TRANSLATIONS ON USSR SCIENCE AND TECHNOLOGY BIOMEDICAL AND BEHAVIORAL SCIENCES (FOUO 8/78) EFFECTS OF NONIONIZING ELECTROMAGNETIC RADIATION

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• Publication Date: February 24, 1978
• Content Type: REPORT

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UIV L Y '
? ,7P RS L/7629
2 4 February 19 7.8
TRANSLATIONS ON USSR SCIENCE AND TECHNOLOGY
BIOMEDICAL AND BEHAVIORAL SCIENCES
CFOUO 8/78)
EFFECTS OF NONIONIZING
ELECTROMAGNETIC RADIATION
U. S. JOINT PU~LIC~-TI~NS RESEARCH SERVICE
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BIBLIOGRAPHIC DATA
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'TRANSLATIONS ON USSR SCIENCE AND TECHNOLOGY
24 Februa 1978
BIOMEDICAL AND BEHAVIORAL SCIENCES (FOLIO 8/78)
6.
Effects of Nonionizing Electromagnetic Radiatio
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 JP RS L/7629
24 February 1978
TRANSLATIONS ON USSR SCIENCE AND TECHNOLOGY
BIOMEDICAL AND BEHAVIORAL SCIENCES
(FOUO S/78)
EFFECTS.OF.NONIONIZING..
ELECTROMAGNETIC RADIATION
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I'OR OF IAL USL OIJLY
This serial publication contains abstracts of articles and news items from
USSR and Eastern Europe scientific and technical journals on the specific
subjects reflected in the table of contents.
Ph otoduplications of foreign-language sources may be obtained from the
Photoduplication Service, Library of Congress, Washington, D. C. 20540.
Requests should provide adequate identification both as to'the source
and the individual article(s) desired.
CONTENTS
PAGE
Reactions To Unperceived Stimuli in the Presence of Functional
Disturbances of Sense Organs
(G. V. Gershuni; SOVREMINNYYE TENDENTSII V NEYROFIZIOIAGII, 1
1977) ........................................................
The Role of Isolation, Environmental Deprivation and
Enrichment in Formation of Behavior
(A. D. Slonim; SOVREi~NNYYE TEDENTSII V NEYROFIZIOIAGII, 1977) 19
_ a _ [III= USSR - 22 S&T FOUO]
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REACTIONS TO UNPERCEIVED STIMULI IN THE PRESENCE OF FUNCTIONAL DISTURBANCES
OF SENSE ORGANS
Leningrad SOVREMINNYYE TENDENTSII V NEYROFIZIOLOGII in Russian 1977 pp 68-81
[Article by G. V. Gershuni, Institute of Evolutionary Physiology and
Biochemistry imeni I. M. Sechenov, USSR Academy of Sciences, Leningrad]
[Text] In this article, I should like to discuss the functions of human
sense organs described by such nonstandard criteria as occurrence of reac-
tions to unperceived stimuli. The appearance of such reactions is very
clearly demonstrable in the presence of some pathological states of the
central nervous system as well as, as we have established, in healthy
individuals who have developed conditioned reactions to stimuli that are
below the threshold of sensations of which they are aware.
Studies of this type of phenomenon were conducted by a team of workers at
the Physiological Institute imeni I. P. Pavlov, USSR Academy of Sciences,
for many years (Gershuni, Alekseyenko et al., 1.945; Gershuni, 1947, 1949,
1955). A brief description of this research was published by L. A. Orbeli
in 1949. The results obtained had not been summarized to this time. In
this article, we shall discuss phenomena observed in the presence of
pathology of of the central nervous system occurring as a result of aerial
concussion, closed skull trauma and mental trauma.
The onset of marked vegetative and other reactions to stimuli delivered to
sense organs, the functional state of which is characterized by the usual
clinical methods as partial or total loss of the relevant type of sensibilit_y___.__
(auditory, visual, tactile, nociceptive, olfactory, gustatory) is not unex-
pected. The descriptions of disorders referable to perception of exogenous
stimuli, observed in the presence of 'closed skull trauma and mental trauma
(i.e., in the presence of excessive stress for man) of wartime and peacetime,
provided by clinicians, have long since indicated this (Veraguth, 1909;
Myasishchev, 1929; Panov, 1933; Astvatsaturov, 1935).
What our studies contributed that is new is determined by the introduction
of a quantitative evaluation of phenomena, based on the choice and record-
ing of a specific set of reactions, in the first place, and development of
measurement procedures for threshold stimuli inducing these reactions, in the
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second place, along with classical methods of determining the thresholds of
perceived sensations in response to the same stimuli. By means of these
procedures, we were able to establish two quantitative criteria characteriz-
ing reactions to unperceived stimuli: 1) The difference between the threshold
levels of stimuli inducing this type of reaction and thresholds of per-
ceived sensations.* This difference, which expresses the range of inten-
sity of stimuli that are not perceived, was referred to as the subsensory
zone (Figures 1 and 2). We have used the term, subsensory (Gershuni,
Alekseyenko, et al., 1945) to refer to the actual reactions that arise to
stimuli below the threshold of perceived sensation. 2) Differences in
characteristics of reactions arising in response to unperceived and perceived
stimuli.
Hearing
Cutaneous
V sensibility
Figure 1.
Thresholds of galvanic skin response
(GSR) and thresholds of sensation in
'patient M. upon sonic and electric
stimulation of the skin.
Vertically: intensity of sonic stimulus
(dB) in relation to normal hearing
threshold (0); intensity of skin stimula-
tion (V) for cutaneous sensibility.
White circles--GSR to stimuli below
sensation threshold; black circles--
sensation thresholds; black circles
with crosses--GSR to perceived stimuli.
Striped area--range of stimuli that are
not perceived (subsensory zone); L--left;.
R--right. ?
These criteria enabled us to observe the dynamics of the pathological pro-
cess in the presence of impaired perception of exogenous stimuli. Thus,
at the stage of profound impairment of perception, the difference between
threshold levels of stimuli inducing vegetative reactions and thresholds of
sensation could reach enormous values, then undergo typical changes in
the course of recovery of function. At the same time, we observed typical
changes in the characteristics of the reactions.
In order to determine the thresholds of stimuli., in our first work, which
dealt with research on functional impairment of sense organs in the presence
of wartime trauma (aerial concussion)(Gershuni, Alekseyenko et ~1., 1945),
we used the reaction of dilatation of the pupil, as well as electroencephalo-
graphic indices, the change in spontaneous rhythm and initial responses
(Gershuni, Klaas et al., 1945). In our subsequent studies, we made extensive
use of the galvanic skin response (Gershuni, 1947).
*The term, "perceived (or overt) sensations," is used to refer to phenomena
demonstrable in standard psychophysical measurements, as opposed to another
group of phenomena referred to, even by I. M. Sechenov, as "sensations in
discrete form" (1863).
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o ~-`
17.01
2.02 7.02
i4 /~~
i ~~} 4
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i
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2 4 6. 8 f0 !2 14 16 1d 20 22 V
Figure 2. Reduction of subsensory zone and recovery of cutaneous
sensibility of patient~at the recovery stage (A) and
magnitude of GSR as function of intensity of stimuli
in and beyond the range of the subsensory zone (B)
In A:
X-axis, day of examination; y-axis, intensity of electric stimulus (V).
White circles, GSR to sublim~.nal stimuli; black circles, perception
thresholds. Striped area, subsensory zone.
In B:
X-axis, intensity of electric stimulus (V); y-axis, magnitude of GSR
(relative units). White circles, stimuli that did not elicit responses;
white circles with crosses, GSR to subsensory stimuli; black circles,
GSR to perceived stimuli. GSR threshold, S V; perception threshold, 14 V
Such vegetative and electroencephalographic reactions were found to be
t~.he most sensitive indicators of activity of the central nervous system
occurring in response to stimuli that were not perceived.
Figure 1 illustrates a typical case of altered threshold values of stimuli
evoking galvanic skin reactions and perception thresholds in a patient with
severe, unilateral hypesthesia of the skin and impaired hearing (on the
left) as a result of brain concussion. It shows that galvanic skin reac-
tions, to both electrocutaneous and sonic stimuli on the left, arise at
threshold levels that are much lower than the perception thresholds for
cutaneous and auditory sensations. Upon stimulation of the right side of
the body, with normal cutaneous sensi.bilir_y, the GSR occurs only in
response to stimuli that reach the threshold of perceived sensations.
Along with an increase i.n sensibility as demonstrated by the thresholds of
perceived sensations, decrease in subsensory zone until~it disappears com-
pletely are inherent in recovery of perceptive function in the presence of
the above-mentioned disturbances. This phenomenon was demonstrable both
in the pupillary reaction test in the course of restoration of hearing
(Gershuni, Alekseyenko et al., 1945) and in the GSR in the course of
recovery of cutaneous sensibility (see Figure 2A).
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The characteristics of vegetative reactions undergo substantial changes
when the stimuli that induce them reach the sensation threshold, i.e.,
when they begin to be perceived. The most typical of these changes are
a reduction in magnitude of reaaction and faster extinction under the in-
fluence of successive stimuli. The former phenomenon emerges distinctly
when measuring the magnitude of reactions as function of intensity of
stimuli that are within and above the subsensory range.
