics have
involved microorganism.~, and primarily bacteria; therefore all data on genetic events
occurring in bacteria contai.n a large amount of information.
Moreover bacterial tests are relatively inexpensive to run, they can be carried out
in large numbers, and therefore they may be used extensively in var~.ous screening
programs. They can be used to detect gene mutations, owing to which rare events
can be revealed in larg~ populations within a short testing time. Point or gene
mutations that sio,:i~ficantly influence the size of the so-called "genetic load"
affect only a small part of a gene (about one or two pairs of a total of about
1,000 pairs of bases making up a gene). Substitution, addition or deletion of. bases
is what usually occurs. Point mutations are usually registered in bacteria as re-
verse mutations, in which case a specific mutant allele m~.y be reverted by substi-
tuting one pair of bases by another, while a mutation associated with shifting of the
"readilig framework" may be reverted by shifting the "reading framework." Two types
of similar cells are used in mutagenicity tests: 1) cells with a normal DNA repair
mechanism and 2) cells in which one or several steps of the damaged DNA restoration
pathway are absent.
The simplest classical test is performed in Petri dishes seeded with test cultures
of Escherichia coZi (44) or SaZmoneZZa typhimurium (12, 14). Z"he substance to be
tested is applied to the center of each of these dishes, thus creating a gradient
in its concentration from the center to the rim. If the substance turns out to be
mutagenic a ring of induced mutants forms about it. In this case predominant death
of strains with a defective repair mechanism would indicate that a reaction had
occurred with cellular DNA, and consequently that the given substance is genetically
active. 11n important prerequisite of these tests is that the substance to be tested
must be able to diffuse in agar. If it does not diffuse, the indicator strains may
be processed with different concentrations of suspensions of the substance. For
practical purposes no difficulties arise with water-soluble compounds.
Since Ames' first works were published, test strains have undergone continual im-
provement through the introduction of additional mutations (for example ones which
can reveal defects in the polysaccharide membrane of a cell that increase its per-
meability, and defects in the repair system) and through introduction of episomes,
use of lysogenic cultures and so on, making them more sensitive than wild strains.
Today over 40 strains of S. typhimurZwn, 40 of coZi and about 10 of BaciZZus
subtiZis are used for genetic screening; however, only the 20 or so strains shown
in the table below have practical significance.
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General Characteristics of Bacterial Strains Used Most Often in.Genetic
Screening (Brusick et al., 1976; 7, 8, 52, e~c.)*
~Ha3eauHe ~2~ AOiIOlIHN- B6IABJIAMtHf TNII
UPf21IH3M8 ]fjTBM\1 TCAb1i2A MyT211110NHOI'O
?I OCI{ODHO~ .MyT811NA 1~3MEH2N11A
~ryrauiii+ (~cpcpexTa)
Sa(uionclla 7'A� 15.'~4 (hys D3os3) u~�r B R- rS
lypliimuiiunt TA-1530 R-BPS
(hys- - hysi ) TA�1535 (hys G~6) rfa, u~�r B R- BPS
TA-1536 R - FS
TA-1538 (hys D~~z) rfa, u~~r B R- ~S
7'A-1950 (hys G~a) uvr B R- IIPS �
- TA� 100 R - BPS
TA-98 R - FS
Cscherichia pol A-
coli WPz R-BPS
- (try--arg F) WPz uvr A- R- BPS
(:+rg--arg+) }~I1 ~~13 ~ I13 FM
- C~~661 R - BPS
Ch161 . R - BPS
Waiio/pai~e . ER
Baci!!us H17 Rec+ NHrN6HposaHNe
s~i6filis IP995 Rec- I I pocTa
. (5) .
*Symbols: hys--histidine; try--tryptophan;
arg--arginine; rfa--mutation increasing cell
wall permeability due to defects in the poly-
saccharide layer; uvr--mutations causing dis-
turbance of excisional repair; R--reverse
mutation; BPS--base pair substitution; FS--
"reading framework" shift; FM--forward muta-
tions; ER--excisional repair; Rec'--recambina-
tion insufficiency.
Key:
1. Organism and principal mutation 4. Revealed type of mutational change
2. Strain (effect)
3. Additional mutation 5. Growth inhibition
Because of the high specificity of the test strains, cases may occur in which mutagens
are incapable of inducing detectable types of reversions, and therefore the need for
_ constantly seeking and expanding the set of strains with different mutation mechanisms
is obvious.
In contrast to mutagenic factors such as ionizing radiation and ultraviolet emi.ssions,
chemical compounds undergo complex metabolic biodegradation on entering the organism.
In this case mutagens may transform into inactive compounds, while on the other hand
genetically inactive substances (indirect mutagens) may acquire mutagenic properties.
The metabolic pathways of xenobiotic substances may vary in some ways, but the main
role in their biological transformation is played by cytochrome F-450, an element of
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ruK urr~~ ~Ai. UJ~ VIVLY
tlie mono-oxygenase system associated with intracellular membranes. This system can
be found in a number of microbial species, insects, amphibians, fish, reptiles, higher
plants, birds and mammals.
Grin and Goldberger (1) emphasized the biochemical universality of the operation of
this system, which is responsible for oxidation of lipid-soluble xenobiotic substanCes
and for their conversion into readily excreted hydrophilic compounds. It is an im-
portant component of more-general systems responsible for imanunochemical homeostasis
(3). Z'herefore metabolic activation of the tested substance, which may be done
in vivo or in v'itro, is a mandatory stage of genetic testing. In the former case the ~
substance to be tested is subjected to metabolism in the body of a mammal (an example
would be the recipient medium method), after which it acts upon test microorganisms
injected into the abdominal cavity, circulatory system or testicles of the recipient.
_ Ir?dicator cells processed in this fashion are removed and analyzed by a particular
method. Indicator microorganisms have also been found useful for revealing the ways
of penetration, dissemination and elimination of mutagens with biological fluids--
the urine and blood of animals processed by premutagens. However, the latter
variant has enjoyed limited acceptance due to the frequent instability of the active
form of inetabolites in the body.
The "recipient medium" method is not broadly accepted owing to the need for admini-
stering large doses of the tested substances, its expensiveness and other reasons
such as nonuniform distribution of the active substance in the body and tissue-specific
activation or deactivation of the mutagen. An infectious process may arise in mammals
given a significant number of test strain bacteria when the recipient is sick; and
so on. But even if the recipient remains outwardly healthy,in this case the bacterial
test strains would be subjected to the strong influence of the host's protective
mechanisms.
The method of inetabolic activation by microsomal enzymes in vitro has enjoyed the
broadest acceptance. Considering the biological universality of the mono-oxygenase
system in relation to metabolism of xenobiotic substances, the appropriate enzymatic
preparation may be obtained from the tissues of mice, rabbits, rats, Chinese hamsters,
dogs, quinea pigs, rhesus monkeys, baboons and man (32,36). Although mono-oxygenase
activity is typical of most organs rich in epithelial cells (stomach, intestine,
kidneys, lungs, skin), the commonly accepted and proven source of enzyme preparation
- is liver tissue, which is responsible for more than 90 percent of xenobiotic meta-
bolism. Liver tissue from sexually mature Wistar line male rats is used most often
for this purpose.
Although microsomal enzymatic activity is inherent to all healthy organisms, it may
b e subjected to additional induction and activation by chemical substances present
in food and the environment or introduced into the organism by artificial means. It
increases when a large number of drugs, food additives, salts of heavy metals, poly-
cyclic hydrocarbons, pesticides and other compounds enter the body. For practical
purposes microsomal oxygenases may be induced successfully with sodium phenobarbital,
aiokhlor, luminal, diazepam and other substances, which are administered intraperi-
toneally or per os for 3-5 days prior to dissection of the animals (18, 31, etc.).
Sampled liver tissue is homogenized at a temperature close to 0�, and the obtained
~reparation is used within the first 2 hours.
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- Substances to be tested may be metabolically activated by immobilizing the microsomal
~,reparation and the necessary cofactors in agar, or by p~ocessing the test strains
with a suspension of the substance to be tested (13,33). There may also be cases
- in which a mutagen cannot be detected (for example a chemical substance may be
metabolized not by liver enzymes but by intestinal microflora; the active metabolite
may be too unstable to achieve an effective concentration; there may be strongly
pronounced metabolic differences between species etc.). On the whole, however, on
the basis of 8 years of experience we can confidently recommend microbial test systems
as a means for revealing direct and potential mutagens--systems which can be used to
study most representatives of all classes of chemical compounds. They have been
found to be convenient and effective in revealing the genetic activity of hydro-
carbons, their derivatives, drugs, nitro derivatives, nitrogenous bases, nucleotides,
nucleosides, salts of heavy metals, i.norganic substances, steroids, low molecular
weight ~rganic substances, pesticides, herbicides, defoliants, complex mutagens,
natural physical factors and so on. They include widespread and rarely encountered
compounds, and commonly accepted and "exotic" substances--foodstuffs, food additives,
cosmetic dyes, meat curing fluids, fungal toxins, meadow grass extracts, alcohols,
waste water, tobacco smoke components, :iarcotics, and even the combustion products
of labdanum and church incense.
W~ may assert from an analysis of 1,015 works that authors have used the following
models with increasing frequency for genetic control: Cytogenic tests in man, tests
run on blood cells, experiments with plants, primary and transplanted animal cell
cultures, D1~A preparations and microorganisms, correspondingly in 5,9, 6.4, 7.5,
15.2, 16.4, 17.2 and 31.3 percent of the works.*
Tlte authors selected microbial test systems from practically a third of all of these
works in order to evaluate their genetic effect, and owing to their simplicity,
,racticability and quickness they were able to evaluate 34, 101, 465 and 855 different
compounds in comparable observations within a short period of time (30,50,43,35).
A tremendous amount of facts were accumulated, facts which can and must become the
basis for making the corresponding conclusions and recommendations on the most
suitable use of bacterial tests and for imposing limitations on the use of these
methods. These issues were discussed specially in 1978 in Arlington (USA) and
Dortmund (FRG), in May 1979 in Banbury (USA), in November 1980 in Ottawa (Canada)
and elsewhere. The resulting recommendations have already found their way into some
national programs (19,24,49,55). ~
The International Commission on Protection from Environmental Mutagens and Carcinogens
(ICPEMC) and the Environmental Mutagen Information Center (EMIC), created in 1977,
are making a major contribution to the creation and standardization of quick screening
test systems. In all recommendations and adopted programs, evaluation of the muta-
~teiiic:it~ of chemical compounds using bacterial test systems is the first and mandatory
stage of analysis. The following are also recommended:
*These percentages are the result of an analysis of the principal aoviet and
foreign publications of 1979-1980, including those in the abstract card index
- "Mutagenic Effects of Chemical Factors" published by the Biology Division of
the All-Union Institute of Scientific and Technical Information.
