JPRS ID: 9722 JAPAN REPORT

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• Content Type: REPORTS

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JPRS L/9968
9 September 1981
USSR Re ort
p
LIFE SClENCES
BIOMEDICAL AND BEHAVIORAL SCIENCES
CFOUO 1 1 /81)
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JPRS L/9968
9 September 1981
USSR REPORT
LIFE $CSENCES
BIOMEDICAL AND BEHAVIORAL SCIENCES
(FOUO 11/81)
CONTENTS ~ ~
BIONICS
Ctiaracteristics of Movement hy Fem~ile Gryllus himaculatus
Crickets To~tard the Source of a Calling Son~, and the
Dependence o~ Orientat3on Accuracp on Signal Spectrum.....~..... 1
Iiandbook of Sensory Physiology 11
BIOTECHNOLOGY
Role of Biological Factors in Formation and Development of Man.... 13
PHXSIOZOGY
Effect of Artificial Gravitatior~ in Space Flig.h_~ on Retention
o� Motor Habits by Albino R.ats 19
Review of Intervuz Collection 'Physiological Bases of Human
Fatigue' 26
RADIATION BIOLOGY
~ Conclusion From Book on Ra3iation Biology 29
HUMAN FACTORS
Simulator Systems 37
- a- [III - USSR - 27.a S&T FOUO]
FnR nFFTf ie r i 1.cF, nN1.Y
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a vn va� r a..ar~u v.aa:, v......
PSYCHOLOGY
.
Meeting of Scientific Council on Psychiatry of USSR Academy of
_ Medical Sciences Presidium 41
Book on Higher Neuron Activitty Republished 45
Socialist Countries Aold Youth Conditioned Reflex Activitq
Seminar 47
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BIONICS
QDC 591.185.5:595.729
CHARACTERISTICS OF MOVEMENTS BY FEMALE GT+yZZt48 bimaeuZatus CRICKETS TOWARD THE �
- SOURCE OF A CALLING SONG, ?,ND THE DEPENDENCE OF ORIENTATION ACCUR~CY ON SIGNAL
SPECTRUM
Leningrad ZHURNAI, EVOLYUTSIOT~I1~i0Y BIOIQ~iIMII I FIZIOLOGII in Russian Vol T7, No 1,
Jan-Feb 81 (manuscript received 23 May 80) pp 25-32
[Article by J. Rheinlaender, V. F. Shuvalov, A. V. Popov and K. Kalmrin~,
Institute of Evolutional Physiolo~y and Biochemistry imeni I. M. Sechenov,
USSR Academy of Sciences, Leningrad, and Ruhr University, Bochum (FRG)]
[Text] The orientational behavior of female crickets in relation
to the source of male acoustic calling songs was studied in a
round arena with a diameter of 140 cm, located in a sound~roof
chamber. The nature of movements made by females depends strongly
on their candition, and it may be irregular or continuous. In
both cases the females periodically deviated from their course
as they moved. These deviations are limited to a�50� sector,
they occur irregularly, and they are corrected by turning motions.
_ Correcting turns are made both after halts and during movement,
such that females could determine the position of the sound source
- during movement as well. The greater the error angle, the greater
the probability of turning in the direction of the source. The
orientation error at the edge of the arena (within 30 cm of the
emitter) does not exceed an average of several degrees.
Experiments with simulated calling songs differing in the
content of high-frequency harmonics showed that presence of
high-frequency components in the signal (10 and 15 kHz) does not
improve orientation accuraay, and it does not generally hasten
phon.otaxis. The characteristics of phonotaxis are defined com-
pletely by the low-frequency component, 5 kHz. The results are
compared with the properties of low-frequency ascending auditory
neurons that transmit information to the brain. The orientation
accuracy discovered in females may be the result of evaluation
of the difference in the activity of symmetrical pairs of these
- neurons.
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ruK vrr~~.~AL u~r. ~ivL~
Introduction
_ Female crickets find males of their species by orienting on the acoustic calling
signals emitted by the latter (1-7). Research on crickets of two species, one of
which uses a calling song consisting of rarely repeating bursts (Scap82pedu8
mczryinatus),while the signal of the other is similar in structure to a trill
(TeZeogzy ZZus oeeanicus), showed that movement of females to the source is obviously
discontinuous--that is, it consists of alternating short, usually straight runs, and
halts lasting 250-400 msec. It is only during halts that females determine the
position of the source, make correcting turns, and then begin their runs. During
their movement, females da not perceive information on the position of the source,
and they cannot purposefully correct their course. Before starting movement toward
the source, females make scanning movements from side to side. Thus the orienta-
tion system of females may be described in general as a system with an open feedback
_ (4,6).
The orientation accuracy of S. marginatus females is astoundingly low. They can
determine only the side on which the emitter is located, but they cannot detezinine
the necessary turning angle. T. oeean2eus females can determine the required caurse
direction angle rather accurately when the source is located in the forward sector
from 0 to 30-40�; however, outside this sector the turning angles are weakly corre-
lated with the error angle. In this case the orientation accuracy does not improve
as sound intensity increases. However, there are grounds for hypothesizing that the
described orientation mechanisms are not universal to all crickets, the signals
of which may possess different properties.
No one has examined ttie dependence of the accuracy and effectiveness of female
orientation on the spectral composition of the signal. But at the same time we
know that cricket calling songs have a harmonic spectrum consisting of several
components (8,9), and that there are several frequency channels in their auditoxy
system which may evaluate these components relatively independently (10-13).
We posed the following objective in our work: 1) to study the nature of orientation
by female GryZZtts b2maeuZatus De Geer crickets, the signal of which consists of
frequently repea~ing bursts; 2) to evaluate the influence of different spectral
components of the signal on orientation accuracy; 3) to compare the available
neurophysiological data with the characteristics of orientational behavior.
Materials and Methods
The experiments were performed with G. bimac�uZatus females from a laboratory popula-
tion in the last larval instar phase. Each female was placed in an individual cage
and raised until sexual maturity and arisal of well-pronounced positive phonotaxis
toward the male calling song. The selection *_nethod was described earlier (4).
The experiments were performed in a round plywood arena with a diam~ter of 140 cm,
located in a soundproof 5x5x5 meter echoless chamber at 20�1�C. ~ao loudspeakers
(MSD-100, Japan) were located 30 cm from the edge of the arena 120� apart (Figure 1).
A cage into which the female was placed before the start of the experiment was
located in t~e center of the arena. Thirty seconds after the female was placed in
the cage, an acoustic signal began to be reproduced through one of the emi.tters.
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The female's movements were recorded by amovie camera mounted above ~he arena. ~he
~ filming rate was two frames per second. The location of the.female and the orienta-
tion of its body relative to the cage-emitter line were measured on each.frame.
