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JPRS L/9986
16 September 1981
Translation
_ MEMORY AND ADAPTATION
By
R. Yu. II'yuchenok
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JPRS L/9986
16 September 1981
MEMORY AND ADAPTATION
Novosibirsk PAMYAT' I ADAPTATSIYA in Russian 1979 (signed to press
23 May 79) pp 2, 3-5, 34-72, 142-150, 158-181, 192
[Annotation, introduction, chapters 2 and 3, conclusion, sections 2,
3 and 4 of the appendix, bibliography and table of contents from book
by Rostislav Yul'yanovich I1'yuchenok, "Memory and Adaptation," edited
by Aleksandr Feodorovich Nikiforov, Izdatel'stvo "Nauka" (Siberian
Division), 4,500 copies, 192 pages, UDC 612.821.2+612.821.6]
CONTENTS
Introduction 1
Qiapter 2. General Physiological Mechanisms Which Are the Basis for
Individual AdaFtation and Which Define the Phasic Nature
of the Process~�.~~~~~~~~~~~~��~~~~~~����~~~~~~~~~~~~~~��~~~~~~���� 3
G`hapter 3. Memory--Zhe Basis for Indiv idual Adaptation~ 13
Conclusion 35
APPendix 42
Section 2. Method for Study of the Activity of the Human Operator in a
System of Compensatory Tracking 42
Section 3. Psychological Methods for Study of Memory in the Adaptation
Process 46
Section 4. Psychophysiological Methods for the Study of Human
Adaptation 50
Bibliography 51
Table of Contents 63
- a - [t - USSR - C FOUO]
F(1R (1FFT~'IAT. i1.CF. nNi.Y
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[TextJ In the monugraph the physiological mechanisms
of adaptation are analyzed as a function of the effec-
tiveness of work of the central regulatory systems and
memory. An analysis of the data obtained during the
adaptation of man to various climatic-geographical
conditions (Pamiro-Alay, Altay, Kuril Islands) allowed
us to separate three phases of the adaptational process,
at the base of which lie various physiologi~cal mechanisms.
Particular attention has been devoted to processes of
memory and the functional asymetry of the brain in dif-
ferent phases of adaptation.
The book is designed for physiologists, psycl-~ologists,
psychiatrists, neurologists, and therapists as well
as students of universities and medical institutes.
Int~oduction
Up to the present time we have amassed extensive data about changes in
different systems of the organism, basically during adaptation to extreme and
near-extreme conditions. All this rich factual data has promoted an under-
standing of many particular mechanisms characteristic for concrete types of
adaptation to defined conditions of the climatic-geographical and production
environment. However on the basis of the material we have it is difficult to
give an interpretation of the universal general mechanisms of individual adap-
tation.
Three expeditioas (1975-1977) carried out by the Department of Central Regulatory
Mechanisms of the Institute of Physiology, Siberian Branch of the Academy of
Medical S~^ieaces USSR with analogous programs of research into the adaptation of
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man to contrasting climatic-geographical conditions allowed us to compare the dynamics
of a large number of reactions occurring in parallel and to show up those of them which
were present both during adaptation to mountainous conditions with different climatic
regimes and during a time shift after transmeridional flight, that is, allowed us to
show up those reactions of the organism which are probably universal for the process
of adaptation.
The investigations were carried out in mountainous conditions of Central Asia and
Siberia (Pamiro-Alay, Altay) in points located roughly at an identical elevation,
but differing in climate, and in conditions of a shift in the time zone by four
hours (Southern Kuril Islands).
The expedition of 1975 on Pamiro-Alay (settlement of Konchech, elevation 2,500 m
above sea level, a hot and dry climate) pursued the goal of clarifying adaptation
to mountainous conditions with an elevated temperature.
In 1977 in Altay (the Aktash mine, at an elevation of 2,600 m above sea level with
a markedly continental climate, similar in characteristics to the climate of Novo-
sibirsk) "purer" adaptation to mountainous conditions was studied since inhabitants
of the Academy City (Novosibirsk) and ad~acent places comprised the group of experi-
mental subjects. On the Kunashir Island (a Southern Kuril settlement, time shift of
four hours, a monsoon climate) in 1966 scientists observed a predominance of desyn-
- chronization elicited by the transmeridional displacement, and adaptation in condi-
tions of a monsoon climate.
The choice of geographical places with subextreme climatic conditions was dictated
on the one hand by a desire to follow the clear cut process of adaptation but
without the superimposition of pathological syndromes and on the other hand by a
desire to compare the shifts of different functions.in contrasting conditions while
attempting to show up the presence of several common changes. Although the main
problem of the investigations was a study of inemory processes during adapati.on, a
series of other psychological and psychophysiological tests were carried out
paralleling this study. These investigations allowed us not only to evaluate th~
background on which one or the other process of inemory took place, but also to
follow the course of the process of adaptation through many parameters, and as was
clarified later, to follow its phaselike quality with the physiological mechanisms
intrinsic to each phase.
Therefore it seems expedient before examining the memory processes to give a descrip-
tion of the adaptational restructuring of the neurological status of the vestibular,
statokinetic, psychological, and psychophysiological reactions; then one can show
the significance of inemory during adaptation, with the help of whose regulation
one may predetermine the paths of intervention in the process of adaptation.
The first successful Pfforts to change the course of inemory trace formation and re-
production by pharmacological substances introduce the hope that this path will
furnish real approaches to the cantrol of inemory. Undoubtedly even by the end of
this book the role of inemory it~ different phases of the adaptational process will
not be revealed because of an insufficiency of data. However, if we succeed in
informing the reader that memory in its widest sense takes nart in the process of
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adaptation and can compose the basis of the formation of adaptive programs which
- allow the organism to ad~ust its activity to the changing conditions of the environ-
ment, if we can succeed in informing the reader that memory is the foundation of
physiological mechanisms providing for the individual adaptation of the organism,
then we may consider that we at::ained our goal.