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1) usirig a single set of test strains--TA-1535, TA-1537, TA-1538, TA-98, TA-100 and
:~cxCmnrteZZa typhimurizon, and periodically verifying their genotype;
2) working with bacterial cultures undergoing logarithmic growth at a density of
1-2�109 cells/ml;
- 3) using the S-9 microsomal fraction from the liver of activated rats, preparing
fresh preparation each time or developing reliable methods of stabilizing prepara-
tions;
4) testing the spontaneous mutability of test strains and perfornung control tests
with known direct and potential mutagens;
5) considering the toxicity of the substance to be tested;
f;) determining the dose-effect curve;
7) in the concluding stage it would also be desirable to document the results in a
computer program, which w;:ald be the formal recording unit and the basis for the
program of further tests (21).
It should be noted that most of these recommendations were taken into account in the
methodological directives "A Test System for Evaluating the Mutagenic Activity of
Environmental Contaminants Using SaZmoneZZa" (8)� Timely publication of these
directives played a major role in standardization and development of work in the
Soviet IInion. The following may be regarded as unsolved problems hindering intro-
duction of bacterial test systems:
the need for creating a single collection of lyophilized test strain cultures;
development and industrial production of dry microsomal fraction preparations;
- prociur.tion of a single standardized outfit containing everything needed for control
of substances using bacterial test systems.
The first of these tasks is being worked on by the Institute of Ecology of Plants
and Animals of the USSR Academy of Sciences Ural Scientific Center, whicli has
developed the processes and created a collection of lyophilized SaZmoneZZa typhi-
muriwn cultures suited to long-term storage, transportation and use anywhere. It
has been demonstrated that lyophilization promotes long-term (more than a year)
preservation of the properties of indicator strains.
Researchers have traveled different paths in their search for a stable microsomal
fraction. The possibility fo r long-term storage of this fraction at superlow temper-
- atures on the order of -193�C has been demonstrated (16). Howe~~er, such long storage
and transportation of the preparation were associated with certain difficultips,
and thus this approach has not enjoyed widespread acceptance. The Institute of
Ecelogy of Plants and Animals of the USSR Academy of Sciences Ural Scientific Center
has developed a semi-industrial method of making lyophilized capsulated microsomal
- Fraction preparation not imposing special requirements on storage and trans~ortation
and maintaining its enzymatic activity for more than a year (the time of observation).
Industrial production of the preparation is planned to begin in 1982.
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Flence we arrive at a possibility for creating a single complex which would include,
hesides lyophilized indicator cultures of test bacteria and dry microsomal pr~para-
tion, a complete set of reagents and standard mutagens necessary for positive con-
trol. Creation of such a preparation outfit and broad introduction of bacterial
Cest systems are among the priority tasks of the genetic control service. .
There are other problems requiring immediate solution as well. As of today more
- thaii 3.5 million different chemical compounds are known, and each year up to 80,000
more are synthesized, of which 30,000 are produced on an industrial scale. A
selective study has shown that about 20 percent of them exhibit some level of
genetic activity (2,54). Naturally we could hardly expect testing of all of them
- in the foreseeable future.
We also know that in natural conditions these substances act upon the bodies of man
aiid animals, plants and microorganisms not in isolation but as various combinations,
with the ingredients present in different concentrations. This means that co-muta-
genic action (activation of potential mutagens by nonmutagenic substances) and the
effects of summation, amplification or attenuation are possible (15,25,27,37,45,56).
A large number of observations have shown that temperature fluctuations, the action
of electric and maqnetic fields, light-induced activation, and the composition of
the medium in which mixtures act have a certain influence on the end effect of
mutageiis (20,22,26,34,41).
f3ence arises a new and great important problem--development of ways for determining
the cumulative action of mutagens in the three principal environments (air, soil,
- water). This would allow us to adopt a new, more sensible approach to organizing
the qenetic control service--primary mapping by means of sLUnmational tests, revela-
tion of zones of high genetic danger and subsequent determination of the concrete
causes behind this danger. For the moment we can cite only a limited number of
studies based on this approach. They include an evaluation of the total mutagenic
activity of s~il water in the Surgut petroleum-gas region and of the waters of the
c4ississippi, and mapping of mutagens in the atmosphere above Norway (4,11). A
similar approach would also be sensible for analyzing the possible mutagenic danger
of multiple-ingredient local mixtures--industrial wastes, gas exhausts and so on.
In the opinion of specialists, as reflected in recommendations of a s~nposium of the
Canadian Society of Geneticists, determination and quantitative evaluation of the
action of individual mutagens or their combinations is not yet enough to allow pre-
diction of the end results of change in the gene poo~. of human, ani.mal, plant and
microorganism populations within the ].imits of a closed ecosystem, and all the more
so an open ecosystem. We would have to agree with the opinion that the influence
of mutagens upon a population should be evaluated on the basis of the size of the
qenetic load and the frequency of arisal of hereditary pathology (38). This is why
itlcreasinqly greater significance has been attached in recent years to developing
a:~ystem of genetic monitoring or surveillance of the dynamics of the genetic load
iii 1luman, animal and plant populations in space and in time (9,10,28,29,39,48,51).'
In the Soviet Union, the Interdepartmental Scientific-Technical Council on Complex
Problems of Environmental Protection and Sensible Use of Natural Resources has~fore-
seen development of genetic monitoring of human populations (the Institute of General
Genetics) and a similar system for natural bacterial populations (the Institute of
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- L�:cology of Plants and Animals of the USSR Academy of Sciences Ural Scientific Center).
The object of the observations is the ubiquitous indicator species E. CoZi Corromtrte�
Research is embracing all of the basic units of the species' natural circulation
(healthy people, drinking water, domestic wastes, open-air water basins) and groups
of clones isolated from industrial wastes containing a high concentration of chemical
- mut;actens, from the cooling ponds of atomic electric power plants and from patients
- subjected to intensive antibiotic and chemotherapy.
The system used to describe the gene pool of natural populations includes the
following genetically determined and ecologically significant characteristics:
a) ~rototrophism, or auxotrophy, with the type of trophic dependence being stated;
b) reproduction dynamics on solid and liquid media, with Kp, td, Mm, Xm, t~ and
YE recorded; c) the possibility of utilizing basic carbohydrates and alcohols for
develo~ment; d) antagonism in relation to accompanying bacterial flora; e) sensi-
tivity to bacteriocins produced by satellite microbes; f) resistance to the principal
antibiotics, activated chlorine, ultraviolet light and so on. The obtained results
are subjected to biometric treatment, systems analysis and computer comparison. In
this case E. coZi eorrum~ne is the indicator microbial species, and periodic analysis
of the gene pool of its natural populations would obviously allow us to evaluate the
resulting action of anthropogenically changing natural factors and of regulatory
- mechanisms within populations (6).
- Summing up the published data and the results of our o~m observations, we should note
at th~~same time that bacterial test systems, as with any other method, have their
limitations. Some researchers (40,42,47) feel it impossible to use this approach to
reveal all genetic damage, to test all antimicrobial agents and to reveal carcinogens
activated by other means, apart from liver microsomes. Full quantitative correlation
is not always revealed in experiments using bacterial and other more-complex systems.
But this does not in any way reduce the value of bacterial test systems, which are
a mandatory stage of screening and the simplest, most convenient and fastest model,
and when combined w.ith select.ive tests run on animal cell cultures (5,23), they allow
us to reveal and quantitatively evaluate the genetic effect of all substances, their
mixtures and other environmental factors. This system has already received inter-
national recognition, and it will enjoy even broader acceptance in the future.
BIBLIOGRAPHY
l. Grin, D. and Goldberger, R., "Molekulyarnyye aspekty zhizni" [The Molecular
Aspects of Life], Moscow, 1968.
Dubinin, N. P. and Pashin, Yu. V., "Mutagenez i okruzhayushchaya sreda"
.[Mutagenesis and the Environment], Moscow, Nauka, 1978, 128 pp�
3. Kovalev, I. Ye. and Ma]enkov, A. G., "The Flow of90~lOlgn Substances: The
Effect on Manlcind," PRIRODA, No 9(781), 1980, pp
4. Kolotov, V. M., Nikolayev, A. G. and Pshenichnov, R. A., "Geomikrobiologiya
poiska i razrabotka neftyanykh mestorozhdeniy" [The Geomicrobiology of Exploring
aiid Developing Oil Deposits], Sverdlovsk, UNTs AN SSSR, 1979, pp 138-140.
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5. Pashin, Yu. V., "Practicability of Modern Test Systems for Evaluating the
t~lutagerii~ity of Environmental Factors," in "Geneticheskiye posledstviya zagrya-
zneniya okruzhayushchey sredy" (Genetic Consequences of Environmental Pollution],
Moscow, 2d Edition, 1977, pp 85-88.
6. Pshenichnov, R. A. and Maslov, Yu. N., "Study of the Heterogenicity of Natural
E. coZi Populations as the First Stage of Biological Monitoring," in "Voprosy
genetiki i selektsii na Urale i v Zaural'ye (inform. materialy)" [Problems of
Genetics and Selection in the Urals and Transurals (Information Materials)],
Sverdlovsk, UNTs AN SSSR, 1979, pp 22-23.
7. Fonshteyn, L. M., Revazova, Yu. A., and Vinkler, G. N., "0 nekotorykh podkhodakh
- k otsenke mutagennoy aktivnosti lekarstvennykh preparatov" [Some Approaches to
Evaluating the Mutagenic Activity of Medicinal Preparations], Moscow, Nauka,
1977, PP 42-47.
8. Fonshteyn, L. M., Kalinina, L. M., Polukhina, G. N., Abilev, S. K. and Shapiro,
A. A., "Test-sistema otsenki mutagennoy aktivnosti zagryazniteley sredy na sal-
monellakh" [A Test System for Evaluating the Mutagenic Activity of Environmental
Contaminants Using SaZmoneZZa~, Moscow, VINITI, 1977, 52 pp.
9. Khil'chevskaya, R. I., "Possibilities of Using Morbidity Data and a Number of
Demographic Population Characteristics for the Purposes of Genetic Monitoring,"
in "Materialy zasedaniya Sektsii geneticheskikh aspektov v probleme 'Chelovek i
biosfera [Proceedings of a Meeting of the Section for Genetic Aspects of the
"Man and Biosphere" Problem], Yerevan, 1979, pp 54-57.
