Founders of world and domestic physiology. History of development of physiology (Harvey, Descartes, Sechenov, Pavlov, Anokhin). Physiology BC

The history of the development of physiology, like other biological sciences, originates in ancient times. Man has always been interested in the structure and functions of the body, the first information about this was summarized and presented in the writings of the "father of medicine" Hippocrates. The structure of the digestive organs, blood vessels was described by the ancient Roman physician anatomist Galen (II century AD). An important role in the study of the beneficial effects of hygienic factors (nutrition, sunlight, air) and the nervous system on the human body was played by the scientist (XI century AD) Abu-Ali-Ibn-Sina (Avicenna).

The founder of experimental physiology and embryology is considered to be the English anatomist and physiologist W. Harvey (1578–1657), who proposed a research technique by dissecting tissues (vivisection). This made it possible to make important discoveries in the functions of the cardiovascular system. Based on his numerous observations, Harvey gave a reasonable idea of ​​the circulation of the blood. It was he who first expressed the idea that "all living things come from an egg."

Subsequently, the doctrine of blood circulation was significantly supplemented by the Italian biologist and physician M. Malpighi, who in 1966 discovered the presence of capillaries.

The founder of experimental physiology in Russia is Professor of Moscow University A.M. Filomafitsky (1807–1849), author of the first textbook on physiology.

The introduction of tissue dissection served as a powerful impetus for the study of various body functions. The first, although largely simplified, ideas about the reflex were formulated by R. Descartes (1596–1650), and later developed by the Czech scientist Georg Prohasko, who introduced the term “reflex” into science.

The French scientist F. Mogendi (1785–1855) discovered the separate presence of sensory and motor nerve fibers in the nerve trunks, which made it possible to better understand the nervous pathways for regulating the functions of organs and body systems. The German naturalist I. Müller is the author of works on the physiology of the central nervous system, the sense organs (vision, hearing), and some endocrine glands.

In 1771, the Italian physicist and anatomist L. Golvani revealed the occurrence of electric currents in the muscles. These studies were continued by Müller's students - the German physiologists Dubois-Reymond (1818-1896), Helmholtz (1821-1894).

Soviet physiologists V.Yu. Chagovets (1873–1941) and A.F. Samoilov (1867–1930) was the first to suggest the chemical mechanism of excitation transmission in synapses and that the origin of currents in tissues is based on a change in the permeability of cell membranes for different ions. In the 40-50s of the twentieth century. this idea served as the basis for the outstanding substantiation of the membrane theory of the occurrence of bioelectric potentials in tissues (A. Hodgkin, A.F. Huxley and B. Katz).


Of considerable interest are the works of the English neurophysiologist Ch.S. Sherringston (1859–1952). Soviet physiologist I.S. Beritashvili (1885-1974) substantiated the position on dendritic inhibition and psycho-nervous activity of a person.

In the field of physiology of visceral systems, the works of the English physiologist W.Kh. Gaskell (1847–1914), devoted to the study of the function of the autonomic nervous system. D.N. Langley (1852-1925) called it "autonomous", thus emphasizing its independence from the higher parts of the nervous system. In contrast to this, Academician K.M. Bykov (1886-1959) revealed the presence of conditioned reflex reactions in the activity of internal organs, showing that autonomic functions are not autonomous and are subject to the influences of the higher parts of the central nervous system.

F. Mogendi, C. Bernard, R. Heidenhain, I.P. Pavlov in numerous experiments on different animals substantiated the idea of ​​the trophic role of the nervous system. I.P. Pavlov believed that the function of each organ is under triple control - neuro-functional, vascular and trophic.

L.A. Orbeli (1882–1958) together with A.G. Ginetsinsky (1895–1962) studied the influence of the sympathetic nervous system on various body functions, which subsequently made it possible for L.A. Orbeli to formulate the doctrine of the adaptive-trophic role of the sympathetic nervous system. K.F. Ludwig (1816–1895), F.W. Ovsyannikov (1827–1906) established the presence of a vasomotor center in the medulla oblongata.

K. Ludwig and I.F. Zion in 1866 discovered the centripetal nerve, which slows down the heart and lowers blood pressure. This nerve was called by them the depressor. In the laboratory of Ludwig, the Zion brothers continued their research on the influence of sympathetic nerves on the work of the heart. In addition, K. Ludwig is the author of the invention of the kymograph and the introduction of a graphical method for recording blood pressure in physiological studies. Subsequently, this method was widely used in the study of many other functions of the body.

As a result of studies on frogs and rabbits, A.P. Walter (1817–1889) and C. Bernard (1813–1878) established that sympathetic nerves constrict the lumen of blood vessels.

The English physiologist E. Starling (1866–1927), studying the dynamics of cardiac activity, noticed that the strength of heart contractions depends on the amount of blood flowing to the heart and the length of its muscle fibers at the time of contraction. An important point in physiology was the discovery of N.A. Mislavsky respiratory center in the medulla oblongata.

Academician P.K. Anokhin (1898–1974) put forward the idea of ​​the functional interaction of the internal organs and systems of the body with the central nervous system on the basis of their feedback, which greatly expanded the previous ideas about the nervous mechanism of regulation of functions.

The founder of physiology in the United States, physician W. Beaumont (1785–1853), conducted many years of observation of gastric digestion in a person who had a non-healing gastric fistula after an injury.

An invaluable contribution to the physiology of digestion processes was made by the studies of K. Bernard, R. Heidenhain, B.K. Babkin. V.A. worked in this direction. Basov, Tiri, Vela, who proposed surgical methods for obtaining juices from various digestive glands.

W. Beilis and E. Starling initiated the study of humoral factors in the regulation of digestion, and I.P. Razenkov (1888–1954) successfully investigated the neurohumoral mechanism of regulation of the digestive organs. A.M. Ugolev (1926-1992) developed the theory of parietal (membrane) digestion.

The works of I.M. Sechenov (1829–1905). He has the honor of discovering inhibition in the central nervous system, which made it possible to consider in a new way the regulatory influence of the nervous system on various functions of the body. He established that the activity of the cerebral cortex is based on a reflex mechanism.

THEM. Sechenov successfully worked in Germany in the laboratories of Dubois-Reymond, Ludwig, Helmholtz. Returning to Russia, he created the Russian physiological school, from which such prominent scientists as V.V. Pashutin, A.F. Samoilov, M.N. Shaternikov, N.E. Vvedensky and others. For outstanding achievements in science I.P. Pavlov named I.M. Sechenov "the father of Russian physiology".

Dealing with the problems of neuromuscular physiology, N.E. Vvedensky (1852-1922) formulated the position on the unity of the processes of excitation and inhibition, proved that under certain conditions the process of excitation can turn into inhibition. Developing Vvedensky's theory of lability and parabiosis, A.A. Ukhtomsky (1875–1942) created a theory about the dominant.

The role and merit in the development of physiology in general and, in particular, the physiology of digestion of academician I.P. Pavlova (1849–1936). It was under his leadership that new original techniques for a number of surgical operations for the imposition of fistulas were improved and developed. The Pavlovian method of the chronic (fistula) experiment made it possible to create a fundamentally new direction in the study of the physiology of the whole organism and its relationship with the external environment.

Works by I.P. Pavlov formed the basis of the physiology of farm animals.

I.P. Pavlov was distinguished by the depth and versatility of research. He devoted his inquisitive and observant mind to the study of the physiology of the cardiovascular system, digestion, the central nervous system and higher nervous activity, and proposed a completely new analytical-synthetic approach in physiology to understanding the essence of physiological processes.

No wonder in 1904 I.P. Pavlov was awarded the Nobel Prize, and in 1935, a year before his death, the International Physiological Congress awarded him the honorary title of "Elder Physiologists of the World."

N.F. Popov, I.A. Baryshnikov, P.F. Soldatenkov, N.V. Kurilov, S.S. Poltyrev, V.V. Savich, N.U. Bazanova devoted their scientific activity to the study of digestion and metabolism in different animal species, A.A. Sysoev - reproduction and lactation, K.R. Viktorov - physiology of respiration and digestion in birds. N.F. Popov worked in the field of physiology of the central nervous system, GNA, physiology of digestion in ruminants and horses. G.I. Azimov carried out studies on the study of GNI, lactation, endocrine glands.

D.Ya. Krinitsyn studied the mechanisms of secretion of digestive juices and the motor function of the digestive organs. A.A. Kudryavtsev - metabolism and energy, GNI, analyzers.

And now A.A. Aliev, N.U. Bazanova, V.I. Georgievsky, A.N. Golikov, S.V. Stoyanovsky, each of which prepared a large number of candidates and doctors of science.

For many years in agricultural universities they study physiology according to the textbooks of K.R. Viktorova, G.I. Azimova, A.A. Sysoeva, A.P. Kostina, A.N. Golikova, N.U. Bazanova, V.I. Georgievsky.

Academician I.A. Bulygin, Professor A.N. Cheredkova, I.K. Slesarev and their numerous students, who devoted their work to the study of the physiology of the nervous system, digestion, and metabolism.

For the development of the physiology of digestion, the works of Professor V.F. Lemesh, who for many years headed the Vitebsk Veterinary Institute. In his multilateral studies, he studied the effectiveness of the use of various feeds and feed mixtures by animals. At the same institute, Professor F.Ya. Bernstein and his students studied the role of minerals in metabolic processes in animals.

Scientists of our republic have made a significant contribution to the study of the physiology of digestion, developed original methods for obtaining digestive juices, proposed new feeds and additives that improve digestive processes. A large number of their works are devoted to the study of the resistance of animals and birds in ontogenesis, the search for the most effective methods her stimulation.

Scientific research by agricultural physiologists has always been aimed at increasing the productivity, safety of animals, and their adaptation to environmental conditions.

William Harvey. Claude Bernard.

Carl Ludwig. THEM. Sechenov.

NOT. Vvedensky. A.F. Samoilov.

F.V. Ovsyannikov. I.P. Pavlov.

The term "physiology" in the meaning of natural science has been used since the 16th century. to designate the science of the animal and plant world. With the accumulation in this field of knowledge, the following independent biological disciplines were identified: botany, zoology and anatomy. The tasks of anatomy included at first a description of the structure and functions and their organs. And only in the XIX century. the doctrine of functions was separated from anatomy, for which the old name "physiology" was adopted.

The first information about the physiological functions of humans and animals was known in antiquity. Even Hippocrates (460-370 BC) knew that bile enters the intestines, and muscles cause movements; monitoring the pulse, he assessed the work of the heart. The human body, according to Hippocrates, contains four "basic juices": blood, yellow bile, black bile and mucus.

Physiology BC

Aristotle (384-322 BC) stated that blood is formed in the liver. He proved that the arteries are branches of the aorta, but attributed to them the function of conducting an airy substance.