The collective of coworkers of the Department of Central Re:gulatory Mechanisms of
the Institute of Physiology, Siberian Branch of the Academy of Medical Sciences USSR
took part in the work. V. P. Leutin, I. V. Vol'f, S. B Tsvetovskiy, and E. I.
Nikolayeva investigated the change of bioelectrical activity of the brain and vege-
tative shifts in people during the formation and reproduction of the memory trace.
L. A. Konstantinovskaya and N. I. Dubrovina carried out psychological examinations
of inemory, attention, and functional state (in the first expedition N. K. Kiyashchenko
and S. R. Chaplygina examined these). M. A. Gilinskiy, I. A. Korsakov, and V. L.
Plyashkevich investigated the activity of the human operator during the adapation
of man to mountainous conditions (Pamiro-Alay, Altay) and on�Kunashir. The head of
the faculty of the Belorussian Institute of Adnanced Medical Studies, professor, doc-
tor of inedical seiences L. S. Gitkina and the he~d of the laboratory of the Institute
of Neurology, Physiotherapy, and Neurosurgery of the Ministry of Public Health of
the Belorussian SSR, professor, doctor of inedical sciences I. A. Sklyut took part in
the expedition. L. S. Gitkina carried out the neurological status examination and
- I. A. Sklyut carried out the examinations of static and dynamic coordination and
vestibular reactions.
In experimental investigations L. V. Loskutova, I. M. Vinnitskiy, N. V. Vol'f, and
S. B. Tsvetovskiy studied the role of different~structures of the brain in the
mechanisms of one session learning. N.I. Dubrovfna studied the neuronal mechanisms
of the interaction of these structures, and M. A. Gilinskiy, G. V. Abuladze, V. I.
Masycheva, and I. A. Pukhov studied the significance of various structures of the
brain in the mechanisms of regulation of inemory tract formation and reproduction.
The author expresses gratitude to all coworkers of the department and participants
of the expeditions who took an active part in the research which lies at the basis
of this book.
Chapter 2. General Physiological Mechanisms which arz the Basis for Individual
Adaptation and which Define the Phasic Nature of the Process
Adaptation is the universal property of the living. The individual life of an
organism.is the process of continual ~daptation to a changing environment with the
goal of achieving an adequate relationship with the external world. Without such
individual adaptation life would be difficult and almost impossible.
The genetic programs formulated during the process of.evolution ensure adapation
to constantly operating factors.
The creation of morphofunctional and biochemical constructions are conditioned by
genetic apparatus and all innovations in these constructions can arise only gradually
by means of random mutations and natural selectinn, enabling adaptation by the species
- to very gradual changes in the environment. These acquired adaptations, built into
morphological and functional systems, ar~e relevant to the external world only within
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the boundaries of a given geographic range. The animal organism is adapted highly
only in a given ecological niche (Shmal'gauzen, 1946).
The "creativ~'role of natural selection does not imply retention of genotypes which
could be useful in the future gi~~en an expansion of the habitation range; individuals
are selected which possess characteristics of increased viability in concrete climato-
geographic conditions. The retention of what is useful and elimination of what is
hazardous is accomplished not in expectation of advantage but in expectation of har-
mony with the reai conditions of existence (Timiryazev, 1895).
If the new ecological zones are markedly distinct in their climato-geographic parameters
and if migration occurs rapidly, then adaptationo,~enesis is passible (in particular,
this is characteristic when people of one generation relocate, when the organism en-
counters repeating but intermittently operative factors or constantly operative ones,
for exam le,
but for not so prolonged an exposure w'hich might alter the genotype, P
changes in tem~erature, pressure, social conditions, etc.). Then, in what fashion
do living beings adapt to altered conditions~of existence? A genetic program within
the defined limits could be beneficial in the process of migration and assimilation
of new ranges; but responses, constructed on the basis of these changes, make it
possible to interact with the external world, but not always in an optimal fashion.
Such reactions must either match, optimize, or reconstruct the same programs.
During the process of evolution of the animal world, we could expect the appearance
of some universal mechanisms which could have enabled adaptation of the living.being
to the continuaily changing conditions of its existence. "Examples of adaptation
which acquired universal signi~icance in the evolution of animals and which determined
group aromorphosis, are the development of the reproductive process and diploidy,
multicellularity and differentiation of tissues and organs, transformation from roe
to the ovum of viviparity, transformation from an independently formed larva to develop-
ment inside a maternal organism (embryonizati~n)a increased protection of the embryo,
nourishment of the young with milk, activation of hormones and more effective use
of food, intensifieation of the oxidizing processes, improvement in thermoregulation,
development of organs for higher nervous activity and improvement in reflex and motor
apparatus, use of individual experience for protection and.training of the young,
adaptation to new situations by individual changes in behavior and development of
behavior of the cognitive type" (Zavadskiy, Kolchinskiy, 1977, pp 14-24).
Data from the I. P. Pavlov school attest to the special role of higher nervous acti-
vity in ensuring balance of the organism with the external environment. As A. N.
Severtsov wrote as early as 1922, it is necessary to point out that in higher animals
and especially ~n man, there is a strong ability for behavioral adaptation to the
_ surrounding environment. When confronted with rapidly approaching changes in surroun-
ding conditions, the anima.l responds with a defined behavioral reaction without re-
construction of its organization and, in the ma3ority of cases, adapts to new con-
ditions. Bedesitheior anism withmadditionalipossibilitiesinwhichucananottonlynexpand,
havior provi g
' but also alter autonomic reflex reactions.