Figure 2B illustrates such data, obtained upon measurement of the galvanic
skin response (GSR) as related to intenstiy of electrical stimulation of
the. skin in a patient with markedly diminished dermal sensibility; the
figure shows that when the stimulus reaches the sensation threshold there is
a sharp decline of GSR. The entire curve for perceived stimuli is shifted
in the direction of higher intensities. Figure 3 illustrates differences in
magnitude of GSR and rate of extinction with delivery of stimuli to areas of
skin with normal and markedly diminished sensibility.
As can be seen in Figure 3, upon stimulation of the hypesthetic skin area,
delivered at intervals of 1 to 1.5 min, GSR of greater amplitude occur
throughout the period of stimulation; on the side with normal sensibility,
GSR. occur only in response to the first stimulus reaching the sensation
threshold; there is no reaction to subsequent stimuli.
We should mention one more distinction in the dynamics of effects of un-
perceived and perceived stimuli; it i~s referable to increased sensibility
(sensitization) under the influence of successive stimuli. This phenomenon,
which was studied in healthy individuals by A. I. Bronshteyn (1946), is
very marked in patients with impaired dermal sensibility (Figure 4).
Figure 4 shows that with successive delivery of stimuli to the skin surface,
which induce GSR, the threshold of perceived sensation is reached with
the ninth stimulus of very great intensity (3 times greater than the
threshold for occurrence of GSR). With continued stimulation, there is
a drop by almost SO% of sensation thresholds (sensitization phenomena);
accordingly, there is a sharp reduction in the subsensory zone. The thresholds
of occurrence of GSR to subsensory stimuli do not undergo apprecialbe changes.
Thus, heighted sensibility is inherent expressly in conditions, under which
conscious perception of exogenous stimuli occurs.
In different cases of impaired conscious perception, different variants of
the above-described phenomena may be observed. The described features of
reactions arising to subsensory stimuli, namely, greater amplitude and
stability in response to a series of stimuli and lack of dynamics typical
of conscious perception (extinction of reactions and sensibilization), are
the typical signs that are demonstrable in studies of diverse forms of
sen.si bi].ity (auditory, cutaneous) and different vegetative reactions
(papillary, galvanocutaneous).
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v
120
24
1
I!
111
NO
22
f00
ZO
4
90
18
~
80
16
4
70
f4
, ~
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0 4
60
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50
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40
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0
0
30
6
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20
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f0
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0
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S f0 15 20 ZS 30 3
S k0 min
Figure 3. GSR to successive stimulation of anesthetized and normal
skin areas
X-axis, time in min; y-axis, intensity of single electric stimuli delivered
to the skin (by condenser discharges)(V) and magnitude of GSR (relative
units). White circles, unperceived stimuli that do not elicit GSR;
white circles with crosses, unperceived stimuli; black circles with crosses,
perceived stimuli associated with GSR; black circles, perceived stimuli not
associated with GSR. Columns, magnitude of GSR as related to stimuli.
Patient So-va (deep anesthesia of both legs, from the toes to the knee). Ef-
ferent electrodes on the left hand, silent electrode on the left foot.
Stimulating electrode placed as follows: I and III on the lower third of
the right leg, anesthetized region; II on the lower third of the thigh on
the same side, area of normal sensibility (according to experiments of
A. M. Alekseyev and A. A. Arapova).
A comprehensive clinicophysiological description of a group of patients
who had sustained air concussion (106 people) was published previously
(Gershuni, Alekseyenko et al., 1945). The data illustrated in Figures 1-4
a're referable to a group of patients who had sustained closed brain
trauma in peacetime (concussion)(studies of Alekseyev and Arapova; Arapova
and Orlova; Arapova, Gershuni and.Orlova). G. V. Gershuni (1947), A. A.
Arapova and G. M. Orlova (1948) published a brief report. All such patients
presented impaired perception with severely marked subsensory reactions (22
cases). In a small group of patients (six people), in the history of which
the effects of mechanical factors inducing trauma could not be established,
similar perception disorders were observed, characterized by marked subsensory
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reactions; the factors inducing these disturbances should have been referred
to psychogenic ones (excessive stress for the individual).
As we know, in the clinical literature such disturbances were often designated
by the term "histerical" or "hysterotraumatic" (Astvatsaturov, 1935). A
similar case in wartime was described in 1945 (Gershuni, Alekseyenko et al.,
1945). In a work published in 1957 (Avakyan et al., 1957) there is a
comprehensive physiological analysis of one peacetime case. Patients with
impaired hearing (Kristostur'yan, 1952; Gershuni et al., 1954) and cutaneous
sensitivity (Arapova and Orlova, 1948) of peripheral origin were also
studied.
In addition, studies were made of patients with impaired cutaneous sensibility
as a result of lesions to different levels of the nervous system (syringo-
myelia, lateral amyotrophic sclerosis, hemorrhages in the region of the
pons varolii, comminuted trauma to the right parietal region), but no
sensory reactions were demonstrable; the GSR thresholds were found to either
coincide or to be above the sensibility threshold. In only one case of a
disorder of vascular origin (cerebrovascular thrombosis) with the main
.focus localized in the right parietal region, accentuated ;GSR were observed
to stimulation of the skin, which were considerably lower than the threshold
of perceived sensation. In this case, it was not possible to rule out
lesions to other structures, including subcortical ones.
With reference to the above-submitted data as a whole, it should be indicated
that the same symptoms, characterized by clinical neurologists (Kryshova,
1945) as subcortical-stem symptoms, are found in patients who have suffered
wartime air concussion and peacetime concussion, with which there is typical
occurrence of subsensory reactions. The obtained data are inadequate for
more precise description of the structures, with injury to which there is
typical occurrence of subsensory reactions. The set of phenomena observed
with the described disorders of perception is quite typical. It can be des-
cribed as the syndrome of unconscious [unperceived] perception or, more
briefly, the subsensory syndrome.
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In research dealing with the effects o.f stimuli below the threshold of per-
ceived sensation, in addition. to vegetative reactions, electroencephalographic
indices were studied. The results of analysis of the electroencephalogram
(F.EG) and changes therein under the influence of stim~ili in patients who
had suffered air concussion revealed that spontaneous (background) activity
(occipital and temporal leads were used) deviated appreciably from normal
and was characterized by the following: 1) instability of the main 8-12-s
alpha rhythm, ready disappearance thereof and change to either a faster
o~r slower rhythm; 2) presence of slow waves, of the order of 1-3 per second
aid spike discharges considerably exceeding the normal EEG variations;
3) impaired electrical activity of tYie cerebral cortex during sleep (Gershuni,
A.l.ekseyenko et al., 1945; Gershuni, Kl.aas et al., 1945).
The responses to exogenous stimuli (sonic, photic, mechanical, olfactory)
are usually manifested by a change in amplitude of dominant EEG rhythm, appear-
ance of relatively fast electric waves and new rhythms at the time the stimuli
are used.
Iii the patient group examined, the reactions to exogenous stimuli were
demonstrable with stimuli below the perception threshold.: Thus, distinct
reactions were demonstrated under the influence of sonic stimuli in indi-
viduals who were totally deaf, as well as in response to stimuli delivered
to the skin in the presence of severe decrease in tactile and nociceptive
sensibility, under the influence of odoriferous substances in cases of total
lack of olfaction.
A comprehensive study of reactions to sonic stimulation revealed that the
electrical responses are the most distinct at specific times after trauma
was sustained. Paradoxical changes were observed in response to relatively
mild stimuli and a significant decrease in such changes was found with
increase in force of the stimuli. The intensity of the electrical responses
diminished in the course of the overall recovery process (Gershuni,
Alekseyenko et al., 1945; Gershuni, IClaas et al., 1945).
The subsensory zone could be established from the difference between thresholds
of electrical response of the cortex and thresholds of auditory perception,
as had been done with respect to thresholds of other reactions (pupillary,
galvanocutaneous).
In the patient group studied, the nature of responses to exogenous stimuli
presented several distinctions, as compared to the normal findings. In addi-
tion to the paradoxical response to mild stimuli, which we have already dis-
cussed, the reaction is quite often manifested by intensification, rather
than depression, of alpha rhythm, which is very similar to the electric
response to exogenous stimuli in the intermediate state between sleeping
ar-d waking.
It is significant that, in the course of recovery of different types of
sensibility and speech, the stable slow rhythms do not demonstrate a
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correlation to these changes, whereas the reactions to exogenous stimuli
change in accordance with recovery of sensibility of a given sense organ.
One of the parameters used for quantitative description of the electrical
reaction of the cortex to sonic stimuli was the latency period of such
reactions. The latency period was defined as the time that elapsed from
the moment sound was delivered to the moment of appearance of EEG changes
(accentuation or attenuation of alpha rhythm, accentuation of beta rhythm,
appearance of initial response). Shorter latency periods were found in a
number of subjects who had sustained air concussion. Increase in the
latency period to close to normal levels occurred concurrently with recovery
of hearing and speech. Shorter latency periods were not observed in
patients with penetrating skull wounds.
T.abl~ 1. Latency periods of EEG responses to sonic stimuli
Nature of Disturbance Subject T.atenc~period
Penetrating wound
M, ~ 0.23
p, 0.45 0.32
I, 0.29
F, 0.29 0.26
G. 0.23
Hearing and speech disorders P. 0.07
following air concussion A. 0.08 0.09
Z, 0.14
T; 0.15
I, 0.20 0.12
~, 0.13
Table 1 lists mean latency periods for normal individuals, patients with
penetrating wounds to the temporoparietal region and patients who suffered
air concussion with hearing and speech disorders at the early posttraumatic
period. The data referable to latency periods at different stages of
recovery of hearing and speech (Table 2) are demonstrative. As can be
seen in Table 2, the latency period increased by 3 times during the
period, within which hearing and speech are restored.