Drawing the trajectory, we used arrows to designa'ce�the orientation of th~ long
axis of the female's body at each moment in time. The effectiveness of feanale
orientation was assessed on the basis of the following indicators: Tp--reaction
, time from the moment of emergence from the box (cage) to the moment the female
reached the edge of the arena; ~8--angle of deviation from the line extending from
the cage to the emitter at the edge of the arena; -maximum angle of deviation
from the course while en route from the cage to the edge of the arena (the first
20 cm of the route from the cage were not counted, because some fem~les wandered
randomly prior to the start of purposeful motion toward the emitter); a~--maximum
error angle between the long ay;~ of the body and the bearing to the emitter
throughout the entire time o~ travel.
~
C
F ' ~ 7oO
_ R ~
; 920 ~
_ ` JO
~
0
N
Figure 1. Experimental Arena and an Example of the Trajectory of a
Female, With the Measured Angles Labeled: Arrows indicate
the trajectory (the bearing of the arrow corresponds to the
orientation of the long axis of the female's body at each
moment in time); H--emi.tter, C--c3ge. Linear dimensions are
given in centimeters. See the text for further explanation.
= In the greatPr part of the experiments the intensity of the signal at the cage was
_ 70 db relative to 0.00002 N/m~. It increased to 82 db at the edge of the arena,
which corresponds to the intensity of a song produced by a male in natural condi-
_ tions from a distance of 0.2-1.0 meters (9). In two series of experiments the
signal intensity was increased by 16 db. In this case the intensity of the signal
at the edge of the arena attained a level corresponding to that of a signal heard
several centimeters away from a singing male.
r
The signals consisted of computer-synthesized sounds with an envelope corresponding
to the envelope of the male's calling song at the same temperature; the intensity
of harmonic components was arbitrari.ly varied. The spectrum of the calling song
contains three harnonic components--4-5, 9-10, and about 15 kHz (Figure 2). At a
signal intensity of 25-40 db the level of the second and third harmonics is below
the level of the main camponent.
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(1)
Q6
40 .
20 �
~ ~ 0.2c~3)
- p 2 4 6 8 10 12 14 K~u(~C~
~ ~ 6
,
Figure 2. Calling Signal of a Male Crieket (a) and the Acoustic Model (b)
Used in the Experiments: A spectral cross section of a calling
song pulse is shown on the graph
Key: ~
1, ~ 3. Sec
2. kHz
Inasmuch as the high frequency components are often below the noise level in the
natural conditions of biotopes experiencing a hiqher noise level (9), one of the
models contained only the 5 kHz component (model 5/0--the digit following the
slash.line indicates attenuation in decibels)� To reveal the role of high fre-
quency components, their leve~. was increased in two other models. The second model
contains the S.and 15 kHz (5/0 and 15/-10) while the third contains all three
_ components (5/0, 10/-20, and 15/-10). The acc~ustic models were recorded and played
back with a Nagra IV-s tape recorder.
Thirty females participated in the experiments. Each female was tested once a day
for 10-14 days.`
Results
The reaction of females to the test models did not differ in general characteristics
fr~m the reaction to a natural calling song. Orientation of females in relation to
the sound source may be divided into three phases: 1) activation phase, 2) initial
o.rientation phase, and 3) phonotaxis phase. The activation phase begins the moment
- the sound is turned on, and it continues until the moment the female exits from the
box. Its duration varies within broad limits, from several seconds to dozens of
seconds, apparently due to differences in motivation of the females.
After leaving the cage, in the ovestivhelming majority of cases the females halted
at the bottom of the chute for several seconds and then.turned in the direction of
the emitter (the initial orientation phase). In this case we never noted "sca.nning
- movements" prior to the start of the purposeful run, as described for Seapsipedus
_ marginatus (4) meaninq that for G. bimacuZatus they are not a mandatory component
of orientation. The initial orientation phase.wa~ vezy short for some females,
which headed for the source almost immediately, while individual specimens wandere~
randomly around the cage on leaving it, resolutely heading toward the emitter only
after this.
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The phonotaxis phase, or the phase of purposeful movement toward the soyarce n~ the
cal.ling song, proceeds differently for different females. Young, timi~i f-~m~-ales,
especially when in the arena for the first time, travel the distance i~? short,
often straiqht runs alternating with halts. After the sound is turned cn~f~, the
nature of the movements does not change, meaning that the halts were x~~t n~cessarily
elicited by a need for course correction. As a rule the direction of mo~e~ent
changed somewhat after a halt even when the f~male was already facin'rg th~e emitter.
These turns far from always occur in the direction of the emitter, and ~onsequently
they are not necessarily correcting turns, bei~g mor.e likely a reflectian of some
sort of perturbations in the movement control system. The larger tlae e~ror angle
at the time of a halt, the greater is the probal~i~.ity of turning in, the direction
~ of the emitter.
Calmer, more-strongly motivated females move signif.icantly more confidently, with
fewer halts or without any at a11; in this case their rate of movemerit gradually
increases as they come closer to the emitter. Runs are not always s~rictly linear.
- On the contraxy the female is frequently seen to periodically deviate from side to
- side in relation to the true course by an insignificaa~t angle. Such oscillations
about the true course are limited to a�30-50� sector (see the va'lues for angle
a~ in the table), while their amplitude is insignificant, inasmu.~c3n as the turns in
different directions alternate with one another quickly. Clwing to these the
oscillations do not have a significant effect oii the fors~~rd tra~j'ectory as a whole,
appearing instead as oscillations in the orientation of tYse b~dy relative to the
principal direction of movement. Course correction occurs in these cases while
on the move or after a halt, meaning that halts are not manda't~ry to a female's
determination of the locai:ion of the sound source. Evidence of this can also be
found in data from experiments in which the sound was produced first from one
emitter and then ~during the time of the female's movement to~ard it) through the
other located opposite the former. In these cases at leasit some of the females
~ turned quickly 180�, without halting their movement or with a few halts. These data
also indicate that females confidently distinguish sounds c~rning from the rear from
sounds coming from the front.
The trajectory of feniales toward the sound source varied within broad limi.ts
(Figure 3a). In most cases females headed toward tl~e sou~ce w~th sufficient accuracy
from the very beginning, wit3.iout noL-iceable deviations to the s~ides (Figure 3b).
Some females occasionally deviated significantly from the course to the emitter,
later correcting their movement by means of one or several turns (Figure 3e).
Certain females ran relatively straight, though at a certain angle to the source,
and then corrected their course at the ~exy edge ot t.~e arena (Figure 3d). The
possibility is not excluded that pathologica~. alterations were present in at least
some of these insects, leading to interaural asxmmetry.
