doi:10.1152/japplphysiol.00435.2001 92:1378-1382, 2002.J Appl PhysiolphysiologistsEarly contributions of Russian stress and exercise

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historical perspectivesEarly contributions of Russian stressand exercise physiologists

ATKO VIRUInstitute of Exercise Biology, University of Tartu, Tartu 51014, Estonia

Viru, Atko. Early contributions of Russian stress and exercise phys-iologists. J Appl Physiol 92: 1378–1382, 2002; 10.1152/japplphysiol.00435.2001.—In Russia, the free development of scientific ideas wassuppressed in 1950 as a result of the actions of the Joint Session of theAcademy of Sciences and the Academy of Medical Sciences of the USSR.Hans Selye’s theory on the general adaptation syndrome was consideredunscientific. From 1956 to 1958, Pjotr Anokhin and Pjotr Gorizontovpresented definitive arguments for having the theory accepted by scien-tists while the significance of hormones in adaptation became a topic ofendocrine studies (Boris Aleshin, Igor Eskin, Vassily Komissarenko,Samuel Leites, and Michael Kolpakov). Later, Felius Meerson madeessential contributions to the adaptive significance of protein synthesisand stress-limiting systems. The area of exercise physiology dealing withacute and chronic adaptation to strong physiological stressors wasfounded by Leon Orbeli and developed by Aleksei Krestovnikov. Signif-icant contributors to this area were Vladimir Farfel, Nikolai Yakovlev,and Nikolai Zimkin. Although the majority of their publications haveremained unknown outside of Russia, it is interesting that many of theirresults have been “rediscovered” by others. Yakovlev also deserves rec-ognition because he was among the founders of contemporary exercisebiochemistry and because his research has provided the foundation forcurrent investigations. Several generations of young scientists have beeninspired by the above-mentioned Russian scientists. Today, however, theresearch activities of scientists are no longer limited by political pres-sures but by financial resources instead.

adaptation; exercise biochemistry; exercise physiology; general adapta-tion syndrome

THE PURPOSE OF THIS PAPER is to recognize the impact ofRussian physiologists who introduced studies on gen-eral adaptation syndrome and who ensured develop-ment of exercise physiology in the Soviet Union.

In the 19th century, several Russian physiologistswere known for their pioneering research, for example,the discovery of the central inhibition by Ivan Sech-enov (26), studies on nerve physiology by Nikolai Vve-denski (30), recognition of inotropic effects by cardiacnerves (22), and establishment of the main principlesof digestion physiology by Ivan Pavlov (23). During thebeginning of the 20th century, Pavlov (24) started his

extended studies on the central nervous system andLeon Orbeli (21) elaborated the theory of adaptive-trophic action of sympathetic innervation.

The scientific research climate was altered after1917. The economic conditions after World War I andthe Russian Revolution restricted opportunities for ex-tended research activities, although Pavlov and a fewother outstanding scientists (e.g., Orbeli) had adequatefacilities and funding. In addition, an increased dis-trust of capitalist countries arose, which meant sup-pressed international contacts for Soviet physiologists,who required approval from Pavlov (as a result of hisNobel Prize) to maintain contact with foreign scien-tists.

The Cold War arising after World War II disruptedwhat remained of international scientific contacts, and

Address for reprint requests and other correspondence: A. Viru,Institute of Exercise Biology, Univ. of Tartu, 18 Ylikooli, Tartu51014, Estonia (E-mail: aluik@ut.ee or viru@ut.ee).

J Appl Physiol 92: 1378–1382, 2002;10.1152/japplphysiol.00435.2001.

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the “iron curtain” separated the Soviet Union andcapitalist countries. Leaders of the Communist partypromoted the attitude that Soviet scientists had betterideas than their foreign colleagues who were lookingfor Soviet ideas to obtain grant funds in their coun-tries. The possibilities to take course work and toparticipate in joint research with foreign universitiesdisappeared; these opportunities did not reappear un-til the 1960s and then only to a limited extent. Forexample, to publish a paper in a foreign journal, scien-tists had to receive special permission and only leadingscientific institutions were permitted to purchase ad-vanced research equipment that was manufacturedabroad.

The development of disciplines such as physiologywas disturbed by political interference into the activi-ties of scientists. In 1950, the Joint Session of theAcademy of Sciences and the Academy of Medical Sci-ences was held (25). The purpose of the session was topromote the use of Pavlov’s findings on scientific inher-itance for the development of physiology. However, thesession actually canonized Pavlov’s concepts and sup-pressed the free development of scientific ideas. Therequirement to follow Pavlov’s ideas was a form ofpolitical control on the research work of physiologistsand other scientists. Because of this, the scientificauthority of Pavlov was discredited, even though Pav-lov was known to be an active fighter for the scientificmethod as well as for the principles of democracy.