10. Shirinyan, G. S., Batikyan, G. G. and Arutyunyan, R. M., "Principles of Popula-
tion Monitoring for the Intensity of Chemical Mutagenesis in the Armenian SSR,"
in "Voprosy molekul~arnoy kletochnoy biologii. Materialy nauch. konf." [Pro-
blems of Molecular Cell Biology. Scientific Conference Proceedings], Yerevan,
1979. PP 81-82.
, 11. Alfheim, I. and Moller, M., "Mutagen1city of Long-Range Transported Atmospheric
Aerosols," SCI. TOTAL ENVIRON., Vol 1, No 3, 1979, pp 275-278.
12. Ames, B. N., "The Detection of Chemical Mutagens With Enteric Bacteria" in
Hollaender, A. (Editor), "Chemical Mutagens, Principles and Methods for Their
Detection," N.Y., Plenum Press, Vol 1, 1971, pp 267-282.
13. 11mes, B. N., Durston, W. E., Yamasaki, E. and Lee, F. D., "Carcinogens are
Mutagens. A Simple Test Combi.ning Liver Homogenates for Activation and Bacteria
_ for Detection," PROC. NAT. ACAD. SCI~ USA, Vol 70, 1973, pp 2,281-2,2$5.
1~. Ames, B. N., McCann, Y. and Yamasaki, E., "Methods for Detecting Carcinogens
and Mutagens With the SaZmoneZZa/mammalian-Microsome Mutagenicity Test,"
MUTATION RES., Vol 31, 1975, pp 347-364.
1`_~. Arisawa, Mikio, Fujiu, Morio, 5uhara, Yasuji and Matuyama, Hiromi B., "Differen-
tial Mutagenicity of Reaction Products of Various Pyrazolones With Nitrite,"
MUTATION RES., Vol 57, No 3, 1978, pp 287-296.
39
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- 16. Ashwood-~mith, M. J., "Stability of Frozen Microsome Preparations for Use in the
Ames SaZmotteZZa Mutagenicity Assay," MUTATIONS RES., Vol 69, No 1, 1980, pp 199-200.
17. Bartelmess, A., "Chemical Mutagenesis in Mamanals and Man," Heidelberg, Springer
Verlag, 1970, pp 69-78.
18. Beije, B., Jenssen, D., Arrhenius, E. and Zetterqvist, M. A., "Isolated Liver
Perfusion--a Tool Mutagenicity Testing for the Evaluation of Carcinogens," CHEM.
BIOL. INTERACT., Vol 27, No 1, 1979, pp 41-57.
19. Braun, R. and Schoneich, J., "The Mutagenicity Testing System Used in the ~
German Democratic Republic," in "Eval. Embryotoxicity, Mutagen and Carcinogen
Risks in New Drugs. Proc. 3d Symp. Toxicol. Test. Safety New Drugs (Prague,
1976)," Prague, 1979, pp 253-256.
20. Chrisman, C. L. and Baumgartner, A. P., "Micronuclei in Bone-Marrow Cells of
Mice Subjected to Hyperthermia," MUTATION RES., Vol 77, No 1, 1980, pp 95-97.
21. Claxton, Larry and Baxter, Richard, "The Computer Assisted Bacterial Test for
Mutagenesis," MUTATION RES., Vol 53, No 3, 1978, pp 345-350.
22. Commoner, Barry, Vithayathil, Antony J., Dolara, Piero, et al., "Formation of
Mur.agens in Beef and Beef Extract During Cooking," SCIENCE, Vol 201, No 4359,
1978, pp 913-916.
23. Hollstein, Monica, McCann, Joyce, Angelosanto, Frank A. and Nichols, Warren W.,
"Short-Term Tests for Carcinogens and Mutagens," MUTATION RES., Vol 65, No 13,
1979, pp 133-226.
24. Ilsie, Abraham W., O'Neill, J. Patrick and McElheny, Victor K., "Quantitative
Mammalian Cell Mutagenesis and Mutagen Screening. A Report on Banbury
Conference II.," MUTATION RES., Vol 64, No 6, 1979, pp 437-440.
25. Isliizawa, W., Utsunomiya, T., Kinoshita, N. and Endo, H., "Formation of Methyl-
nitrosocyanamide From Methylquanidine and Sodium Nitrite in Simulated Gastric
Juice and in Stomachs of Rats: Quantitative Esti.mation by a Mutagenicity Assay,"
J. NAT. CANCER INST., Vol 62, No 1, 1979, pp 71-77.
26. Kada, Tsuneo, Morita, Kaziyoshil and Inoue, Tadashi, "Anti-Mutagenic Action
of Vegetable Factor (S) on the Mutagenic Principle of Tryptophan Pyrolysate,"
MUTATIC~N RES., Vol 53, No 3, 1978, pp 351-353.
27. Kada, Tsuneo and Kanematsu, Nobutake, "Reduction of N-Methyl-N~-Nitro-N-Nitro-
soquanidine Induced Mutations by Cobalt Chloride in Esc12e2yich2a coZ2.," PROC.
JAP. ACAD., Vol E54, No 5, 1978, pp 234-237.
28. Kale, P. Y. and Baum, J. W., "Sensitivity of DrosophiZa meZanogaster to Low
Concentrations of Gaseous Mutagens," MUTATION RES., Vol 68, No 1, 1979, pp 59-68.
29. Klekowski, E. J. Jr., "Screening Aquatic Ecosystems for Mutagens With Fern
Bioassays," ENVIRON. HEALTH PERSPECT., Vol 28, 1978, pp 99-102.
40 ~
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30. Lijinsky, W. and Andrews, A. W. ,"The Mutagenicity of Nitrosami~es in SaZmoneZZa
~typhi~mcriwn," MUTATION RES., Vol 68, No 1, 1979, pp 1-8-
31. Magnusson, J., Hallstrom, I. and Ramel, C., "Metabolic Activution of Vinyl
Chloride in DrosophiZa meZanogaster After Pretreatment with Inducers of Micro-
= somal Detoxication Enzymes," MUTATION RES., Vol 64, No 2, 1979, pp 138-139.
3'l. Maier, P., Philphot, R. M., Nbhn, G. R., and Malling, H. V., "Influence.of Sub-
cellular Fractions of Mam~?alian Testes on the Mutagenic Activity of Nitrofurans
Toward ESCYC22�ZCY12Gi COZZ. Presence of a Co-Mutagen-Like Factor," MUTATION RES
Vol 63, No 2, 1979, pp 223-243.
33. Malling, H. V., "Comparison Between in 'Uitro and in vivo Activation," in "In
vitr~o Metabl. Act. Mutagenesis Test. Proc. Symp.," 1978, pp 143-156.
34. McCoy, E. C., Hyman, J. and Rosenkranz, H. S., "Conversion of Environmental
Pollutant to Mutagens by Visible Light," BIOCHEM. AND BIOPHYS. RES. COMMUN.,
Vol 89, No 2, 1979, pp 729-734.
- 35. McMahon, Robert E., Cline, John C. and Thompson, Christina, Z., "Assay of
855 Test Chemical in Ten Tester Strains Using a New Modification of the Ames
Test for Bacterial Mutagens," CANCER RES., Vol 39, No 3, 1979, pp 682-693.
36. Muller, D., Nelles, J., Deparade, E. and Arni, P., "Studies of the Activity of
S9-Liver Fractions From Various Species in the Ames Test," MUTATION RES., Vol 64,
No 1979, pp 104-105.
37. Nagao, Minako, Yahagi, Takie and Sugimura, Takashi, "Differences in Effects
of Narharman With Various Classes of Chemical Mutagens and Amounts of S-9,"
BIOCHEM. AND BIOPHYS. RES. COMMUN., Vol 83, No 2, 1978, pp 373-378.
38. Newcombe, Howard B., "Problems of Assessing the Genetic Impact of Mutagens on
Man," CAN. J. GENET. AND CYTOL., Vol 20, No 4, 1978, pp 459-470.
39. Nilan, R. A., "Potential of Plant Genetic Systems for Monitoring and Screening
Mutagens," ENVIRON. HEALTH PERSTECT., Vol 27, 1979, pp 181-196.
40. Pini, L. A., Fabio, lJ. and Della, Cass G., "Considerzioni critiche e valuta-
zione di un test microbiologico quale metodo di screening per sostanze poten-
zialmente cancerogene," SECURITAS, Vol 63, No 5, 1978, pp 257-272�
41. Pool, B. L., "Nachweis der mutagegen Wirkung von 5-Methoxypsoralen (Bergapten),"
- ARZTL. KOSNiETOL., Vol 9, No 6, 1979, pp 349-350, 352, 355.
~12. Rinkus, S. and Legator, M., "Deficiencies in the Microbial Activation System for
Mutagenic-Carcinogenic Screening," MUTATION RES., Vol 53, No 2, 1978, p 254.
43. Rinkus, St. L. and Legator, M. S., "Chemical Characterization of 465 Known or
Suspected Carcinogens and Their Correlation With Mutagenic A~tivity in the
ScxZmoneZZa tz~phirrntriwn System," CANCER F~;S., Vol ~9, No 9, 1979, pp 3,289-3,318.
41
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44. Rasenkranz, H. S., "Bacteriological Detection of Carcinogens," UMSCHAU, Vol 74,
1974, pp 745-747.
45. Rosin, M. P. and Stich, H. F., "Inhibitory Effect of Reducing Agents on
N-Acetoxy- and N-Hydroxy-2-Acetylaminofluorens-Induced Mutagenesis," CANCER RES.,
Vol 38, No 5, 1978, pp 1,307-1,310.
46. Sadaie, Y. and Kada, T., "Mapping of Rec-Genes in Bae2ZZus subt2Zis," ANN. REP.
NATL. INST. GENET., Vol 23, 1972, pp 74-75.
47. Serres, Frederik, J. de., "9trength and Weaknesses of Microbial Test R~sults for
Predicting Human Response," J. ENVIRON. PATHOL. AND TOXICOL., Vol 1, No 2, 1977,
pp 43-48.
48. Serres, Frederik, J. de., "Introduction to Utilization of Higher Plant Systems
_ as Monitors of Environmental Mutagens," ENVIRON. HEALTH PERSPECT., Vol 27, 1978,
pp 3-6.
49. Serres, Frederik, J. de. and Shelby, Michael, D., "Recommendations on Data
Production and Analysis Usinq the SaZmoneZZa/Microsome Mutagenicity Assay,"
MUTATION RES., Vol 64, No 3, 1979, pp 159-165.
5U. Simmon, V. E., "In Vitro Mutagenicity Assays of Chemical Carcinogens and Related
Compounds With SaZmoneZZa typhimur?iwn," J. NAT. CANCER INST., Vol 62, No 4,
1979, PP 893-899.