Physiological concepts reached their greatest development in the writings of the Roman physician Claudius Galen (129-201 AD). He was the ancestor of the autopsy (vivisection) of animals (monkeys and pigs). Galen described the periosteum, the vocal apparatus, distinguished seven pairs of cranial nerves. Using vivisection, he proved that the blood moves not only through the veins, but also through the arteries, found out the participation of the intercostal muscles and the diaphragm in respiratory movements. Proved the presence of sensory and motor nerves. So he can be considered the first physiologist-experimenter. The basis of human life, according to Galen, is the soul, which is part of the universal soul - pneuma.

Despite some erroneous ideas and statements of ancient doctors and thinkers, they paved the way for the emergence of physiological science.

Physiology in the Renaissance

During the Middle Ages, the development of science slowed down sharply, and only in the Renaissance began its renewal. Carried out in the 16th century studies of the founders of anatomy A. Vesalius (1514-1564), M. Servetus (1509 or 1511-1553) and G. Fallopius (1523-1562) paved the way for physiological discoveries, in particular the systemic circulation of blood flow. For the first time the correct opinion about blood circulation was expressed by Servetus, he also discovered a small circle of blood flow. The English physician W. Harvey (1578-1657) proved in 1628 that blood moves from the heart through the arteries, and to the heart through the veins, and the constant flow of blood is due to contractions of the heart. Therefore, 1628 is considered the year when man and animals arose. Harvey did not know how the blood from the arteries passes into the veins. This question was solved by the Italian scientist M. Malpighi (1628-1694), who discovered blood capillaries, described blood erythrocytes, and studied the structure of the skin, kidneys, and lungs.

Iatrophysics and iatrochemistry

In science XVII-XVIII Art. the descriptive-anatomical direction prevailed, but even then attempts were made to introduce the methods of physics and chemistry into physiology. In the 17th century In medicine, two directions were formed: iatrophysical and iatrochemical. Iatrochemists tried to explain physiological processes from the standpoint of chemistry, and iatrophysicists from the standpoint of physics and mechanics.

The iatrophic direction is based at the University of Padua. The representative of this school was G. Borelli (1608-1679), who considered the human body as a machine, equated the movements of the limbs with levers, and applied the laws of hydrodynamics to explain the movement of blood. In 1643, K. Scheiner (1575-1650) showed that the refraction of light in the lens of the eye is carried out according to the laws of optics and the retina of the eye plays a role in the appearance of visual sensations. From the point of view of mechanics, R. Descartes (1596-1650) described in 1644 a reflex act, although the term reflex was proposed later by I. Prohaska. For the first time in 1733 blood pressure (by direct method) was measured by the English scientist S. Gales (1677-1761).

The origins of iatrochemistry are associated with the name of Paracelsus (1493-1541), who believed that all processes in the body are chemical in nature. This idea was further developed at the University of Leiden (Netherlands), where J. B. van Helmont (1579-1644), who believed that not a single process in the body is possible without the participation of enzymes, defended it. He found acid in the stomach, sea salt in the blood and urine. However, F. Silvius (1614-1672) is considered the real creator of the school of iatrochemistry, who claimed that there are saliva and pancreatic juice that convert one substance into another. At the same time, Sylvius paid much attention to the study of the anatomy of the brain. R. de Graaf (1641-1673), who studied the anatomy and physiology of the pancreas, was a student of Sylvius.

Iatrophysicists and iatrochemists were representatives of the extreme trends in medicine. Along with this, some scientists understood that neither with the participation of physics with mechanics, nor with the participation of chemistry, it is impossible to explain all the complex processes that occur as in a healthy person; as well as in a sick body.

For the 18th century such facts in the development of physiology are also characteristic. The Russian scientist M. V. Lomonosov (1711-1765) formulated in 1748 the law of conservation of matter and energy. The Italian physician L. Galvani (1737-1798) discovered in 1791 bioelectric phenomena. The Czech scientist I. Prohaska (1779-1820) described the main properties of reflexes (1794). The first textbook and eight-volume guide to physiology was written in 1755-1766. Swiss scientist A. von Haller (1708-1777). Since 1738, physiology began to be taught at the Academic University of St. Petersburg.

Physiology in the 19th century

In the 19th century there was a separation of physiology from anatomy and histology. She achieved significant success and began to teach her as a separate science. In many countries, physiological schools were created and developed, the basis of which was the performance of precise experiments. The most famous representatives of such schools were: in Germany - J. Müller (1801-1858), G. Helmholtz (1821-1894), E. Dubois-Rsimon (1818-1896), R. Heidsengain (1834-1897), K. Ludwig (1816-1885), in France - F. Magendie (1783-1855), C. Bernard (1813-1878), in England - C. Bell (1774-1842), J. Langley (1852-1925), Ch Sherrington (1857-1952), in Russia - I. M. Sechenov (1829-1905), M. E. Vvsdinsky (1852-1922). I. P. Pavlov (1849-1936), in Ukraine - V. Yu. Chagovets (1873-1941), V. Ya. Danilevsky (1852-1939), in the USA - W. Cannon (1871-1945).

Johannes Müller studied the reflex activity of the spinal cord and medulla oblongata, developed problems of sensory physiology, studied the microscopic structure of the connective tissue, the kidneys, and described the early stages of development of the human embryo. Wrote one of the most authoritative textbooks on physiology.

His students were G. Helmholtz and E. Dubois-Reymond. Helmholtz is known as a physicist, mathematician, physiologist and psychologist. His main works in the field of physiology are devoted to muscle contraction and sensory systems. He measured the duration of a single contraction, the speed of propagation of a nerve impulse, proposed the theory of tetanic contraction of skeletal muscles, the theory of accommodation of the eye, the resonant theory of hearing, and the doctrine of color vision.

Emile Dubois-Reymond investigated animal electricity, the presence of which he proved in the muscles, nerves, glands, skin, retina of the eye. He discovered the physical electrotone, formulated the first theory of the origin of bioelectric potentials (electromotive molecules), began electrophysiological research using an induction coil and electrodes.

Rudolf Heidenhain recorded the release of heat during single muscle contractions, established the role of the renal epithelium in the formation of urine, proposed the use of the isolated ventricle method to study gastric secretion, and proved that pepsin and perchloric acid are secreted by various cells of the gastric glands. He laid the foundations of knowledge about the secretory process, wrote a guide to physiology.

Karl Ludwig introduced graphic registration of processes with the help of a kymograph and the method of perfusion of isolated organs into physiology, proposed the filtration theory, discovered the secretory nerves of the salivary glands, and wrote a manual on human physiology.

The main scientific works of Ch. Bell devoted to the anatomy and physiology of the nervous system. He was the first to suggest (1811) that the anterior spinal roots are motor, and the posterior roots are sensitive. In 1822, this was experimentally confirmed by F. Magendie.

F. Magendie's scientific research concerns the physiology of the nervous system. He studied movements after removal of the cerebral hemispheres and cerebellum, demonstrated the trophic influence of the nervous system on organs and muscles, and proved the motor functions of the anterior and sensory functions of the posterior spinal roots.

For some time, Bernard worked in Magendie's laboratory, who studied the structure and functions of the glands of the gastrointestinal tract, the action of digestive juices, the metabolism of carbohydrates, and the vasoconstrictive functions of the sympathetic nerves. He is considered one of the founders of the doctrine of homeostasis.

The founder of the doctrine of the autonomic nervous system is J. Langley. He described the general plan of the structure of the autonomic nervous system, established the exit points of the autonomic nerve fibers from the central nervous system.

The English scientist Charles Sherrington made a great contribution to the development of the physiology of the central nervous system. He investigated the features of the conduction of excitation along the reflex arc, established the one-sidedness of the conduction and the presence of a synaptic delay. Introduced the concept of "synapse" and "neuron" into science. He discovered the phenomena of relief, convergence, occlusion, described decerebral rigidity, explained the development of spinal shock, studied inhibitions in the spinal cord. In 1932, he was awarded the Nobel Prize for these studies.

I. M. Sechenov is considered the father of Russian physiology. After graduating from Moscow University in 1856, in 1860 he improved his skills in the laboratories of C. Bernard, G. Helmholtz, C. Ludwig, E. Dubois-Reymond. Sechenov enriched science with facts and concepts of fundamental importance: he created the doctrine of blood gases, explained the respiratory function of blood, discovered carbhemoglobin, as well as the phenomena of summation of excitations and inhibitions in the central nervous system, formulated the central nervous theory of fatigue, introduced the concept of active rest, formulated the position that the activity of the brain is based on reflexes, substantiated the reflex nature of human mental activity. Sechenov lectured on electrophysiology, he is considered the founder of labor physiology. Sechenov's students were N. E. Vvedensky (1852-1922). B. F. Verigo (1860-1925), N. P. Kravkov (1865-1924), A. P. Samoilov (1867-1930), M. M. Shaternikov (1870-1939), V. V. Pashutin ( 1845-1901).

The Russian scientist N. E. Vvedensky worked in the field of physiology of excitable tissues. With the help of a signal amplifier, he studied excitation impulses in the nerve and muscle, discovered the phenomena of optimum and pessimum, formulated the concept of functional mobility, or lability, and analyzed the phenomenon of nerve fatigue.

The work of I. P. Pavlov, who in 1904 was awarded the Nobel Prize for his work in the field of digestion, had a particularly great influence on the development of physiology. Main directions scientific activity Pavlova - physiology of blood circulation, digestion and higher nervous activity. He created the doctrine of the trophic function of the nervous system, developed and improved the methods of surgical operations on the digestive organs, introduced a chronic experiment into physiology, discovered the secretory nerve of the stomach and pancreas, and also the new kind reflex reactions - conditioned reflexes, created the doctrine of the types of higher nervous activity, two signal systems and a dynamic stereotype, formulated the concepts of the analytical and synthetic activity of the cerebral cortex. Pavlov trained a large number of students, among whom were B. P. Babkin (1877-1950), L. A. Orbeli (1882-1958), K. M. Bykov (1886-1959).

The American physiologist W. Kennon entered the history of physiology as one of the founders of the theory of homeostasis and the sympathetic-adrenal system. He investigated the role of adrenaline as a mediator, found that during stimulation of sympathetic nerve fibers, sympathin is released - a substance similar to adrenaline, suggested the presence of two types of sympathin.

V. Yu. Chagovets began his scientific activity as a third-year student in the laboratory of I. R. Tarkhanov. In 1896 he published an article on the application of S. Arrhenius' theory of dissociation to electromotive phenomena in living tissues. So, he was the first to use the physicochemical approach to solving physiological problems and formulated the ionic theory of the origin of bioelectric potentials and the capacitor theory of excitation. Together with his students, he studied the electrogastrogram. V. V. Pravdich-Neminsky (1879-1952), A. I. Venchik, L. L. Gidzheu became his students.