A. D. Slonim (1976) examined certain forms of homeostatic behavior, that is, behavior
directed lationPoftsurroundingeenvironmentaliconditi ns and specializedrformsnof~ ac-
tive regu
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activity, directed at creating the optimal conditions of existence in nature. Example~
include beh3vior thermoregulation (Milner, 1973), construction of nests, search for
shelter and wetting of the fur with saliva when it is hot. The ability of animals to
adapt to new conditions solely by means of behavior is shown graphically in training
of rats by sub~ecting them ta cold, th~rmal lamps, activated by depression af a lever,
(Wiess, Laties, 1961), cold ;~howers, or moving them into a heated cage (Epstein,
Milestone, 1968). Similar analysis of work on the homeostatic behavior of organisms
is provided in the studies of A. D.Sbnim and his co-workers (Slorii~ 1976).
In man, the development of higher nervous aetivity achieved such a high level that
behavior became a determining factor in his adaptation. Adaptation of man to different
conditions in the external environment is based primarily on distinct forms of behavior,
including artistic and technical abilities, by which he is able to exist in conditions
which are unsupportable for other organisms (Hensel, Hilderbradt, 1964). This fact
suggests that "adaptation, in the broad sense, is a process of a3aptation by man to
, conditions in his living environment which, to a large extent, he creates by trans-
formation of nature, directed at the preservation,.development and achievement of
his main goal: human progress" (Kazancheyev, Lozovoy, 1974, p 4).
Thus, man is adapted not only to conditions of habitation, but also, to a la.rger
extent, he adapts the external environment to hi~ biological abilities, creating an
artificial environment--an environment of culture and civilization, by which he
adapts to any conditions of existence. Of all living beings, man ~ossesses the
greatest ability for adaptation (Sevetsov, 1942; Deryapa, et al., 1975; and others).
However, examination of the social aspects of adaptation by man falls outside the
focus of this book, in which a detailed analysis of only the physiological mechanisms
of adaptation is presented.
The ab~lity of different systems in an organism, which guarantee homeostasis, to adapt
effectively its activity to changing conditions of the surrounding environment is de-
termined primarily by the function of central regulatory mechanisms. Creation, in
the process of evolution, of the regulatory systems with the correspanding morpho-
functional substratum led to the appearance of a potential for more precise reaction
to the external environment, an increase in the ~ange of adaptability without radical
morphological and biochemical reconstruction of tissues, adaptation of physiological
mechanisms by means of changes in function--that is, "ad~ustment" or optimization of
responses.
All normal processes of life activity have an adapri.ve character, that is, all physio-
logical reactions can be either adaptive to concrete "stationary" conditions of the
envir.~nment, that is previous processes of adaptation, or nonadaptive ones, found in
the process of adaptation. Consequently, both during the process of adapation and
in the final stage of adaptation, it is important to represent the degree ef parti-
cipation of various physioZogical mechanisms and the role which they play in the
creation of optimal interaction of the organism with the external environment. The
basic burden in certain phases of the adaptational process fall on these mechanisms.
The process of adaptation is a function of time. Consequently, at different stages
of it different physiological mechanisms can be involved. We assume that at the
outset of the proeess of adaptation; in special complex situations when the organism,
because of indadequate information, can not ad~ust appropriately or completely
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adequately to specific forms of adaptational reactions nonspecific mechanisms de-
teriorate such as emotion and Selye's adaptation syndrome by which the organism,
expending a large amount of energy, adapts its activity to specially changing
conditions. The participation of this syndrome in the mechanism of adaptation is
referred to by the author's name, G. Selye (Selye, 1973). He asserts that, w3thou~t
stress, life does not exist. He considers that peasants working intensely duxing
harvest, administrators undertaking a difficult problem, sportsmen striving for
victory, a man sitting in a vehicle with hi~ dying wife--all experience stress.
Stress, according to the definition formulated at the International Congress on~
Stress, i~ viewed as a r?onspecific reaction to any influence brought to bear on an
organism. This, as well as distress is "bad" stress but eustress is "good" stress.
It is interesting that in both forms of stress, nonspecific sympThusarstressecanibes
(increase in excretion of adrenalin) are similar (Levi, 1972~1�
without suffering, without grief, without unpleasantness and d~isease; it can arise
from any stress--physical, emotional or intellectual; essentia~ly, the degree of
stress also limits the potentials of an individual.
At the present time, in the literature and in natural phenomena there are s~ufficient
data, obtained in expeditionary conditions, to explain the initialinvolved~intthe
process of adaptation, including the special adaptive mechanismss,
adaptation syndrome of Selye. Thus, data are cited on the signif.t~1915�~/andtothers).
responses of an organism to extreme cold (Slonim, 1964; Maustrakh, ,
For S. N. Kukishev (1972), the inversican of circadian rhythm with the phenomenon of
~ decreased chronognosis is considered to be a stress reaction. The firs'~ period of
adaptation to inversion of sleep rhythm--wakefulness--is viewed by +these authors as
a stage of fear. However, it is inappropriate in this sense to consider this period
before 10 days time and furthermore, as an example of the first stage of stress
according to Selye --an anxiety reaction is delayed for 24-48 hours. In this period,
a decline in many functions is noted. These effects are not related to specific
forms of act~vity--nervous emotional stress can impaioints totthetinhibitionuoffunc-
tions. Thus, as illustrations, A. D. Slonim (1964) p
autonomic reactions; salivation, gastric secretions, respiration?, cardiac activity,
etc.