The EEG data obtained on patients who suffered peacetime concussion presented
the same feature in common, appearance of reactions to stimuli below the
threshold of conscious perception. To illustrate this, we have submitted
in Figure 5 the results of EEG tests on the same patient, whose GSR was
studied comprehensively (see Figure 1). This patient has a distinct alpha
rhythm. Sonic and cutaneous stimuli (Frey's bristles and hairs) delivered
to the hypesthetic half of the body induced subsensory reactions of de-
pression of alpha rhythm. A comparison of these reactions to those occur-
ring~in response to stimulation of the other half of the body, with normal
cutaneous and auditory sensibility, showed significant differences in
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latency periods and duration of alpha rhythm depression. Thus, the latency
period for subsensory sonic stimuli (on the left) is considerably shorter
(0.18 s) than upon subliminal (perceptible) stimulation of the right half of
the body (0.45 s), The reaction to subsensory stimuli (0.1 s) was found to
last much less time than in response to stimuli above the sensibility
threshold (2.4 s) with the same physical intensity of these stimuli. The
differences in duration of alpha rhythm depression, as can be seen well on
the oscillograms, are due to the after-effect, which is several times greater
with stimuli that elicit conscious sensations. The phenomenon is equally
marked under the influence of both sonic and cutaneous stimuli.
Table 2. Changes in latency periods of EEG responses (according to Gershuni,
Klaas et al., 1945) and at different stages of recovery of hearing
and speech
Latency
Patient Date examined Condition of hearing and speech period(s)
A-v, trauma 19 Sep 43 Hearing and speech absent 0.08
on 15 Aug 43 25 Sep 43 Appearance of hearing at all fre-
quencies on the left, no speech 0.16
12 Nov 43 Appearance of hearing on the right,
speaks in a distinct whisper 0.30
F'-f, trauma 14 Sep 43 No speech or hearing 0.07
on 21 Aug 43 15 Oct 43 Appearance of hearing at all fre-
quencies in both ears; stutters 0.12
22 Oct 43 Speaks more distinctly 0.13
21 Nov 43 Hearing improved, speaks freely 0.22
The foregoing data characterized perception disturbances that could be demon-
strated by specific procedures of psychophysical measurement. These pro-
cedures involved the use of simple physical stimuli (for example; pure
tones for hearing) and examination of reactions that did not require preli-
minary experimental development. These conditions facilitated significantly
measurement of thresholds and determination of range of stimuli that were
nut consciously perceived, However, such a study was not sufficient to
characterize perception disturbances; in the first place, it was necessary
to obtain data on how more complex stimuli, including those used in real
life (the sounds of speech, for hearing), are perceived; in the second place,
we had to determine the extent to which it is possible to learn to perform
specific activity in response to stimuli that are not perceived. Experi-
mentally, this was a question of. developing conditioned reactions to sub-
sensory stimuli.
Data pertaining io perception of speech sounds and development of conditioned
reactions to subsensory stimuli had already been obtained in the first in-
vestigation (Gershuni, Alekseyenko et al., 1945). We submit data below,
which pertain to perception of speech sounds described in this work.
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Figure 5. Reactions of depression of alpha rhythm in the EEG of
patient M. Left-sided decrease in tactile, nociceptive
and auditory sensibility following concussion. Occipital
lead.
'Top, stimulation mark; bottom, time (1 and 0.2 s), From top to bottom:
tactile stimulation of. dorsal surfaces of left arm (subliminal stimulus);
tactile stimulation of the same area on the right (supraliminal stimulus);
sonic stimulation on the left (1000 Hz; stimulus is 20 dB below sensibility
threshold); sonic stimulation on the right, same intensity as on the left
(stimulus is 5 dB above threshold)
Perception of speech sounds: Studies of thresholds of auditory sensibility
using sinusoidal oscillations of a specific frequency (pure tones) revealed
a consistent course of restoration of hearing at different frequencies.
In some patients, there was some instability to the degree of decline; however:,
these fluctuations of threshold did not exceed 10-12 dB. Under specific condi-
tions, speech sound stimuli elicited changes of a very different nature,
which could be roughly described as constituting 30-50 dB. The conditions,
under which these phenomena could be observed, ensued from consideration of
a very interesting procedure for testing hearing, described by L. B.
Perelman (1943) in a study of similar patients. This procedure, which the
author called "combined test," is based on concurrent delivery of verbal and
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written signals, fixing the attention on the latter: questions are posed to
the patient in writing, and he answers them in writing,or veibally (if
he can speak). Concurrently with writing the phrase, the experimenter
pronounces it at a certain volume; several questions, which follow one
another, are written more and more indiscernably. But the subject continues
to give the correct answers, and this could only be the result of a reaction
to the sounds of speech. We were able to completely corroborate the pheno-
menon described by L. B. Perelman, having observed it distinctly in a number
of patients (10 out of the 35 studied). By varying the volume of speech
sounds, it is possible, in some, cases, to determine how the reaction
threshold changes on the basis only of fixing the eyes on the pencil moving
along the paper. Of course, this is a rough determination, but this does not
diminish its basic significance. Let us consider the two most vivid cases.
1) Patient S-ov lost his hearing and speech after an aerial bomb explosion.
At the time of the study, he had regained'speech; he had no hearing on the
left and could hear loud speech close to the concha on the right. He could
respond to relatively low speech, the intensity of which was at least 30-40 dB
lower than Che level to which he usually reacted, by fixing his eyes on a
moving pencil (nothing is written on the paper). The test was repeated many
times; each time, fixing the eyes on the pencil elicited a distinct response
to low speech. 2) Patient Ya-uk lost his speech and hearing after an aerial
bomb explosion. At the time of the study he stuttered. When addressed
directly, he could not hear loud speech uttered at close range, near the
concha, or the sounds of tuning forks. He could respond to moderately,
loud speech at a distance of 1 meter when fixing his eyes on a moving pencil,
but did not respond to low speech. Thus, fixing the eyes on a moving pencil
elicits a reaction to speech sounds, the intensity of which is at least 30 dB
lower than the intensity of sounds that are inaudible to the patient when
delivered at the concha directly. After 4 days, the patient suddenly began
to hear a loud shout near the concha.; audiometry revealed a decline of
audibility threshold on the order of 100 dB at all frequencies. ,After 2 more
days, he perceived moderately loud speech addressed to him. Audiograms
showed typical decline at moderate and high frequencies (of the order of 60 dB).
We sY-all submit several facts referable to the study of development of con-
ditioned reactions to subsensory stimuli and stimuli beyond this range. We
tried to develop conditioned reflexes in response to sonic signals, in the
presence of external signs of deafness. In our experiments, we used primarily
the method of verbal (written) reinforcement, according to A. G.
Ivanov-Smolenskiy. Upon appearance on a screen of the written signal to
"depress," the patient squeezed a rubber bulb. A metronome began to tick
a few seconds (2-3) prior to appearance of the instruction. 'Depression
of the bulb at the sound of the metronome, before the writing appeared,
served as an indication of formation of a conditioned reaction. In several
tests we used electric stimulation of the skin rather than verbal reinforce-
ment. The patient had to perk his hand away from the electrode during
passage of current. The clicking metronome preceded delivery of current.
Experiments on 29 patients who were totally deaf or had a severe hearing
impairment (over 80 dB) revealed that it may not be possible to develop
a conditioned reflex in response to the metronome (intensity level of the
11
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order of 40-50 dB). In this respect, the experimental results were quite
similar. As a control, we conducted tests on the same group of patients,
using contact with the skin as a stimulus rather than the metronome. Test-
ing of 21 patients revealed that 16 of them could develop a conditioned reflex
to tactile stimulation as a result of several combinations. Finally, on
:mother group of patients (15 people) whose hearing had been restored to
the e:ctent of perception of the sound of the metronome, the conditioned
refle:c could be readily formed in 13 cases.
The above facts indicated, on the one hand, that the impossibility of forming
conditioned reflexes to inaudible metronome sounds could not be the result
of sorue methodological flaws that generally prevented formation of conditioned
associations under our experimental set-up; on the other hand, they showed
that conditioned reflexes to sonic stimuli can be readily formed in res-
ponse to sounds above the audibility threshold.
We altered the experimental set-up to determine the significance of audibility
threshold to development of conditioned reactions. Thus, we used a generator
of electric oscillations and a telephone as sonic stimuli. We determined
the audibility threshold and range of the subsensory zone according to the
pupillary reaction at a frequency of 512 Hz. Table 3 illustrates the
results of these studies; it contains the typical data obtained on one of the
patients who had suffered air concussion. These data show rather clearly
that formation of a conditioned motor reflex (movement of the fingers) can
he distinctly demonstrated only with sounds that are slightly above the
audibility threshold. No conditioned reactions were developed to sounds of
considerable intensity, 43 and 50 dB above the threshold for the pupillary
dilatation reaction, i.e., stimuli that definitely elicited a flow of
afferent impulsation in the acoustic tract.
Table 3. Development of conditioned reflex with the use of different
intensities of sound, in patient Sh.