In the 19th century physiology was enriched, moreover, by such new facts and discoveries. The German physiologist E. Pflüger (1859) formulated the patterns of stimulation by direct electric current, which were supplemented by B. F. Verigo. N. A. Mislavsky (1885) established the location of the respiratory center, and F. V. Ovsyannikov (1871) located the vasomotor center. AI Babukhin (1877) proved the ability of nerve fibers for bilateral conduction of excitation. IR Tarkhanov (1889) described the galvanic skin reflex. E. Marey designed a device for pneumatic recording of movements (Mare's capsule), and A. Mosso - a plethysmograph (for studying the blood filling of organs) and an ergograph (for studying fatigue). In 1836, the first reference books on physiology in Russia were published at the same time: in St. Petersburg - by D. M. Vellansky, in Moscow - by A. M. Filomafitsky.

Physiology in the 20th century

The development of human and animal physiology in the 20th century is characterized primarily by the attempts of physiologists to understand the processes of vital activity at the level of molecular interaction. Therefore, the concept of "life processes" acquire a clear and understandable content, cease to be mysterious and enigmatic. At the same time, physiologists do not confine themselves to studying the activity of individual organs, but study the functioning of integral organisms, find out the mechanisms for combining and coordinating the totality of life processes.

The directions of research started earlier are being further developed and new ones are being formed. There has been a quantitative increase in research and researchers. If at the end of the XIX century. the number of annual physiological publications worldwide does not exceed 700, then in the 70s of the XX century. it reached 60,000. Thus, the development of physiology in the 20th century should be considered in terms of areas of research.

The electrophysiology and physiology of excitable cells begin to develop especially rapidly. The German physiologist Julius Bernstein formulated in 1902-1912. membrane theory of bioelectric potentials, J. Loeb (1910) studied the effect of ions on the functional state of tissues. P. P. Lazarev (1923) developed the ion theory of excitation generation, A. Hodgkin and E. Huxley (1952) formulated the modern membrane theory of bioelectric potentials and excitation. Significant progress has been made in the field of electrophysiology of nerve cells. J. Erlanger and G. Gasser (1937) studied the conduction of nerve fibers, I. Tasaki (1957) substantiated the saltatory conduction of excitation, J. Eccles (1966) and B. Katz (1968) thoroughly studied the mechanisms of synaptic transmission of excitation. PG Kostyuk (1986) found out the functional role of Ca 2+ in the activity of neurons.

Recently, electrophysiological studies have focused on the study of the ion channels of the plasma membrane of various cells (B. Hille, 1975; E. Neer, B. Sakkman, 1987). The Nobel Prize winners were J. Erlanger and G. Gasser (1944), J. Eccles, A. Hodgkin, E. Huxley (1963), B. Katz (1970).

The study of the nervous system was not limited to the study of only electrophysiological methods at the cellular level. In 1912, V. V. Pravdich-Neminsky registered the electroencephalogram of a dog, and in 1929 G. Berger, an electroencephalogram of a person.

The study of the reflex function of the central nervous system was continued by IP Pavlov and C. Sherrington. Sherrington brought up a large galaxy of neurophysiologists, the most famous of which are R. Magnus, J. Eccles, R. Granit, V. Penfield and others.

A new direction in the study of the physiology of the central nervous system was started by the Dutch scientist R. Magnus, who discovered static and stato-kinetic reflexes (1924), with the participation of which the body's position in space is maintained.

In the 40s of the XX century. G. Magun, R. Rainis, J. Mruzzi investigated the functional role of the reticular formation in the regulation of excitability and tone of all parts of the central nervous system.

A great achievement of the physiology of the XX century. consider the emergence of the doctrine of mediators, which provide chemical transmission of impulses in synapses. The Austrian pharmacologist A. Levi (1921) became the founder of this doctrine. The chemical transmission of a nerve impulse was substantiated by A. P. Samoilov (1924), A. V. Kibyakov (1933), A. G. Ginetsinsky (1935).

In close connection with the physiology of the nervous system, the physiology of sensory systems also developed. One of the methods of physiological study of sensory systems was the method of conditioned reflexes, which was used to determine the sensitivity of the sense organs, the boundaries of the perception of stimuli, and the localization of sensory zones in the cortex. Electrophysiological studies of receptor cells were successfully carried out by E. Adrian (Nobel Prize, 1932). The discovery of the electroretinogram belongs to F. Gotch (1903). In 1930, E Weaver and C. Bray discovered the mic curl effect. G. Bekesy (Nobel Prize, 1961) electrophysiologically confirmed the resonator theory of hearing by G. Helmholtz.

Physiological studies of muscles have developed in several directions: excitability and excitation of muscle fibers, the relationship between excitation and contraction, the mechanism and energy of contraction. In 1907, V. Fletcher and F. Hopkins found that when a muscle contracts, lactic acid is formed in it. A. Hill and A. Meyerhof (Nobel Prize, 1922) came to the conclusion that lactic acid reacts with muscle and this leads to a change in their mechanical properties.

As early as 1930, E Lundsgaard found that by suppressing glycolysis with monoiodine acetate, the muscle can contract for some time, although lactic acid is not formed. It can contract as long as it contains creatine phosphate (discovered in 1927), the breakdown of which was considered as the initial reaction in the energetics of contraction. In 1929, K. Loman discovered ATP, which was recognized as a direct source of energy for contraction. A. Szent-Györgyi (1939-1946) proved that "muscle protein" consists of myosin and actin. In 1939, V. A. Engelgardt and M. M. Lyubimova established that ATPase activity is characteristic of myosin.

On the basis of electron microscopic and x-ray studies, E. Huxley (1957) proposed a theory of contraction, according to which it occurs due to sliding and convergence of actin and myosin protofibrils. This theory is being detailed and deepened even today. In 1965, A. Sandow elucidated the role of Ca 2+ in electromechanical communication.

At the end of XIX - early. 20th century made important discoveries in the physiology of blood circulation. In 1893, V. Gis described a bundle of muscle fibers of the heart, which was named after him. In 1906, S. Tavara discovered the atrioventricular node, and soon A. Kos and M. Fleck described the sinoatrial node. Electrocardiography dates back to 1903, when W. Einthoven (Nobel Prize, 1924) standardized the conditions for recording electrocardiograms. A. P. Samoilov made a significant contribution to the theory and practice of electrocardiography. In 1914, E Starling concluded that the mechanical strength of the heart muscle depends on the length of its fibers.

In the 20s of the XX century. K. Wiggers divided the cardiac cycle into separate phases: systole and diastole. German scientists N. Goering (1924) and K. Heimans (Nobel Prize, 1939) found out the role of mechano- and chemoreceptors of reflexogenic zones in the regulation of the heart and vascular tone. A. Krog (Nobel Prize, 1920) proved that the number of functioning capillaries increases during the activity of skeletal muscles.

Research on respiration was aimed mainly at elucidating the mechanisms of its regulation and the transport of gases by the blood. Chemoreceptors of the carotid body, the irritation of which causes a change in respiration, was discovered by K. Geymans (1928). The pneumotaxic center was discovered by T. Lumsden (1923), and the fact that gas exchange in the lungs is carried out by diffusion was established by A. Krogh (1910) and J. Barcroft (1914).

At the beginning of the XX century. Digestion studies were carried out by students of I. P. Pavlov (B. P. Babkin, L. A. Orbeli, I. P. Razenkov, K. M. Bykov). In 1902 V. Beilis and E. Starling discovered secretin, in 1906 D. Adkins discovered gastrin, and in 1943 A. Harper and H. Reiper discovered pancreozymin. In 1958, A. M. Ugolev (1926-1991) discovered membrane digestion.

A significant contribution to the physiology of digestion was made by P. G. Bogach (1918-1981). who studied the central and peripheral mechanisms of regulation of the activity of the digestive organs, the electrophysiological properties of smooth muscle and secretory cells of the digestive tract, discovered hypothalamic mechanisms for regulating bile formation and bile secretion. The electrical connection between the smooth muscle cells of the gastrointestinal tract was discovered by M.F. Shuba (1928-2007). He also investigated the ionic nature of their resting membrane potential and electrical activity, the ionic mechanisms of the action of acetylcholine and norepinephrine on them as mediators of the autonomic nervous system.

In 1917, A. Keshni proposed the filtration-reabsorption theory of urine formation, which was further developed by A. Richards, G. Smith and others. And also the 20th century. characterized by the discovery of hormones (I. Takamine and T. Aldrich, 1901) and vitamins (K. Funk, 1912). These discoveries were of great importance for medicine and veterinary medicine.

Conclusion

In the development of physiology today, its further differentiation and specialization (cosmic physiology, neurophysiology), the use of accurate quantitative research methods at all levels of the organization of the living, using computer technology, theory, and automatic control are observed. The analytical approach to the study of the vital activity of organisms is combined with a synthetic approach, which makes it possible to determine the functional integrity of organisms, the spatio-temporal organization of physiological processes, and complex acts of human and animal behavior.

Already in ancient times, elementary ideas about the activities of the human body were formulated. Hippocrates (460-377 BC) represented the human body as a unity of liquid media and the mental make-up of a person. In the Middle Ages, ideas based on the postulates of the Roman anatomist Galen dominated.

The official date of the emergence of physiology can be considered 1628, when the English physician, anatomist and physiologist William Harvey published his treatise Anatomical Study of the Movement of the Heart and Blood in Animals. In it, he first presented experimental data on the presence of large and small circles of blood circulation, as well as on the effect of the heart on blood circulation.

In the 17th century scientists conducted a number of studies on the physiology of muscles, respiration, and metabolism. But the experimental data obtained were explained at that time from the positions of anatomy, chemistry and physics.

In the XVIII century. the doctrine of "animal electricity", discovered by the Italian scientist L. Galvani, arose. The principle of reflex activity is further developed (I. Prohaska, 1749-1820).

First tutorial in Physiology was published by the German scientist A. Haller in the middle of the 18th century.

Physiological science received further development in the 19th century. This period is associated with advances in organic chemistry (F. Weller synthesized urea); in histology - by the discovery of the cell (T. Schwann); in physiology - the creation of a reflex theory of nervous activity (I.M. Sechenov).

An important milestone in the development of experimental physiology was the invention of the kymograph and the development of a method for graphic recording of blood pressure by the German scientist K. Ludwig in 1847.

A significant contribution to many areas of physiology during this period was made by the famous French scientist C. Bernard (1813-1878). His research concerned the functions of the spinal cord, carbohydrate metabolism, the activity of digestive enzymes, and the role of endocrine glands.

Interesting discoveries in the field of physiology in the middle and end of the 19th century. were made in the field of regulation of the activity of the heart and blood vessels [K. Ludwig (1816-1895), I.F. Zion (1842-1912), C. Bernard (1813-1878), F.W. Ovsyanikov (1827-1906)].