On the 2nd-3rd day of residence in Southern Kuril we noted a decrease ~n accuracy
and an increase in the number of errors in comparison with control standards in cor-
rective tests; there was a sharp increase on these days in variability of indices
for the number of errors (fig 24). On subsequent days, the ~ndices for number of
errors and accuracy were normalized. We noted on th~e 2nd-3rd day of the adaptation
process a decrease in activity (iig 25) evaluated sub~ecti.vely by the test subjects
in experiments conducted by SAS [Siberian Academy af Sciences]. The subjects
experienced an improvement in their sense of well-b~ing and their test performances
were within the highest level of control nuII?bers. ~.'he number of erro~rs in tracking,
that is, operator activity declined. Distu~bances in the work capacity of the test
subjects became evident in analysis of prod~ctivity (the quantity of reviewed letters,
accuracy of work) in the first27ndandcofd~hemlevelSOfnstability oftreproductionaof
after hourly experiments (fig )
words throughout the course of the experimen,t (fig 28)�
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:,~1 1 i~.~ o,n.~
t, ~ ~
iiiiiiii~iiii~iiirti~iii~itiiirrii~rrtirir "
:~i ~2 i n f~~,,,. ~3~
(1) ~,r,:.;.::: ..I _ ~i~:,,,.�,;u ~t~?f~~~~~,r.~~
1~~~,�. :'I Il:~nii~ui�inia llnruc~~cnu nuca::ari~:icii ~iurna onn~fiuit i~
~;u~i~iri;r~'~iuuii n~~unu.
Figure 24. Changes in Variability of Indices for the Number
of Errors in Corrective Tests
Key:
1. Novosibirsk
2. Southern Kuril
3. Days
~1~
0
~,~i _c-,-_--_._..~ -
y-~_
ti ~~,c~ ~
~~.:r
u,u~
,
i~iiiiiiiiiiri~i~~ii~irrii~iiiiriii~ii~~i
U
~ , 11,U1
`.1 ~ _
4 ' ~o
?
II~IIIIIIIIII~II~III~II11~IlI1i11111~1'I11111
�i !1 :'i I �7 !i .'l/(Inl ~w~
~2~~.,,~,~�:,~.~,~,. - ~g~;n~,,,~,-nyf~,,,,~.-�.,~I
l'~~r Il:~airnrunr uut;a:~:ni~nrii :u;iuuunr�ru (u) n n:ir~r~~urmut
Figure 25. Changes in Indices for Activity (a) and Adjustment (b)
Key:
1. Standard units 3. Southern Kuril
2. Novosibirsk 4. Days
7
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:r
. ~1~r.,
~ ; ~2
Prr~�. _f;. li:tni~~uouiti+ ~ut~'-
u~ nu unrnGuu n~in nocnpu-
"o f' ICtltl!14QIII1lt C]IUII It llO0-
3- - J . i~~~ccc a~~~urruitF~u. ~ .
J V .
o G
m ~,-r- m - r -a-
~2 4 ~3> ~~r ~4~
~~a~~n
A.~rnuii _
~�ubu~~~.x
: I
~ Figure 26. Changes in the Number of Errors in Reproduction of
Words during the Process of Adaptation
Key:
1. Number of errors in reproduction 3. Altay
2. Novosibirsk 4. Days

~ ~v:,u- - [.__7 ~ .
~
~'u,�. ~ir. ISu~r~~r.i:�rv~~u~isi upu
~ia. 1'nrrurpani~~a ~c~n+encnun
~.~~~~nuii.uuc~rn paiGuri.t upu ~i II ~y
a)~~~nra~lm( u rupax If~innipo- 1,UUU ri 1 .c~ `
,
:~~i:ui u�ri~muc:.~no ~?u~~:i~i ~J ~im ^
n~~nrniur~iruui~~~ :sti;ir.ou u�ru�
puii nunuuun~.[ u~~oGi.i ~c nep- :
nuii. . r
i - - ~~;~~~,~n~~, s - ~wnc~~ o~~~,~ ra. c,.,_�~ ~ _ i__. - ~---r
D? 3 9 fl 21 n~
Figure 27. Carrective Tests. Histogram of ChangeS in Stability of
Work given Adaptation to the Mountains of Pamiro-Alay--
_ The Relationship of Numbers of Examined Indices during the
Second Half of the Tests to the First Half. 1--Beginning
2--End of Experiment ~
Key:
1. Day
During these days in mountainous conditions, a decline in attention, disturbance
in stability of work according to corrective tests and dynamic coordinaCion of
random indices~occurred. On the 2nd day of residence in the mountains, we noted
a worsening of the sub~ective condition, detected by the methods used by SAS; in-
crease in variability of indices for motor memory of the right hand and maximum
~ scatter of indices for motor memory of the left hand; a decline in reaction to
moving ob3ects and sensory-motor response, as well as an increase in the variability
of their indices. .
It is doubtful that a sudden contrasting change in climatic conditions causes a
stress reaction without emotional overtones. But:emot3on, perhaps, should be
viewed as an adaptive reaction to generalized nature. Even Charles Darwin (1872)
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assigned significance to the adaptational character of emotion. It is doubtful that
this adaptive sign can be viewed as gradual loss in the process of evolution of its
adaptational significance. On the contrary, and not ina~vertently, the greater the
degree of evolution that has occurred in an organism the richer it is, not only in
number of reactions for expression of emotion but also, perhaps, in its ability for
emotional survival. Man of all living beings possesses the greatest potential for
adaptation to multiformed changes in the external environment and in man emotions
are the most deveZoped. It is difficult to verify that this is merely an apparent
parallelism and not a fundamental relationship. The specific reactians can be
depressed when man is introducted to new conditions. The majority of reactions turn
out to be nonadaptational ones and only interfere with the interaction of the organism
with the external environment. Ideally, reactions should be more generalized. Simi-
. lar phenomena are observed at different phases in the development of the conditioned
reflex when the absence of a specialized reaction is compensated for by the appearance
of emotion and the animal is able to adequately achieve, at least in part, the r.e-
quired goal. .