On the whole the orientation accuracy of females, evaluated oa~ the basis of their
location at the end of the route (beside the.edqe of the arena), was sufficiently
high. The averages for angle ~p do not usually exceed 6-7�. When the emitter was
brought close to *he edge of the arena, a11 females approached it,, correcting small
deviations in the final leg of the r~ute.
Statistical treatment of the obta.ined material (see table and Fic1ure 4) showed that
presence of high-fre~uency compone~~ts in the signal does not significantly improve
orientation accuracy, and it does not accelerate phonotaxis. The contrazy is more
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run w�raa..~na, voa: v~~a..a
Effectiveness of Orientation by Female Crickets
During Movement Toward the Source uf a Calling Song
M~del T(sec) ~m~~ a~ N,/sec� � r~
5/0 20.65�13.33 6.86�6.07 25.28�18.44 44.61�12.41 5.35�3.66 44
5/0 15.-10 16.21�7.38 T0.36�7.36 27.76�20.92 54.31�18.15 4.49�1.05 39
5/0 10/-20 15/-10 26.6t23.8 6.42t7.46 22.22�17.73 42.02�15.85 4.84�1.27 53
5/0+16 ~ 15.72�5.48 6.0�5.36 15.33�9.41 39.00�11.0 5.17�0.78 14
5/0 15/-10+16 db 38.25�15.73 33.00�25.03 47.75�25.16 86.67�40.1 2.52�1.56 5
Note: The values given for each variable are x�6; N--arbitrarily selected
measure of the effectiveness of phonotaxis, which is a function of orienta-
tion accuracy (~p) and running time (Tp). N= ln 1/Tp��p. The larger N is,
the more effective is phonotaxis; n--number of ineasurements. See text for
further explanations.
a
v d ~
c
6 0 ,-.~..-f 4
1 ~ ~
% ~ .
.
~ ~ 1
~ . .
f ~ !
.
~ ,
. . ~
~ ~ ~
Figure 3. Trajectoxy of Females During Phonotaxis: see text for explanation
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o c
N e H
!5 ~SPo~=b.42*7.46 ~s ~~001=6.86!6.01~
I ~
~v ~ ~o ~
s s
0 f0 20 90 40 50 0 ~0 20 30 I~a l�
N ' 6
15 ~ amQ=I = 42.02: 15 . 85' ~ ~ a~~~ ~44.61; 72 .4!'
tp !0
.
~ S S
0 20 40 60 60 f00 ~ a��~� 0 20 40 6a 8Y) ~00 ~rs~~~�
Figure 4. Distribution of Angles ~p and a in th~ ~our.s~~af Phonotaxis
Toward Models 5/0, 10/-20, 15/-10 (a,~v) a~d 5~/0 (e,d) :
a--n = 53, D~p = 2�; b--n = 50, ~am~ =10� j e-�-rc = 44,
p= 2�; d--n = 42, ~am~ =10 N--numbex~ ca f runs
likely true. Angle a is significantly greater with m~odel 5/0,15/-10 than with
- the other models. The differences in the rest of ~t'~e parameters are insignificant.
The accuracy of orientation on model 5/0 does not Eliffer from that of orientation
on a natural signal.
Raising signal intcnsity by 16 db significantly increased the effeativenesS of
phonotaxis toward model 5/0+16 db and, on the other hand, significantly decreased-~
~ the accuracy of orientation c+n the m4de1 containing the high-frequency component
(5/0 and 15/-10+16 db). In the latter case the females began rushing from side to
side, as if searching for the source. This was accompanied by an increase in all
orientation indicators (Tp, ~p, ~~X, a; see table).
Discussion
The obtained data shaw that the orientation behavior of G. bimaeuZatus females
differs dramatically from behavior described in the literature for S. marginatus
~ and T. oceanicus (4,6) in having the capability for evaluating the position of the .
source and correcting course n~t only during halts but also while moving, and by
absence of scanning movements before the start of phonotaxis. Wendlar et al. (7)
recently came to similar conclusions in research on the orientation behavior of
the field cricket, GryZZus eampestris, the calling song of which is very similar
in temporal organization to the signal produced by G. bim~zculatus (8,9)� The
oscillatory deviations of the long body axis and the trajectory from the direction
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rvn vra�a~.~hi, a~oc vi~L.i
to the sound source may be the result of random shifts or noises in the systein con-
trolling the movement of right and left legs. The randomness of these deviations
- and their significant individual variability argue in favor of such an interpretation.
Yn any case, inasmuch as these deviations were altogether absent from many femalee, ~
it may be hypothesized that oscillations are not a mandatory component of the guidance
mechanism. Course correction compensating for random deviations of the long axis of
the body from the direction to the target is apparently achie~ed by feec~ack signals.
These signals may be dependent on the error angles. Obviously, the greater the
error angle, the larger is the probability of a correcting instruction and its in-
tensity; this probability approaches 1 as the error angle reaches its critical value.
Our data showed that the maximum error angle for phonotaxis does not usually exceed
30-50�. Within the �30-50� sector, the intensity of the correcting signal may be
commensurate with the level of noises within the system, and this may explain the
randomness and unpredictability of "minor" turns along the course to the goal.
Deflection of the long axis of the body from the direction to the emitter is
accompanied by appearance of asymmetry in stimulation of the right and left auditory
channels. Information on sound in different frequency bands is processed separately
in the auditory system of crickets, beginning at the peripheral level and ending in
the highest associative center in. the protocerebrum. The calling signal is perceived
by G. bimacuZatus by means of receptors narrowly tuned to a frequency of 5 kHz and by
second-order ascending neurons (LF-neurons), which transmit information on the signal
_ to the protocerebrum and apparently play the decisive role in control of pkionotaxis.
. The functional properties of these elements have been described earlier (11-14).
There is one LF-neuron on each side, and the relationship between the discharges
contains information on the position of the sound source in s~ace. Each of the
LF-neurons is maximally sensitive to sound coming from tY~e ipsilateral side (Figure 5).
The difference in their discharges is obviously a function of the error angle. We
can see from Figure 6 that the difference in th s number of impulses produced in re-
sponse to a calling song burst grows dramatically as the source shifts from foxward
position (0�) up to 80-90� to either side. At the critical error angle for phono-
taxis (50�), tl~e difference in the number of impulses contained in a discharge by
these neurons in response to a calling song burst is 15-20 at average and high
intensity. This ccrresponds to a difference in stimulation intensity on the order
of 18-25 db, which greatly exceeds the differential intensity threshold of hearing
which, judging from behavioral experiments, is about 2-3 db for G. bimacuZatus (157.
A difference in the number of impulses on the order of 2-4 per burst, which is what
arises at error angles on the order of 3-7�, corresponds to the differential intensity
_ threshold. This amount is well consistent with angle ~p (about 6�). At larger
deviation angles a feedback signal should arise, the intensity of which is proportional
to an error angle within up to 80-90� (if its intensity is governec~ by the difference
in stimulation of LF-neurons).