In the second half of the 19th and early 20th centuries Physiological research has also received significant development in Russia thanks to the research of I.M. Sechenov (1829-1905), I.P. Pavlov (1849-1936) and other Russian scientists.

An important merit in physiology belongs to I.M. Sechenov, who first discovered the presence of inhibition processes in the central nervous system and, on the basis of this, created the doctrine of the reflex activity of the body. His work "Reflexes of the brain" served as the basis for the formation of the doctrine of nervism. In this work, he suggested that the various manifestations of human mental activity ultimately come down to muscle movement. Ideas I.M. Sechenov was later successfully developed by the famous Russian physiologist I.P. Pavlov.

On the basis of an objective study of behavioral reactions, he created a new direction in science - the physiology of higher nervous activity. The teachings of I.P. Pavlov on the higher nervous activity of man and animals made it possible to deepen the theory of the reflex activity of the brain.

In addition, he made many other discoveries in physiology. He discovered the presence of a sympathetic nerve that enhances the contraction of the heart (1881). Created the doctrine of the trophic influence of the nervous system (1920). For many years he studied the physiology of digestion and developed methods for imposing a permanent fistula of the pancreas, forming an isolated ventricle, determined the main patterns of the secretory activity of the digestive glands, the role of sympathetic and parasympathetic nerves in the reflex regulation of this activity. I.P. Pavlov published two major works: "Lectures on the work of the main digestive glands" (1897) and "Physiological surgery of the digestive tract" (1902), which were of great importance in the development of world physiology. For research in the field of physiology of digestion, Academician I.P. Pavlov received the Nobel Prize in 1904.

I.P. Pavlov founded the school of Russian physiologists, which made a great contribution to world science. His students were Academicians P.K. Anokhin, K.M. Bykov, L.A. Orbeli and many other scientists.

Academician N.E. Vvedensky (1884-1886).

The works of A.A. Ukhtomsky. He formulated the principle of dominance.

Academician K.M. Bykov carried out various studies on the role of the cerebral cortex in the activity of internal organs.

L.A. Orbeli developed the teachings of I.P. Pavlova on the trophic influence of the nervous system.

In the 30s of the XX century. the chemical mechanism of nerve impulse transmission in synapses was proved (O. Levy and G. Dale).

The development of the membrane theory of bioelectric potentials in living cells (A.L. Hodgkin, E.F. Huxley, B. Katz) was of great importance.

The twentieth century was rich in discoveries in the field of endocrine glands and the physiology of digestion. For example, A.M. Ugolev (1926-1992) discovered membrane intestinal digestion.

Designed by I.M. Sechenov and I.P. Pavlov, the principles and methods of physiological research formed the basis for the development of the physiology of farm animals. Under the editorship of A.V. Leontovich in Russia in 1916, the first domestic textbook, “Physiology of Domestic Animals,” was published. Professors A.V. Leontovich and K.R. Viktorov conducted in-depth research in the field of digestion in birds.

Research in the field of physiology of lactation in animals was carried out by Professor G.I. Asimov and his school.

A significant contribution to the field of studying the physiology of digestion in animals was made by the studies of N.V. Kurilova, A.D. Sineshchekova, V.I. Georgievsky, A.A. Kudryavtsev.

Domestic researchers made a great contribution to the study of metabolism in animals: A.A. Aliev, N.A. Shmanenkov, D.K. Kalnitsky, N.S. Shevelev and many others.

Significant progress in the physiology of excretion in animals was made by V.F. Lysov, A.I. Kuznetsov, and in the physiology of the endocrine glands - V.I. Maksimov, V.P. Radchenkov and many other scientists.

Significant results in the field of physiology of reproduction of domestic animals were achieved by domestic scientists I.I. Ivanov, V.K. Milovanov, A.I. Lopyrin.

Research in the field of animal physiology continues at the present time in various educational and research organizations.

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Physiology (from the Greek physis - nature, logos - teaching) is a science that studies the patterns of functioning of animal organisms, their individual systems, organs, tissues and cells. The totality of physiological knowledge is subdivided into a number of separate but interrelated areas - general, particular and applied physiology. General physiology includes information regarding the nature of the main life processes, general manifestations of vital activity, such as the metabolism of organs and tissues, general patterns of the response of the body and its structures to environmental influences - irritability. This also includes features due to the level of structural organization, different conditions of existence. Consequently, general physiology describes those qualitatively unique phenomena that distinguish the living from the non-living. Particular physiology studies the properties of individual tissues and organs, the patterns of combining them into systems, as well as the physiology of individual classes, groups and species of animals. Applied physiology studies the patterns of manifestations of the activity of an organism, especially a person, in connection with special tasks and conditions. These sections include the physiology of labor, sports, nutrition, environmental physiology. Physiology is also conventionally divided into normal and pathological. The emergence of physiology occurred in antiquity in connection with the needs of medicine, the best representatives of which clearly understood that it was possible to help the patient only by knowing about the structure of the body. Hippocrates, the father of medicine, laid the foundations for understanding the role of individual systems and functions of the body as a whole. Similar views were held by another famous physician of antiquity, the Roman anatomist Galen, who for the first time in history introduced an experiment into the practice of medicine. His experiments formed the basis for theories that lasted almost 14 centuries without any significant changes. The origin of physiology as a science that studies the processes occurring in the body and combines them on the basis of observations and experiments, refers mainly to the second half of the 16th - the beginning of the 18th century. At the same time, the anatomist Andreas Vesalius was the first to correctly describe the structural features of the human body, and also created the first manual on animals. The most important stage in the development of physiology is considered to be 1628, when the English physician and physiologist William Harvey published his immortal book Anatomical Studies on the Movement of the Heart and Blood in Animals, in which he outlined the foundations of his great discovery - the existence circulation. The discovery of blood circulation became possible due to the fact that Harvey introduced a new technique into the practice of scientific research - vivisection or vivisection. This technique provides for the exposure of the covers and tissues of certain organs of animals through certain incisions, which creates the possibility of direct observation of the work of these organs. In addition, experiments were carried out using various influences on the process under study. The correctness of ideas about the presence of a closed circulatory system was confirmed by the Italian biologist Marcello Malpighi (1628-1694). He owns the discovery of blood cells, the alveolar structure of the lungs, as well as the connection of arteries with veins through capillaries. Among the most important achievements of the XVII-XVIII centuries. refers to the idea formulated by the French philosopher, mathematician, physicist and physiologist Rene Descartes about the "reflected activity of the body." Descartes, using such facts as blinking naturally occurring when touching the cornea, put forward the concept of reflex. By the first half of the XVIII century. refers to the beginning of the development of physiology in Russia. I. M. Sechenov entered the history of science as the "father of Russian physiology", a thinker who for the first time dared to subject to experimental analysis the most complex area of ​​nature - the phenomenon consciousness. The scientific activity of I. M. Sechenov consisted of several stages. He was the first who succeeded in extracting and analyzing the gases dissolved in the blood, establishing the relative effectiveness of the influence of various ions on the physicochemical processes in a living organism, and discovering the phenomenon of summation in the central nervous system. He also became the founder of a new direction in physiology - labor physiology. The greatest glory to Russian science was brought by the discovery by I. M. Sechenov (1862) inhibition in the central nervous system. The development of domestic and world physiology was greatly influenced by the works of I.P. Pavlov, an outstanding representative of natural science, the creator of the doctrine of higher nervous activity animals and humans. Pavlov established the existence of special nerves, some of which enhance, others delay the work of the heart, and others are able to change the strength of heart contractions without changing their frequency. IP Pavlov explained this phenomenon by the property of these nerves to change the functional state of the cardiac muscles, reducing its trophism. Thus the foundation was laid theories about trophic innervation of tissues. Simultaneously with the study of the cardiovascular system, IP Pavlov investigated the physiology of digestion. By developing and applying a number of subtle surgical techniques, he essentially re-created the physiology of digestion. Studying the dynamics of the secretory process of the gastric, pancreatic and salivary glands, the work of the liver when eating different foods, IP Pavlov showed their ability to adapt to the nature of excitatory secretion. These works were based on the idea nervism, by which I. P. Pavlov understood “a physiological direction that seeks to extend the influence of the nervous system to as many activities of the body as possible. At the beginning of the 20th century, V. M. Bekhterev established the role of subcortical structures in the formation of emotional and motor reactions animals and humans; the nuclei and pathways of the brain are open; the functional-anatomical basis of balance and orientation in space was revealed; thalamus functions; centers of movement and secretion of internal organs were identified in the cerebral cortex; it has been proved that the motor fields of the cerebral cortex are the basis of individually acquired movements. Freud formulated the idea of the prevailing importance of instincts, the dominant value of unconscious mental processes. A. A. Ukhtomsky formulated the leading principle of the brain - dominant revealed its characteristic features - an increase in excitability in the dominant center, the stability of this excitation over time, the possibility of its summation, the inertia of excitation and inhibition of other reflex mechanisms that are not involved in the dominant reaction. Currently, the dominant is recognized as one of the main mechanisms of brain activity. In the current century, a great contribution has been made to the study functional relationships between the cerebral cortex and internal organs. K. M. Bykov, studying the regulatory influence of the cerebral cortex on the work of internal organs, showed the possibility of changing their activity by a conditioned reflex. Thanks to the study by V. N. Chernigovsky of the problems of sensitivity of internal organs, relationships with the cerebral cortex, as well as the determination of projections of afferent systems of internal organs in the cerebral cortex, thalamus, cerebellum, reticular formation, a detailed study of the unconditional reflex activity of these organs during stimulation of interoceptors by mechanical, chemical and other agents opened a new chapter of physiology - interoception.

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Subject, tasks of age physiology and its connection with other sciences

Age physiology is a science that studies the features of the life process of an organism at different stages of ontogenesis.

It is an independent branch of human and animal physiology, the subject of which is the study of the patterns of formation and development of the physiological functions of the body throughout its life path from fertilization to the end of life.

Depending on what age period the age-related physiology studies, there are: age-related neurophysiology, age-related endocrinology, age-related physiology of muscle activity and motor function; age-related physiology of metabolic processes, cardiovascular and respiratory systems, the digestive and excretory systems, the physiology of embryonic development, the physiology of infants, the physiology of children and adolescents, the physiology of adulthood, gerontology (the science of aging).

The main objectives of the study of age physiology are as follows:

The study of the features of the functioning of various organs, systems and the body as a whole;

Identification of exogenous and endogenous factors that determine the features of the functioning of the body in different age periods;

Determination of objective age criteria (age standards);

Establishment of patterns of individual development.