, r~� ~r,U:~ _
,"0
~
-�l Puc. `LS. 1TaMen~~uue c~r:t-
\
(iuii,uui:�ru iwcnpon;~isc~to~u~~i
M, \ cnou n xu~~o aiccuepa~icirra
u upoi~~~~�;�o n~~nmri~lu?~ (u~r- i
~iuntt~mui ~incn~ nncnpon:r i
ut~;l!'wiii n�ropoii uunonuur~t
U,'~l i~ ' UIII~I'i'il 1: IIC~IL(111~. i
~,1,~~,,,~,,,,,,,,,~,:
.1 ; i ~1
~lr~�~~~ j (2) (3 I
~
ruo � A~~~" . - i
' ~ ,
_ Figure 28. Changes in the Stability of Word Reproduction in the
Course of the Experiment on the Adaptational Process
(Relationship of Numbers of Words Reproduced during
the Second Half of the Experiment to Numbers Reproduced
during the First Half)
- Key.
1. Novosibirsk
2. Southern Kuril
3. Days
However, it is doubtful that the role of emotion is limited merely to those external ~
phenomena and mobilization of energetic reserves. The maximum output ot power is
achieved by mobilization of a wide range of adaptive programs in a short interval
of time during critical situations for the organism with the aim of utilizing already
existing programs for special adaptation. The physiological meaning of emtoions is
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explained well by the hypothesis of P. V. Simonov (1970) in which emotions are con-
sidered to be reflections of the dimensions of requirements and the likelihood of
their satisfaction at a given moment. In particular, emotions can guarantee satis-
faction of the needs of the organism in conditions of an information deficit that,
undoubtedly, occur at the onset of residence by Che organism in new conditions.
In recent years, the role of emotion in adaptive reactions of man and animals has
been underscored by thorough analysis (Shingarov, 1971; Kruglikov, "1971; Khananashvili,
1972; Simonov, 1975; Fress, Piazhe, 1975; Ivanitskiy, 1976; Val'dman et al., 1976;
and others). Some authors even give fundamental significance to the emotional state
in the mechanisms of adaptation of man and animals (Banshchikov et al., 1975). Un-
doubtedly, emotion superimposes its own imprint on the subsequent processes for the
formation of new adaptive programs as the perception of external signals is altered
(Gerken, Neff, 1963; Hall, Mark, 1967; Gasanov, 1972; Kostandov, 1977) and the pro-
cess of formation of their imprints (Whitly, 1962; I1'yuchenok, Yeliseyeva, 1966;
Smythies, 1967; Beritashvili, 1968; Veyn, Kamenetskaya, 1968; Latash, 1968; Khananash-
vili, 1972; Il'yuchenok, 1972, 1977; Brady, I975; Gromova, 1976) takes place.
After nervous emotional stress disappears masked signs of these states, characteristic
for activation of the regulatory systems, are developed. Changes in external condi-
tions require "adjustment" of physiologic reactions. On the basis of the existing
program, genetically complicated during the process o� evolution or created previously
in the process of earlier adaptations, regulatory mechanisms are responsible for al-
tering the level of regulation and switching over to another program. Although these
programs for a new living environment are, on tt~e whole, nonspecific, they are also
fixed in their range of possible deviation, thus making their utilization feasible.
One can assume that the fundamental burden on regulatory mechanisms occurs at initial
stages in the process of adaptation. The range of adaptation which is guaranteed by
- the central regulatory mechanisms depends on the limits of their potentials, reliability
and reserves of resistance. Of special importance in a given situat1974iSKaznacherev,
of stress on the regulatory system (Kaznacheyev, 1973; Vasilevskiy, , Y
Bayevskiy, 1974). A similar anal}isis of the resistance of the regulatory system in
the process of adaptation was conducted by N. N. Vasilovskiy (1974, 1976) and S. I.
Soroko (1975).
It is important to point out that the participation of the regulatory system is
most apparent at the onset of the process of adaptation when the response is still
based on old programs. This is illustrated by data, obtained in our laboratory,
both for adaptation in mountains and in conditions of other zonal times. In this
period, the resistance of function in different systems in the organism is decreased
and the variability of indices for many physiological functions is increased. Changes
in manifestation of orientational and cardiac reactions (fig 29), indices for dynamic
coordination, reactions to moving objects (see fig 7b), sensory motor reactions and
errors in evaluation of the proper time are increased. Reconstruction of the organism
on a new level of function goes through a phase of destabilization (Miller, 1975;
Bekhtereva et al., 1977).
Increase in variability leads to expansion of the range of response which improves
the potential for correlation of important life-related information with the multi-
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_ formed reactions of the organism. Change in the homeostatic regimes activate condi-
tionally the regulatory systems. The fundamental task at this level of regulation
is the search for the optimal regime for function (Breynes, Svechinskiy, 1963; cita-
tion from Kuritskiy, 1969; Ktiritskiq, 1969). A higher level of regulation adds to
the continuously active process of regulation of new "laws" and changes not only the
limits of significance for parameters which characterize the regime for systemic
function in the organism, but also the regime for this very sear~h. It is unlikely
that a new resistant state of function in the systems of the organism is achieved only
when a switch over to a new program for interaction of the organism with the external
environment occurs. Reconstruction of different systems in the organism occurs to
accommodate a new regime for function, thereby the maximum level of reconstruction
_ and the duration of it are dissimilar in different systems.