51. Sram, R. J. and Kulesov, N. P., "Monitorovani geneticlych ucinku factori zivotniho
prostzedi v lidske populaci," BIOL. LISTY, Vol 44, No 2, 1979, pp 81-103.
- 52. Tazima, Y., Kada, T. and Murakami, A., "Mutagenicity of Nitrofuran Derivatives
Including Furylfuramide, a Food Preservative," MUTATION RES., Vol 32, 1975,
pp 55-80.
53. "The Foundation of the International Commission for Protection Against Environ-
mental Mutagens and Carcinogens," BIOL. ZBL., Vol 96, No 6, 1977, pp 735-736.
54. Verschaeve, L., "Problemen van mutagenese," TIJDSCHR. GENEESK., Vol 35, No 20;
1979, pp 1,289-1,295.
55. Wiekramasinghe, R. H., ''Short-Ternn Mutagenicity Test Systems for Detecting
Carcinogens. Report of an International Symposium in Dortmund," IND. ARCH.
OCCUP. AND ENVIRON. HEALTH, Vol 44, No 1, 1979, pp 61-64.
56. Yoshida, D., Matsumoto, T. and Okamoto, H., "Interaction Between Amino-Z-Carboline
and, Amino-Y-Carboline an Mutagenicity in SaZmoneZZa typhimux~iwn," MUTATION.
RE;S., Vol 68, No 2, 1979, pp 175-178.
COPYRIGHT: Izdatel'stvo "Nauka", "Ekologiya", 1981
11004
CSO: 1840/90
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~ UDC 581.5+577.391
90Sr AND 137Cs ACCUMULATION BY SOME LOWER PLANTS IN VICINITY OF BELOY.ARSKAYA
ATOMIC POWER PLANT IN THE URALS
Sverdlovsk EKOLOGIYA in Russian No 6, P?ov-Dec 81 (manuscript received 27 Mar 81)
pp 94-97
[Article by M. G. Nifontova and N. V. Kulikov, Institute of. Ecology of Plants
and Aninals, Ural Scientif ic Center, t~SSR Academy of Sciences]
[Text] Considering the future of atomic power, the law~ governing migratir,n and
accumulation of artificial radionuclides in components of natural ecosystems at the
locations of industrial atomic electric power plants (AES) are an extremely important
object of study. Such components may include the lower plants, representatives of
which (the lichens for example) are capable of greater accumulation of radioactive
and stable nuclides (3,4,8).
This paper presents data on the concentration of y~Sr and 137Cs in the thalluses of
the leafy epiphytic lichen Nt~pogymnia phz~sodes (L.) Nyl. and in a number of widespread
species of edible fungi growing in forested areas of the health protection zone of .
the Beloyarskaya AES in the Central Urals: agaric--RusBUla et~cr.xcmtha (Schw.) Fr.,
birch mus~~room--Leceinum scabrum (Fr.) S. F. Gray, pepper mushroom--Lactarius resitmts
Fr., and the valuy [transliteration]--RusBUZa foetens (Fr.) Fr.
The plant samples and their substrates (birch, pine and brushwood bark for lichens
and the upper layer of forest litter and soil to a depth of 0-5 cm for fungi) were
taken from 50x50-meter sampling areas in three to six repetitions in July-August 1978-
1979. Then the material was dried and reduced to ash at 450-50�C. 90Sr was deter-
_ mined* radiochemically by the presence of daughter 90Y~ 137Cs was determined
with an AI-128-2 scintillation gaimna-spectrometer with an 80x80 uun NaI(T1) crystal
and a 24x40 mm socket. The statistical error of the analyses did not exceed 10-15
percent.
As we can see from Figure 1, the concentration of y~Sr in epiphytic lichen growing
_ on various substrates varies from 5.8 to 9.4 nanocuries, while that of 137Cs varies
- from 9.5 to 17.9 nannocuries per kilogram dry weight. These magnitudes do not exceed
the concentration of radionuclides in sfmilar groups of lichens from other regions
of the country subjected to worldwide radioactive atmospheric fallout (4). The con-
ceiitration of 90Sr and 13~Cs in different substrates is practically the same, and
it is several times lower than in lichens growing on these substrates. In this case
we did not note any dependence between accumulation of radionuclides in lichens and
*The authors express their deep gratefulness to S. Kh. Fakhrutdinova and S. I.
Rossal~nko for their assistance in radiometric processing of the preparations.
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rvec vrr~~-~~si, vor_ v~v~.i
tY~Pir concentration in the substrates. The latter indirectly confirms that the
principal source of 90Sr and 127Cs taken up by epiphytic lichens is not the substrate
upon which they grow but atmospheric precipitation and water draining from tree
crowns, enriched by radionuclides to a certain degree.
~ 20
~
~16
~
~12
x
~ 8
~
' , ~
m 4
~
0
1 2 3 4 5 6
Figure 1. Concentration of 90Sr (Empty Columns) and '`~~Cs (Cross-Hatched
Columns) in Thalluses of Hypo~mnia ph~s~:~e~ and Substrates:
- 1-3--lichens (1--from birch, 2--from pine, 3--from brushwood);
4-6--substrates (4--birch, 5--pine, 6--brushwood)
Key:
1. Concentration, nanocuries/kg dry weight
The concentration of radionuclides in edible mushrooms was about an order of magnitude
lower than in lichens (Figure 2), which is explained.mainly by the life span of the
objects studied. Epiphytic lichens usually live several decades, and consequently
they accumulate radionuclides in their thalluses for many years. The life span of the
fruiting bodies of mushrooms is limited to several days (5). Moreover, as was noted
above, lichens accumulate radionuclides from atmospheric precipitation and from water
draininq off of tree crowns; the principal source of radioactive products in the
fungal mycelium is the substrate itself--that is, soil and litter, in which 9~Sr and
137Cs exist predominantly in a bound form with low mobility (2). Figure 2 also
shows that these radionuclides are not readily available to the fruiting bodies of
' mushrooms. We can see that the concenicration of radionuclides in fungi is signifi-
cantly lower than in the substrate. In this case the accumulation coefficients
arc~ only 0.07-0.09 for 9~Sr and 0.18-0.22 for 137Cs. Consequently, given uptake
, ~f both nuclides by the fruiting bodies of mushrooms, they are noticeably enriched
by radioactive cesium in comparison with 90Sr. The concentration of 9~Sr and
1;~Cs in the caps of the fruiting bodies of some mushroom species is somewhat
t~iyher than in the stems (Figure 3). But on the whole the concentration of these
isotopes in edible mushrooms living in the health protection zone uf the Be~~oyarskaya
AT;S does not exceed amounts explainable by worldwide radioactive fallout, inasmuch
as the same concentrations of radionuclides were noted for mushrooms in the Lithu-
- anian SSR (1), the FRG (6) and a number of other countries.(7).
44
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F()R OMFI('IA1, ll~M: ON1.Y
80I
1
El),
~ 2,8
~ '
v 2,4
~
~
i ~
)a
~ 1,6
: $
~ 1,2
0
Q8 -
Q4 -
0 .
1 2 3 4 S
Fi~Ture 2. Concentration of 90Sr (Empty Columns) and 137Cs (Cross-Hatched
Columns) in Fruiting Bodies of Agaric Mushrooms and in Sub-
strate: 1--RussuZa cr~axantha; 2--Leccinum scabrwn; 3--Laet,ar?ius
resimus; 4--RussuZa foetens; 5--soil
Key: .
1. Concentration, nanocuries/kg dry weight
~ 2,0 - n'CS
v
~ 1,6
~
~l?~ 1,2 -
~ .
a
_ ~ 0~8 - .
Q
~ Q4
~
0
0
1 ~ 3 1 2 3
- Figure 3. Concentration of 90Sr and 137Cs in the Caps (Empty Coltunns) and
- Stems (Cross-Hatched Columns) of Mushroom Fruiting Bodies:
1--Russula foetens; 2--RussuZa cn~axantha; 3--Lactarius resimus
Key:
1. Concentration, nanocuries~kg dry weight
45
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rvn vrriwMa, a,oc v~~i.~
Tlius tYie data presented here permit the assumption that the concentration of 90Sr
and 1j~Cs in epiphytic lichens and edi.hle agaric mushrooms in the vicinity of the
Beloyarskaya AES does not exceed the background concentration of these radionuclides
typical of similar plants from other habitats outside the possible influence of the
AE5.
BIBLIOGRAPHY
1. Dushauskene-Duzh, R. F. and Urbonas, V. A., "Migration of 210Pb and 90Sr in
Terrestrial Ecosystems," EKOLOGIYA, No 5, 1978, pp 97-100.
"l. Kulikov, N. V. and Molchanova, I. V., "Kontinental'naya radioekelogiya"
[Continental Radioecology], Moscow, Nauka, 1975, 184 pp.
3. Moiseyev, A. A. and Ramzayev, T'. V., "Tseziy-137 v biosfere" [Cesium-137 in the
Biosphere], Moscow, Atomizdat, 1975, pp 152-166.
4. Nifontova, M. G. and Kulikov, N. V., "Accumulation of Strontium-90 and
Cesium-137 by Lichens in Nature," EKOLOGIYA, No 3, 1977, pp 93-96.
5. Serzhanina, G. I. and Zmitrovich, I. I., "Makromitsety" [Macromycetes], Minsk,
Vysheyshaya shkola, 1978, 190 pp.
6. Grueter, H., "Product 137Cs in Mushrooms in W. Germany During 1963-70," HEALTH
PHYS., Vol 20, No 6, 1971, pp 655-656.
7. Haselwandter, K., "Accumulation of the Radioactive Nuclide 137Cs in Fruiting
Bodies of Basidiomycetes," HEALTH PHYS., Vol 34, No 6, 1978, pp 713-715.
8. Mattsson, S., "Radionuclides in Lichen, Reindeer and Man," Lund., 1972, 48 pp.
COPYRIGHT: Izdatel'stvo "Nauka", "Ekologiya", 1981
11004
CSO: 1840/90
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UDC: 502.7
COLLABORATION OF U.S. AND SOVIET SCIENTISTS IN LEGAL PROTFCTIQN
OF THE ENVIRONMENT
- Moscow VESTNIK AKADEMII NAUK SSSR in Russian No 8, Aug 81 (signed to press 4 Aug
81) pp 102-109
(Article by 0. S. Kolbasov and M. I. Kozyr', doctors of law]
[Text] In the second half of the 20th century, environmental protection has
become one of the most serious global problems, the solution of which requires
coordinated effort on the part of all countries of the world.
Enormous and valuable knowhow has been accumulated in the Soviet Union in the
area of environmental protection; relevant laws have been elaborated and
successfully implemented. The communist party and Soviet government attribute
much importance to further intensification of ineasures for protection of the
environment and devote their constant attention to this matter.