As a consequence of the Joint Session, several out-standing physiologists such as Orbeli, Pjotr Anokhin,and Aleksandr Ginetsinski were removed from theirleading positions in institutes. In addition, severalresearch directions were officially labeled as being un-desirable, which meant that they were forbidden. Dur-ing the latter portion of the 1950s, Soviet physiologybegan to be gradually released from political suppres-sion. Unfortunately, the consequences lasted muchlonger, as research work was seriously disarrangedand competent progressive scientists were replaced bythose who were politically loyal but had limited scien-tific ability. However, a number of moral and scientif-ically respected physiologists survived this period (e.g.,Anokhin, Farfel, Ginetsinski, Pjotr Gorizontov, Yakov-lev, Zimkin, to name a few). Later, they became leadersin restoring previous scientific traditions and the prac-tice of the scientific method. They were soon joined bymany young scientists who strived to become ac-quainted with the results of foreign physiological re-search and to have the opportunity to study abroad.

STRESS PHYSIOLOGY

The theory of Hans Selye on the general adaptationsyndrome was among the forbidden research themes.Two scientists who introduced the stress theory intoscientific understanding in the Soviet Union were Anok-hin (2) and Gorizontov (9).

Pjotr Anokhin (1898–1974). Anokhin graduated fromthe Leningrad (Saint Petersburg) Medical Institute. In1921, being a degree candidate, he began his research

work at the Institute of the Brain under the supervi-sion of Vladimir Bekhterev. Later, he became a post-graduate student of Pavlov. In 1930, he was recom-mended by Pavlov to be professor of physiology at theUniversity of Nizhni Novgorod. In 1935, he moved toMoscow and became the Director of the Institute ofPhysiology of the Academy of Medical Sciences in 1946.After the Joint Session of the Academies, he was as-signed to the Rjazan Medical Institute (�200 kmsoutheast of Moscow). Although the opportunity forexperimental research was limited at Rjazan, he re-mained scientifically active and focused on theoreticalgeneralizations. In 1953, he returned to Moscow; threeyears later, he was elected to be professor of physiologyat the Moscow First Medical Institute, which meantthat his “political rehabilitation” was complete. Later,he founded the Research Institute of Normal andPathological Physiology.

Anokhin was a neurophysiologist whose main scien-tific contribution was developing integrative physiol-ogy. In 1935, he published the first outlines of thistheory. Further development made it a fundamentalgeneralization of functional integration having adap-tive and behavioral aims (4). Unlike the widespreadapproach, he analyzed the relationships between Pav-lov’s theory on conditioned reflexes and recent ad-vances of neurophysiology (see Ref. 4). This integralapproach guided him to the analysis of emotionalstates and the theory of stress as a general adaptiveresponse. In 1958, Anokhin (2) described the essence ofSelye’s stress theory: that specific physiological pro-cesses ensured the body’s adaptation to unfavorableconditions and that these made the body more resis-tant. This was a courageous scientific statement be-cause, in the late 1950s, the political control of sciencepronouncements was still in existence. For that time,the positive evaluation of the stress theory by a leadingscientist was highly significant.

In 1962, Anokhin published a profound analysis ofadaptation processes and their relationships to ho-meostasis (3). He discriminated between 1) plastic con-stants of the body, which were changeable to a greatextent; and 2) rigid constants, which have a very nar-row range of fluctuation between the resting and activ-ity levels from which deviations cannot be associatedwith maintenance of life. However, the main contribu-tion of Anokhin for the adaptation processes was histheory of integrative physiology. The further develop-ment of this theory was made by Sudakov in 1984 (29).

Pjotr D. Gorizontov (1902–1987). Gorizontov gradu-ated from Omsk Medical Institute and in 1962 becamethe director of in the Institute of Biophysics in Moscow.In the Joint Session of the Academies, he exhibitedgreat courage by defending the banned scientists. In1956, Gorizontov published (9) a review paper on thegeneral adaptation syndrome and diseases of adapta-tion in which he criticized the approach that attemptedto explain all findings in clinical and experimentalpathology by actions of the nervous system, an ap-proach that ignored the responses of endocrine glands.

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He later emphasized that the unspecific nature ofstress responses has only a relative significance be-cause these responses depend on the initial state of thebody. Gorizontov considered that a limitation in Selye’sapproach was that Selye only dealt with pathogenicmechanisms of diseases and not the etiological mech-anisms (11). In 1976, Gorizontov (10) edited a largevolume on the significance of homeostatic regulation inadaptation.

Two books by Selye were published in Russian in1960 (28) and 1961 (27). The second book was intro-duced by Vassily Parin, a leading cardiologist and aninitiator of space physiology in the USSR.