_ LF-neurons react almost identically to saund from the front and from the back, and
consequently there should be additional elements in the auditory system of crickets
having difFerential sensitivity toward the back and toward the front. Similar elements
have been discovered in locusts only in the protocerebrum (16). The acuity of their
directional sensitivity grows as sound intensity increases. If crickets also have
such elements, then our data indicating growth in orientation accuracy with increasing
sound intensity receive a fully natur.al explanation.
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n D'
~l~ 32 r~y
/I NyMeupoH ~r'' ~ lTHY�HeupoM
~ ~ Z4 ~
- - ~ i �
- /-10db~~~ 16 ~ -lOd6
Od6 I ~
~ ~ . Od6
/ i 1 8 ~ ~ (3)
/ ~ ) ~
0
270 ~ 90
~ \ \
~ ~ \
` ` _ i s
~ _ ~
~
~ '
~ e ~
_ \ /
1~~ _ _ / '
180� .
Figure 5. Characteristics of the Directional Sensitivity of a Symmetrical
Pair of LF-Neurons of the Cri~ket's Ventral Nerve Chain in
Response to the Action of a Calling Song Containing Bursts of
Five Pulses: n--number of impulses contained in neuron responses
to a calling song burst, recorded at the level of the protocerebrum
from axons of LF-neurons (from experiments conducted in Bochum,
the basic results of which were published earlier (11)). An
intensity of 0 db corresponds to a signal intensity of 95 db
relative to 0.00002 N/m2. Sc.e text for further explanation.
Key:
1. Left LF-neuron 3, cm
2. Right LF-neuran
dn
30 ~ - ~O,q 6
.
~ 20 ~ ~ . C1~ .
C ~
90 ' O,q6
~
0 ~ ~
. ,
~ ~ , .
90 ~ / .
C ~
v .
C ZO ~ ~
` ~ i
. ~
J
L 1 I ~ 1~I ~ I I ~ I I I
180� 270� 0� 90� 180~
Figure 6. Dependence of the Difference in Responses (~yy) by the Same Pair
of LF-Neurons on the Er:.or Angle, Given the Same Signal Intensity
as in Figure 5(Evaluated on the Basis of the Number of Impulses
in Response to a Calling Song Burst): �~ee text for further
explanation
Key:
1. db 9
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. �'V~\ Vl'l ~~.IAV v?V VL\1.~�
Experiments with models having different spectral compositions showed that the
characteristics of phonotaxis depend entirely on the first low-frequency maximum of
the signal (5 kHz). High-frequency components influence neither the effectiveness
of orientation nor the rate of phonotaxis, even if their level is significantly higher
than in the natural signal. Sharp disturbance of orientation in the presence of a
high-intensity signal with high-frequency components may be a reflection of a transi-
tion by the female from phonotaxis to a search for the male. In natural conditions
the high-frequency components achieve their highest level only in direct proximity to
a singing male.
The cricket orientation system may be of interest from the standpoint of bionics
as an example of a guidance system used to assist movement.
BIBLIOGRAPHY ~
� 1. Regen, I., PFLUGERS ARCH., Vol 155, 1913, p 193.
2. Regan, I., S.B. AKAD. WISS. WIEN., Vol 132, 1923, p 81.
3. Walker, T. J., ANN. ENTOMOL. SOC. AMER., Vol 50, 1957, p 626.
- 4. Murphey, R. K. and Zaretsky, M. D., J. EXP. BIOL., Vol 56, 1972, p 335.
5. Popov, A. V. and Shuvalov, V. F., J. CON~. PHYSIOL., Vol 119, 1977, p 111.
. 6. Bailey, W. J. and Thompson, P., J. EXP. BIOL., Vol 67, 1977, p 61.
7. Wendler, G., Dambach, M., Schmitz, B. and Scharstein, H., NATURWISS, Vol 67,
1980, p 99.
, 8. Popov, A. V., ENTOMOL. OBOZR., Vol 51, 1972, p 17.
9. Popov, A. V., Shuvalov, V. F., Knyazev, A. N. and Klar-Spasovskaya, N. A.,
ENTOMOL. OBOZR., Vol 53, 1974, p 258.
10. Popov, A. V., ZH. EVOL. BIOICHIM. I FIZIOL., Vol 9, 1973, p 265.
11. Rheinlaender, J., Kalmring, K., Popov, A. V. and Rehbein, H. G., J. COI~.
PHYSIOL., Vol 110, 1976, p 251.
12. Elsner, N. and Popov, A. V., ADV. INSECT PHYSIOL., Vol 13, 1978, p 229.
13. Popov, A. V., "The Insect Acot~stic Communication System," Dissertation Abstract,
Leningrad, 1979.
14. Shuvalov, V. F. and Popc~v, A. V., ZH. EVOL. BIOKHIM. I FIZIOL., Vol 9, 1973, p 177.
- 15. Shuvalov, V. F. and Popov, A. V., DOKL. AN SSSR, Vol 246, 1979, p 247.
16. Adam, L. J., ZOOL. JB. PHYSIOL. [sic] Vol 81, 1977, p 250.
COPYRIGHT: Izdatel'stvo "Nauka" "Zhurnal evolyutsionnoy biokhimii i fiziologii", 1981
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HANDBOOK 0~' SENSORY PHXSIOLOGY ~
Leningrad ZHUTINAL EVOLYUTSIONNOi~ BIOKHIMII I FI.ZIOLOGII in Russian Vol 17, No 1,
Jan-Feb 81 pp 103-104 ~ ,
[Review by F. Gribakin of book "Rukovodstvo po fiziologii organov chuvstv" [Handbook
of Sensory Physiol~gy], Volume VII/6A, "Vision in Invertebrates", P~rt A: "Inverte-
brate Photoreceptors", edited by Prof Kh. Autrum, Springer-Verlag, Berlin-Heidelberg-
New York, 1979, 729 pages, 344 figures] ~
� [Text] This book is the first part (A) of the next volume in a handbook of 5ensory
physiology known the world over, containing basic information on the general physio-
logical mechanisms of invertebrate photoreception. Special emphasis is placed on the
fact, sometimes unfor�:unately disregarded, that invertebrate photoreceptors not only
serve as models for studying the basi,c photoreception processes, but also owing to
numerous special adaptations they have significant value to expanding our knowledge
of ecology and evolution. This is reflected in the full title of the volume--"Compara-
tive Physiology and Evoluticn of Invertebrate Vision".