Article 18 of the USSR Constitution states: "In the interests of the present
and future generations, the necessary steps are being taken in the USSR to
protect and make wise use, with scientific validation, of the land and its
mineral resources, water resources, the plant and animal kingdom, to preserve
the purity of air and water, implement reproduction of natural resources and
improve man's environment." There is a special section in "The Main Directions
of Economic and Social Development of the USSR in 1981-1985 and for the Period
up to 1990," which were approved at the 26th CPSU Congress, concerning environ-
mental protection, and in the section listing the main objectives of economic
- and social development of our nation for the future peri~d it is stated: "There
must be intensification of protection of nature, the land and its mineral re-
sources, atmospheric air, water reservoirs, animal and plant kingdom. Wise
use and reproduction of natural resources must be implemented."
Tlie Soviet Union is actively in favor of continued deve~.opment and deepening
of international collaboration in the area of environmental protection. Con-
structive proposals on this subject are constantly offered at party congresses,
sessions of the USSR Supreme Soviet, in speeches of Soviet representatives in
various international forums. ~
~ Mucl? work on environmental protection is being pursued in the United States of
America; considerable knowhow has been accumulated there, particularly with re-
gard to refinement of laws and management in this area.
47
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1?? view oF the ~~normous importance and acuity of the problem, as well as the
need tu combine forces for. environmental protection, the governments of the
USSR and United States signed an agreement on collaboration in the area of
environmental protection on 23 May 1972, in Moscow.
Soviet-American collaboration is directed at solving the main aspects of the
- problem of the environment and developing bases for regulating the effects of
human endeavor on nature with, of course, consideration of the differences in
socioeconomic and political systems of these two countries. The main aspects
of this collaboration are: prevention of air and water (including the marine
environment) pollution; protection of the environment against the adverse
effects of agriculture and other industries, improvement of the ecological en-
vironment of cities, organization of reservations, forecasting ea rthquakes,
study of biological and genetic consequences of environmental pollution,
regulation of the effects of human endeavor on climate, investigation of the
_ distinctions of arctic and subarctic ecological systems, analysis of various
legal and administrative measures to preserve the quality of the environment.
One or several joint projects and collaborating organizations were assigned
for each of the above problems. The forms of work were defined: exchange of
scientists, specialists and trainees, as well as scientific and technical
information, documentation and results of research, holding joint symposiums,
conferences and meetings of experts, joint development of specific scientific
programs, etc. The plans for such work are approved at annual meeting~ of
the Soviet-American commission, which isjointly headed by the chairman of
the USSR State Committee for Hydrometeorology and Control of the Environm ent
and director of the Environmental Protection Agency (EPA) of the United
States.
Iri view of the fact that work on legal and administrative measures for environ-
mental protection touches upon many common problems of government and law, the
Institute of Government and Law, USSR Academy of Sciences, was name3 as the
chief organization for the Soviet side (since 1972, there has been a sector
at this institute working on legal problems of environmental protection).
Ttie Council on Environmental Quality under the U. S. president is the chi.ef
organization on the American side.
Since this collaboration started, there have been five 2-week conferences of
experts (in 1973, 1976 and 1980 in the United States; in 1974 and 1978 in the
US5R); two American specialists spent 5 months in the Soviet Union in training,
there was regular exchange of literature and information on legislation and
practical use thereof, consultations were offered on different questions of
law and management in the area of environmental protection, assistance was
given in establishing direct contact between public environment protection
organizations of the USSR and the United States. Of course, personal meetings
of specialists and expert conf erences were also very important.
At tlie meetings of specialists, there was discussion of such important topics
as ttie correlation between international and national law in environmental
protection, correlat~on between law and management on different levels (all-
Union--federal, republic--state, etc., down to the level of individual enter-
~~rises); legal forms of participation of the USSR and United States community
in environmental protection.
48
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Exchange uf opinions, knowhow and information on these pressing problems,
joint work on theoretical conceptions in the area of law and administration
are instrumental in mutual enrichment of theory and practice on both sides,
dissemination over the entire world of knowhow in environmental protection of
the two large industrial countries, strengthening the spirit of mutual under-
standing of the Soviet and American peoples.
While visiting the country in which a meeting of experts is convened, the
scientists of ttie other side have the opportunity to learn about environmental
protection in the host country, the nature of activities of environment pro-
tection organizations, the work of reservations and national parks.
Thus, American specialists, who came to the Soviet Union in 1974 and 1978
learned about the wo~k of state agencies and public organizations concerned
with environmental protection in Moscow and Moscow Oblast, Kazakh SSR, Kiev,
Baku, Tashkent, Vladimirskaya and Rostovskaya Oblasts. In particular, they
visited the Botanical Garden of the Institute of Botany imeni V. L. Komarov,
Azerbaijan Academy of Sciences, the Kanevskiy Preserve and ma.rine oil wells in
Baku. They talked with the staff of the Presidium of the USSR Supreme Soviet,
chairman of the Commission for Environmental Protection of the RSFSR Supreme
Soviet and members of the Presiditun of the Supreme Soviet of Azerbaijan SSR,
representatives of the USSR Supreme Court, USSR Mi.nistry of Justice, RSFSR
Ministry of Agriculture, Vladmirskaya and Rostovskaya oblispolkoms, agencies
for protection of fish reserves and supervision of. hunting, administrators of
tt?e All-Russian and Kazakh environmental protection societies, as well as
Rostovskaya Oblast organization of the All-Russian Society for Environmental
Protection, etc. In 1978, an agreement was made for establishment of direct
contact between the All-Russian Society for Environmental Protection and the
American Sierra Club, and in accordance with this agreement, American public
figures concerned with environmental protection made their first visit to the
- USSR in 1979.

~
American specialists and public figures who visited the USSR praised the work
conducted in our country in the area of environmental protection.~ Such a
rating was reflected in the speech of (J. A. Basterud}, a member o~ the
Council on Environmental Quality, at the Internal Expo--74 exhibition
in Spokane. Prof T.(Shoenbaum), who studied for several months the organiz~~
tional and ~legal measures for the protection of reserves in our country,
published an article entitled "Protected Preserve Regions in the Soviet Union
and United States. Comparative Aspect," in the AMERICAN JOURNAL OF COMPARATIVE
LAW, in 1976, where he shed light on our achievements in this area and re-
commended that some elements of Soviet knowhow in legal regulation of protec-
tion of reserves be used in the United States. Representatives of American
public organizations who visited the USSR in 1979 gave a high rating to the
statutes in the USSR Constitution pertaining to environmental protection.
In turn, Soviet specialists had the opportunity of learning about organization .
and content of the work of the Council on Envirotunental Quality, EPA
and some of its peripheral departments, the work of the U. S. comgreas perman-
ent commissions, the Departments of Justice. The Interior, Agriculture, the
Atomic Energy Commission, the ii. S. Supreme Court and certain other regional and
st~ite courts, environmental protection organizations in Washington, New York,
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Boston, San Francisco and Los Angeles. They visited Grand Teton, Yellowstone
and Grand Canyon National Parksr learned about the work on redeveloping land
in the coal mines of the (Deker) Mine Company (Montana), protection of water
and atmospheric air at the Kennecott Copper copper and molybdenum enterprise
(Utah), and participated in seminars in Denver (Colorado) and Salt Lake City
(Utah).
The fifth conference of Soviet and American specialists on legal and adminis-
trative aspects of environmental protection was held in December 1980, in the
United States (Washington, Miami, Atlanta and New York). The American delega-
tion was headed by F. Knight, who performed the duties of the chief jurist of the
host organization, the Council on Quality Environmental. The delegation con-
~ sisted of prominent specialists: N. Yost, director of the president's working
team for preparation of a global ecological forecast up to the year 2000, J.
Moorman, head of the Land and Natural Resources Division, U. S. Department of
Justice, G. (Widman), deputy director of the ~uridical service of the Dept.
of the Interior, E..Greenberg, head of the juridical service of the National
Oceanic and Atmospheric Admin. of the Dept. of Commerce, (V. Smith), chief
economist of the EPA, N. Robinson, member of the board of directors of the
Sierra Club, 0. Hook, vice-pres~dent of the National Wildlife Federation,
(V. Futrell), president of the Institute of Environmental Law, members of
different departments and agencies, K. (Hall), B. (Bramble), G. (Osherenko)
_ and D. (Shay).
The Soviet delegation included 0. S. Kolbasov (head), M. I. Kozyr~, Yu. S.
Shemshuchenko, A. M. Kaverin and Yu. A. Starikov.
In his opening remarks at the conference, F. Knight stressed the fact that
collaboration of the United States and the USSR on a very difficult problem
affecting the interests of all c:ountries of the world--enviranmental protection--
was quite fruitful.
0. S. Kolbasov noted the importance of Soviet-American collaboration in rhfl
area of environmental protection, indicating its place and role in the system
of ineasures to preserve peace, improve the general political climate, as w~ll
as living conditions. He informed the audience of ineasures impleme~eted in
recent years in the USSR to protect the environment and stressed that a very
important place is given to environmental protection in the draf t of the "Main
Directions of Economic and Social Development of the USSR in 1981-1985 and
for the Period up to 1990."
Uur.ing the meeting, Soviet and American specialists discussed issues related to
tlie need and mandatory nature of considering ecological aspects and questions
pertaining to the environment in elaborating construction plans, with analysis
of tlie role in this process of the U. S. EPA and analogous state organizations.
In particiilar, determination was made of the extent to which this agency takes
into consideration economic factors in making decisions related to environmental
protection. We sensed a concern on the part of the agency with the fact that
the supposedly strict laws, as well as EPA rules for environmental protection,
which supposedly held back development of the economy were submitted to ex-
cessive criticism during the period of the pre-election campaign in the United
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Seaces in 1980, and the opinion was voiced that it was necessary to strictly
"straighten out" the EPA officials and reduce expenses for environmental
protection.
During the meeting there was also comprehensive discussion of global environmental
protection, international mechanisms of assessing the ~~nvironment and problems
of environmental protection in the entire world.
American specialists acquainted the Soviet scientists with an abstract published
in July 1980 of a report to the president of the United States, which contained
an ecological forecast for the year 2000.