Developmental physiology is closely related to many branches of physiological science and makes extensive use of data from many other biological sciences. Thus, in order to understand the patterns of formation of functions in the process of individual development of a person, data from such physiological sciences as cell physiology, comparative and evolutionary physiology, the physiology of individual organs and systems: the heart, liver, kidneys, blood, respiration, nervous system, etc. are needed.

At the same time, the patterns and laws discovered by age physiology are based on data from various biological sciences: embryology, genetics, anatomy, cytology, histology, biophysics, biochemistry, etc. Finally, age physiology data, in turn, can be used to develop various scientific disciplines. For example, age physiology is of great importance for the development of pediatrics, pediatric traumatology and surgery, anthropology and gerontology, hygiene, developmental psychology and pedagogy.

History and main stages of development of age physiology

The scientific study of the age characteristics of the child's body began relatively recently - in the second half of the 19th century. Shortly after the discovery of the law of conservation of energy, physiologists discovered that a child consumes slightly less energy during the day than an adult, although the size of a child's body is much smaller. This fact required a rational explanation. In search of this explanation, the German physiologist Max Rubner studied the rate of energy metabolism in dogs of different sizes and found that larger animals consume much less energy per 1 kg of body weight than smaller ones. Having calculated the surface area of ​​the body, Rubner made sure that the ratio of the amount of energy consumed is proportional to the size of the body surface - and this is not surprising: after all, all the energy consumed by the body must be released into the environment in the form of heat, i.e. the energy flux depends on the heat transfer surface. Rubner explained the difference in the intensity of energy metabolism between large and small animals, and at the same time between adults and children, precisely by differences in the ratio of mass and surface of the body. Rubner's "surface rule" was one of the first fundamental generalizations in developmental and environmental physiology.

This rule explained not only the differences in the magnitude of heat production, but also in the frequency of heart contractions and respiratory cycles, pulmonary ventilation and volume of blood flow, as well as in other indicators of the activity of autonomic functions. In all these cases, the intensity of physiological processes in a child's body is significantly higher than in an adult's body.

Such a purely quantitative approach is characteristic of the German physiological school of the 19th century, consecrated by the names of prominent physiologists E.F. Pfluger, G.L. Helmholtz and others. Through their labors, physiology was raised to the level of the natural sciences, standing on a par with physics and chemistry. However, the Russian physiological school, although rooted in the German one, has always been distinguished by an increased interest in qualitative features and regularities.

An outstanding representative of the Russian pediatric school, Dr. Nikolai Petrovich Gundobin, at the very beginning of the 20th century.

argued that a child is not just small, he is also in many ways not the same as an adult. His body is arranged and works differently, and at each stage of its development, the child's body is perfectly adapted to the specific conditions that it has to face in real life.

These ideas were shared and developed by the remarkable Russian physiologist, teacher and hygienist Pyotr Frantsevich Lesgaft, who laid the foundations for school hygiene and physical education for children and adolescents. He considered it necessary to deeply study the child's body, its physiological capabilities.

The central problem of developmental physiology was most clearly formulated in the 20s of the XX century. German physician and physiologist E. Helmreich. He argued that the differences between an adult and a child lie on two planes, which must be considered as independently as possible, as two independent aspects: the child as a small organism and the child as a developing organism. In this sense, Rubner's "surface rule" considers the child in only one aspect - namely, as a small organism. Much more interesting are those features of the child that characterize him as a developing organism.

One of these fundamental features is the uneven development of the sympathetic and parasympathetic influences of the nervous system on all the most important functions of the child's body, discovered by Ilya Arkadyevich Arshavsky at the end of the 1930s. I.A. Arshavsky proved that sympathotonic mechanisms mature much earlier, and this creates an important qualitative originality of the functional state of the child's body. The sympathetic division of the autonomic nervous system stimulates the activity of the cardiovascular and respiratory systems, as well as metabolic processes in the body.

Such stimulation is quite adequate for an early age, when the body needs an increased intensity of metabolic processes necessary to ensure the processes of growth and development. As the child's body matures, parasympathetic, inhibitory influences intensify.

Chapter 1. History of Physiology. Methods of physiological research

As a result, the pulse rate, respiratory rate, and the relative intensity of energy production decrease.

The problem of uneven heterochrony (time difference) in the development of organs and systems has become the central object of research by the outstanding physiologist Academician Pyotr Kuzmich Anokhin and his scientific school.

In the 1940s, he formulated the concept of systemogenesis, according to which the sequence of events unfolding in the body is built in such a way as to satisfy the changing needs of the body in the course of development. At the same time, P.K. Anokhin for the first time moved from consideration of anatomically integral systems to the study and analysis of functional relationships in the body.

Another outstanding physiologist, Nikolai Aleksandrovich Bernshtein, showed how algorithms for controlling voluntary movements are gradually formed and become more complex in ontogenesis, how the mechanisms of higher movement control spread with age from the most evolutionarily ancient subcortical structures of the brain to newer ones, reaching an ever higher level of “building movements”. In the works of N.A. Bernshtein, it was shown for the first time that the direction of ontogenetic progress in the control of physiological functions clearly coincides with the direction of phylogenetic progress. Thus, on the basis of physiological material, the concept of E. Haeckel and A.N. Severtsov that individual development (ontogeny) is an accelerated evolutionary development (phylogenesis).

Academician Ivan Ivanovich Shmalgauzen, a prominent specialist in the field of the theory of evolution, also dealt with ontogeny for many years. The material on which I.I. Shmalgauzen drew his conclusions rarely had a direct bearing on the physiology of development, but the conclusions from his works on the alternation of stages of growth and differentiation, as well as methodological work in the field of studying the dynamics of growth processes, carried out in the 30s , and are still of great importance for understanding the most important patterns of age-related development.

In the 1960s, the physiologist Akop Artashesovich Markosyan put forward the concept of biological reliability as one of the factors of ontogeny. She relied on numerous facts that testified that the reliability of functional systems increases significantly as the body grows older. This was confirmed by data on the development of the blood coagulation system, immunity, and the functional organization of brain activity.

In recent decades, many new facts have accumulated that confirm the main provisions of A.A. Markosyan's concept of biological reliability.

At the present stage of development of biomedical science, research in the field of age physiology is also continuing, already using modern methods research.

Thus, physiological science currently has at its disposal considerable multifaceted information concerning the functional activity of any physiological system of the child's organism and its activity as a whole.

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Main article: History of physiology

In Russia, physiology began to develop in the 18th century. From the very beginning, Russian physiology showed the greatest interest in the study of the physiology of the nervous system.

Efrem Osipovich Mukhin (1766-1850), professor of anatomy and physiology at the Medical-Surgical Academy of Moscow University, can be considered the founder of the physiology of the nervous system.

In the 19th century In Russia, a brilliant group of physiologists came to the fore, among whom I. M. Sechenov stood out in particular. Almost simultaneously with Sechenov, or a little later, V. Ya. Danilevsky worked in Kharkov and I. A. Mislavsky in Kazan.

Formulated by Russian physiology, starting from Mukhin, then by Sechenov, Pavlov and others, the reflex theory also includes the activity of the cerebral cortex. This does not leave room for the assumption that any functions of the cortex can occur spontaneously, without stimuli from outside or inside.

Mukhin E. O.

In 1800, E. O. Mukhin defended his dissertation on stimuli that excite the human body, and received a doctorate in medicine and surgery. The main direction of all his scientific activity was the study of the function of the nervous system, the elucidation of the meaning of irritations that cause actions and determine all the phenomena of life. He believed that external and internal factors serve as stimuli, that all body functions are determined. At the same time, he pointed out that the state of the organism, its reactivity, also matters. Irritations, in his opinion, can lead both to actions and to the cessation of actions (that is, to inhibition), a struggle between stimuli can occur in the body, moreover, a stronger irritation overcomes a weaker one; he considered the brain to be the first place of sensations; excitation, he pointed out, quickly spreads through the nerves of the whole body, like electric current; the transition of excitation from one half of the body to the other occurs in the medulla oblongata, in the pons of Varoli, in the commissure of the hemispheres. Mukhin insisted that the work of the nervous system makes the body integral and that, thanks to its ability to respond to changes in the external environment, it merges with it together.

The high merits of this outstanding and undeservedly half-forgotten Russian physiologist are evident from the fact that even now, after a century and a half, we can hardly change anything in the list of his statements, he penetrated so deeply into the functions of the nervous system even when there was not even good research methodology.

Sechenov I. M.

Of greatest importance are the works of Ivan Mikhailovich Sechenov, who is rightly considered the founder of Russian physiology. He was a versatile scientist. He carried out research on the physiology of blood and developed a method for obtaining gases from the blood. IM Sechenov worked extensively on the physiology of respiration and metabolism.

BRIEF HISTORY OF THE DEVELOPMENT OF PHYSIOLOGY

However, his most important work concerns the physiology of the nervous system, where he made classical discoveries on the question of inhibition in the nervous system and on the functions of the cerebral cortex. Working a lot and fruitfully on the mechanism of reflexes, their ways and summation of excitation and the brain, he came to the conclusion about the predominant role of the cerebral cortex in the nervous system of higher animals. The cerebral cortex receives stimuli from all parts of the body and sends excitations to them. Sechenov developed the most important thesis in the physiology of the cerebral cortex, which consists in recognizing that reflex mechanisms underlie the activity of the cortex.

Danilevsky V. Ya.

Danilevsky was interested in electrophysiology, discovered electric currents in the cerebral cortex, studied the muscular system and metabolism in it.

Mislavsky I. A.

Mislavsky studied the cerebral cortex a lot, observing the effects of direct stimulation of it at different points. But his most important merit was the discovery of the location of the respiratory center with its exact localization in the medulla oblongata. Mislavsky's school also studied the innervation of the glands, especially the endocrine glands.

Vvedensky I. E.

In the end. 19th century in Russian physiology, a prominent place was occupied by I. E. Vvedensky (Petersburg), who worked on general issues arousal. Studying the phenomena of nerve death on a neuromuscular preparation, he discovered the patterns of changing the process of excitation by the process of inhibition, known as parabiosis. It is remarkable that the regularities established by him are applicable to all manifestations of excitation in the nervous system and in other excitable formations. Material from the site http://wiki-med.com

Pavlov I.P.

From the end of the 19th century The development of physiology in Russia is associated, first of all, with the activities of the outstanding researcher and versatile experimenter Ivan Petrovich Pavlov (St. Petersburg). His outstanding work focused on two large areas of physiology. This is the study of the process of digestion, where Pavlov gave a wonderful technique for imposing fistulas on different parts of the digestive canal, which allowed him to directly observe the processes in deep-lying organs. He developed this area of ​​physiology with such perfection that he received the Nobel Prize for these works.