S~
- N
6' -
~
. , .
_ o
..~i iiiiiiii~riiriiiniri�~
l~ur. :..1. Ilar.~enciiu~~ ltnrucpi�nii ~1~'.s ' .s�~ 7J ;>>/~~~u ~3~
~ ~'I';U,~~n~.is ~~t~~iiii(nii u u~~ui~~~r. llu~n, (2~
rr ~~)~~urr:u~ini. ~�riGu~~~�n ~ //~~,~~upo Annii . ..I
Figure 29. Changes in Variability of Cardiac Reactions in the
Process of Adaptation
Key:
1. Novosibirsk
2. Pamiro-Alay
3. Days
The results obtainedduring three expeditions showed that in conditions at a height of
2,600 m and shifts in zonal time by 4 hours, reconstruction of the neurological (L. S.
Gitkina), vestibular, statokinetic (I. A. Sklyut) and immunologic reactions (L. V.
Devoyno, L. S. Eliseyeva, M. A. Cheydo et al.) resolved, on the whole, by the 2nd
week of adaptation.
Autonomic guarantee of reactions by an organism given various forn~s of adaptation can
vary; some are characteristic for a given form of adaptation with predominance of either
parasympathetic or sympathetic components. We assume that stimulation of both systems
occurs because of activation of the reticular formation in the brain stem. However,
in this form of adaptation and in its different phases, one system predominates.
_ Thus, in adaptation to mountainous conditions, the activity of the sympathetic system
predominated. This was characteristic also for the initial period of cosmic flight
with a subsequent short-term strengthening in the tonus of the parasympathetic section
of the nervous system (Vorob'ev et at., 1976). In adaptation to shifts in zonal time
the activity ~f the parasympathetic system predominated.
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In our experiments, the period of activation of the majority of regulat~ry systems
occurred on the 3rd-7th day of adaptation. Total use of the existing program in an
organism, given the need for future maintenance of activity in functional systems
within the assigned limits, lead~~to stress of the regulatory system. In special
conditions, this period is prolonged because for these conditions none of the adap-
tational programs can be optimal. However, the system functions within the limits
of regulation and in the future in conditions of its disruption.
' Given long term changes in external conditions stress on the regulatory system can
be weakened or eliminated by including other, more adaptive programs for reaction,
but given conditions where all program reserves are exhausted, this is possible only
after f~rmation of new programs on the basis of imprints of informatipn concerning
the changing surrounding environment; that is, use of inemory. Those reactions, pre-
determined on the basis of genetic mechanisms, do not lend themselves well to "training"
(they do not allow formation of fundamentally new programs for interaction with the .
external environment) because they either reached previously the limits of their re-
training capacity or their adequateness in external conditions is ensured only in
the course of the whole process of adaptation by maintenance of stress on the regula-
tory system, possibly, by transfer to other genetic programs for response. Recently,
the process of adaptation for a number of~functions can be viewed not as normaliza-~
tion of variability of indices, but as a decrease in them as we see, for example, in
the dynamic coordination reaction to moving ob~ects, the time of hand reaction given
- presentation of numbers and others.
Alleviation of the process of regulation with retention or even increase in the effec-
tiveness of responses and decrease in expenditure of energy can be accomplished only
by formation of new programs for adequate interaction of the organism with a changing
external environment, and this ensures the possibility of registration and preserva-
tion of imprin*_s of information regarding these changes. The more sharply distinct~
the dimensions of the parameters of the situation in which it is necessary to adapt,
the greater the stress on the regulatory system; but the mora diversity in the
changing situation., the more need for time to completely fix all of its elements.
With sharp differences and diverse factors in the new situation, inopportune and
inadequate fixation of an imprint can not weaken the limited stress on the regula-
tory system and interruption of regulation in separate functional systems can set
in before the completiong of the adaptational process. It is possible that, as a
in the first days of
result of such dissociation experienced in high al~itudes,
adaptation disturbances in higher nervous activity, diffuse general inhibition and
~ a number of other pathological phenomenon (Sirotinin, 1957; and others) can occur.
Shortening the transition period and the resulting stress on the regulatory system
can enccursaalsorwith accelerationbofifixationeofnimprintsaof~changingSexternal
This o
influences and the creation of new adaptive programs.
All the above mentioned considerations facilitate examination of the formation of
adaptive states as a dynamic process with resulting transitory phases on which are
builti physiological mechanisms. It is likely that these principles must serve as
the bases for a targeted program of study of the physiological mechanisms which
lie at the foundation of the process of adaptation (see the program in the Appendix).
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Chapter 3. Memory--The Basis for Individual Adaptation
Memory, arising at the earliest stages of evolutionary development, has universal sig-
nificance for the living world: on its basis rudimenXary progress became possible
for the organization of resporises in an ever more complex environment of habitation
by the use of accumulated experience and creation of new programs on its basis. This
characteristic not only retained its effectiveness in the most highly complex living
- systems, but also acquired more cardinal and universal significance for forms, existing
at the highest levels of evolution because it consistuted the basis for individual
behavior, attaining its more complete form in man in the form of behavior of the cog-
nitive type.
The phenomenon of inemory includes the process of transmission of information at the
moment of its entry in the brain through selection for storage, the mechanisms for
its retention and reproduction before return to the external world in the form of
behavior of the animal or man. The essence of adaptation consists of the ability of
an organism, after obtaining information from the external world, to respond adequately
on the basis of formed adaptive programs of re~ction to the environment.
Memory must be considered in the broadest sense as fixation of imprints to form pro-
gramsf~r the most varied responses, both behaviorally and autonomically, and as the
basis for all physiological reactions of biochemical transformation. However, the
most important factor in individual adaptation of higher animals and man to changes
in the external environment is the ability to register and retain i~?formation about
these changes in order to alter behavior in relation to acquired experience (I1'yuchenol+
1972).