In the opinion of the authors of the report, in the year 2000 earth's
population will be 6.35 billion, and 90% of the population growth will
occur in developing countries. By that time, foodstuff prcduction
will almost double; however, the increase per capita will be less
than 15%. There will be an even greater discrepancy between the
economic welfare of the people of developed and developing countries:
industrially developed countries (one-quarter of the worZd's popula-
tion) will, as before, consume three-quarters of the worldwide
mineral resources. A water shortage will increase in some parts of
tlie world. There will be a 50% decrease (per capita) in stock of
growing timber of commercial size; in developing countries, about 40%
of the remaining forests will be removed. In view of the serious
changes in living cond~~ions, perhaps up to 20% of the existing species
of animals and plants will disappear forever. It is expected that the
atmospheric concentration of nitrogen dioxide and other chemicals that
destroy ozone will increase at such a rate that they will alter appre-~
ciably earth's cli~nate by the year 2050. In spite of the fact that
an increasing effort is being applied at the present time to protect
the environment, in the opinion of the authors of this report this
i~ obviously insufficient and cannot alter the negative ecological
trends. At the same time, the authors did not take into considera-
tion the real economic anci political diff iculties that prevent people
from forming their relations with nature more wisely, and this in-
creases even more the seriousness of the looming problems.
No concrete plan oF actions was advanc~d in this report to correct the ecologi-
cal situation (work is not yet finished on the third volume thereof, which
- contains recommendations on models of behavior of U. S. management in implementa-
Cioii of ecological policies). It merely offers general recommendations: to
- cii5play more initiative in solving ecological problems, elaborate global
cum~lex and long-term programs of environmental protection that should "inaugurate
an era of unprecedented collaboration and selflessness of all peoples of the
- wor.ld. The authors of this report maintain that, at the present time, all
of the opportunities are available, as well as weighty grounds for the United
StaCes to become a world leader in the movement for environmental ~rotection.
Soviet specialists found the report quite interesting and considered that its
- material merited carefu.l investigation. At the same time, they stressed the
f.act that so long as there is a real threat of thermonuclear war because of
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Chuse wlw oppose detente, and there is intensification of the arms race, it is �
difficult and unrealistic to speak about"unprecedented international collabora-
tion" in the area of environmental protection. As for leadership of the
United States in tl->e international environmental protection movement, it could
be accepted by the international community only if it is implemented with no
detriment to and not at the expense of other nations, if it becomes a leader-
sfiip through good examples and assistance to needy nations.
_ 'fhe scientific papers delivered by members of the Soviet delegation offered in-
depth and comprehensive analysis of different aspects of existing Soviet
legislation for environmental protection, including preventive measures, and
it was noted that questions of environmental protection are taken into compre-
liensive consideration in our country in preparing construction plans, and that
a thorough ecological expert evaluation is made of such plans. The speakers
~;ave detailed information on the content of new Soviet laws on protection of at-
mosptieric air, protection and use of the animal kingdom; they told about the
pr~cedures for planning wise use of natural resources and environmental protec-
tion in the area of agriculture, the juridical guarantees for land protec-
tion and improvement of soil fertility.
Ttie American participants displayed much interest in the information about deve-
lopment and use of laws for environmental protection in the USSR, and voiced
tlieir interest in strengthening Soviet-American collaboration in the area of
environmental protection.
During their stay in the United States, the Soviet specialists had comprehensive
tallcs with American specialists dealing with legal arid administrative problems
of env?.ronmental protection in the USSR and United States, the EPA, U.S. Depart--
ments ot The Interior, Commerce and Justice, as well as at the Kennan Institute
and Institute of Environmental Law in Washington. They discussed with M. (Wilkie),
a member of the Federal Court of Appeals of the District of Columbia, the question
of the role of U. S. courts in the area of environmental law. They had talks
with ~overnment officials of several states and some local agencies on the
effectiveness of using federal and state laws, rules and procedures for environ-
~ mental protection, with representatives of U. S. public organizations concerned
wit}i environmental protection, in particular, with activists of the Sierra Club
and National Wildlife Federation, as well as prominent practicing
jurists concerned with legal problems of environmental protection and professors
on the faculty of law at Pace University.
'Itie members of the Soviet delegation visited the national Everglades preserve,
a farm near Miami (Florida), a heat and electric power plant and regional
department of the EPA in Atlanta, the department for environmental protection
ot the state of ~iew Yurk, a gas plant near Calvert Cliffs (Maryland) and were
received the Mayor of Greenburg, L~ew York.
During these meetings, American figures in the area of environmental protection
~ displayed much interest in Soviet knowhow and praised it highly. Thus, J. W.
Moorman, assistant attorney general; Land and Natural Resources Division, Dept.
of Justice, voiced his admiration to Soviet specialists for setting up well the job
of protecting the environment in the USSR; in several of_ his speeches Prof
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ri.A. Robinson rated highly the collaboration between the Soviet Union and the United
States, with reference to legal and administrative problems of environmental
protection, repeatedly citing articles 18 and 58 of the USSR Constitution as a
model of responsible attitude toward environmentat protection, calling for
further expansion of collaboration in this area. The major American special-
ists, N. Yost and 0. Hook, made similar statements.
How can one assess the current status of American law and government, and their
- role in environmental protection?
- In ~he opinion of Soviet specialists, some progress in legal protection of
atmospheric air and coastal zones of adjacent seas has been made in the United
- States in the last decade (in previous years, there were mainly measures to
protect waters and regulate land use on the basis of territorial zoning).
Worthy of especially careful study is, in our opinion, the effective use in the
United States of a new legal institution, "Declarations on Effects on the
F.nvironment," referable to construction of various industrial pro3ects and
facilities (gas and pipe lines, dams, hydroelectric power stations, irriga-
tion canals, etc.), which is a rather powerful means of preventive control
[inspection] that American specialists believe helped increase significantly
the ecological validation of management decisions avoid mistal~es and improve
effectiveness of environmental protection. The i~merican specialists recommend
the use of this form by other countries, as well as in international relations.
This system of preliminary ecological control is implemented in industry in
accordance with the Law on National Policy in the Area of Environmental
Yrotection (January 1970) and analogous laws in certain states. The
laws stipulate that, at specific times prior to construction and remodeling of
enterprises and other economic facilities, the entrepreneurs and state agencies
must make a public "declaration on effects on the environment" of these instal-
lations. The contents of the declaration are analyzed for a specific time by
state inspection [control] agencies, public organizations and the local public,
who can voice objections for ecological reasons against a particular project
as a whole, propose alternative variants or corrections to the plan. Entre-
preneurs and state agencies must examine the corrections and, within a specific
time, announce whether they are accepted or rejected. If the authors of the
comments and proposals are not satisfied with the results of this examination,
they can tile suit to force implementation of their suggestions. To date,
several tens of thousands of declarations on effects on the environment have
been examined on different levels of the American governmental structure; up to
5i of them have been disputed in the courts.
H~wever, by far not all of the important aspects of legal environmental protec-
tion in tlie United States could be studied during the years of collaboration.
'ffiere is still a need for deeper investigation of the system, functions and
i?iteraction in environmental protection of American state agencies, which
include, in particular, the U. S. Department of Justice, Council on Environmental
Quality under the U. S. president and the Environmental Protection
Agency.
TI~~~ system of preventive environmental protective measures in U. S. agriculture,
~~s w~ll ~s the adverse effect of industrialization and particularly use of
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ctiemistry in agriculture on the environment have not been adequately studied
either.
It is of ~;reat in~erest to Soviet jurists--scientists and practitioners--to
study the practice of examining environmental protection cases in the American
courts and to analyze measures of liability [as to property] for the detriment
to the environment (for example, fines of $25,000 for each day of violation).
International environmental protection measures of the United States, their
place in the general policy of this state, as it applies to different parts of
the world as a whole, but particularly as it relates to food aid for develop-
ing countries, also merits investigation. Finally, we must also study ttie as-
pects of American theory and practice that are related to development of inter-
national environmental laws.
As for assessment of the current status of Soviet-American collaboration in
law and administrative problems of environmental protection, it was formulated
as follows in the summary minutes of the meeting: "Both sides observed that
the fifth meeting was successful and that there was satisfactory exchange of
information and knowhow. The specialists conceded that there is a need to
continue with the exchange of opinions, information and documents, and observed
ttiat collaboration on environmental issues on a worldwid,e scale is very
important and merits increasing attention."
The minutes also formulated the main directions of collaboration in 1981-198~.
As before, there will be continued exchange of information and opinions concern-
~ ing elaboration and use of legislation on environmental protection in the USSR
and United States. There will be continued exchange of information about
research and evaluation of effects on the environment, both within each nation
and on earth as a whole, about comparison of economic factors to environmental
factors, implementation of laws pertaining to the environment. E~:change of
scientists and students concerned with legal and administrative aspects of
environmental protection will be continued.
It was also decided to examine the question of the means of future collaboration
of the USSR and United States in the area of long-term problems of environmental
protection and wise use of natural resources on an international scale.
It is planned to discuss all tliese problems at the sixth meeting of Soviet and
American specialists, which is to be held in the USSR in 1982.
Th~: participants at the meeting agreed to explore the possibility of joint
preparation and publication in both countries of a collection of articles
dealing with evaluation of environmental protection on the national and inter-
national levels, use of legislation, as well as analysis of administrative
. measures in the area of environmental protection that are being implemented
in the USSR and United States.
With reference to the results of 9-year Soviet-American collaboration on legal
aiid administrative problems of environmental protection, it should be noted
tli:?t the main actiievement was deeper knowledge of theory and practice of legal
rey;ulation in the area of environmental protection, in both countries and in
ttie system of international law.
~ 54
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Aside from its direct professional usefulness, collaboration was instrumental
in maintaining a spirit of trust and mutual understanding between the Soviet
and American people.
COPYRIGHT: Izdatel'stvo "Nauka", "Vestnik Akademii nauk SSSR", 1981 .
10,657
CSO: 1840/120
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MEDICINE
UDC: 613.644
LOW-FREQUENCY ACOUSTIC VIBRATIONS IN INDUSTRY
Moscow NIZKOCHASTOTNYYE AKUSTICHESKIYE KOLEBANIYA NA PROIZVODSTVE in Russian
1981 (signed to press 22 Aug 80) pp 2-5, 190-191
[Annotation � introduction and table of contents from book "Low-Frequency
Acoustic Vibrations in Industry" by Nadezhda Ivanovna Karpova and Eduard
Nikolayevich Malyshev, Izdatel'stvo "Meditsina", 1266 copies, 192 pages]
(Text] This monograph is concerned with hygienic evaluation of infrasonic
- waves in industry, their biological effects on man and animals, and prevention
of their deleterious effects. For the first time, information is systematized
concerning the physical nature and distinctions of infrasonic waves. Techniques
are described for measurement and esti.mation of infrasound in industYy. Methods
are discussed .for conducting tests on man and experiments on animals to study
reactions to low-frequency vibrations. The authors' own data, as well as
those of the staff of the Department of Industrial HygienP, Leningrad Medical
Institute of Sanitation and Hygiene, obtained using modern investigative
methods, which describe the influence of industri.al parameters of subsonic
vibration on the subjective state of man and functional changes occurring in
the central nervous, cardiovascular and respiratory systems, as well as analyzers
and viscera are presented. Special attention is given to the approach to setting
standards of subsonic vibrations in industry. In conclus3on~ measures are listed
for the prevention ot the deleterious eff ects of industrial infrasonic waves of
specific parameters and medical preventive measures. This~monograph fills a gap in
- the literature# since there are no sucb w~orks eitfier in our country or abroad. It
is intended for industrial hygienists, occupational pathologists, industrial
health inspectors [physiciansJ, engineering and technical workers, as well as
uther specialists concerned with the study of industrial noise, control of noise
and infrasonic waves. ~
Introduction
The decisions of the 25th CPSU Congress provide for further improvement of work-
ing conditions that would be instrumental in lowering morbidity and increasing
labor productivity in all sectors of industry.