Intensity of sound Intensity of sound Number
(512 Hz) above the (512 Hz) in relation Formation of of Level of
normal audibility to patient's audibility conditioned combi= conditioned
threshold (dB) threshold (dB) reflex nations reflex
92 _7 No 30 --
99 0 (threshold) No 37 --
106 +7 Yes 4 mild
114 +15 Yes 4 strong
'.~~;~~~
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The patient had suffered air concussion; he was totally deaf on the left,
8U dB threshold of cochleopupillary reflex; diminished hearing on the
right, 100 dB, 50 dB threshold of cochleopupillary reflex. subsensory
zone, 50 dB. The telephone was put to the right ear.
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. FOR OFFICIAL USE ONLY
nditioned reactions to subsensory stimuli
ment of CO ated in
uestion of develop exception was investig an et al.,
The q impaired auditory P Avaky
in the presence of uent study in our laboratory ( deaf in both
greater depth in the subseq female]), who was totally 18 times) in
act (patient M? ~ times
1957). The subj trauma, was tested many during an experimental
ears as a result of mental She regained her hearing
5 months: we used the galvanic skin reaction
the course of 1? reliminary development. This
To test the absolute threshol s, the extinction
session. without any P
demonstrated in numerous case ,
response, which occurred
ically intensities at three fre-
was stereotyP marked. The threshold
did not differ by over 10-12 dB from the
phenomenon was minima11y4000 Hz)
quencies (200, 1000 and i.e.~ they were within the normala~cogrding
be normal, clinical audiometry. Thus,
levels considered to led in e of intensity of
according to the criteria adop
the subsensory zone involved the entire rang
L-o the GSR, .
sonic frequencies used.
conditioned reactions thesmethod
es of reactions to develoblinki.ng reaction using
We used two tyP range 1)
stimuli within the subsensory which permits measurement of absaevelop~
Avakyan (1955), during
in individuals with normal hear n ,
developed by R. V.
ential thresholds voluntary motor reactions based
differ onses to sounds; b) G, Ivanov-Smolesnkiy
meet of conditioned resp the method of A? s of. movements
on verbal or written instructions by o ra~hic recording ~~
sin "lift your finger.
(1933)? We made mechanical earancecof theg g when appearance or this
onse to aPP ed rapidly
performed in resp
Normally, conditionedaneaudlonsignailop
sign was preceded by the intensity of
reactions to sounds, range)
Development of conditioned ollSkagg(i?e?, within the subsensoryh it was
ed from 90 as compared to normal, althoug
which was Chang 1 altered,
fixing of conditioned blinking re-
was found to.be sharp y articularly slow: about 200 combinations
possible. Formation and p was very Even
flexes in response to low intensities
for this, i.e., about 20 timesmm~~e f differentiations,
an
were required develop
greater differences were observed in
'ndicative of significant intensification of success ve
and this was i
extinction inhibition.
development of. conditioned reflex finger
the blinking reaction, ust like the conditioned motor reac-
Un].ike could not be
lifting with written reinforcement, ~ ment of conditioned motor
the finger away with electric reinforcement,
lion of jerking id develop hotic stimuli.
There was rap in response to p
demonstrated at all? e in this patient,
reactions of this tYP exception disorders that can
The above facts are conditioned reflex method.P
be detected by the our studies,
occurred in this patient during
of hearing, which earance of a marked condition rwhich
henomenon,
e of aPP
ery
i
hi
s p
m
Recov
with the t
sound. T
coincided exactly onse to a erimenter during a test,
the finger in resp the exP
lion of articular interest, was detected by
was of p
13
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 --:t._ ~:.;5..++T'sz-_,??= u,---~-. _ .. .. _~?.z~~aw?;:.~.~ tr:G~._._.:r3~~~.=-W ~r,~i?^-~ ~ -'~r~
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on the basis of developing reduction of latency .periods of motor reactions.
Thus, with delivery of the 18th stimulus, the motor reaction recorded on
the electromyogram, occurred 0.1 s before appearance of the sign "lift your
finger." Thus, we see that there is development of a conditioned motor
response to sound. The appearance of distinct, conditioned finger movement,
which could not be demonstrated in any of the preceding tests, compelled the
experimenter to open the door and enter the chamber, in which the patient
was situated. The patient was excited and her face was very flushed; when
the experimenter appeared she said: "Last time, I heard something before
the sign appeared; before, only the sign appeared. Before this, I felt
something like a blow to the head." In answer to the question, "Do you
hear ine?" the patient exclaimed "I do hear, I hear." She was very excited,
tearful and exclaimed "I can hear," and hugged everyone there. The test
was interrupted.
Continuation of the study after a break revealed that, with each delivery
of the sonic signal there was a conditioned motor reaction, in the form of
lifting a finger, and after being instructed by the experimenter to respond
verbally to the stimulus, the verbal report of "sound" appeared in response
to the stimulus.
A comparison of the signs of restoration of hearing in this patient to the
reactions recorded on oscillograms shows that appearance of the first
marked conditioned reaction of lifting the finger in response to a sonic
stimulus coincides exactly with the same stimulus, with which the patient
first reported auditory perception preceding appearance of the sign.
Let us try to evaluate the obtained facts. According to the measurements
of absolute thresholds according to the GSR, the sensibility of this
patient's auditory system was close to normal. Hence, at sound intensities
that were used to develop conditioned motor re~.ctions of finger lifting
(50 dB), the flux of afferent impulsation in the auditory tract should have
been quite significant. However, this flux could not be used to develop
the motor act, which is referable to a complex system of movements referred
to as voluntary, in just the same way as for construction of .conscious per-
ception of the signal.
However, development of conditioned reactions could occur on the basis of a
motor reaction of very specialized significance (blinking reflex); in this
case development of the reflex was slow and distorted.
The foregoing indicates that the subsensory syndrome is characterized by
rather profound and, at the same time, differentiated disturbances of central.
nervous system function with complete preservation of afferent flux. Very
generally, these disturbances can be described as impaired use of information.
contained in the afferent flux in specific sensory pathways.
lip
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What facts can we offer to describe the changes that occur in the central
nervous system i.n the presence of the subsensory syndrome: First of all,
let us discuss the characteristics of generalized vegetative and electro-
encephalographic reactions.
The following were typical of vegetative reactions: a) greater amplitude of
reactions; b) considerably greater stereotypy of responses in the case of
multiple repetition of stimuli.
The following are typical electroencephalographic reactions: 1) shorter
latency period and shorter inhibition of. alpha rhythm (when this rhythm is
pronounced); b) shorter latency period of other reactions (appearance of
alpha rhythm in the early responses); c) distorted (accentuated) reactions
to mild stimuli.
It should be stressed that the above changes in characteristics of reactions
are typical of stimuli in the subsensory range. Such findings are parti=
cularly distinct in studies of the process of recovery of perception func-
tion. This warrants consideration of the above-described changes as
indications of changes in some common elements of the chain of events occur-
ring in the central nervous system, needed for conscious perception of an
external signal, as well as for development of conditioned motor reactions
related to the system of voluntary movements, rather than concomitant pheno-
mena.
What is the significance of these changes? On this score, several hypotheses
can be expounded, which are based on the attempts to interpret the aggregate
of observed changes as an expression of impairment of some general biological.
reactions, in particular, the orienting, waking up reactions, the set of
emotional reactions determined by subcortical structures. Very plausible
assumptions of this kind were voiced in the work of B. D. Asafov, per-
forined in the laboratory of A. M. Zimkina (Asafov, 1965) and T. N.
Reshchikova, performed in the laboratory of E. A. Kostandov (Reshchikova,
1969), as well as Zakharova (1973), in describing subsensory reactions ob-
served in patients with lesions to deep structures of the brain.
It is quite significant that, both in our studies and the above-mentioned,
there is a link between the subsensory syndrome and disturbances referable
to iieep structures of the brain. However, it is not deemed, possible at
this time, on the basis of the available data, to describe .these structures
more precisely and, accordingly, to expound hypotheses with sufficient sub-
stantiation concerning the role of various regulatory systems in the
observed functional disturbance.
However, we could use another approach to this question, which has found ex-
perimental expression in the form of studies of formation of conditioned
reaction to subsensory stimuli in healthy individuals. It is only after
reporting the results of this type of study, with which our team has been
concerned for about 10 years, that it is possible to try to discuss the
mairi question, which we can again formulate as follows: How are we to
15
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interpret the internal correlation between such phenomena as awareness of
perception of an exogenous signal (onset of conscious perception) and
conditioned motor reactions related to the system of voluntary movements?
1. Avakyan, R. V. "Measurement of Threshold Intensities of Sounds and
Differential Frequency Thresholds Using Conditioned Blink Reflexes
of Man," in: "Fiziologicheskaya akustika" [Physiological Acoustics],.
Leningrad, 1955, pp 52-59.
2. Avakyan, R. V.; Gershuni, G. V.; and Ratenberg, M. A: "Investigation
of Function of Auditory Analyzers in Patients With Signs of Hysterical
Deafness," ZHURN. VYSSH. NERVN. DEYAT. [Journal of Higher Nervous
Activity], Vol 7, Vyp 3, 1957, pp 325-334.
3. Arapova, A. A., and Orlova, G. M. "Correlation Between Thresholds of
the Galvanic Skin Reflex and Perception Thresholds in .the Presence of
Disorders Referable to Cutaneous and Auditory Sensibility," in "Tez.
Dokl. na XIII soveshch. po fiziol. probl." [Summaries of Papers Delivered
at the 13th Conference on Physiological Problems], Leningrad, 1948,
pp 8-10.