All 12 chapters are written by top-class specialists. Readers of our journal would
be interested to learn that the "Introduction" (Chapter 1, Prof IQz. Autrum) discusses,
in particular, Ya. A. Vinnikov's idea that microvillous receptors are phylogenetically
older than ciliary receptors; of course the editor of the volume himself feels that
this supposition must be supported by further research. Chapter 2(B. D'yen) is
devoted to the physiological mechanisms of photoreception in the Protista. Chapter 3
(U. Miller) exami.nes the optic properties of the eye's filtering media (aromatic amino
acids, shielding pigments, and so on). Chapter 4, which is devoted to the physiology
of invertebrate vi.sual pigments, is extr.emely interesting. This chapter was written
- by Prof K. Khamdorf, who discovered that photoregeneration is the principal means of
regeneration of rhodopsin in cephalopod mollusks and arthropods, while rhodopsin in
vertebrates is regenerated by enzymatic mechanisms. Chapter 5, "The Physics of Vision
in the Compound Eye", contains all of the basic information on the optics of photo-
reception by the invertebrate eye. The author of this chapter, prFSently a professor
of applied mathematics at the University of Australia, A. U. Snayder, is essentially
the "father" of photoreceptor optics, which grew up into an independent branch of
knowledge in the last 15 years. Chapter 6, "The Receptor Potential of Invertebrate
- Visual Cells" (M. Yarvilekhto), contains contemporary data on the regeneration
mechanisms of ear~:y and late receptor po~ential in arthropod photoreceptors. It goes
on to examine the physiological mechanisms behind arisal of a false pupil, or a
pseudopupil, in the compound eye. A purely optical phenomenon, the pseudopupil is
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usually seen as a dark spot formed out of omanatidia oriented strai~ht at the
observer, and consequently not reflecting and not scattering ~ight in this direction;
the possibilities for capitalizing on this phenomenon as a valuable tool for
studying the optic qualities of the ele and the properties of its photoreceptors
and pigment cells are demonstrated (Chapter 7, D. Stavenga). Modern ideas are
presented on the superposition and apposition of compound eyes (P. Kuntse, Chapter 8);
the first subdivisions of the chapter rather extensively explain the classical
viewpoints of Johannes Muller and Sigmund Ecksner. Chapter 9, "Spectral Sensitivity
and Color Vision of Invertebrates" (R. Mentsel')r i.s a most extensive review summar-
izing, in one place, all that is known today about the spectral sensitivity of the
photoreceptors of invertebrates, beginning with the Protista, and about the central
mechanisms af color information processing, to include the behavioral aspects of
the problem. Chapters 10 (M. Ioshida) and 11 (Miriazn Bennett) are devoted to
extraocular photoreception and its role in triggerinq and maintaining diurnal rhythms.
Finally Chapter 12, "A Genetic Approach to the Visual System" (M. Geyzenberg), deals
basically with research on vision of a classical object in genetics, mainly various
eye mutants of DrosophiZa, since in the author's opinion only research on
DrosophiZa fits within the framework of the term "genetic approach", even though
the genetic literature on vision Presentlyincludes organisms from halobacteria to
the white tiger.
Evaluating the content of the volume as a whole, it should be stated that it is
a superior source of contemporary data having fundamental significance today. Never-
theless a number of problems still rema.in outside the field of vision of the author
collective (the ion mechanisms of generation of receptor potential, the genetic
approach to studying ,.vision in the honeybee, analysis of the electric field of a
compound eye, and so on). We would hope that these aspects of the physiology of
invertebrate vision would be reflected in the second part of Volume VII/6.
which is now being prepared for publication. Unfortunately, the iridex of cited works
by Soviet authors is not very large: As a rule only those works which are published
in English in international journals are cited, which once again emphasizes the need
for publishing in fo~eign sources.
COPYRIGHT: Izdatel'stvo "Nauka" "Zhurnal evolyutsionnoy biokhimii i fiziologii", 1981
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BIOTECI~TOLOGY
ROLE OF BIOLOGICAL FACTORS IN FORMATiON AND DEVELOPMENT OF MAN
Mosc~w VESTNIK AKADEMII NAUK SSSR in Russian No 6, Jun 81 pp 42-47
[Text of statements made by Academician A. D. Aleksandrov and Academician
N. P. Dubinin]
[Text] As was already reported on the pages of VESTNIK AN SSSR
[Vestnik of the USSR Academy of Sciences] (No 4, 1981, p 107),
there was a session of the General Meeting of the USSR Academy of
Sciences on 21 November 1980. At this session, Academician A. D.
Aleksandrov spoke on the role of biological factors in the
formation and development of man. Academician A. P. Aleksandrov,
- president of the USSR Academy of Sciences, then gave the floor to
Academician N. P. Dubinin to respond to his points. In accordance
with the r~quest of the speakers, the text of their speeches is
published below.
Academician A. D. Aleksandrov
- Man is, to use the terms of philosophy, the carrier of two forms of movement of
matter, "biological" and "social." But it is expressly the latter that distinguishes
man. Within the same biological species and even the same race, people demonstrate
an immense diversity of forms of a.ctivity, which they create and transmit to one
another and to subsequent generations by means of direct and indirect communication.
Since the brain is the source of the programs of these forms of endeavor, we can
refer to the cerebral evolution of man in contrast to genetic evolution.
The enormous area of what is specifically human, human history, develops, of course,
- on the basis of genetically determined human distinctions and, first of all, the
_ distinctions of the human brain. There, the same chief law common to all living
organisms is in effect, according to which genes ~'-etermine the possibility of
development of traits of an organism, but this possibility is realized depending
on conditions. However, ~specially man genetically represents merely a possibility
~ that changes into reality through interaction between the genotype and conditions, ~
mainly social ones. These conditions themselves are created and changed by the
activities of people. For this reason, it can be stated that man (in the collective
' sense~ cxeated a~d continues to create himself on *_he basis of his specific genotype.
~ TYius, three factors affect the formation of man: genotype, living conditions and
tY~e t~ansfo~zning, creative endeavors of man himself The extretnely complex
*'~Creati;ve endeavor" is used to construe any activity that is not performed accord-
ing to a pxeset program. ~
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_
correlation between them constitutes a fundamental and difficult problem about
which the most varied v~iews are expressed. Representatives of different special
fields often exaggerate expressly what is closer to them, disregard{ng what is
"alien" to them. Geneticists tend to underestimdte the social factor, while
sociologists do so for the genetic factor, as well as the creative fdctor in
~ some cases, interpreting, for example, the personality as the passive product of
existing social conditions. ~ ~
The colossal extent, to which geneticists sometimes disregard the social factor, is
indicated, for example, by an article by the well-known American geneticist,
re:cipient of the Nobel prize, J. Lede~berg, which was published at the time in
LITERATURNAYA GAZETA [Literary Gazette] (No 20, 1969). There, among others, the
following question was discussed: "Do the poor have proper [perfect] genes?" The
answer being: "Perhaps genes do play some part in man's economic failures. For
this reason, the excessive proliferation of the stratum of the poor could be
eugenically harmful. But birth control among the poor will not yield results
very soon.:.. Of course, it would be easiest to suggest another solution: let not
- the poor have any children at all. But if we were to take this route, we could
go even farther: fight for eradication of poverty by means of physical extermination
of all the poor. Although, perhaps, this would be ~nore humane than to doom them
- to a hungry death."
