EFFECT OF MUSCULAR ACTIVITY ON CONDITIONED SECRETARY REFLEXES IN MAN
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CIA-RDP81-00280R001300170021-8
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RIPPUB
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U
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20
Document Creation Date:
December 22, 2016
Document Release Date:
June 8, 2011
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21
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Publication Date:
October 25, 1956
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STAT
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Fisiclogl a nerve kh V. A. Novi, Candidate
protsessov [Physiology in Medical Sciences,
of Nerve Processes), Laboratory of the Higher
1953, Kiev, Pages .38-449 Nervous Activity and
Nerve Trophies, Institute
of Physiology iteni
A. A. Bogouolots, Academy
of Science-- Ukrainian SSHH
The study of the higher nervous activity of aaicxils was
placed on an objective foundation by the method of conditioned
sali'?ary railexos originated by I. P. Pavlov. No other ciethod
aspiring to objectivity can compare to it in this regard. There-
fore, while Tavlov was still alive Lents and Smirnov used the
method of conditioned secretory reflexes to study the higher
nervous acts, ty of Healthy human adults as well. Despite the
fact t.tat Pavi,iv did not regard this absolute imitation of animal
experiments as fully valid, many physiologists have employed and
still today employ his classic method in the study of the laws of
nerve proLuGi in the child, as well as in the healthy and sick
adult.
This c.%n perhaps be explained by the fact that to this day
there is no universally accepted method for studying the higher
nervous activity in man. On the other hand, while the fundazsental
mechanisms of the higher nervous activity, discovered in studios
of conditioned reflexes in animals, also apply to tan, Pavlov al-
ways warned against simply transferring data from the animal to the
human sphere. In this regard, K. U. Hykov has written chat 'human
activity involves a number of specific :actors which cannot bv
t,iven due eonaldoration in a si:.plified analysts based solely on
STAT ~
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has yet buca devised. Therefore, objective and properly-grounded
Conclusions with re atd to va_ious phenomena of human cranial
ac[i%ity cat: be obtained only if this judgment is based on the
study of nunan cortical processes not merely by one, but by a
n:neber of c.ut;tods.
This methodologically multi.aceted approuch to the study of
any ,,1ven manifestation of higher nervous activity in man uust
employ as one of its means the investigation of conditioned secre-
tory reflexes.
G. V. Fol'bort and associates have determined the basic laws
of the occurrence and interaction of the processes of exhaustion
and recovery fundamental to the processes of fatigue and rest.
The development of this direction of study is currently in the
stage of the investigation of the processes of exhaustion and re-
covery in the cerebral cortex. Under Fol'bort's 1.idauco we are
presently doing research on the interrelation betwe,n the processes
of excitation and inhibition in human muscular activity, and toward
that end we are employing a number of methods.
In full consciousness of the fact that an effort to apply
to man the results of research on the conditioned reflexes of
dogs canno..be successful, we nonetheless decided to supplement
the nateriai we had previously collected by undertaking a study of
the conditioned reflexes of man in muscular activity.
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Conditioned secretory reflexes were investigated in a
study of the cranial activity of man in muscular activity. Saliva
from the parotid gland was withdrawn in a ploxit;lass capsule (de-
signed by Krasnogorskiy, Levin, and Klorin). The amount of saliva
secreted was determined by the Ganiko-Kupalov method (passage
throui;h water and air, one drop corresponding to 3 gradations on
the scale, and 4U drops to one cu cm). The nuuber of drops was
recorded by kymo;,raph, whereby a drop falling on the Gaaike in-
strument closed an electric circuit and was recorded on a smoked
tape by an electrically-operated needle.
unco,x:itionod stimulus. The unconditioned stimulus was applied
for 30 seconds, during which tine the subject drank 8 to lU cros
of cranberry Juice. The positive conditioned reflexes employed
included a quiet bell, the lighting of a .0-w electric bulb, the
lighting-up of a red square, a motronone at 180 beats to the vin-
ute (this stimulus was used for differentiation with 2 of the sub-
jects), and the conditioned stimulus "I an giving you the cranberry
Juice." The differentiation stimuli were the li,,htin,-up of a
~.reen square, and a metronome at 90 beats to the utlnuto (this
stimulus woes positive with 2 subjocte).