4. Asafov, B. D. "Functional Organization of the Orienting Reflex in the
Presence of Lesion to the Deep Structures of the Human Brain," in:
"Rol' glubokikh struktur golovnogo mozga cheloveka v mekhanizmakh
patologicheskikh reaktsiy" [The Role of Deep Structures of the Human
Brain in Mechanisms of Pathological Reactions], Leningrad, 1965, pp 18-20.
5. Astvatsaturov, M. I. "Nervous Diseases," Leningrad, 1935,.432 pp.
6. Bronshteyn, A. I. "Sensitization of Sense Organs,'.' Leningrad, 1946,
133 pp. '
7. Gershuni, G. V. "Investigation of Subsensory Reactions in Sense Organ
Function," FIZIOL. ZHURN. SSSR [Physiological Journal of the USSR],
Vol 33, Vyp 3, 1947, pp 393-412.
8. Idem, "Reflex Reactions to Delivery of Exogenous Stimuli to Human Sense
Organs as Related to Sensations," Ibid, Vol 35, Vyp 5, 1949, pp 511-560.
9. Idem, "Distinctions of Conditioned Galvanic Skin Reactions and Reactions
pf Inhibition of Alpha Rhythm Occurring Under the Influence of Subliminal
and Supraliminal SOI1iC Stimuli in Man," ZHURN. WSSH. NERVN. DEYAT.,
Vol 5, Vyp 5, 1955, pp 665-676.
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10. Gersl~~uni, G. V.; Alekseyenko, N. Yu.; Arapova, A. A.; Klaas, Yu. A.;
Maruseva, A. t1.; Obraztsova, G. A.; and Solovtsova, A. P, "Impairment
of Activity of Sense Organs and Certain Other Nervous Functions in the
Presence of 'Air Concussion'," in: "Voyei:no-meditsinskiy sbornik"
[Military Medical Collection], Moscow--Leningrad, Vol 2, 1945, pp 98-192.
11. Gershuni, G. V.; Klaas, Yu. A.; Liva.nov, rS. N.; and Maruseva, A. M.
"Electrical Activity of the Brain in the Presence of Hearing and Speech
Disorders Occurring as a Result of 'Air Concussion'," in: "Tr. Fiziolog.
in-ta AN SSSR" [Works of the Physiology Institute of the USSR Academy
of Sciences], Moscow--Leningrad, Vol 1, 1945, pp 115-128.
12. Gershuni, G. V.; Kozhevnikov, V. A.; and Matyatova, Ye. S. "Investiga-
tion of Some Manifestations of Activity of the Human Auditory Analyzer
Using Conditioned Galvanic Skin Reflexes," VESTN. OTORINOLAR. [Vestnik
of Otorhinolaryngology], Vol 16, Vyp 4, 1954, pp 14-20.
13. Zakharova, N, N. "Distinctions of Perception of Sensory Stimuli in the
Presence of Emotional Stress and Posttraumatic Pyschopathoid Syndrome,"
ZHURN. NEVROPATOL. PSIKHIATR. [Journal of Neuropathology and Psychiatry],
Vol 73, Vyp 3, 1973, pp 401-406.
14. Ivanov-Smolenskiy, A. G. "A Method of Studying Conditioned Reflexes
of Man," Moscow, 1933, 104 pp.
15. Kristostur'yan, S. G. "Use of Conditioned Galvanic Skin Reflexes in
the Presence of Pathology of the Auditory System," VESTN. OTORINOLAR,
Vol 14, Vyp 2, 1952, pp 11-15.
16. Kryshova, N. A. quoted by Gershuni, Alekseyenko et al., 1945.
17. Myasishchev, V. N. "The So-Called Psychogalvanic Reflex and Its Signi-
ficance in Personality Studies," in: "Novoye v refleksologii i fiziologii
nervnoy sistemy" [News in Reflexology and Physiology of the Nervous
System], Leningrad, Vol 3, 1929, pp 233-255.
18. Orbeli, L. A. "Problems of Higher Nervous Activity," Moscow--Leningrad,
1949, 499 pp.
19. Panov, A. G. "Experimental Analysis of Hearing Disorders in the Presence
of Nervous System Pathology," in: "Psikhofiziologicheskiy eksperiment
v klinike nervnykh i dushevnykh bolezney" [Psychophysiological Experi-
mentation in Symptomatology of Neurological and Mental Disease],
Leningrad, 1933, pp 6-23.
?_0. Perelman, L. B. "Reactive Postcontusion Surdomutism, Detection and
Treatment Thereof," Moscow, 1943, 56 pp.
21. Reshchikova, T. N. "Investigation of Subsensory Reactions of Patients
With Long-Term Sequelae of Closed Cerebrocranial Trauma," author abstract
of candidatorial dissertation, Moscow, 1969.
17
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22. Sechenov, I. M. "Reflexes of the Brain," in: "Izbr. proizv." [Selected
Works], Moscow, Vol 1 (1863), 1952, pp 7-127.
23. Veraguth, 0. "Das Psychogalvanische Reflexphenomenon," Berlin, 1909.
COPYRIGHT: Izdatel'stvo "Nauka", 1.977
10,657
CSO: 8144
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THE ROLE OF ISOLATION, ENVIRONMENTAL DEPRIVATION AND ENRICHMENT IN FORMATION
OF BEHAVIOR
Leningrad SOVREMENNYYE TENDENTSII V NEYROFIZ~IOLOGII in Russian 1977 pp 304-316
[Article by A. D. Slonim, Institute of Physiology and Experimental Pathology
of High Altitudes, Kirgiz Academy of Sciences, Frunze]
[Text] The problem of separating complex forms of behavior into phenotypic
and genotypic elements compelled researchers to develop a number of experiments
with animals at different stages of postnatal and prenatal ontogenesis. Addi-
tion to the environment of new factors, unusual for the organism, or elimina-
tion of some of the customary ones were used extensively in recent years to
determine the genesis of certain elements (patterns) of reactions of the
organism and its systems.
Environmental changes not only have effects based on the plus-minus inter-
action principle, they also disrupt sharply the entire process of formation
of behavior as an integral system. This prompted researchers to investigate
r.he effects of a so-called enriched or deprived environment, as well as to
evaluate the significance of this interaction, which occurs in accordance
with the feedback principle. Reverse afferentation [feedback], postulated
by P. K. Anokhin (1968) as an acceptor of action acquries exceptional import-
ance in many animal species at a specific stage of development, and it is
a most important mechanism (for example, in some passerines.) that adjusts
vocal reactions.
At the same time, elimination of some types of afferentation and reduction
of overall information from the environment lead to prevalence of intero-
ceptive influences on the central nervous system and could create a new
level of physiological state, sleep. Isolation, with regard to certain
types of food, sharply alters some types of appetite (Ugolev, Kassil', 1965)
and even leads to energy balance disturbances (Chernigovskiy, 1962).
At the present time, researchers have accumulated considerable material dealing
with the effect of isolation and environmental deprivationaon the developing
organism. These studies can be arbitrarily divided into the following three
groups: 1) experiments involving isolation, with regard to some environmental
factors; 2) experiments involving isolation of an animal of a given species
lg
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from its usual habitat (and development); 3) experiments involving exclusion
of sensory systems (sensory deprivation).
Historically, it happened that the method of isolation of the developing
organism from the external environment turned out to be the simplest and,
it would appear, most promising method. The history of this method goes
back to the legendary experiment of Lycurgus, which has retained its signifi-
cance to this day.
The famous legislator of ancient Sparta, Lycurgus, took two puppies from the
same litter, keeping one of them in a ditch in isolation from the environment
and the other under ordinary conditions, among people and animals. When the
puppies grew up, they were released to chase a hare. The dog that was
raised in isolating hid from fright; the one raised in liberty dashed after
the hare and choked it. This experiment served as the basis for education
theory, which stipulated that it was imperative to overcome vital difficulties
at an early stage of development (childhood) in order to develop positive
personality traits in the adult. The experiment of Lycurgus was repeated in
the laboratory of I. P. Pavlov, and it was found that the dogs raised in
isolation were notable for cowardice and poor adjustment to changing living
conditions (Vyrzhikovskiy, Mayorov, 1933).
Isolation may refer to elimination of specific environmental factors, specific
types of food, other animals of the same kind, specific forms of activity,
physical environmental factors (for example, light), chemical environmental
factors (change in air composition), habitat, etc. The isolation method
is used at all stages of ontogenesis, but it is particularly important at
the early stages of development of an organism, when different forma of
behavior and activity are formed.
In the experiment of Cuvier (1842), a young beaver was nursed by a woman.
When it switched to plant feed, it would first stack part of the willow
twigs in a corner of its cage after having torn the bark off,. When some
earth was brought into the cage, the little beaver, which had never seen
beaver dams, tamped the earth with its tail and stuck the willow twigs into
it. Evidently, the experiment of Cuvier is. the first reliable experiment
on isolated rearing of an animal in order to detect his natural instincts,
and it defined the significance of isolation to formation of genetically
programmed behavior. patterns. This was confirmed by Eibl-Eibesfeldt (1961).
Spalding (1965) kept young swallows in small cages, that precluded any
possibility of flying or exercising wing movements. In spite of this,
when the swallows were rel.eased.at the stage of development when they should
normally fly, they flew just as well as other birds of the same age.
In the experiments of Grohman (1938), pigeons were kept in cardboard tubes
that restricted wing movement. Control pigeons of the same .age were able
to move their wings without restriction in their nest. The experimental
birds were then released from the tubes and their capacity for flying was
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then compared to that of control birds in their first flight. Grohman
failed to demonstrate a difference between the two groups.