This opinion is extremely instructive in many respects, and it should be discussed
in detail, but unfortunately, we do not have space here to do it. We shall merely
call attention to the blindness of this authoritative scientist who does not see
_ that poverty is a social phenomenon and that for this reason it is simply absurd to
discuss "eradication of poverty by means of physical extermination of all the
poor" (incidentally, ~et us recall that Schubert and Van Gogh, for example, were
paupers and, one would think, their poverty was related expressly to their genius,
which compelled them to direct their efforts to an activity that yielded no
economic success).
There is also a very obvious ethical aspect to such ~udgments. An important dis-
tinction of all sciences dealing with man, be it psychology or human genetics, is
that it applies, in principle, to everyone, not excluding the researcher himself,
and for this reason, in the sciences dealing with man the scientific position is
particularly closely related to the ethical [moral] position. Of course, this
does not mean that science must subordinate its objectivity to ethical goals.
Marx wrote: a man who strives to adapt science to such a point of view
that is not gleaned from science itself (:no mazfe~ Rqw ~quch t~e la~,tex ~a~ in exro~'~
but from the outside, toward a point of view that ts dictated by i~te~+e~fis that
are alien to science, endogenous in relation ta it�-~T ca~,7, such a-man R~i~~erio~'
But one must clearly understand the human meaning of what is said and done in
the science of man and there must be a warning signal in front of everyone who
talks about man, parttcul,a�r~y tt~ose who do so on behal~ o~ science; "Be c~ze~ult
people! Be ob~ective to the utmost!" This app~ies in particular tp statetqen~s
made in the broad press.
*K. Marx and F. Engels, "Soch." [Works], Vol 26, Pt 2, p 125.
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But here we read the article of Academician N. P. Dubinin, "Inheritance of Biologi-
cal and Social Factors,"* where this warning is not heeded at all.
N. B. Dubinin wishes to develop the idea of Marx concerning the social essence of
man,and he criticizes geneticists who exaggerate the role of the biological factor
in man to the detriment of the social one. But he himself goes to an utter
extreme.
His own idea is that it is absurd to search for an explanation of normal mental
phenomena in any biological functions. He decisively denies that biological,
genetic heredity has any influence on man's normal psyche and, in particular,
writes: "Vn2gar b fologizational interpretation of inentai functions as genetically
_ determined traits constitutes idealism in comprehension of the problem of man.
Authors who relate the human psyche directly to genes, attributing the mental
properties of man to physiological functions of neurons of the brain, vi~w
themselves as materialists of the highest order. However, the human psyche cannot
be found in ei.ther the function of genes or the function of neurons, for the simple
reason that its level has gone beyond the boundaries of biology. To search for
an explanation of inental phenomena in any biological functions is tantamount to
attributing spiritual properties of all matter" (p 72).
But we know, for example, that sex is genetically determined, that the psyche of
normal men and women is not the same, and that such mental differences are attri-
butable, at least in part, to biological functions. ThnsL_the main thesis of the
article, and one stated so categorically, is an overt contradiction to obvious
facts.
- The real problem is to examine which mental features, how and to what extent they
depend on heredity or social conditions. But N. P. Dubinin closes this issue for
normal people, leaving it to medical genetics only with regard to abnormal people.
However, it is beneficial to note that the very concepts of normal genotype, normal
brain, normal psyche, that Academician N. P. Dubinin uses with such confidence,
are not completely definite and imply demonstration of various aspects and gradations
of normalcy and abnormality, which depend on the same genetic heredity. For this
reason, the desire of Academician N. P. Dubinin to limit the influence of genes
- only to an "abnormal" psyche is invalid.
Further, we see from the statements quoted above that Academician N. P. Dubinin
decisively denies a direct relationship of the psyche to neuronal functions in
the brain. But, for example, the well-known effect of alcohol on the mind is
- apparently related to its effect on neuronal function in the brain, so that here
too there is no validity to the idea of Academician N. P. Dubinin.
Indeed, the problem consists of investigating physiological, biochemical, physical
structures and mechanisms that determine mental phenomena. N. B. Dubinin "closes"
this problem also.
Let us recall, however, higher nervous activity, the types thereof established by
I. P. Pavlov. He made the direct statement that certain .temperaments correspond
to these types, and that they are determined by the genotype. So that Pavlov
- See: KOMMUNIST, No 11, 1980.
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maintains there is a direci. relationship of the psyche to the genotype and brain
neuron functions and, thus, is among the authors who "directly relate the psyche
to genes. Let us assume that Pavlov was m~.staken, but then he should be refuted
with scientific arguments, and not disposed of in such a manner. Incidentally,
it would appear that Pavlov was not mistaken here....
The problems that are directly related to man require utmost objectivity of judg-
ments, utmost caution of conclusions and utmost attention in research. Questions
of heredity, congenital or acquired traits, abilities or defects, questions of
man's capabilities--they all emerge acutely in life, they affect living people,
sometimes they create difficult situations and they merit the most serious dis-
, cussion. This applies fully to the problem of abilities.
A socialist society is based on the principle of "from each according to his
abilities," and for this reason the question of abilities is of basic importance
to us, not only theoretically but practically, both on the level of society as a .
whole and of each man: how to best demonstrate and develop his abilities, what
can be expected and demanded of him. N. P. Dubinin answers the question rather
simply: "All normal people are capable of virtually unlimited spiritual develop-
ment" (p 68). "Giftedness is the effecti~ve development of essential human traits
combining a normal genotype with beneficial conditions for development thereof
(i.e., chiefly the "'acquired nature' thereof)" (p 73). In other words, apparently
with a normal genotype, the beneficial conditions can make a L~monosov, Marx,
Newton, Beethoven, Rafael, or some~ne else out of any child by providing for
"virtually unlimited spiritual development." And if there are very few even
less gifted people who develop from the multitude of children, it is the fault of .
their parents who did not provide the same conditions for them as were present for
Lomonosov.
Incidentally, Academician N. P. Dubinin stipulated: "i.e., chiefly the 'acquired
nature'." And this apparently means that "the inborn factor" also plays a part,
if not the leading one. If this is so, does not N. P. Dubinin find himself
among the "authors who related the human psyche directly to genes," at least to
an insignificant degree? The question of being gifted requires investigation. But
N. P. Dubinin, proceeding from his views, sweeps away in-depth studies of gifted-
ness, whose authors try to demonstrate the potential abilities of man. 5uch
studies have not yet advanced enough, but to abandon work in this direction would
be extremely detrimental to the development of our society, as would abandonment
in general of all three problems that N. P. Dubinin "closes."