Mien the reflexes had been elaborated, and differentiation
fixed, the effect of muscular activity thereon was tested. In
other instances a study was made of the effect of muscular activity
on the procass of elaborating conditioned reflexes. The effect of
muscular work was also tested in cases in which, despite the consid-
erable number of cocbinarions, the conditioned reflexes retained in-
rons,ant.
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The conditioned swallowing reflex was studied simultaneously
with the conditioned secretory reflexes. The method of air trans-
mission developed by X. I. [rasnogorskiy was employed to record
swallowing.
The muscular activity consisted of pedalling a stationary
bicycle carrying a load differint, for each subject. The difficulty
of the work was determined by taking the pulse and measuring res-
piration and perspiration. The subjective attitude of the subject
to the work was determined, the units of of fort applied were cal-
culated, and aeroenergographlc data were taken into account. The
method of acroenorgographic recording has been described by hhur-
avlev and Kudryavtsev. (The subject is told to squeeze a rubber
bulb, the ccmprossion being recorded by a kycio;raph operated by
the air in the bulb. A Richardson bulb and pressure _..tui;e are
used to ostablish the desired pressure in the system, ti,us provid-
ing the resistance needed for compression by i;and). We asked the
subjects to squeeze the bulb with utmost stren;,th and frequency
until co longer able to do so. The subject was asked to do this
before pedalling the stationary bicycle, a,;ain i,asaodiately after
finishint; that work, and subsequent to rest thereafter. In view
of the results of the experiments of Zhuravlev and Ktuiryavtsov,
who noted a rise in work capacity after limited work, and a decline
after much work, we believed that these seroonergoirams would help
us determine the difficulty of the work performed.
On the basis of the various indices listed above we clas-
sifted the work performed by the subject as light, tedium, and
:Davy, or the individual concerned. Our subjects were students,
acttool pupils. and technical personnel, lu is nuuber. A tutul of
We were convinced ourselves at the very outset of the dif-
ficulty of elaborating conditioned secretory reflexes in rsan. Our
first experiments, with subjects 0. t7. and L. D., produced no condi-
tioned reflexes even after 100 or none combinations of stinulA. Los-
ing hope of elaborating conditioned reflexes in the usual way, we
asked one of our subjects to try to think about the conditioned re-
Flax when it appeared, givint; attention to its taste and color.
This did produce reflexes on 2 or 3 occasions, but as they failed
to become constant, we temporarily dropped the experiments as
vacation time had come. e.hen work with these subjects was renewed
the latter showed precise reflexes for a short period, but there-
after the reflexes appeared and disappeared irregularly.
Developing conditioned reflexes in subject L. N. proved
equally difficult. In her case, it took more than :0 combinations.
With the rest of the subjects, conditioned reflexes appeared after
lu to 14 combinations. With certain individuals the conditioned
reflex first appeared to light stinali, and then to sound (P.V.),
while with others it was just the opposite (S. 1.). The condi-
tioned reflex appeared more rapidly to the oral stimulus ("I am
giving you the cranberry juice") than to the others. With the
subjects who developed conditioned reflexes rapidly, relative
r.tability of these reflexes was evident from the seventh or eighth
experiment, while in subject 0. G. this stability was obtained
only in the thirtieth experiment, her reflexes being clearly do-
fined from the thirtieth to the thirty-seventh experiments,
thereafter tarot thus to the and of the study) alternately appear-
ing and failing to appear. Conditioned secretion in subjects
L. N. and L. D. was likewise alternating in its appearance. With
regard to the remaining subjects, it is also impossible to speak
of clearly-defined conditioned reflexes, although home degree of
constancy was nonetheless to be observed. True, toward the and
of the observations, even relative constancy was lacking in virtu-
ally all the subjects, and the reflexes were lacking in a number
of cases.