This experiment showed that the main motor elements of flying in pigeons
could develop without prior experience and formation of proprioceptive feed-
back. However, there was no quantitative evaluation of flying performance.
Analysis of fine adaptations developing in the course of exercise is very
difficult to make. Petersen et al. (1957) studied development of flying
in cabbage butterflies (Pieris). Butterflies that had just emerged from
their cocoons were let out from a brightly lit area to determine the height
to which they could fly. With age, they ascended higher and higher. Three
groups of butterflies were allowed different periods of flying time in
order to investigate the role of practicing flying in this refinement. One
group was tested from the 1st to 5th day of life, 4 times a day, at 20-min
intervals. The second group was kept inactive for the first 4 days, cooling
them in darkness to 18?~, then tested just like the first group, on the 5th
and 10th days of life. The third group was tested similarly, but was allowed
20 additional "practice" flights before each mandatory test in the first 5
and on the 10th day of life.
Flying characteristics improved with age in all groups, regardless of practice.
This improvement was the result of increased rigidity of the wings. The
height reached by representatives of all groups of the same age was the
same, which rules out the significance of practice. Similar results were
obtained in a study of formation of motor activity in an aquatic environment,
while swimming.
Carmicheal (1927) restricted tadpole movement that usually takes place in the
eggs by submerging them in urethane solution. The tadpoles developing in a
motionless state swam quite normally after the anesthetic solution was
washed off. Carmichael's data were not completely corroborated by D. A.
Sakharov (1957) in the laboratory of Kh. S. Koshtoyants, who observed
impaired swimming movements in clean water, after removal of the anesthetic,
in frog larvae (Rana pipiens) raised under anesthesia.
Considerable data have also been gathered on formation of specialized reactions
in animals using the isolation method. Even L. Morgan (1899) had described
a case, when baby squirrels, removed from the nest before they could see,
were raised in a room and took nuts, put them on the rug and made "bsrrowing"
movements. After a certain number of such movements, the squirrel would
take another nut and repeat the whole procedure.
In studies of the effects of isolation on development of behavior, much
attention was devoted to sexual and parental behavior, which are related to
preservation of the species. Thus, animals on different phylogenetic levels,
which were raised in.isolation and encountered the relevant stimuli for
t11e first time, manifested sexual activity. Male leaping spiders. go through
a very complex courting ritual, which is usually also manifested in specimens
raised in isolation (Drees, 1952). True, there are data to the effect tha t
the sexual behavior of guinea pigs is impaired as a result of being raised .
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in isolation (Young , 1957): significantly after isolating animals (males)
at the age of 2 days and less so when isolated at the age of 10 days (Gerall,
1965). However, according to the~data of Harper (1968), when isolated
immediately after birth and tested between the ages of 80 and 90 days, the
sexual. behavior of male and female guinea pigs did not differ from that of
control animals. There is reason to believe that these differences are
related to the incomplete physical development of animals as a result of
ruling out play activity when they .are kept alone in opaque cages. This
fact confirms our conclusions (Slonim, 1961, 1971) that play is a programmed
element of development and has distinct quantitative characteristics refera ble
to specific stages of development (Ponugayeva, 1964, 1968).
Precise quantitative characteristics are very difficult Co obtain with
regard to the behavior of inexperienced mammalian mothers. For example, rab-
bits build each successive burrow better than the preceding one (Ross et al.,
1956). Such behavioral changes, elicited by activity, are not very
noticeable.
A more variegated set of findings is observed in experiments with lower
monkeys. Singh (1969) demonstrated that wild specimens of Macaca rhesus
and monkeys from Indian temples did not differ from animals raised in partial
isolation, with respect to their ability to solve problems in a problem cage.
Moreover, the monkeys raised under artificial conditions solved the problems
faster than their wild relatives. At the same time, raising Macaca rhesus
in isolation led to impairment of "social" and sexual .behavior of these
animals. In particular, maternal rejection of its offspring and aggressive
behavior toward offspring of the same age were observed (Harlow et al., 1971).
This problem, that of development of aggressiveness in isolation, is being
worked on very intensively in several laboratories. It was investigated
the most systematically on laboratory mice (Valzelli, 1969x, 1969b, 1973),
in which a distinctive "isolation syndrome" was described, consisting of
several peripheral, behavioral and neurochemical changes in the organism.
They include increased muscle tones, corneal and skin reflexes, piloerection
and tremor. There is an increase in spontaneous motor activity, hyper-
reactivity and development of hypertension. Isolated aggressive mice pre-
sented an increase in weight of the adrenals and very accentuated orienting
reaction. Progression of. these symptoms can be observed when the animals
are kept in isolation for 1 to 4 weeks. This effect is observed only in
males, since females are not aggressive. Aggressiveness does not develop
if over S mice are kept in the cage, but it develops to a mild degree when
there are 2 and 3 animals per cage. Castration prevents development of ag-
gressive behavior; however, if it had developed prior to this operation,
the latter did not eliminate it (Burge, Edwards, 1971). Adrenalectomy
attenuates aggressiveness but does not prevent development of the phenomenon
(Brain et al., 1971).
The above-described phenomena should be classified as the consequences of
"social" isolation, when environmental deprivation involves primarily iso-
lation from specimens of the same species.
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Vocal reactions of animals are another exampJ_e of the role of experience in
development of motor patterns of behavior. In birds, there is a distinct
difference between a song, which is a more or less complex sonic expression
and calls, which are shorter and simpler sounds. Most calls develop nor-
mally in birds raised in isolation from adult specimens of the same species,
regardless of whether or not they had heard the sounds of other species.
Thus, normal vocal reactions were observed in roosters and chickens raised
in an incubator (Schjelderup-Ebbe, 1923). Experiments with small Passeri-
formes, warblers and black thrushes, raised in isolation, revealed that
their calls develop normally (Sauer, 1954; Messmer, Messmer,r-1956; Thielcke-
Poltz, Thielcke, 1960). The calls were exactly the same as in wild specimens
of the same species in all 25 thrushes raised in isolation. The young of
many other species of nesting Passeriformes birds were removed from their
nest at different times between hatching and acquiring their plumage, and
they were hand-fed in varying degrees of isolation from their own and other
species. The authors believe that experience in perception of sounds during
the period prior to isolation could not have a direct effect on subsequent
development of calls, aJ.though this possibility cannot be entirely ruled out.
The calls of small Passeriformes, skylarks and pied flycatchers (Canyon, 1957;
Curio, 1959), developed Normally in almost all cases with isolation. A wild
hybrid from a cross between the pied flycatcher and collared flycatcher
developed an intermediate call, differing appreciably from the calls of
both species (Haartman, Lohrl, 1950; Lohrl, 1950). Consequently, the calls
of passerine Passeriformes raised in isolation did not differ from calls of
birds in their natural habitat (Miller, 1921; Nice, 1943; Marler, 1956).
Ttiis confirms the conclusions in old studies involving hand-feeding of nesting
birds (Heinroth, 1924) to the effect that, in general, development of inter-
specific differences in bird calls is relative and unrelated to the acoustic
influences of other birds (Stadler, 1929). Normal development of singing was
demonstrated in handed baby swallows and sand martins, wrens, pikas, star-
lings, orioles, reed and corn buntings, as well as bullfinches (in the
surveys o.f Heinroth, 1924; Stadler, 1929; Thorpe, 1961; Marler, 19.63). A. N.
Promptov (7.944, 1956), A. N. Promptov and Ye. V. Lukina (1945) demonstrated
the great importance of imitation in development of singing in birds, but the
authors also devoted much attention to coordinations in the vocal effector
apparatus and differences that arise here. According to A. N. Promptov,
vocal reactions as a form of congenital behavior are largely determined by
morphological distinctions and function of vocal muscles .of the lower larynx.
Attention is devoted to both imitation proper, i.e., the capacity for imita-
tion which is congenital, and conditioned reflex changes in focal reactions
acid motor behavior of birds.
At the present time, it may be considered established that imitation and
learning play different roles in formation of vocal reactions (calls and
singing) in birds. In this respect, all birds can be divided into three
groups (Galambos, Worden, 1972): first group, birds (for example, domestic
ones) that retain species-specific vocalization, even if~they were deprived
of hearing immediately after hatching, i.e., before exposure to calls.
This implies the existence of genetically programmed vocal reproduction.
In these species, there is no process of comparison of the genetic program
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to acoustic information. The second group refers to birds (passerine
Passeriformes) that can develop normal singing, even if raised in isolation;
if thl~y are deaf at birth, normal. singing does not develop, and this suggests
that acoustical feedback (their own voice and singing) plays an important
role, apparently in relation to the genetic sound code; the third group of
birds refers to those for which not only auditory feedback but an external
"mode:l." of species-specific singing must exist for development of voice and
singing. For example, in chaffinches raised in isolation singing is impaired,
and there are many structures, which are related to imitation in the course
of deVelopment~ that are wanting.
There are very few data concerning voice development in mammals, with the
exception of man. Most important is the study of Boutan (1913) of the be-
havior ofa hand-fed gibbon, in whom all 30 studied cries developed normally,
although it was not known to cahat extent this animal was familiar with the
sounds of its wild relatives. This question often arises in primate studies.