Declaring loudly that he is speaking from the vantage points of Marxism, N.'P.
Dubinin considers this as grounds, from the very first lines of his article, to
discredit people of a different mind stating that they are "undertaking an
attempt at revision," and even trying to "abolish the Marxist teaching on the
integral social essence of man, ad~�ancing the thesis that the genetic program
- supposedly controls the social behavior of people" (pp 62, 63).
He mentions the late Academician B. L. Astaurov as the first example of those "who
did not escape this craze," and quotes an extensive passage where B. L. Astaurov
writes about the significance of the law, according to which all of the proper-
ties of an adult organism, not excluding man and his psyche, depend on heredity
and the environment. But N. P. Dubinin interprets recognition of this law as a
"thesis," that supposedly the "genetic program controls the social behavior of
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people." Confusing the concepts of "to be rel.ated jor dependent] and "to be
controlled," he attributes to B. I,. Astaurov sometfiing he did not say and, at
~ the same time, involvement in efforts to "aboJ.ish Marxist teaching on the integ�-
ral social essence of man."
IncidentaLly, N.P. Dubinin also attributes to Marx things that he did not write at
all. It is expressly in the "Theses about Feurbach" that Marx states: the
essence of man is not an abstraction inherent to a separate individual. In reality,
it is the aggregate of all social relations."* N. P. Dubinin writes, with
reference to man, that "in the words of K. Marx, he is the aggregate of all social
relations" (p 62). But is it not clear that "man" and "essence of man," and in
general any object and its essence are ratner different concepts, so that to
replace one with the other is to distort the author~s thought. Let us also mention
that Marx writes about the essence of man in its reality, of man in the collective
sense, rather than an individual; let us compare this to the thesis, according to
which genes determine the possibility of development of an individual's traits....
It is obvious that the contradiction that N.P. Dubinin wishes to see does not exist
here.
However, it is not the inaccurate quotation or inadequate comprehension o~ some o~
the judgments of Marx that is the main point. The main point is that the article
of Academician N.P.Dubin~n, with all its claims of a Marxist position, is in
reality something that is quite the opposite of Marxism.
The spirit of Marxism lies in systematic scientific attitude, concrete analysis,
in-depth penetration into the entire aggregate of available facts referable to the
problem under study. But in the article in question, only declarations are made
with regard to the psyche of a normal man. ~
The spirit of Marxism lies in dialectics and, consequently, consideration of all
associations, mediations and mutual transitions, to the extent of the unity of
opposites. But in the article under discussion, the very thoughtof interaction
between biological and social factors with reference to the psyche of normal man
is re~ected.
The spirit of Marxism lies in unconditional, specific humanism. But in the article
in question, concrete analysis of human problems relatedt~i-heredity (with a"normal"
genotype) is replaced with abstract statements as to the unlimited abilities of man.
Furthermore, there is the question of principles of scientific research and scienti-
fic polemics. The main point is not referable to specific mistakes and absurdities
per se. The main point is for something alien to science by virtue of inethods of
reasoning and approach to the problem not to replace research and scientific -
discussion of the most serious problems.
Academician N. P. Dubinin
I never denied the importance of problems of human genetics. Science has offered
good proof of the effect of biological heredity on physical, neurodynamic and psycho-
dynamic distinctions of man. At the present time~ more than 2300 Mendelian traits
have been studied. Moreover, questions of damage to biological heredity are being
*K. Marx and F. Engles, "Soch.," Vol 3, p 3.
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. . _
raised with exceptional acuity at the present time. According to data for 1977-1980,
10.5% of the children in the world are born with genetic defects. Of these, 3%
(according to the UN O.r.ganization) are born with inheri~ed metital deficiency. ~
There is a question of dynamic growth of the genetic burden under the influence of
mutagens, which pollute the environment, and other facY.ors. Much work is being
done on this subject. I myself am a participant of many All-Union programs and,
in particular, I head the section of "Genetic Aspects of the Problem of 'Man and
the Biosphere of the USSR State Committee for Science and Technology, and am
involved in the work of international committees dealing with the problem of the
biological factor in man.
And, in the mentioned article, "Inheritance of Biological and Social Factors," in
the ~ournal KOMNIUNIST (No 11, 1980), the signif icance of the biological factor is
very clearly stated. Of course, each man is a distinctive genetic individual with
his own properties, his own anatomical, physiological and biochemical features,
with numerous distinctions, such as elements of the psyche, endocrine system,
speed of neurodynamic processes, etc. But the point is how this is related to
man's social essence: do variations of normal genotypes inherited from parents have
a direct, adequate effect on social and human properties, on man's spiritual world?
Efforts are being made to relate them in a fatal way. It has even been said that
creation of a new man would be impossible without altering the genetic prerequi-
sites of modern people. '
~tatements are being made that one can supposedly "predict" all of the potential
spiritual abilities of each neonate on the basis of a study of his genes. It
was written in LITERATURNAYA GAZETA (4 July 1980) that one can identify a.future
member of the Writers' Union at the embryonic stage, according to the set of
"writer" genes received in the zygote after fertilization. . Such statements are
beyond the boundaries of science. In the presence of dialectics of social and
biological factors, it is imperative to draw a distinct line between them, since
the biological part of man is merely the lowest form of movement of matter. The
essence of man is social. This idea is the basis for my article in KOMMUNIST.
I am deeply convinced that my point of view discloses real possibilities for the
spiritual, social and industrial development of each man, while the vew of fatal
genetic predetermination precludes these possibilities.
Moreover, the latter point of view is scientifically incorrect: we have no data to
the effect that man~s spiritual world, his work activities are biologically de- .
termined; in fact the specifics of man are created as the product of all social
relations.
For this reason, we shall fight for the view that offers our people the opportunity
of enormous social development on the basis of normal human heredity. The biological
basis, being the lowest form of movement of matter in man, serves anl.y as the pre-
requisite for him to enter into the social form of movemant of matter. Conscious-
ness, the spiritual world and ability to work do not appear in a child from his
biology, but as a result of his active entrance into the world of social practice.
Herein lie the enormous capabilities of pedagogics and psychology, that chief
factor that should serve as the foundation of formation of new man by the condi-
tions of the social environment.
COPYRIGHT: Izdatel'stvo "Nauka", "Vestnik Akademii nauk SSSR", 1981
10,657 .