Intensive scientif ic and technological progress is leading to development of
hig}i-power equipment and machines. However, operation of such equipment is
not infrequently associated with considerable noise. Infrasound, which did
not attract the attention of scientists for a long time, is a constant companion
- of noise.
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Infrasound varying in in+tensity and frequency is always present in our environ-
ment, particularly in large cities with well-developed industry and intensive
transportation. Man may be exposed to low-frequency sonic vibrations at work
- and at rest.
There are natural sources of infrasound. Many natural phenomena, such as earth-
quakes, eruptions of volcanoes, thunder, landslides, hurricanes, storms at sea,
large waterfalls, aurora borealis, generate infrasonic waves. In addition to
natural sources, more and more often man is exposed to increasing levels of
infrasound occurring in industry and transportation as a concomitant factor of
technological processes.
Infrasonic waves are formed during operation of compressors, turbines, diesel
engines, electric locomotives, air-conditioning systems, commercial fan~ and
other large machines and equipment.
With the development of technology, machines are appearing that are large in
size, operating at high speed and high power, whose function is associated with
excitation of intensive low-frequency sonic waves. Modern technology is
increasing the noise background in industry, which has a broad spectrum and
increasing intensity.
Various forms of transportation are also artificial sources of infrasound:
seafaring, railroad, aviation, including jet aircraft and the high-power
engines used in space flights (R. W. Stephens, 1969, 1974; W. Tempest, 1971).
Man's exploration of space generates noise during launchings that has maximum
energy in the low-frequency part of the spectrum, in the range,,of 1 to 100 Hz.
Infrasonic vibrations are not only a component, but in many cases the pre-
vailing part of spectra of industrial noises. At the present time, the effects
of industrial noise in the acoustic range of frequencies on man have been studied
rather well.
There is extremely little information in the Soviet and foreign literature about
the biological effects of low-frequency vibrations. The data in the literature
are disparate and often contradictory. There are few sources with systematized
data on the biological eff ects of infrasound.
Studies have revealed that man is sensitive to low-frequency vibrations and
compel us to admit that the problem of studying the effect of infrasound on
man merits the very closest attention.
Infrasound may be one of the deleterious factors of the industrial environment.
Certain levels oL infrasound may be deleterious to human health. It affects
tlie entire human body, man's health and work capacity.
Most workers in modern industry and transportation are exposed to such a~little-
studied factor as infrasound. This puts to industrial hygienists the task of
com~~rehensive investigation of the distinctions of its biological action in order
to determine the degree of harm of low-frequency sonic vibrations, set hygienic
standards and elaborate measures to eliminate the deleterious effects af some
- parameters of low-frequency vibrations on workers.
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liecause of the distinctions of propagation of infrasonic waves and their dif-
ference from audible vibrations, the study of biological effects of infrasound
requires new methodological approaches to set the scientif ically substantiated
ran~;c: of permissible levels of infrasonic pressure and develop optimum amelio-
r~tive measures. It is very important to determine the threshold intensity of
infrasound with acute and chronic exposure to it, as well as to demonstrate the
physiological mechanisms of the body's reactions to infrasound.
Tlie objective of this book is to acquaint the reader with infrasound as a del~-
terious factor of the industrial environment; as well as experimental data
from studies of biological effects of infrasound of industrial parameters on
animals and man.
This work summarizes the knowhow of Soviet and foreign authors in the area of
studying the biological effects of low-frequency vibrations, as well as the
results of studies conducted by t~e authors and staff inembers of the Department of
Industrial Hygiene of the Leningrad Medical Institute of Sanitation and Hygiene
. and Department of Industrial Safety of the Leningrad Institute of Railroad
Engineers.
It is imperative to widen the �ront of scientific research in this interesting
brarich of science.
CONTENTS Page
Introduction 3
Chapter 1. Physical Characteristics and Distinctions of Low-Frequency
Acoustic Vibrations 6 ~
Concept of infrasound 6
Piechanism of formation of infrasonic vibrations 8
Pliysical characteristics of infrasound 10
Distinctions of propagation of infrasonic vibrations 12
Rate of propagation of infrasound 15
Chapter 2. Methods for Measuring and Evaluating Low-Frequency
Acoustic Vibrations in Industry 17
~ Analysis of existing methods of ineasuring noise and possible use
thereof to record noise in the infrasonic range of frequencies 17
Equipment and methods of ineasuring noise in the infrasonic range 19
Chapter 3. Hygienic Characteristics of Industrial Sources of
Inf rasonic Waves 2~
Investigation of infrasonic waves in industry 27
Evaluatior. of infrasonic vibratiQn of compressors 33
Evaluatic,n of infrasound of vibrating surfaces 38
Infrasonic vibration of transportation.vehicles 42
Cha~ter 4. ;~~o?.ogical Effects of Low-Frequency Vibrations in the
Infrasonic Range 48
Chapter 5. Generation of Infrasonic Waves Under Experimental
Conditions 63
Chapter 6. Methodological Bases of Conducting Research for the .
Study of Effects of Industrial Infrasonic Vibrations 72
Experimental complex for bielogical studies and simulation of
infrasonic field occurring in industrial premises 72
Methods of studying the biological effects of infrasound 75
58 ~
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(;ha~[~r 7. Investigation of Lffects of Infrasonic Vibrations on
Experimental Animals 80
Behavioral reactions of animals 80
Investigation of functional state of cells of the central nervous
system and viscera by the method of intravital tissue stain 81
Investigation of functional state of the ra~bit's central nervous
and cardiovascular systems 82
Investigation of microcirculation in the pYa mater of white rats 9U
Amounts and distribution of nucleic acids in receptor cells of
Corti's organ 92
Investigation of enzymatic and metabolic changes in experimental
animals 94
Chapter 8. Histological and Electron Microscopic Changes in Various
Organs and Systems Under the Influence of Infrasound 108
Morphological changes in viscera 108
Reactive changes in brain cells 115
Investigation of effects of infrasonic vibrations on biological
objects 122
Chapter 9. Investigation of Effects of Infrasonic Vibrations on
Human Physiological Functions 126
Subjective perception of infrasonic vibrations 126
Effect of infrasonic vibrations on functional state of the
central nervous system 132
Effect of intrasonic vibrations on the cardiovascular system 134
Effect of infrasonic vibrations on the respiratory system 139
Condition of some analyzer systems 140
Changes in the auditory analyzer 140
Changes in the vestibular analyzer 145
Changes in skin's sensitivity to vibration and temperature 147
Physiological study of the effects of infrasound in itidustry 149
Investigation of eff ects of infrasonic vibrations on man 158
Maximum permissible levels of infrasound 161
Chapter 10. Means of Prevention of Deleterious Effects of Low-
Frequency Acoustic Vibrations on the Organism . 166
Medical preventive measures 166
Technical methods of controlling low-frequency sonic vibr~ations 166
Absorption of infrasound 171
Ultrasonic insultation 175
Low-frequency dampers for attenuation of infrasonic waves
generated by aerodynamic processes in industrial installations 177
llesign proposals for reduction of noise from vibrating surfaces 181
13i.b] iogr.aphy 183
COPI'I:iCHT: Izdatel'stvo "Meditsina", Moskva, 1981 ~
10,657
CSU: 1840/119
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HUMAN FACTORS
UDC: 629.7.05
ANALYSIS AND OPTIMIZATION OF CONTROL SYSTEMS FOR MANNED AIRCRAFT
Moscow ANALIZ I OPTIMIZATSIYA SISTEM UPRAVLENIYA PILOTIRUYEMYKH LETATEL'NYKH
APPARATOV in Russian 1981 (signed to press 2 Jun 81) pp 2-8, 200
[Annotation, introduction and table of contents from book "Analysis and
Opti.mization of Control Systems for Manned Aircraft", by Grigoriy Illarionovich
Ryl'skiy,.Izdatel~stvo "Mashinostroyeniye", 1240 copies, 200 pages, illus-
trated]
[Text]. This book analyzes the performance of operators in control systems (CS)
of manned aircraft, as well as the possibility of describing it mathematically.
A classification is proposed and analysis made of modern mathematical models
that describe certain forms of human work in aircraf t control systems. The
advantages are shown of models that make use of queueing theory, and new
engineering methods were developed on the basis of these models for demon-
~ strating the characteristics of human operators. This book is 3ntended for
engineers involved with the design and operation of a broad class of man-
machine control systems.
Intr~duction
The cybernetic problem of ref ining control of various systems (engineering,
biological, economic, management [organization], man-machine, etc.) is a
rather pressing oneo The most important element of this problem is to find
optimuin variants of organization of structure and processes of control in
such systems. The concept of system emerged as a result of development of en-
gineering disciplines, since it is expressly the increasing complexity of
various technical systems that made it necessary to elaborate methodology that
would help integrate all parts of such systems. At the present time, there
are quite a few def initions of systemo However, there is still no conven-
tional [generally used] definition. Each researcher gives this concept his
own special content and singles out essentially the element of the system that
he submits to investigation (see, for example, the works of foreign [2, 58,
79, 92, 101, 107, 117, 121, 129, 135, 136] and Soviet [7, 43, 77, 78, 109, 113,
119, 125, 130, 146, 147] authors).
In this book, system is used to refer to the aggregate of elements that are
interlinked into a specific structure. With such an approach, industrial,
educational and research organizations, man-machine systems, equipment, etc.,
can be considerQd to be a system.
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Systems with infinite diversity can be classif ied according to a number of ,
features: purpose, complexity, level of automation and others.