Nevertheless, special study of the processes of inemory in the conditions of adaptation
has been devoted only to separate elements�of function. Thus, in experiments on dogs
subjected to prolonged residence at altitudes of 1,650 m above sea level (Avadkhara,
the Caucusus), improvement in performance of ext~~ded dynamic stereotypes and acceler-
ation of consolidation of temporal cont~ections became apparAnt (Kipiani et al., 1974).
Improvement in the formulation of inemory imprints was noted here and in experiments
on mice to observe performance of conditioned reflexes for active avoidance based
on adaptation to an altitude of 3,200 m above sea level (the m~untain pass of Tuya-
Ashu) (Meyerson et al., 1971). Acceleration in the performance of positive conditioned
reflexes in rats for adaptation to mountainous conditions (4,200 m, Minkush) was
apparent also in the experiments of M. A. Aliyev and A. K. Kasymova (1970). In these
studies, in contras~ to others, an acceleration in performance of inhibitory condi-
tioned reflexes was noted also. Interruption of the influence of hypoxia in condi-
tions of a decompression chamber (daily for 5-6 hours at an altitude of 5,000-7,000
- m above sea level) resulted in an improvement in rentention of the defensive condi-
tioned reflexes of passive avoidance and an increase in resistance of the process of
fixation of conditioned reflexes to~the effect of electroshock given activation of
nucleic acid and protein synthesis (Meyerson et al., 1970, 1976).
A smaller scale investigation of inemory in conditions of adaptation was conducted in
humans. Using psychological testing, scientigts noted a worsening of inemory in the
first days of adaptation (Aydaraliyev et al., 1977) and an improvement in immedl.~te
word reproduction on a later date in the process of adaptation to high altitudes
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(Mirrakhimov et al., 1975) and in the extreme North (Kust, 1974). However, these data
on improvement of inemory did not lead authors to assume that memory constitutes the
basis for the process of individual adaptation.
Recently, the problem of interaction of inemory and adaptation in the process of its
formulation has attracted more attention (I1'yuchenok, 1972, 1977a; KiFiani et al.,
1974; Mirrakhimov et al., 1975; Slonim, 1976; Vasilevskiy, Trubachev, 1977; Aliyev,
~ 1977). The importance of a multifaceted approach to the study of this specif ic
problem has been stressed (Bekhtereva, 1974) because memory ensures maintenance of
the resistant state of health (homeostasis) and adaptive reconstruction (Bekhtereva,
1976) .
Adaptation is a long-term process, making the role of inemory at various stages of
it ambiguous. Therefore, it is necessary to elucidate the character of changes in
memory at different phases of the adaptational process. It would be incorrect to
expect that these changes would be great in the collective sense, but, more correctly,
they would Ue of long duration. In order that these changes would not be confused
as pathological alteration in the function of different functional systems of the
organism in extreme and close to extreme conditions, scientists found it advisable
to analyze them given ambiguously expressed changes in the climato-geographic environ-
ment (altitude of 2,500 m and changes in zonal time by 4 hours).
V. P. Leutin, Ye. I. Nikolayeva, N. V. Vol'f and S. B. Tsvetovskiy conducted poly-
graphic recordings by electroencephalogram, electrocardiogram and dermo-galvanic
_ reaction in expeditionary conditions.
The highly informative nature of electroencephalographic and autonomic indices was
revealed during analysis of inechanisms for concentration of attention, perception,
formulation of short-term and long-term memory (Pollen, Trachtenberg, 1972; Rusalov,
Mekachchi, 1973; Klinger et al., 1973; Martidale, Armstrong, 1974). Using EEG methods,
the complex regulation of space-time organization in brain processes in man in the
normal state and in the pathological one, given formulation of associations and
different forms of inemory, was detected (Livanov, 197~; Voronin, Konovalov, 1976; and
others). However, similar analysis was of no practical importance in understanding
the effect of different climato-geographical conditions on these mechanisms.
With the completion of psychological tests, changes in the bioelectrical activity of
the brain, dermo-galvanic reactions and the dynamics of cardiac rhythm occurred. Con-
trol studies at Novosibirsk showed that autonomic shifts are part of the phasic dermo-
galvanic reaction which accompanies memorization and reproduction of words. Also seen
was an increase in the frequency of cardiac contraction and the appearan~e of reactions
for EEG activation (fig 30). .
In the initial period of residence by the subjects in Southern Kuril (2nd-3rd day),
against a background of increase in motor activity, spontaneous dermo-galvanic reaction
and slowing of the cardiac rhythm, V. P. Leutin, N. V. Vol'f, S. B. Tsvetovskiy and
Ye. I. Nikolayeva detected a decrease in the immediate reproduction of words given
an increase in the manifestation of accompanying EEG activation and a decrease in the
dimensions of recorded dermo-galvanic and cardiac reactions. A decline in word re-
production of the last words on the list occurred (fig 31). Immediately after
passage to the mountains of Altay, testing showed that a decline in word reproduction
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did not occur but the number of errors increased (fig 32). In addition, an increase
was noted in the manifestation of dermo-galvanic and cardiac reaction which accompanied
word reproduction with retiention of manifested EEG aetivation within normal limits
- (Fig 33). In the experiments of L. A. Konstantinovskiy, on the 2nd day of residence
in Altay conditions, a decline in verbal and form-space memory was not noted. In
Pamiro in this period, similar studies were not conducted. With regard to motor memory,
by the 2nd day, the indices for left hand function showed an increase in the amount of
deviation in the reproduced height from the preset one; in the indices for right hand
function, no changes were observed on the 2nd day.
a~ _ r_-_T-~----,
b ~ w, ~~.-+..-�-}..~.r.,,r..,..,,,...---....~-w,~..-a
c� rrYt~trvpWPYih1hYYYil~tYtYi~i~MY'r4Wly{~YPfhi~}ihYi~?t~rrl~rwrtrwra~trr*PrtMrwt+�fr~rYiv~~
~ ! ~ . \
t--- .