The magnitude of the reflex was small in all subjects,
even those in whom the reflex was distinct. In the majority of
the subjects the conditioned reflex represented 3 to ) gradations
on the scale, reaching 10 to 15 only in the case of S. T. In ad-
dition, the unconditioned stimulus produced an adequate secretory
effect.
Figure 1 presents kymographic records of the conditioned
reflex. Here (subject Kh. 1t.), the conditioned secretion con-
sisted of 3 drops of saliva, preceded somewhat by a motor reaction,
swallowing. A combined study of the swallowing reflex and the se-
cretory reaction showed inconstancy in this reflex reaction as well.
In a number of canes this reflex was clearly defined, in which case
the swallowing motions preceded the conditioned reflex, but in the
majority of experiments, the conditioned reflex for swallowing was
absent. Krasnogorskiy and Shastin believe that the motor rer.ctions
izust be retarded only as supplementary, in view of their lack of
constancy.
requested them to take neither dinner nor supper before the ex-
periment* The rule was for the subjects to participate in the
experiment before breakfast. However, if the experiment were to
be performed curing the second half of the day, the last intake of
food was not to be less than 5 hours before the experiment. After
the experiment the subject was given breakfast in the laboratory.
Our obs.'rvations of these subjects showed more constant conditioned
reflexes than in others, but this constancy was disturbed there-
after, and the reflexes began to disappear. Questions to the sub-
jects elicited the information that, despite the artificial induc-
tion of hunger, they had never thought about food, although other
thoughts had often occupied them.
Our preliminary work in the elaboration of conditioned hu-
man secretory reflexes may be summed up as follows. In man, con-
ditioned, artificially-induced reflexes are not stable and easily
disappear on repetition. The magnitude of the conditioned reac-
tion is low and irregular. Only a large amount of experimental
data makes possible the discovery of a relationship between the
rtiagni it!do of the reaction and the usa of any given stimulus. Dif-
ferentiation was easier to obtain and was more constant. S. T.
was an exception In this regard; she failed to reveal differentia-
tion.
On the basis of the foregoing standard, which is not fully
precise, we nonetheless proceeded to study conditioned reflexes of
secretion during muscular activity. In 2 Instances we began our
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Our observations showed that muscular activity facilitates a
more precise ranifestation of conditioned reflexes and differentia-
tions, and more rapid formation thereof. But this aeries of ob-
servations was likewise characterized by lack of constancy in the
manifestation of conditioned reactions. In a number of experiments
very precise results were observed, but o.i other occasions the find-
ings, under identical conditions and with the same subjects, were
different, and we could thus not be confident of the conclusion
drawn on the basis cf th prior experiment.
table, In compiling it, we distinguished, to begin with, the find-
ings resulting from different types of work: with ti(;ht, avorat;o,
and muscular loads respectively. Then, in each series of obsorva-
Lions ""ud for each experiment, we determined strengthoninj;, weaken-
ing, r failu.-. of the conditioned reflex to show chant,e. The
same procedure was followed, as concerns differontiation.
EFFECT OF MUSCULAR ACTIVITY ON COMITIONED SECRETORY
REZ IEXES IN MAN
Conditioned reaction
Differentiation
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Of 28 experiments with _ight muscle load and different sub-
jects, the data i.t the table silow that there was an increase in the
conditioned reflexes in 18 exptorinionts, and no change in the re-
maini_.i; lo. wi noted no weakening of the conditioned reflexes to
light wo.%. :irht muscular activity disorganized differentiation
in 10 experiments. In 17 experiments the reaction remained un-
changed, and in one it was strengthened.
It, work of medium difficulty, the conditioned reflex also
showed an increase in the majority of cases (23 of 40 experiments),
while differentiation was weakened in 13 experiments. On the
other hand, in 8 experiments of :iedi%im difficulty, the conditioned
reflexes were weakened.
In 20 of 37 experiments involving severe muscular activity
the table shows that the result was an increase in the conditioned
secretion only nt the beginning of the observation, while at the
end the reflexes declined. In 4 experiments with heavy muscular
activity a weakening in the conditioned ref loxes was observed
throughout the .ntiro experiment, and in 2 cases the reflexes in-
creased throu;,:.out the experiment. On heavy work, differentiation
was disorgan.t..-.kk in 13 experiments, while in 6 it increased.