Because of the restrictions imposed by the JointSession, endocrine studies were possible only if theycontributed to the neural regulation of endocrine func-tions. However, in 1955, a journal on problems ofendocrinology (Problemo Endokrinologij) was founded.Subsequently, hormones in adaptation became a topicof study by many investigators [Aleshin (1), Eskin (6),Kolpakov (13), Komissarenko (14), and Leites (17)],and several research centers were established thatproduced relevant research in endocrinology. Signifi-cant contributions were made by Yudayev (36) and histeam on the biosynthesis and metabolism of steroidhormones.

Meerson made essential contributions to short- andlong-term adaptations to stress. In 1965 (19), he indi-cated that a molecular mechanism exists that linkscellular function with its genetic apparatus. Throughthis mechanism, the synthesis of proteins is enhanced.He implicated his findings in the analysis of cardiolog-ical problems (20) and elaborated a concept of stress-limiting systems (antioxidant mechanism and endoge-neous opioid responses).

EXERCISE PHYSIOLOGY AND ITS CONTRIBUTIONTO STRESS PHYSIOLOGY

Exercise physiology deals with acute and chronicadaptations to strong physiological stressors. In Rus-sia, the development of exercise physiology is attrib-uted to activities of Krestovnikov (16), Orbeli (21), andZimkin (37), as well as of Farfel (8) and Yakovlev (33,34). The first three were pupils of Pavlov and profes-sors of exercise physiology in the Leningrad (SaintPetersburg) Leshaft Institute of Physical Education(Orbeli from 1919 to 1927, Krestovnikov from 1927 to1955, and Zimkin from 1961 to 1975). Farfel (8) initi-ated strong centers of exercise physiology in Moscow.Yakovlev (33, 34) made the Leningrad Research Insti-tute of Physical Culture well known by his research inexercise biochemistry.

Aleksei N. Krestovnikov (1985–1955). Krestovnikovgraduated from Saint Petersburg University (1912)and from Petrograd Medical Institute (1923). In 1912,he worked in Pavlov’s laboratory and later becamesenior assistant to Orbeli in the Department of Physi-ology, Leningrad Leshaft Institute of Physical Educa-tion. Orbeli demonstrated the increase of contractionamplitude of fatigued muscle when sympathetic nerves

were stimulated (21). Krestovnikov, his collaborator,found an increased oxidation rate of muscle when sym-pathetic nerves innervating the muscle were stimu-lated (15). In 1927, Krestovnikov replaced Orbeli asprofessor of physiology in the Leshaft Institute andintroduced studies on the physiological changes duringexercise and training. Noteworthy were his studies onthe reflexes evoked by movements of the head. He useda special plaster cast to exclude any movement of head.Even in highly qualified athletes, the result was dis-turbed motor skills. The same effect occurred whenperipheral but not central vision was excluded. Inregard to exercise training, he found conditioned re-flexes were necessary for increased performance. Hisresearch has been summarized in two monographsthat were edited in 1939 and 1951 (16).

Nikolai N. Yakovlev (1911–1992). Yakovlev gradu-ated from the Leningrad Medical Institute in 1928 andjoined the Laboratory of Physiological Chemistry at theLeshaft Research Institute of Natural Sciences. Later,he moved to the Laboratory of Pathological Physiology,Leningrad Research Institute of Physical Culture. Hiswork concerned the role of vitamins in the trainingprocess and the contribution of hormones from thepancreas during exercise. After World War II, his re-sponsibility was to establish a laboratory of exercisebiochemistry at the Leningrad Research Institute ofPhysical Culture.

He discovered (1949–1959) the phenomenon of “su-percompensation” of muscle and liver glycogen andmuscle phosphocreatine during postexercise recoveryand with training (32, 34). In 1955, his work on thesignificance of anaerobic resynthesis of ATP in inten-sive exercise was published (32). His research on me-tabolism involved several areas: 1) the training effecton the mitochondrion; 2) ATP, ATD, and AMP concen-trational changes in muscles with various exercises; 3)lipid and phosphorous changes in brain tissue; 4) therelationship between GABA and fatigue (brain GABAcontent increased in rats with fatigue); 5) the role ofornithine and polyamines in muscle hypertrophy; 6)cAMP changes and the action of hormones; 7) antici-patory changes in blood of athletes before competition;8) dependence of carbohydrate and phosphorous me-tabolism on the state of the central nervous systemduring exercise; 9) exercise-induced metabolic re-sponses in myocardium and liver; and 10) several prob-lems related to nutrition of athletes (for reviews, seeRefs. 33 and 34). Beginning in 1949 and continuing to1976, he founded the concept of the biochemical speci-ficity of tissue responses to training, which indicatedthat the biochemical response of a tissue depended onthe type of training that was performed (31, 34). Dur-ing 1973–1975, he found that the training-inducedincrease in tissue sensitivity to epinephrine was typi-cal for the first period of training and related thechange to an increased activity of adenylate cyclase. Healso showed (35) that this was followed by an increasein AMP-phosphodiesterase and a normalization in thesensitivity to epinephrine.