For example, up to 32 different vocal signals can develop in hand-fed chim-
panzees (Yerkes, Learned, 1925), but there are no data pertaining to com-
parison thereof to the calls of wild ani.ma.ls. In spite of persistent attempts
to tench lower monkeys new sounds, including human speech, they either failed,
or else the results were very insignificant (Furness, 1916; Yerkes, Yerkes,
1929):
Riggs et al. (1972) tried to investigate the effect of lack of hearing on
vocalization of adult Saimiri sciureus monkeys. They demonstrated a lack
of visible differences in vocalization of normal and hearing-deprived animals.
After their birth, the mothers raised their young in an environment deprived
of species-specific sounds. One of the offspring was deprived of hearing
surgically 5 days after birth. Two otF-ers were raised under normal conditions,
i.e., they were exposed to species-specific vocalization. Additional data
were obtained on 6 other offspring: 4 were raised normally, 2 isolated from
their mothers. The sonic spectrograms were studied for 6 months on those
raised in isolation and for up to 17 months on the normal ones. Samples of
this spectrographic material were analyzed (form of calls and quantitative
criteria, i.e., duration, frequency characteristics of squeals and cries).
No differences could be demonstrated between experimental and control animals.
The sounds emitted by babies and adult animals also failed to differ with
regard to these parameters (Winter et al., 1973). Perhaps, there is normal
development of vocalization in lower and even higher primates, gibbons and
orangutans in isolation.
Seitz (1955) raised a group of raccoon dogs (Nyctereutes procyonoides) in
isolation and observed that six calls and variants thereof developed normally;
the same was observed in two domestic cats raised in isolation (Weiss, 1952)
and in Duplicidentata isolated after the first month of life (Severaid, 1958).
In our laboratory, a large series of studies was conducted at'different times
on ma~ranals isolated from the environment, related to formation of specialized
(species-specific) alimentary and defense reactions.
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In the studies of E. R. Uzhdavi.ni (1958a, 1958b), salivary duct fistulae were
created in puppies immediately after birth; he found that, at the age of
20-21 days, the puppies raised on a milk diet presented a positive reaction
to meat, consisting of movement toward meat and salivation.
Ttris fact indicates that the inborn reflex to the odor of meat appears in
dogs, as in predators, at a specific period of life, at the time when ani-
mals switch from milk to a mixed diet. If this reflex is not reinforced
with meat, it is extremely unstable and no longer demonstrable at the age
of 8-9 months. It is totally replaced by natural conditioned reflexes,
which developed in the course of being on a milk diet. Similar findings
ware made on puppies isolated from their mothers and raised on bottles.
The reaction of newborn Ungulata to shading .[dark] above the head, which is
particularly vivid on the 1st-3d day of life, disappears when the animal
is fed from a bottle, i.e., when this darkness is not reinforced by food
(Slonim, 1961). In puppy dogs, a positive reaction to stimulation of the
snout with fur is particularly marked on the 2d day, rather than immediately
after birth, even if the animals are fed from special droppers. Then, on
the 3d-5th day, this reaction disappears, failing to be reinforced by
contact with the nursing female.
The studies of A. M. Ugolev (1953) performed with kittens revealed that
there is no salivation in response to the sight and odor of meat while they
.ar.e hunting for live prey, just as .is the case in response to the sight
and odor of living prey, mice and birds. It was found that this is specific
for a predator with the type of nutrition involving stalking of the prey
and prolonged tracking thereof; in the kittens that 11ad never hunted, there
were the usual, natural conditioned alimentary salivary reflexes, even
though they were on a mixed diet. The transformation of kittens into
"hunters of prey," i.e., the transition to independent searching for food
(feeding on live prey) is associated with an abrupt change in the entire sys-
tem of unconditioned and conditioned alimentary reflexes. The natural
salivary reflexes disappear entirely during the period of ,stalking the
prey or prior to eating, as is the case in adult cats. This unique "trans-
formation" of the kitten into a hunting animal, a predator, occurs at a
certain age which, however, fluctuates over a rather wide range.
It was also established that if a kitten, which had not yet hunted for food,
is given cut pieces of dead mouse, then a live mouse but with some cuts
in it, this transformation can be accelerated due to development of con-
ditioned reflexes to the mouse as a source of food. The abruptness of this
transformation in predators is of enormous interest, since it permits
determination of the direct effect of the environment on rate of formation
of unconditioned reflexes, which are thus far from absolutely independent
of. environmental factors that have immediate biological significance.
However, the isolated maintenance of kittens in many homes does not prevent
them from changing into predators and catching mice and rats; consequently,
~j
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neither imitation nor development of special reflexes determine formation
of the predator reaction to live prey. According to the data of A. M.
Ugoley, in cats that do not hunt for mice, there is no inhibition of saliva-
tion at the sight and odor of food and live prey, which they had never caught.
Formation of alimentary reflexes in predators was confirmed in a newborn
lion cub raised on a milk diet to the age of 50 days. It was found that,
starting at the age of 30-31 days, it developed a stable, positive motor
reaction, without extinction, to the sight and odor of meat. It appeared
concurrently with an eye movement reaction to a moving object. Consequently,
inborn reactions to meat develop independently of feeding conditions
(lJzhdavini, Shepeleva, 1966).
In the studies of A. I. Shcheglova (1959, 1961), young rodents were nursed
by their mothers or females of another species: an albino rat nursed a
great gerbil, a great gerbil nursed an albino rat, a gray rat nursed a
gerbil, etc. In a special series of experiments, a group of gerbils and rats
was raised in a warm bag and fed milk from a piece of cotton and a dropper.
1'he animals were tested, starting on their 2d day of life, for 10 min in
a special chamber, which contained a digging object, i.e., dry roasted sand,
and gnawing objects, wooden sticks. It was found that gnawing and digging
began and developed in all gerbils at the same time, regardless of whether
they had been nursed by a gerbil, white rat or with a dropper. Digging
activity was much less marked in the gray rat than the great gerbil. It
appeared on the 18th-19th day of life; gnawing was observed on the 22d day t
and burrowing on the 14th day. A1]. these reactions developed at the above-
mentipned times, regardless of conditions under which the animals were
reared.
In th~~ experiments of A. G. 1'onugayeva (1960) who studied the milk-hoarding
reaction in young golden hamsters, the offspring were raised with their
mother but fed a .liquid mixture that these animals could not store. In
another series of experiments, young golden hamsters were taken from the
mother on the 24th day and also fed a liquid diet. These experiments re-
vealed that formation of the feed-storing reaction in the golden hamster
is unrelated to rearing conditions.
However, by far not all alimentary reactions develop in isolation, as was
shown in the above examples. K. Rakhimov (1958, 1959) found that there
was no attempt to graze by hungry lambs and kids raised up to the age of
5 months on a milk diet in strict isolation from others of their species,
as well as from pasturage and rought plant feed, when out in pasture. They
developed the ability to graze only after being with the herd for a few
days. The act of grazing, which is in essence a conditioned reflex, is
formed in the nature of an imitation reflex. It is important to mention
this, since V. I. Klimova (1956, 1958) established that neonate rabbits,
kept on a milk diet, presented an inborn reaction to green feed: The
numerous rxirer.i.mente of: K. Rakhimov failed to demonstrate this unconditioned
reflex to feed of plane origin at any of the phases of development of neo-
nate Ungulata raised on a milk diet. These differences in formation of
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unconditioned alimentary reflexes in rabbits and ungulates could be related
to the difference in ecological conditions under which the young develop,
a burrow for the rabbits and herd fior the ungulates, with the enormous oppor-
tunities for forming behavior based on imitation.
The above material warrants the conclusion and the isolation method is, so to
speak, the antithesis of the method of development of artificial conditioned
reflexes and experimental elimination of some forms of conditioned reflex
activity. This route of research, largely substantiated by I. P. Pavlov,
presents difficulties, with respect to interpretation of the results, since
it requires mandatory comparison to development of inborn behavior:
All of the studies lead to the same conclusion, that the alimentary act and
associated salivation reaction are stimulated by the sight and odor of
food, and that these reflexes should be referred to the natural conditioned
reflexes. If an animal is denied the food inherent in adult specimens of
its species from the day it is born, these reflexes are not manifested.
Thus, we also find an explanation for the numerous findings of disappearance
of predator's reflexes when raised with other species (for example, wolf
cubs and lambs, etc.).
However, when interpreting these facts, it is imperative to bear in mind
that the transition from milk to a mixed diet, then to independent feeding
occurs within specific ontogenetic time frames, and that it: is possible
to demonstrate the appearance of specialized reflexes to specific foods
only in these periods of development of an animal (Kossobutskiy, 1951;
Ugolev, 1953; Slonim, 1955; Klimova, 1956, 1958; Rakhimov, 1958, 1959;
Uzdavini, 1958a, b, and many others).
With reference to the .results of experiments involving isolation from some
types of food, we cannot fail to call attention to the fact that appearance
and disappearance of a positive reflex to the odor of food is inherent in
the ontogenetic process. In these experiments, it is difficult to imagine
(in spite of all the precautions taken by the authors: isolated feeding,
special uniforms for the service personnel, etc.) that there was total
elimination of odors of meat, which had never been reinforced and therefore
became an inhibitory stimulus. A special experiment conducted in our labo-
ratory with puppies kept in the vivarium building but in separate cages
where olfactory stimuli referable to the odor of meat were not ruled out,
and where they were kept on a diet of milk alone, confirmed this thesis.
Such puppies did not differ in any way from those kept on a milk diet in
an isolated dog house.