The di!',culty of muscular work was determined in accord-
ance with a nun_ or of indices, but it was always difficult to
make accurate allowance for such a factor as the attitude under
which the work was performed. Therefore, in a number of instances
the possibility that work of medium difficulty could become heavy,
and vice versa, cannot be excluded. It is possible that it is
specifically this which explains the fact that in s oxpori:rents
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Having than grouped our data in a table, and having found
lo u majority of instances definable principles, we believed it pos-
sible to advance selected experiments which would confirm the
principle inherent in each, although the number of those precise
experiments constitute only a third of our total number of obsorvu-
tions.
Figure 2 presents an experiment, subject L. N. The illustra-
tion shows clearly the strengthening of conditioned reflex activity
in lic.ht muscular activity. Before this work, her reaction to the
metronome, at 90 beats, was one drop of saliva at the sixty-first
combination, and 2 drops in response to the rod square at the
twenty-ninth try. Her conditioned secretion increased during the
performance of work. Four drops were secreted in response to the
lighting of the lamp on the first try, 3 drops on the thirty-sec-
end try to the red rectangle, and 2 to the 90-boat metronome at
the sixty-second combination of stimuli. After rest, the condi-
tioned secretory reaction again showed a decline, one drop of
saliva being secreted in response to the 9U-beat metronome at the
sixty-third combination.
Jn a number of cases the conditioned swallowing reflex was
manifested clearly, in which situation it pret.eded the secretory
reaction. In the majority of cases, however, the swallowing reac-
tion sot in only in response to the unconditioned stimulus.
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anee of work, the conditioned reaction to light, metronome, and
bell was 3 gradations on the scale, and differentiation was com-
plete. During work of medium difficulty the conditioned secre-
tion increased 2- or 3-fold (8 gradations on the scale to the
thirteenth combination of stimuli including the lamp, 12 ;;radations
at the fourteenth, and 9 gradations at the fifteenth. Difierenti-
ation was markedly disordered (saliva secretion to 180 beats of
the metronome being , radations at the sixteenth combination or
stimuli, and 10 gradations at the seventeenth). After 13 ^inutes
of rest, the magnitude of the conditioned reflexes declined, but
not to its initial value.
In the majority of cases heavy muscular activity elevated
the secretory reaction only at the beginning of the experiment,
its magnitude declining thereafter. Figure 4, showing Experiment
8 with suuject V. Kh., shows this effect. Prior to work, the
nineteenth combination of stimuli including the lamp showed the
c(adltioned secretory reaction to be 7 gradations on the scale.
After 6 minutes of work, secretion of saliva to that stimulus
constituted 8 gradations on the scale, but only 2 after 3:i minutes,
and 3 after 4o minutes. The 180-boat metronome yielded a .-grada-
tion reaction before work, 5.2 after 11 minutes of work, 7.5 after
19 minutes, and 2 after 49 minutes.
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method of condtt:onee secruto.-p ,:. mats is use..- in
certain manifestations of the higher nervous activity In man. Con-
ditioned ref locus of secretion are often developed, it is true,
with great difficulty, the stren;;th of the inhibitory process thus
making itself felt.
One is struck by the irregularity of the conditioned reflex,
and by its disappearance, for which no explanation Immediately
presents itself. We were unable to determine any precise connec-
tion between theme phenomena and either happenings it, the lives
of the subjects, or their state of mind, or even with :ood excita-
bility, although in a number of instances such a relationship
clearly existed. Experiments set up against the background of the
uninterrupted reading of books by the subjects presented interest-
ing resultr. In these cases the conditioned secretory reaction
omer4,od more clearly. The conditioned reaction also appeared with
greater clarity on performance of light muscular activity.
As long ago as when Lents demonstrated the first condi-
tioned secretory reflexes, it was observed by V. Yu. Cha ovets
that the major obstacle with respect to the observation of condi-
tivne