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Yakovlev was among the founders of contemporaryexercise biochemistry, and his work influenced re-search in other laboratories. Although biochemical re-search techniques have been vastly improved since histime, contemporary studies have confirmed his find-ings and continue to be an inspiration to contemporaryresearchers.

Nikolai V. Zimkin (1899–1990). Zimkin graduatedfrom the Academy of Military Medicine in 1926. Hestarted his scientific career under the supervision ofPavlov. By 1945, he was the head of a Laboratory in theInstitute of Evolutionary Physiology (Leningrad). In1947, he received a professorship in the Military Insti-tute of Physical Education (in Leningrad) where heconducted longitudinal experiments on training of var-ious motor abilities (37). Because most of his mono-graphs and articles were unknown in other countries,many of his results have been “rediscovered” by others(e.g., cross-training and detraining effects; positive andnegative interaction in improvement of strength,speed, and endurance; mastery of motor skills). It isnoteworthy that his studies on training effects on bodyresistance to heat, poisons, hypoxia, and radioactiveirradiation provided the first experimental evidence ofchanges in body adaptivity with chronic exercise (38).These results showed increased adaptation under theinfluence of exercise training. However, if the trainingregime was too intense, survival of the experimentalrats was decreased.

Other meaningful results from his scientific researchwere the concept of “fatigue focus” and the demonstra-tion of extrapolation in athletes’ motor skills (rapidadjustments of precise motor skills in changeable sit-uations). According to the concept of fatigue focus,published in 1962, the type of exercise, the externaland internal conditions, and the individual peculiari-ties of the subject determine a critical complex ofchanges, disturbing the activity of neural centers or ofmuscle metabolism and causing a decrease in workcapacity. This concept is in accord with the catastrophetheory of fatigue (see Ref. 5). According to Zimkin,latent fatigue has to be recognized. This is manifestedby a feeling of tiredness or discoordination of functionswithout any decrease in work quantity.

Vladimir S. Farfel (1904–1979). Farfel was educatedas a physiologist at Leningrad University. His mentorwas Aleksei Ukhtomski who was an expert in centralnervous system physiology. Farfel became the head ofthe Physiological Laboratory of the Central ResearchInstitute of Physical Culture (1934–1947) and profes-sor in the Pedagogical Institute of Moscow’s Region(1947–1958). Later, he directed a laboratory in theInstitute of Ontogenetic Physiology, Academy of Peda-gogical Sciences (1958–1960), and finished his careeras a professor of physiology in the Central Institute ofPhysical Education (1960–1972). At every institution,he established a strong team, conducted meaningfulresearch, and was chiefly responsible for making hislaboratory well known in the USSR.

Farfel became recognized for his systematic physio-logical classification of exercise. In following the prac-

tice of Archibald Hill (12), he analyzed track events bylogarithmically plotting the running velocity (intensi-ty) against exercise duration and found that fourstraight lines were present. Consequently, he was ableto discriminate four different types of cyclic exercise bytheir intensity (7). He concluded from his studies(1939–1940) on elite athletes during important com-petitive events that trained subjects during maximalexercise have more pronounced functional cardiovas-cular and respiratory manifestations than untrainedsubjects, even though their responses are less withsubmaximal exercise.

Farfel conducted an extensive study on endurancethat is seldom cited. In 1949, a member of his researchteam demonstrated that glucose administration justbefore the start of a long distance event will causehypoglycemia to occur during the early stages (18).Obviously, it was related to glucose effect on insulinsecretion.

Several generations of young scientists were in-spired by these four Russian “giants.” Because theirresults have been published predominantly within theRussian literature, a language barrier greatly limitedthe distribution and awareness of their findings. Dur-ing the 1980s, the leading scientists were Yakov Kots(neural mechanisms of movement), Viktor Karpman(cardiovascular studies), Sergei Kuckin (respiration),Erlena Matlina (catecholamine metabolism), ViktorRogozkin (mechanism of induction of protein synthe-sis), Jelena Sologub (EEG studies), Vitali Thorevski(circulation), Nikolai Volkov (muscle energetics), andVladimir Zatsiorsky (biomechanical mechanisms).

In summary, an educated and inquisitive mind canfind opportunities for creative activity despite restric-tive and oppressive conditions. Fortunately, politicallydriven restrictions for scientific contacts and ex-changes between Russia and other countries are nolonger present as they were before and during the ColdWar. Today, the effectiveness of research work is lim-ited only by the availability of funding. However, newgenerations of scientists remain in debt to the impor-tant research work of the above-mentioned “greats.”

The author gratefully acknowledges Dr. C. M. Tipton’s idea toconsider Russian stress and exercise physiologists who are essen-tially unknown to their American colleagues.

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