2; . . _ . _~l~~.l~J.l1..~1.lAl~~.?l/.NJ~A.I~IR. 1~ . -
~..s..e~s ~wr.~.......~rwu.m
g' ~ - -
l'ur. :fl1. Aluucoi;anani.nai~~ ~~ernr,i�p~u~n?i ~Z~nanwtorn~iccuux ~~~yuiu~uii
ni�.ni~iTyCniurn.
~i - +ior~~u upen~cnn, 10 c; f - oeCltTpo;)Ilqi�~~~a1J1oP~lA~1MA; ~r ~JIPIfTp111t~IpJ~llq-
I ~~nnlMil; .t - Itq)Ifllll-17111L071111i9~SC1lilll ~1L�:Iltllll~l; v-- ~10111/1'~IJ~iMA (IICIIACCItIf 110-
~~u:nu~~�n�~i n ntm~ruTi~ uocu~u~~cwi�Acuun cnnutt nuirnnru~uuum t~ upu ycru~~tx
~rrurrax urni~ryeniuro); c-~ini~nu nr,pec~+venuti auy~uin :):)1'; ~k-uTeicrim
upurpuMn~ia u~n,rra.
Figure 30. Multichannel Recording of Physiological Function of
Test Sub3ects. a--Time Markers, 10 s; b--Electroencephalo-
gram; c--Electrocardiogram; d---Dermo-Galvanic Reaction; e--
Phonogram (Spikes Appear at the Moment of Word Reproduction
on Tape Recorder and given Oral Responses by Sub~ect); f--
Number of Intersections at "Zero" on EEG; g--Control Points
of Experimental Program.
~ a " ~ ~,,.._b
0
ti'; v
~
� i ;
r
t' ~ l'~ir. :!l. I~nrl~~u~ii,tn~~;~~~
Q ~uir IU niir,qi~:Uuu (u) n
~ I I11 Il~~~lltbl\ (~i) ~'.~Illlt ~�II~I ~ -
~ ~ ~ 1'Gil.
I ~ i:- , i
. t y' 1. ' 1 J
1 I ~ :
M. ~A n ~
. . ~ r(3) r~ M 3
~t ,y _ Q ~ " ~ ( )
..f t,~') , . ~ ~'(2)'~:'~'',.
Figure 31. Reproduction of Last 10 (a) and First 10 (b) Words on List
Key:
1. Novosibirsk 3. Days
2. Southern Kuril
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;i;
Id
i; ~ f�
- r~ ~
!'~N�. ~i;.~. I1:~~I1�IIPIIUt' 'tll~'llil 1111111-
imi, ~i~/tl nui�n~iuli:utr14r1111n r:inu .,~a
~S
n u~~ui~r~�~�u a~~;inr~ii~un.
t; ~i~~~iiiiii~iiiii~ii -
",S ~1 11 lf~.~i
- I ( 3 )
- ~1~ , - (2)
Figure 32. Changes in Number of Errors in Word Reproduction during
the Process of Adaptation
Key:
1. Novosibirsk 3. Days
- 2. A1Cay _
o ,
�
.
i~~
-
i,
l.' ~lrl'
r' u,ul (~r11 ~
n, - r.t .
rt �
' i~
0
ii ~~~~~i~~~~~~~i~i~~~i ~ ii i~~~~i~i~i~iii~~i~~~i -
;~,s a n ; ~ � %~~'~3~
- 2 ~~~~,~;~M~ ~2~ --I
1 r:,f,~,. � n,:.�:a� ~ ~ -
( 7
I'ur. ;f~f. ll:i~iruruuo ni,i~u~;r,~~iuu~ri~~ c~~~~,~ryuuii ~io~i~t~~~nt (n) n ~~r~~i;
i~nu J;~1' ~~ri�uii~punuanii,ini u~n~ itucu~~on:?ue/~~win ~v~uu.
Figure 33. Changes in Manifestation of Cardiac Reaction (a) and
EEG Desynchronization (b) durii~.g Word Reproduction
Key:
1. Novosibirsk 3. Days
2. Altay
In the next phase of the adaptation process (3rd-4th day), an increased level of
spontaneous dermo-galvanic reaction continued; in Southern Kuril, cardiac rhythm
slowed down; in the mountains, frequency of cardiac rhythm increased with a normal
dermo-galvanic reaction; orientational reactions increased and motor activity was
normalized.
- In this period, memorization of words did not differ from cantrol data for total
memorization of the list. The number of test and control studies wa~ the same.
On the basis of a number of indices, on the 3rd day immediate repruduction decreased
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and by the 3rd-6th day, delayed word reproduction was noted in Southern K~uril. EEG
activation reaction, accompanying word reproduction, diminished to the lower limits
of the control. Dermo-galvanic and cardi.ac reactions remained below control levels
although their increase had begun by the 4th day. In the mountains of Altay, on the
3rd-4th day, immediate and delayed word reproduction were somewhat lower than the
control, but in another test, the indices exceeded the control. Levels of the
accompanying EEG activation, dermo-galvanic and cardiac reactions were within the
limits of normal. Verbal memory for syllables from the 4th day, and form-space
memory from the 3rd d~y showed definite improvement (fig 34).
~,