Studying the processes of digestion, IP Pavlov paid special attention to the role of the nervous system in general and the cerebral cortex in particular in these processes. In connection with this, Pavlov developed the doctrine of conditioned reflexes, which later became the main direction of his scientific activity. Using conditioned reflexes, Pavlov was able to penetrate into the most intimate physiological processes in the cerebral cortex. The development of these questions continues even now with great success.

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The formation of physiology as a science

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The birth of physiology as a science is associated with the name of the outstanding English physician, physiologist and embryologist William Harvey. (Harvey, Wiliiam, 1578-1657) (Fig. 90), who is credited with creating a coherent theory of blood circulation.

At the age of 21, W. Harvey graduated from the University of Cambridge. At the age of 24 he became a doctor of medicine in Padua. Returning to his homeland, Harvey became a professor of anatomy, physiology and surgery in London.

Based on the achievements of his predecessors - Galen, Vesalius, Colombo, Fabricius - Harvey mathematically calculated and experimentally substantiated the theory of blood circulation, according to which blood returns to the heart in small and large circles. Due to the fact that during the life of Harvey, a microscope was not yet used in physiology, he could not see the capillaries - they were discovered by Marcello Malpighi (Malpighi, Marcello, 1628-1694) four years after Harvey's death. According to Harvey, blood passed from arteries to veins through anastomoses and through tissue pores.

After many years of testing in the experiment, W. Harvey outlined his theory in the fundamental work “Anatomical Study of the Movement of the Heart and Blood in Animals” (“Exercitatio anatomica de motu cordis et sangvinis in animalibus”, 1628) and was immediately subjected to fierce attacks from the church and many scientists. R. Descartes was the first to recognize Harvey's theory, then G. Galileo, S. Santorio, A. Borelli. I.P. Pavlov defined it as not only “the fruit of his mind of rare value, but also a feat of his courage and selflessness.”

The activities of the outstanding English philosopher Francis Bacon (Bacon, Francis, 1561-1626) had a great influence on the development of natural science (and physiology in particular). Not being a doctor, Bacon largely determined the paths for the further development of medicine. In his work "On the Dignity and Improvement of the Sciences", he formulated three main tasks of medicine: "the first is to maintain health, the second is to cure diseases, the third is to continue life." Being engaged in experimental work in the field of physiology, Bacon posed several specific questions for medicine: the study of the anatomy of not only a healthy, but also a diseased organism, the introduction of anesthesia, the use of natural factors in the treatment of diseases and the development of balneology. The solution of these and many other problems put forward by F. Bacon took centuries.

A contemporary of Francis Bacon, the outstanding French scientist Rene Descartes (Descartes, Rene, 1596-1650) developed the reflex arc scheme in its simplest form. He divided all the nerves into centripetal, through which signals enter the brain, and centrifugal, through which signals from the brain move to the organs. Descartes believed that vital actions are of a reflex nature and obey mechanical laws.

R. Descartes was a typical representative iatrophysics - a direction in natural science and medicine, which considered wildlife from the standpoint of physics. Compared with medieval scholasticism, the metaphysical thinking of the 17th century. was a progressive phenomenon, and the mechanistic views of Descartes had a positive impact on further development philosophy and natural science in the era of modern times. However, along with the materialistic understanding of the world, Descartes interpreted phenomena idealistically in a number of issues. So, he believed that thinking is a faculty of the soul, not of the body.

Another direction in natural science was iatromechanics. Its main provisions are clearly stated in the essay “On the Movement of Animals” (Fig.

History of development of physiology.

91) Italian anatomist and physiologist Giovanni Alfonso Borelli (Borelli, Giovanni Alfonso, 1608-1679), one of the founders of biomechanics. From the standpoint of iatromechanics, a living organism is like a machine in which all processes can be explained using mathematics and mechanics.

Among the outstanding achievements of the Renaissance, which were related to both physics and medicine, is the invention at the end of the 16th century. thermometer (more precisely, an air thermoscope). Its author is one of the titans of the Renaissance, the Italian scientist Galileo Galilei (Galilei, Galileo, 1564-1642), who confirmed and developed the heliocentric theory of N. Copernicus (1543). Many of his precious manuscripts were burned by the Inquisition. But in those that have survived, they found: drawings of the first thermoscope. Unlike a modern thermometer, it expanded air, not mercury. Almost simultaneously with Galileo, a professor at the University of Padua Santorio (Santorius, 1561-1636), a doctor, anatomist and physiologist, created his own device, with which he measured the heat of the human body (Fig. 92). The device was quite bulky. Santorio installed it in his backyard for all to see. The heat of various parts of the body was determined during ten pulse strokes by changing the level of the liquid in the tube, the scale of which was arbitrary.

At the beginning of the XVII century. many original thermometers were made in Europe. The first thermometer, the readings of which did not depend on changes in atmospheric pressure, was created in 1641 at the court of Ferdinand II, Emperor of the Holy Roman Empire, who was not only known as the patron of the arts, but also was the author of a number of physical instruments. With his participation, thermometers, funny in form, similar to small frogs, were created. They were intended to measure the warmth of the human body and were easily attached to the skin with a plaster. The cavity of the “frogs” was filled with a liquid in which colored balls of various densities floated. As the liquid warmed up, its volume increased and its density decreased, and some of the balls sank to the bottom of the apparatus. The patient's body heat was determined according to the number of multi-colored balls remaining on the surface: the fewer of them, the higher the body heat of the test subject.

The development of a single scale of degrees stretched over a century. The last word on this issue belongs to the Swedish astronomer and physicist Anders Celsius (Celsius, Anders, 1701-1744), who in 1742 proposed a centigrade scale: for 0 ° he took the boiling point of water, and the melting point of ice corresponded to 100 °. Subsequently, this scale was inverted, making 0 ° the point of melting ice and the starting point. In this form, the Celsius scale has reached our days, having won the widest popularity.

In medical practice, thermometry began to be used much later - only in the second half of the 19th century. The active introduction of this method in Russia in 1860 is associated with the name of the outstanding Russian clinician S. P. Botkin (see p. 270).

Iatrochemistry and medicine

Along with iatrophysics and iatromechanics, iatrochemistry, a direction in medicine associated with the advances in chemistry, was widely developed during the Renaissance. Iatrochemists believed that the processes occurring in the body are chemical, therefore, both the study of these processes and the treatment of diseases should be associated with chemistry.

One of the founders of iatrochemistry is the outstanding physician and chemist of the early Renaissance, Philippus Aureolus Theophrastus Bombastus von - Paracelsus, 1493-1541. Swiss by birth, he was educated at the University of Ferrara (Italy) and subsequently lectured at the University of Basel in his native German instead of the accepted Latin in the scientific world.

Paracelsus was one of the founders of the experimental method in science. The doctor's theory is experience. No one can become a doctor without science and experience,” he argued.

At the time of Paracelsus, surgery in Europe was not considered a field of medicine and was not taught at universities (artisans were engaged in it), and Paracelsus insisted on combining surgery and medicine (i.e. therapy) into one science, because both of them come from the same root. He himself proudly called himself "doctor of both medicines." His books "Small Surgery" ("Chirurgia minor", 1528), "Great Surgery" ("Chirurgia magna", 1536) and others were very popular (Fig. 93).

With Paracelsus begins a radical restructuring of chemistry in its application to medicine: from the search for ways to obtain gold - to the preparation of medicines. According to Paracelsus, health is associated with the normal content of three principles in the human body: sulfur, mercury and salt; violation of their correct ratios leads to illness. That is why doctors and pharmacists of the Renaissance attached great importance to medicines containing sulfur, mercury and various salts, and often smelted them themselves from natural ores. Paracelsus proudly wrote that he and his students "have a rest in the laboratory, stick their fingers into coals and garbage and all sorts of dirt, and not into golden rings, and are like blacksmiths and smoked coal miners."

In his writings, he also wrote about the diseases of miners and foundry workers associated with poisoning with sulfur, lead, mercury, antimony, and thus laid the foundations for the future science of occupational diseases. Georg Bauer, a contemporary of Paracelsus, also known under the pseudonym Agracola (Agricola, Georg, 1493-1541), also wrote about the diseases of miners and their prevention in his essay “On Mining and Metallurgy” (“De re metallica.”, 1556).

The development of medicinal chemistry during the Renaissance led to the expansion of the pharmaceutical business. The pharmacy as an independent institution arose in the second half of the 8th century. in the Middle East. (The first pharmacy in the Near and Middle East was opened in 754 in the capital of the Caliphate - Baghdad.) In Europe, the first pharmacies appeared in the 11th century. in the Spanish cities of Toledo and Cordoba. By the 15th century they spread widely across the continent.

During the Renaissance, the size of pharmacy shops increased significantly: from simple shops of the developed Middle Ages, when the entire pharmacy was located in one room, they turned into large pharmaceutical laboratories, which included a room for receiving visitors, pantries where medicines and raw materials were crushed and stored , and the actual laboratory with an oven and a distillation apparatus (Fig. 94).

Starting from the XV century. with special diligence cultivated pharmaceutical botanical gardens; they were also called health gardens - Hortus sanitatis. From this Latin name came the Russian - Vertograd (i.e. garden, flower garden). In the XVI-XVII centuries. Vertograds spread widely in Russia. Mineral substances and parts of animals were also used as medicinal raw materials. Overseas travels were of great importance, from which foreign medicines were brought.

Ideas about the therapeutic effect of many medicines at that time were often far from the truth. So, for almost two millennia (from the 1st to the 20th century), there was an opinion that theriac is a universal remedy against all diseases. It was composed by the doctors themselves with a large crowd of people from more than 70 components, and then kept for six months: the teriac prepared in Venice was especially famous.

Renaissance pharmacists, like other professionals, made a great contribution to the formation of the culture of their time. They occupied a high position in society, but their activities were regulated by the state. In the middle of the XVI century. the first pharmacopeias began to appear, listing the medicines used in a given city or state, their composition, use and cost. This was the beginning of the official regulation of prices for medicines in Europe.

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Ticket 4. The role of domestic scientists in the development of physiology.

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The first Russian physiologist and doctor of medical sciences was one of the outstanding associates of Peter I P.

The formation of physiology as a science. History of development of physiology.

V. Posnikov (born in 1676). P. V. Posnikov set himself the task of experimentally studying the cause of death.

The famous Russian scientist M. V. Lomonosov (1711-1765) did a lot for the development of physiology. He not only for the first time formulated the law of conservation of matter and the transformation of energy, but also developed the scientific foundations of the oxidation process. Later, his conclusions were confirmed by the French chemist Lavoisier, who discovered oxygen. The ideas of M. V. Lomonosov were later taken as the basis of the doctrine of respiration. M. V. Lomonosov was the first to formulate a three-component theory of color vision, gave a classification of taste sensations, and suggested that the body is a source of heat generation.