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PHYSIOLOGY
UDC 612.821.6+613.693
' EFFECT OF ARTIFICIAL GRAVITATION IN SPACE FLIQiT ON RETENTION OF MOTOR HABITS BY
ALBINO RATS
Moscow ZHURN~IL VYSSHEY NERVNOY DEYA~Z'NOSTI IMENI I. P. PAVLOVA in Russian Vol 31~
No 2, Mar-Apr 81 (manuscript received 6 Mar 80) pp 261-266
[Article by N. N. Livshits, Z. I. Apanaserilco, M. A. Kuznetsova and Ye. S. Meyzerov]
[Text] The goal of this project was to s~cudy the influence of artifical gravita-
tional forces on some indicators of tk~e 2~igher nervous activity of albino rats in
space flight. Certain changes in thc behavioral reactions of animals following
their exposure to space flight factors in biosatellites were noted by us earlier
(3,4). Accelerations are known to ha~~e an effect on the central nervous system
on earth (6,7, etc.), but there is no information in the literature on the action
of these factors during orbital flight.
We studied behavior of animals in the first 2 weeks after the landing of a bio-
satellite aboard which artificial gravitation was created.
Methods
The conditions under which animals were maintained in a vivarium and aboard the
biosatellite were published earlier, as was the complete program of clinical-bio-
logical research (1).
In addition to groups of animals studied in accorclance with the complete program,
there were a group of rats for which only higher nervous activity was investigated,
and a group of rats studied in accordance with an abbreviated program containing
only a third of the procedures included in the complete program (Table 1).
A maze devised by Ya. Dombrovskaya was used to study rat higher nervous activity.
We recorded the number of refusals to run the maze, the time taken by the rat to
move from the starting ccmpartment to the feed compartment, and the mi.stakes made
en route. We kept count of the number of times neurotic states arose, and of the
number of disturbances in the normal dynamics of mistakes in the course of experi-
ments. The research methods were described earlier (3). It should be pointed out
- that we attached special significance to the number of times rats refused to run
the maze in our analysis of the results. Disturbances in higher nervous activity
deduced from other indicators could be assessed at less than their true importance
in the event of a significant increase in the number of refusals, because this would
19
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Table 1. G'haracteristics of Experimental Conditions
No. of Angular Volume of
Group Animals Accelera- Velocity, Experimental
Exposure Conditions S ols in Group tion, g Radians Research
Aboard b~io- Ddotionless IlH3 5 0 0 Complete (all
satellite BIOS [not procedures
further
identified]
5 1 5.3 "
ICosmos-936 Centrifuge ~2
radius ~
320 mm
Mock-up Motionless CH3 5 1 0
on earth BIOS
Centrifuge C~2 4 1.4 5.3 Reduced (only .
radius higher ner-
320 mm vous activity)
Centrifuge I~~ 4 l.l 5.3 Complete (all
radius procedures)
98 mm
Vivariums Ma~ n group ffiC 6 1 , ~
Additional ffiC~ 18 1 0 Partial
- group (1/3 proce-
dures)
Intact ~3 9 1 0 Reduced (only
animals , higher ner-
_ group vous activity)
(with no
sensors
implanted)
mean that information about those animals which experienced the most significant
- changes in higher nervous activity would be excluded from the analyzed material. ~
Concurrently with recording the state of higher nervous activity, we used a five-
point system to score the feeding activity of animals in the feed compartment of
the maze. The lowest point score, l, meant total absence of a reaction to the
feeder. The highest, 5, signified fast eating of all of the feed. Intermediate
poin't scores (2,3,4) corresponded to different degrees of taeakening of the feeding
reaction (not finishing all of the feed, slow feeding).
We began training the rats to find the feed in the rnaze 2 weeks before the flight,
and stopped the training 5 days before the flight. The animals were distributed
such that each rat in the experimental group had a partner in the control group
with similar higher nervous activity indicators. The symbols used to refer to the
different groups and descriptions of the effects to which they were exposed are
shown in Table 1.
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60
CN~ '
~ ~
x ~
- 40 -x x-x- x
~
, X\ ~u~
X-x-x-x
~~flX
~
~
\
20 nNJ
\
8ifa
~ ~ o
~ u2
.
O B~" ~ ~-r ~ ~i � �~,~-n~s!?
:
: _
3 ,5 7 . 9 f1 >3
Figure 1. Refusals to Run the Maze in the Recovery Period: abscissa--
days after landing; ordinate --refusals to run the mazes, percent.
Letters beside the curves are the abbreviated names of the animal
groups. See text for e~splanation.
dt,~
~20 ~ '
I~
~00
I
~1)
6i~adc ea 80
� 6-
60
4 ~
UO
2 ?0
- ~ 8x,~ Qlf BH~ r,y~ q; nH~ nu2 ~
ex,~ el( BRd CNJ uz OHJ lIUZ
Figure 2. Mistakes in Recovery Period: Figure 3. Maze Running Time in the
abscissa--animal groups (see tPxt and Recovery Period: ordinate--time, sec.
Table 1); ordinate--difference between See Figure 2 for other symbols.
group average for indicator under analy-
sis in the testing period and two addi-
tional experiments performed subsequently
Key:
1. Absolute units
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Body temperature sensors were implanted into the abdominal cavity of all animals
- (except group HKg) 2 weeks before the flight.
The maze tests were resumed 3 days after the biosatellite landed. For 16 days we
studied the animals' capability for restoring and reinforcing the previously developed
feed-seeking habit.
An experiment with a heightened functional load was performed on the 17th-18th days
in order to study tiring of the higher divisions of the central nervous system.
During this experiment the number of maze runs was increased to 16, and to avoid
saturation of the animal, the number of individual reinforcement feedings was re-
duced corresporidingly.
The results were treated statistically using algorithms for comparison of two
processes, calling for dispersion analysis (5), and, in some cases, analysis on the
basis of the "phi" and inversion tests.
Research Results and Discussion
The results of research on rats maintained in a vivarium showed that the behavioral
habit previously developed in the maze was easily restored by intact animals, and
with somewhat greater difficulty by rats subjected to surgery (implantation of
sensors in the abdominal cavity). In this case the greatest difficulties were noted
among animals tested in accordance with the complete program. This can be distinctly
seen from change in the number of refusals (Figure 1), the maximum increase of which
was recorded in group BK, and the minimum of which was recorded in ffiCH (P < 0.05).
Changes in the number of refusals in group ffiCA were close to the indicators for
group HKx; however, the rest of the parameters underaent significantly greater
- alteration than in the intact control (HiCH). Differences in the number of mistakes
_ (Figure 2) and time (Figure 3) were significant (P < 0.05).
Similar relationships in the change in the number of refusals recorded for control
groups subjected to comparison ~aere also revealed in the experiment with the
heightened functional load.
Research conducted on animals maintained in a m4ck-up of the biosatellite also re-
veals certain difficulties in restoring the maze habit. All indicators for animals
_ in group CH3 changed for the worse, without exception, in comparison with the same
indicators for animals maintained in vivariums (BK). Differences between the groups
were significant in relation to both the refusal criterion (P