_ Control systems (CS) are an extremely important class of systems, which are used
to control other systems (aircraft, plant, etc.). Consequently, such system9
must have the purpose of functioning and, moreover, must have the capacity
to alter the parameters (characteristics) and structures of elements and of the
entire system in accordance to this functioning purpose. Control systems, in
which man is an element, are called automated, ergatic, anthropotechnical, or-
ganizational systems or man-equipment (man-machine) systems. It must be noted
that all of these definitions refer chiefly to man-machine control systems
(MMCS), i.e., in which man (or a group) interacts with technical systems and
devices in the control process. We are not dealing here with control of social
and economic processes, in which man does not necessarily interact with tech-
nical systems.
In modern CS of manned ilight vehicles [aircraft] (MFV), whose complexity and cost
are constantly rising, man's role is increasing more and more ar?d his duties
are growing more and more complex. However, because of the limited psycho-
physiological and intellectual capabilities of man, he is of ten unable to per-
form all of the necessary functions for the control of complex systems and
thereby lowers drastically the efficiency of these systems. Thus, according to
the data of American specialists, of the total number of malfunctions occurring
in the control of complex systems, 40% in the rocket industry, 63.6% in the
navy and up to 70% in aviation occur because of errors made by a numan operator
(HO) while controlling these systems [90]. For this reason, when developing and
operating modern man-machine control systems, there is a pressing need to con-
sider human factors. In this respect, aircraf t and particularly MFV CS are the
~ first and foremost objects of scientific work on ergonomic problems of design,
development and operation of MMCS.
Thus, analysis of Soviet and foreign~experience in the time that has elapsed
since the start of space exploration [55, 59, 110] enables us to derive the
following conclusions from the standpoint of ergonomics: man can not only live
in space for a long time, but perform efficiently a large volume of diversif ied
work; the crew of manned spacecraft must participate actively in implementing
the space flight program, assuring structural redundancy of onboard systems~
and play the role of the adapting element in the general circuit of onboard CS; .
it is desirable to use man in cases where his presence aboard the spacecraft
provides more effective solution of the target (multitarget) problem, as com-
pared to machines, aids in maintaining spacecraft function on a high level of
reliability and successful performance of the program as a whole, lowers the
cost of running these programs, as compared to entirely automatic spacecraft
(with equal effectiveness of problem solving); the designs of spacecraf t and
their onboard systems are such as to aid in making utmost use of man's capa-
bilities, using the rich knowhow accumulated by aviation; the scope and import-
ance of tasks put to man are increasing constantly, and special ergonomic
studies must be pursued for them to be performed well.
There are rather detailed but often far from complete data on many traits and
cliaracteristics of the H0, which can be considered in developing various
elements and equipment of MMCS. Rather extensive recommendations have been
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elaborated fo~ designing information display systems (IDS), indicators and '
signals; development of control levers, consoles and stations; development of
life support systems and systems to maintain high levels of work capacity.
Use of already existing results of research in the f ields of physiology, medi-
cine, psychology, cybernetics, engineering psychology and ergonomics makes ~t
possible to drastically improve the efficiency of MMCS and objects of control,
often without any special industrial and economic expenditures. For example,
according to 0. K. Antonov [8], use of the ergonomic approach to designing
aircraf t cockpits, as compared to previously developed ones, resulted in 20-
40% reduction of work load for the crew with regard to equipment and improved
distribution thereof among crew members; it increased by 30-60% the relative
time available to the crew for piloting the aircraft and scanning the surround-
ings [outside the aircraf t], with concurrent improvement of conditions (visi-
bility, force applied to stick, etc.); it reduced the probability of mistakes
I and increased reliability of work done by the crew to correct accident
[emergency] situations (AS) in flight; it increased by 15-20% the operation-
- ality, readiness of the aircraft for flight by reducing the time required for
the crew to make preparations and taxi to the take-off position.
In our opinion, all MMCS can be arbitrarily divided into three types (unlike
the four types of operator work proposed in [54]), according to nature and .
degree of man's involvement in the control process. The first type refers to
systems, in which man performs his functions directly, observing the object
(process) of control and affecting it by his motor organs or using the most ~
elementary equipment. In such systems (which are often called monoergatic--
with one person), man's functions can be called technological and the operator
could be called the technologist (according to V. P. Zinchenko [54]) or
~ executor (according to Yu. G. Fokin [133]). The second type refers to MMCS,
in which the work done by a man (or group) to control an object (process)
is performed by means of controls and other means on the basis of using
information models. In such systems (often called polyergatic--with several
operators) the man's (group's) work is called operator work and the man is
called an operator (according to B. F. Lomov [83]). We can list as this
type of human work the functions of an operator-observer and operator-
researcher according to the classification proposed in [54]. The third type
refers to systems, in which a team of people participates in the control pro-
cess, some of whom, called administrators [supervisorsJ, do not directly
interact with equipment, but are concerned with organizing operator work. Such
systems are usually called organizational. In this book, we shall deal with
problems referable to MMCS of only these three types.
The complexity of modern systems and, first of all, systems of purposeful con-
trol made it necessary to use the so-called systems approach and general
theory of ~ystems (GTS) to develop them.
According to current conceptions (see, for example, [78]), GTS refers to the
scientific direction related to elaboration of the aggregate of philosophical,
methodological, concretely scientific and applied problems of analysis and
synthesis of complex systems of an arbitrary nature. GTS intierently strives
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toward being interdisciplinary on the basis of creating a single scientific ~
platform. This unity can be achieved by means of analogy (isomorphism) of pro-
cesses taking place in various (engineering, tiological and other) systems.
Strictly demonstrative isomorphism of systems differing in nature permits
"transf er" of knowledge from one discipline tu another. Moreover, GTS as an
area of scientific knowledge implies that it is possible to study the behavior,
including purposeful behavior, of systems whatever their complexity and
purpose.
' The formal organization of GTS is diverse, just as the object of its study,
systems. For example, the following have been developed and used in a number of
Soviet works for mathematical descriptions of real complex systems: machine-
algebraic methods (V. M. Glushkov [37]), test theory (S. V. Yablonskiy [147]),
operator algorithms (A. P. Yershov [45]), causal networks (A. A. Markov [86]),
growing machines (Ya. M. Barzdin' [9]), theory of aggregate systems and
computer modeling methods (N. P. Buslenko, I. N. Kovalenko [21]), algorithmic,
information and network [critical-path?] methods (G. S. Pospelov, V. A.
Ir ikov [ 109 ] ) .
A. I. Kukhtenko [77, 78], who analyzed works dealing with mathematics and GTS,
proposed the following classification of levels of abstract description of
different systems: symbolic or linguistic (semiotics); theoretical=set ["multiple"];
abstract-algebraic; topological; logical-mathematical; theoretical-informa-
tional; dynamic; heuristic. Each of these levels has its own definition of
system. Thus, according to M. Mesarovich [92j, on the symboiic or linguistic
level of abstract description, system refers to a set of correct statements, ~
whereas on the theoretical=set level system is the reflection S:X-~Y of abstract
s et X in abstract set Y, where X and Y are set of inputs and set of outputs,
respectively. According to N. Burbaki [18], on the theoretical-set level,
system refers to the set made up of elements that have certain properties and
certain relations with one another and with elements of other sets.
V. V. Chavchanidze, who pointed to the ineffectiveness of the proposed software
and absence of really common elements in the approaches of different authors,
proposed a new approach [137] to the choice of a language that would describe
the most fully systems and structures of arbitrary complexity. Analyzing the
concepts developed in theory of artificial intelligence (AI) and using the
principle of potential isomorphism of the world and cognitive intelligence, he
proposed that a description be chosen in the form of concept~models directed
at man and completely simulating the process of formation of concepts by a
natural int~lligence. Studies of the mechanism of information storage in the
brain, which are presently being conducted under the guidance of V. V. .
~ Chavchanidze, revealed [138] that there are special forms of information
storage by means of elements~of neuron ensembles ("memorons"). This form makes
it possible to issue information in the form of pictures, including some that
were not presented in the learning process. Several of the works of V. V.
Chavchanidze deal with validation of the need and desirability of the conceptual
_ approach to development of GTS [139-141].
- At tlie present time, there is a considerable number of mathematical methods for
analysis and optimization (according to different criteria) of cqntrol systems
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- and processes, and a rather comprehensive analysis of their current status was
made in, for example, [36, 72, 89, 95, 96]. ~n these methods, the f irst and
foremost objective of the researcher was to completely.formalize the problem,
and then only to choose a criterion and develop methods of optimizing .the con-
trol sy'stem or processes. -
Analysis of man's performance in CS shows that the current level o~ development
oF GTS and various special mathematical methods, as well as the inadequate know-
ledge of psychophysiological and intellectual char.acter.istics,. and behaviorai
motivations of man during such activity, do not permit proposing a universal
~ meth~d'of formalization at the present time, that would,adequately describe all
oF these processes in the work of H0. However,.the.need to consider human fac-
tors led to the use for formalization of different forms of HO activity in CS,
at first known methods (for example, automatic control, information. theories),
and then new and modif ied mathematical methods (theories of queueing--QT,
reliability, etc.): With these methods, which make use of certain anthropometric
data and psychophysiological characteristics of man, a number of ergonomic prob-
lems were solved with regard to developing controls, information display systems
and even some problems of optimum distribution of functions,be,tween man and
equipment. In particular, determination has been.made of optimum form and
~ arrangement of controls and consules, illumination characteristics,.form,,loca-
tion, means of presenting and coding information on znf ormation display.systems
for individual and ~group u5e, etc.
It is known that the objective of optimizing any piece of equipment is to find
its parameters (analysis) and structure (synthesis) that will;provide the given .
quality (or efficiency). As it applies to MMCS,,one of the main,purposes of
optimization is wise (or best possible) distribution of;functions between HO
and equipment assuring maximum (or close to it) efficiency of the entire CS., A
unified formalized apparatus is needed, which could be used to describe the
main properties of an MMCS for quantitative analysis and optimization of this
system.
In this book, analysis is made, on the basis of consideration of .some types of
_ HO activity in modern MMCS, of the possibility of formalizing this activity
by means of mathematical and cybernetic methods. In view of the relative
simplicity of obtaining base information and running algorithms.of.mathematical
description of HO function using QT methods, an engineering method has been
proposed and developed for defining the characteristics of an HO in the case of
his performance with and without mistakes. ,
Since all MMCS parameters ~including HO characteristics) can, in most cases, be
divided into additive, multiplicative and logical, we have proposed here the
choic~~ of a"global" criterion of optimization in both a determined and stochas-
tic formulation. Quantitative methods and algorithms for comparative analysis
and optimization of a wide class of MMCS are offered, which are based on the
~ use of mathematical (dynamic, linear and nonlinear) programming methods. ,
The main purpose of this book was to offer engineering methods and equipment for
optimizing MMCS, whi~ch could increase their efficiency,and improve (accelerate
and ].ower the cost) processes of designing, developing and operating them.
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Tl~e