The founder of experimental physiology is Professor of Moscow University A. M. Filomafitsky (1802-1849), who studied issues related to the physiology of respiration, blood transfusion, and the use of anesthesia. A. M. Filomafitsky wrote the first Russian textbook on physiology:

The beginning of the operative-surgical method of studying the processes of digestion was laid by the surgeon V. A. Basov. A great contribution to the development of Russian physiology was also made by A. T. Babukhin, who established the bilateral conduction of excitation along the nerve fiber, V. F. Ovsyannikov, who described the vasomotor center in the medulla oblongata, N. A. Mislavsky, who studied the peculiarities of the location of the respiratory center, V. Ya. Danilevsky, who discovered the presence of electrical oscillations in the central nervous system, V. Yu. Chagovets, who formulated the basic principles of the ionic theory of excitation.

The works of the revolutionary democrats of the 60s of the 19th century N. G. Chernyshevsky, A. I. Herzen, V. G. Belinsky, N. A. Dobrolyubov, D. I. Pisarev had a huge influence on the formation of materialistic traditions in Russian physiology. In their works, they developed democratic ideas, ardently propagated the achievements of the natural sciences and the materialistic worldview. Among the physiologists-materialists who accepted the ideas of the Russian enlighteners-democrats, I. M. Sechenov and I. P. Pavlov should be put in the first place. studying the relationship between the processes of excitation and inhibition in the nervous system.

The study of the physiology of the central nervous system led I. M. Sechenov to the discovery of the phenomenon of summation of nerve impulses. He discovered the periodicity of electrical oscillations in the medulla oblongata.

The immediate successor of I. M. Sechenov’s research was his student N. E. Vvedensky (1852-1922), professor at St. Petersburg University. NE Vvedensky developed a new method of telephonic recording of electrical phenomena in living tissues. Using this method, he showed that the excitation process depends not only on the stimulus, but also on the state of the excitable tissue. N. E. Vvedensky experimentally proved the low fatigability of nerve fibers. He established the unity of the processes of excitation and inhibition, their inseparable connection. N. E. Vvedensky developed the doctrine of parabiosis - a universal reaction of living tissue to damaging effects.

The ideas of N. E. Vvedensky continued to be developed by his student and successor in the Department of Physiology of the Leningrad University, A. A. Ukhtomsky (1875-1942). He created the doctrine of the dominant - the dominant focus of excitation in the central nervous system under certain conditions.

I. P. Pavlov (1849-1936) played an outstanding role in the development of domestic and world physiological science. 1901) - the physiology of digestion, the third (1901-1936) - the higher nervous activity of animals and humans.

The study of the functions of the higher parts of the central nervous system of animals made it possible to come close to revealing the laws of the activity of the human brain. IP Pavlov created the doctrine of the types of higher nervous activity, which has not only theoretical, but also practical significance.

The pinnacle of I. P. Pavlov's creativity is his doctrine of the signaling systems of the cerebral cortex. IP Pavlov showed the qualitative features of the higher nervous activity of a person, studied and described the mechanisms by which abstract thinking is carried out, which is inherent only in man.

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Brief history of physiology

Physiology owes its origin to the needs of medicine, as well as the desire of a person to know himself, the essence and manifestations of life at various levels of its organization. The need to preserve human life was at all stages of its development, and already in ancient times, elementary ideas about the activities of the human body were formed, being a generalization of the accumulated experience of mankind. The father of medicine, Hippocrates (460-377 BC) represented the human body as a kind of unity of liquid media and the mental make-up of the individual, emphasized the connection of a person with the environment and that movement is the main form of this connection. This determined his approach to the complex treatment of the patient. An approach similar in principle was characteristic of physicians in ancient China, India, the Middle East and Europe.

In the Middle Ages, ideas far from reality, based on the postulates of the Roman anatomist Galen, dominated, and the dominance of the church determined an indefinable barrier between body and soul.

The Renaissance (XVI-XVII centuries), with its increased needs of social production, awakened science and culture to life, and the undoubted successes of physics and chemistry, the appeal of doctors to them determined the desire to explain the activity of the human body on the basis of the chemical (iatrochemistry) and physical ( iatrophysics) processes. However, the level of knowledge of the sciences of that time, of course, could not form any complete and adequate idea of ​​physiological functions.

At the same time, the invention of the microscope and the deepening of knowledge about the microscopic structure of animal tissues prompts the study of the functional purpose of the structures being discovered. The successes of chemistry and the study of the circulation of substances in nature direct the interests of man to the fate of the substances entering his body, which becomes the subject of research interest. The improvement of the exact sciences, natural science in general and philosophy determines the appeal of human thought to the mechanisms of movement. So, R. Descartes (1596-1650) formulates the reflex principle of organization of movements, which is based on the stimulus that induces them.

A special place in the science of man was played by the discovery of blood circulation by the English physician W. Harvey (1578-1657). Possessing extensive anatomical knowledge, V. Harvey conducted experimental studies on animals and observations on humans, founded physiology as a science, the main method of which is experiment. The official date for the emergence of human and animal physiology as a science is 1628 - the year of the publication of W. Harvey's treatise "Anatomical study of the movement of the heart and blood in animals." This work served as an incentive to study the activity of the organism in animal experiments as the main objective source of knowledge.

In the 17th century, a number of studies were carried out on the physiology of muscles, respiration, and metabolism. In Europe in the 18th century, the doctrine of "animal electricity" (L. Galvani, 1737-1798) arose, which grew into one of the leading sections of modern science - electrophysiology. The principle of reflex activity is being further developed (I. Prohaska, 1749-1820). A lot of value is being introduced into the understanding of the activity of the circulatory systems (S. Hels, 1667-1761), respiration (D. Priestley, 1733-1804), metabolism (A. Lavoisier, 1743-1794).

During this period, the Russian Academy of Sciences was opened (1724), where D. Bernoulli performed the first experimental studies in Russia on the movement of blood through blood vessels. In Russia, solid physiological discoveries were made by M. V. Lomonosov (1711-1765).

The 19th century is the heyday of analytical physiology, when outstanding discoveries were made in almost all physiological systems. This happened simultaneously with the rapid growth of natural science, the acquisition of fundamental knowledge about nature: the discovery of the law of conservation of energy, the cellular structure of organisms, the formation of the foundations of the doctrine of the evolution of life on Earth. Of particular importance in the development of physiology were new methodological approaches and inventions of outstanding physiologists of that time, as discussed in the previous section. All this determined the separation of physiology into an independent science in the middle of the 19th century. Physiological laboratories are being set up at the universities of Russia and England, and physiological research is being intensified in Europe.

In the second half of the 19th century - the beginning of the 20th century, physiology in Russia became one of the most advanced in world science, in which an outstanding role was played by metropolitan schools I. M. Sechenov (1829-1905), I. P. Pavlov (1849-1936), famous schools of Kazan, Kiev, Odessa, Tomsk, Yekaterinburg. Russian science, for all its originality and methodological originality, maintained the closest creative ties with the leading physiological schools. Western Europe and then America.

The 20th century is a period of integration and specialization of sciences, and physiology did not bypass the greatest discoveries. In the 40-50s, the membrane theory of bioelectric potentials was approved (A.L. Hodgkin, E.F. Huxley, B. Katz). The role of this theory in establishing the ionic mechanisms of excitation of neurons in 1963 is noted Nobel Prize(D. K. Eccles, E. F. Huxley, A. L. Hodgkin). Fundamental discoveries are being made in the field of cytophysiology and cytochemistry.

The end of the 19th and the beginning of the 20th century was a period of decisive successes in the field of the physiology of nerves and muscles as excitable tissues (Dubois-Reymond, E.F. Pfluger, P.G. Heidenhain, Yu. Bernshtein, G.L. Helmholtz). In Russia, especially notable research in this branch of science is carried out by N. E. Vvedensky (1852-1922),

A. I. Babukhin (1835-1891), B. F. Verigo (1860-1925),

V. Ya. Danilevsky (1852-1939), V. Yu. Chagovets (1873-1941). A. V. Hill (1886-1977) and O. F. Meyerhof (1884-1951) were awarded the Nobel Prize for discoveries of heat generation in muscles. The achievement of the 20th century, marked by the Nobel Prize in 1936, was the discovery of the chemical mechanism of nerve impulse transmission in synapses by O. Levy (1873-1961) and G. X. Dale (1875-1968). The development of this direction in the works of W. Euler, D. Axelrod and B. Katz was awarded the Nobel Prize in 1970. A. D. Erlanger and G. Gasser were awarded the same prize in 1944 for their success in studying the conduction of impulses along nerve fibres. Soviet physiologists A. A. Ukhtomsky (1875-1942), A. F. Samoilov (1867-1930), D. S. Vorontsov (1886-1965) also made a significant contribution to the solution of the problem of excitation of nerves and muscles during this period.

The 19th and 20th centuries were marked by many significant advances in the study of brain function.

An outstanding role in the study of brain functions belongs to I. M. Sechenov (1829-1905), who in 1862 discovered the phenomenon of inhibition in the central nervous system, which largely determined the subsequent success in studies of the coordination of reflex activity. The ideas set forth by I. M. Sechenov in the book “Reflexes of the Brain” (1863) determined that mental phenomena were attributed to reflex acts, introduced new ideas into the mechanisms of brain activity, and outlined fundamentally new approaches to its further research. At the same time, the scientist emphasized the decisive role of the external environment in the reflex activity of the brain.

I. P. Pavlov (1849-1936) brought the theory of reflex activity of the brain to a qualitatively new level, having created the doctrine of the higher nervous activity (behavior) of humans and animals, its physiology and pathology. IP Pavlov founded the school of Russian physiologists, which made an outstanding contribution to world science.

Among the students and followers of I. P. Pavlov are academicians P. K. Anokhin, E. A. Astratyan, K. M. Bykov, L. A. Orbeli and many others who created domestic physiological scientific schools.

The ideas of I.P. Pavlov about the reflex activity of the brain were further developed in the theory of functional systems by P.K. Anokhin (1898-1974), which are the basis for organizing complex forms of behavioral activity and ensuring homeostasis of the human and animal organisms. It is difficult to overestimate the contribution to the physiology of the nervous system by I. S. Beritashvili (1885-1975), who discovered fundamental patterns in brain activity and created a number of original theories about its organization.

E. A. Astratyan (1903-1981) - the author of a number of fundamental works, in which he developed the main provisions of I. P. Pavlov on higher nervous activity. K. M. Bykov (1887-1959) founded the doctrine of the two-way connection of the cerebral cortex with internal organs, of cortico-visceral pathology. His student V. N. Chernigovsky (1907-1981) enriched science with the doctrine of interoception of visceral organs, regulation of the blood system.

L. A. Orbeli (1882-1958) founded the doctrine of the adaptive-trophic influences of the sympathetic nervous system on the somatic and autonomic functions of the body, and was one of the founders of evolutionary physiology.