This is indicative of tYie special role of formation of inhibitory conditioned
reflexes when using the isolation method. Isolation from different environ-
mental factors not only precludes the possibility of development of natural
conditioned reflexes in response to signal-value components of these factors,
but could lead to formation of inhibitory reflexes, to development of signs
of habituation. This applies, first of all, to olfactory and sonic stimuli,
the sensibility thresholds for which ar.e exceptionally high in many animals.
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Disruption of the living stereotype is the second circumstance that is
rather important to evaluation of the results of such research. Expressly
exposure to a stimulus that was never uged before leads to inhibition of
reflex activity. For this reason, for example, stow eating of meat by
"milk" puppies and lack of physiological reactions adequate to this can be
interpreted as the influence of anew alimentary stimulus, unusual for the
animal.
F:ina].ly, use of the isolation method compels us to pay attention to the
fact that isolation from environmental factors and impoverishment of the
latter affect not only formation of~certain natural conditioned reflexes,
but the entire dynamics of cortical processes. This thesis gained experi-
mental confirmation i.n the studies of staff members of the University of
California (Rosenzweig et al., 1972). Studies were pursued of a number of
changes in the brain (occipital cortex) of rats kept in an "enriched" and
"deprived" environment for 25 to 100 days. The authors proceeded from the
fact that, in the laboratory, 3 rats are usually kept per cage. In the
"deprived environment," only one rat was kept in each cage. In the
"enri.ched environment," 12 rats were kept together in a large cage that
contained toys which were changed daily. Food and drink were available
in abundance in all of the cages. Biochemical, histochemical and morpholo-
gical. changes were demonstrated in the occipital cortex of rats from the
"deprived" and "enriched" environment. Almost all of the changes were reli-
able. We were impressed by the sharp increase in thickness of the occipital
cortex of "enriched" rats, as well as ,the increase~in number of anatomical
synaptic connections per unit area (by 50%) and decrease in their size.
A team of Czech physiologists conducted a large series of studies of this
type; they "impoverished" not only the external environment but the diet,
and i.n particular its protein componen~ Animals raised on a low-protein
diet revealed slower myelinization of nerve fibers, development of pyramidal
neurons in the sensomotor cortex, an increase in latency period of optical
and acoustical evoked potentials (Myslivecek, 1970; Safanda et al., 1971;
Myslivecek et al., 1974). Thus, a wide range of phenomena related to
nutrition, on the one hand, and delivery of stimuli from the environment, on
the other, was demonstrated. It is important to note that both endogenous
and exogenous influences determine not only general development of the organ-
ism, but primarily formation of the central nervous system, i.e., the brain.
If we scrutinize the method of isolation in detail, in connection with the
study of ontogenetic development of functions, we cannot overlook the
influence of afferentati.on on the genetically predetermined process of
development. In this respect, it does not matter whether these influences
are considered from the standpoint of action acceptor or construction of some
other system with feedback. Experimental research involving elimination of
different analyzers during the period of postnatal ontogenesis.enables us to
investigate this question and detect disturbances in the natural course of
development, which occur with any form of isolation or restriction of condi-
tions inherent in a given species. In this regard, some interesting data
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are given in the survey of Fox (1970). Fox et al. (1968), having deprived
a dog of sight, found normal development and myelinization of the visual
cortex, i.e., maturation thereof regardless of control and stimulation. In
the in vitro studies of Crain et al. (1970), they demonstrated that blocking
agents of the curare type did not affect development of nerve tissue; there
was normal development of bioelectr.ic activity at the usual rate. It may be
assumed that self-stimulation via proprioceptive feedback after a single
movement or during continuous activity of embryos may be significant for'
normal development of effectors, i.e., the skeletomuscular system, rather
than for. organization of motor acts. Evidently, this feedback develops after
spontaneous mobility is demonstrated, when reflex or evoked activity can be
elicited by mild (cutaneous) or more profound (proprioceptive) external
stimulation of the embryo. After this phase of effector-affector integration
on the segmental level of the spinal cord, there is subsequent integration
on higher levels of the developing nervous system until complex coordinated
action appears. Hamburger (].963) discussed the possible role of these
phenomena in embryogenesis of activity of chickens. Experiments conducted in
his .laboratory (Hamburger et al., 1965) revealed that spontaneous activity
develops before evoked activity, even in isolated segments of the locomotor
system, and that extensive generalization, involving the entire nervous sys-
r_em and observed in normally developing mammals after birth, appears on
about the 17th day of incubation (Volokhov, 1951; Voyno-Yasenetskiy, 1974;
Fox, 1966).
Held and Hein (1963) showed convincingly that deprivation of kittens of
all nonvisual sensibility impairs severely their subsequent exploratory
behavior that is controlled by sight. Rats blinded at an early age were
notable for, poor spatial auditory discrimination, as compared to those
that had the experience of vision (Spigelman, Bryden, 1967); but rats deprived
of sight at an even earlier age coped well with simple problems of auditory
discrimination. On the basis of these comparisons, Fox concludes that it is
difficult to separate the role of maturation processes from influences of
prior experience in development of a specific branch of the nervous system
or behavior pattern. In such cases, experiments involving isolation or.
restriction often induce effects in the form of excessive excitation after
isolation and lead to paradoxical phenomena. Thus, Lindsley et al. (1964)
discovered that monkeys raised in darkness reacted to light just like
monkeys raised under normal conditions react to the light being switched
off. Increased excitation in dogs after isolation disrupts their ability
to solve problems (Fox, Stelzner, 1.966) and to respond to nociceptive
stimuli (Melzack, Scott, 1957). Other "nonspecific" influences that could
distort efforts to examine a given sensory system or behavior pattern
include masking or depression of certain responses due to lack of condi-
tioning or reinforcement; these responses may simply disappear or become
"overmature," or else they may be related to unusual stimuli. Kovach and
Kling (1967), for example, found that kittens fed from a pipette subse-
quently refused to suckle their mother. Isolation may disrupt not only
maturation of a specific branch of the nervous system (or behavior pattern),
but structural and functional integration of one system (or element of
interaction) with another. For example, Held and Bauer (1967); Hein and
29
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and Held (1967) showed that if a collar is put on the neck of a newborn ,
monkey or kitten, which prevented them from seeing their forelegs, it
retarded development of association of the eye and front limbs and, after
the color was removed, there was no sight-controlled movement of these
limb's. Thus, some adjustments must be made to the isolation method. However,
for the time being, it is the only means of eliminating learning and memory
phenomena and thereby of separating genetically heterogeneous elements of
complex behavior of an organism into different stages of formation of
functional correlations.
1. Anokhin, P. K. "Biology and Neurophysiology of Conditioned Reflexes,"
Moscow, 1968, 546 pp.
2. Voyno-Yasenetskiy, A. V. "Pr.imary Rhythms of Excitation in Ontogenesis,"
Leningrad, 1974, 147 pp.
3. Vol.okhov, A. A. "Ontogenetic }?atterns of Nervous Activity," Moscow--
Leningrad, 1951, 301 pp.
4. Vyrzhikovskiy, S. N., and Mayorov, F. P. "Data on the Effects of.
Rearing on Type of Higher Nervous Activity in Dogs," TR. FIZIOL.
LABOR. AKAD. I. P. PAVLOVA [Works of the Physiology Laboratory of
Academician I. P. P avlov], Vol 5, 1933, p 171.
5. Klimova, V. I. "Alimentary Reflexes and Natural Stimuli in Ontogenesis
of Rabbits and Dogs," in "Soveshch. po vopr. evol. fiziol. Tez. i ref.
dok7.." [Conference on Problems of Evolutionary Physiology. Summaries
and Abstracts of Papers], Leningrad, 1956, p 82.
6. Tdem, "Alimentary Reflexes to Natural Stimuli in Ontogenesis of Rabbits
and Dogs," in "Problemy sravnitel'noy fiziologii nervnoy,deyatel'nosti"
[Problems of Comparative Physiology of Nervous Activity]; Leningrad,
1958, p 54.
7. Kossobutskiy, V. I. "Analysis of Development of Food-Hunting and
Defense Instincts in Ontogenesis of Some Predators," author abstract of
dissertation, Moscow, 1951.
8. P9organ, L. "Habbit and Instinct," St. Petersburg, 1899, 315 pp.
9. Ponugayeva, A. G. "Physiological Studies of Animal Instincts,"
Moscow--Leningrad, 1960, 180 pp.
10. Idem, "Prolonged Effect of Ambient Temperature on Formation and Per-
formance of Voluntary and Muscular. Activity in Ontogenesis," in:
"Teploobrazovaniye v organizme" [Thermogenesis.in the Organism], Kiev,
1964, pp 168-169.
30
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 II.iL~---_.._w.e_?4~ e~t.v r--~-~_vs-~_t~t.._.._ _.. ..._~i~li~.. _-' ~~.:~~~7:tY~+'~'i';~.+~u;.S,~
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1.1. Ponugayeva, A. G. "Research on Play Activity of Rats and Golden
Hamsters," in: "Sravnitel'naya i vozrastnaya fiziologiya" [Comparative
and Developmental Physiology], Leningrad, 1968, pp 104-117.
12. Promptov, A. N. "Vocal Imitation in Passeriformes Birds as One of the
Specific Properties of Their Higher Nervous Activity," DAN SSSR
[Reports of the USSR Academy of Sciences], Vol 45, No 6, 1944, pp 278-
281.
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COPYRIGHT: Izdatel'stvo "Nauka", 1977
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CSO: 8144 END
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