L. S. Stern (1878-1968) created the doctrine of the blood-brain and histo-hematogenous barriers that provide homeostatic functions in humans and animals.

Great is the merit of A. A. Ukhtomsky (1875-1942) in the study of the physiology of the central nervous system. His doctrine of the dominant - the "basic principle of activity" of the brain, still feeds the ideas of organizing the purposeful activity of humans and animals.

There is no doubt that the contribution of Russian physiologists to the world science of the brain is original and generally recognized, much has been done in the study of the localization of functions in the brain (V. M. Bekhterev, M. A. Mislavsky, F. V. Ovsyannikov, etc.), in the development of methods his study.

At the end of the 19th and in the 20th century, brain physiology was successfully developing in Europe and America. To a large extent, this is due to the creation of a neural theory of reflex activity of the brain on the basis of his histological study by C. Golgi (1844-1926) and S. Ramon y Cajal (18512-1934), who were awarded the Nobel Prize in 1906, and then Lorente de No.

An outstanding role in the study of the functions of the central nervous system was played by Ch. S. Sherrington (1856-1952), who developed and formulated the basic principles of the coordination activity of the brain. These works were awarded the Nobel Prize in 1932. The award was also received by the electrophysiologist

E. D. Adrian (1889-1977), who also made a significant contribution to modern ideas about the activity of the brain. The merit of Ch. S. Sherrington is that he brought up a galaxy of physiologists to whom science owes many outstanding discoveries (R. Granit, R. Magnus, W. Penfield, J. Eccles, and others).

R. Magnus (1873-1927) owes science to the doctrine of the adjusting reflexes that distribute the tone of skeletal muscles. R. Granit, X. K. Hartlainen and D. Wald in 1967, and D. Hubel and T. Wiesel in 1981 were awarded the Nobel Prize for their work on the physiology and biochemistry of the visual analyzer. Russian scientists P. P. Lazarev (1878-1942) and V. S. Kravkov (1893-1951) also made a worthy contribution to this branch of science.

The modern physiology of the reticular formation of the brain was created by the experimental studies of G. Magun and D. Moruzzi. It should be emphasized that the results of the scientific work of I. M. Sechenov and V. M. Bekhterev served as the basis for these studies.

Of course, the functions of the brain have attracted and continue to attract the attention of many prominent scientists in the world, and successful searches continue in this area. Their main results are described in the corresponding chapters of the textbook, with the names of living physiologists also mentioned.

The physiology of visceral organs has occupied a very prominent place in the history of science from the time of the emergence of physiology to the present day. The 19th and 20th centuries were marked by major discoveries on the mechanisms of regulation of the activity of the heart and blood vessels: K. Ludwig (1816-1895), I.F. Zion (1842-1912), K. Bernard (1813-1878), F.V. Ovsyannikov (1827-1906), V. Einthovei (1860-1927), E. G. Sterling (1866-1927) and others.

A. Krogh (1874-1949) was awarded the Nobel Prize in 1920 for studies of capillary circulation. V Soviet time A major scientific contribution to the physiology of the cardiovascular system was made by V. V. Parin (1903-1971), V. N. Chernigovsky, A. M. Chernukh and others.

The 20th century is rich in successes in the field of physiology of respiration, especially its regulation (N. A. Mislavsky, K. Heimans, D. S. Haldane). For work in this area, K. Heimans (1892-1968) received the Nobel Prize in 1939. Major discoveries were made in the biochemistry of gas exchange and cellular respiration (A. Krogh, D. Barcroft), and O. G. Warburg (1883-1970 ) for the discovery of the enzymatic mechanism of cellular respiration, the Nobel Prize was awarded in 1931. M. V. Sergievsky (1898-1982) made a great contribution to the physiology of the respiratory center.

Outstanding physiologists of Europe and America (K. Ludwig, K. Bernard, R. Hedenhain, E. Starling, etc.) were engaged in the physiology of digestion at different times, but “recreated the physiology of digestion” (as stated in the diploma of the Nobel laureate in 1904) and P. Pavlov is the first among the physiologists of the world and the first Russian scientist to be awarded this high title.

History of the development of physiology

The work of another Nobel laureate, I. I. Mechnikov (1845-1916), was devoted to intracellular digestion. E. S. London, I. P. Razenkov, G. V. Folbort, B. P. Babkin and others worked in the laboratory of I. P. Pavlov, who continued the glorious traditions of pioneers in the field of digestive physiology. An outstanding role in this field of science was played by A. M. Ugolev (1926-1992), who owns the honor of discovering membrane intestinal digestion and determining its place in the digestive conveyor, modern concepts of the endocrine activity of the gastrointestinal tract, the evolution of secretory processes, the theory of adequate nutrition and other original theories and hypotheses in physiology.

In the physiology of visceral systems, the basic concepts of the functional organization of the autonomic (vegetative) nervous system were formed. These pages of the history of physiology are described in sufficient detail in section 4.3 of the textbook.

The 20th century is rich in discoveries in the field of studying the activity of the endocrine glands. In 1923, the Nobel Prize was awarded to F. G. Banting (1891-1941). D. McLeod (1876-1935) and C. G. Best (1899-1978) for their work on insulin. This prize was awarded in 1947 to B. A. Usai (1887-1971) for his discoveries in the field of the physiology of the pituitary gland. Work on the study of the function of this gland was also noted in 1977 by R. Guillemin, E. V. Schally and R. S. Yalou. In 1950, the Nobel Prize for the study of adrenal function was awarded to F. Sh. Hench (1896-1965), E. K. Kendall (1886-1972) and T. Reichstein (b. in 1897).

In 1971, E. W. Sutherland (1915-1974) became the Nobel laureate, who discovered the role of AMP in the regulation of metabolism, showed its importance as a mediator in the hormonal effect on metabolism.

Domestic physiologists have priority in creating an artificial heart (A. A. Bryukhonenko), EEG recording (V. V. Pravdich-Neminsky), creating such important and new areas in science as space physiology, labor physiology, sports physiology, and the study of physiological mechanisms adaptation, regulation of mechanisms for the implementation of many physiological functions. These and many other studies are of paramount importance for medicine.

THE SUBJECT OF PHYSIOLOGY, ITS RELATIONSHIP WITH OTHER SCIENCES AND SIGNIFICANCE FOR PHYSIOLOGY AND SPORT

Physiology is the science of the functions and mechanisms of activity of cells, tissues, organs, systems and the whole organism as a whole. Physiological function- this is a manifestation of vital activity, which has an adaptive value.

physiology as a science is inextricably linked with other disciplines. It is based on the knowledge of physics, biophysics and biomechanics, chemistry and biochemistry, general biology, genetics, histology, cybernetics, anatomy. In turn, physiology is the basis of medicine, psychology, pedagogy, sociology, theory and methodology of physical education. In the process of the development of physiological science, various particular sections of it emerged from general physiology. physiology of labor, physiology of sports, aerospace physiology, physiology of underwater labor, age physiology, psychophysiology, etc.

General physiology is the theoretical foundation of sports physiology. It describes the basic regularities of the activity of the body of people of different ages and gender, various functional states, the mechanisms of operation of individual organs and systems of the body and their interaction. Its practical significance lies in the scientific substantiation of the age stages of development of the human body, the individual characteristics of individuals, the mechanisms for the manifestation of their physical and mental abilities,

features of control and possibilities of managing the functional state of the body. Physiology reveals the consequences of bad habits in humans, substantiates ways to prevent functional disorders and maintain health. Knowledge of physiology helps the teacher and coach in the processes of sports selection and sports orientation, in predicting the success of the competitive activity of an athlete, in the rational construction of the training process, in ensuring the individualization of physical activity and open up the possibility of using the functional reserves of the body.

PHYSIOLOGICAL RESEARCH METHODS

Physiology is an experimental science. Knowledge about the functions and mechanisms of the body's activity is based on experiments conducted on animals, clinical observations, and examinations of healthy people under various experimental conditions. At the same time, in relation to a healthy person, methods are required that are not associated with damage to his tissues and penetration into the body - the so-called non-invasive methods.

In a general form, physiology uses three methodological methods of research: observation or the "black box" method, acute experience and chronic experiment.

The classical research methods were methods of removal and methods of irritation of individual parts or entire organs, mainly used in experiments on animals or during operations in the clinic. They gave an approximate idea of ​​the functions of removed or irritated organs and tissues of the body. In this regard, the method of conditioned reflexes developed by IP Pavlov was a progressive method for studying the whole organism.

In modern conditions, the most common are electrophysiological methods that allow recording electrical processes without changing the current activity of the organs under study and without damaging the integumentary tissues - for example, electrocardiography, electromyography, electroencephalography (registration of the electrical activity of the heart, muscles and brain). The development of radio telemetry makes it possible to transmit these received records over considerable distances, and Computer techologies and special programs - provide a fine analysis of physiological data. The use of infrared photography (thermal imaging) allows you to identify the hottest or coldest areas of the body observed at rest or as a result of activity. With the help of the so-called computed tomography, not

opening the brain, you can see its morphofunctional changes at different depths. New data on the functioning of the brain and individual parts of the body are provided by the study of magnetic oscillations.

A BRIEF HISTORY OF PHYSIOLOGY

Observations of the vital activity of the organism have been made since time immemorial. For 14-15 centuries BC. in ancient Egypt, in the manufacture of mummies, people were well acquainted with the internal organs of a person. In the tomb of the pharaoh Unas, ancient medical instruments are depicted. In ancient China, up to 400 diseases were surprisingly subtly distinguished only by the pulse. In the IV-U century BC. e. there, the doctrine of functionally important points of the body was developed, which at present became the basis for modern developments in reflexology and acupuncture, Su-Jok therapy, testing the functional state of the athlete's skeletal muscles by the magnitude of the electric field strength of the skin in bioelectrically active points above them. Ancient India became famous for its special herbal recipes, the impact on the body of yoga exercises and breathing exercises. In ancient Greece, the first ideas about the functions of the brain and heart were expressed in the 4th-5th centuries BC. e. Hippocrates (460-377 BC) and Aristotle (384-322 BC), and in Ancient Rome in the 11th century BC, the physician Galen (201-131 BC). e.).

However, as an experimental science, physiology arose in the 17th century AD, when the English physician W. Harvey discovered the circles of blood circulation. In the same period, the French scientist R. Descartes introduced the concept of reflex (reflection), describing the path of external information to the brain and the return path of the motor response. The work of the brilliant Russian scientist M. V. Lomonosov and the German physicist G. Helmholtz on the three-component nature of color vision, the treatise of the Czech G. Prochazka on the functions of the nervous system, and the observations of the Italian L. Galvani on animal electricity in nerves and muscles marked the 18th century. In the 19th century, the ideas of the English physiologist C. Sherrington on integrative processes in the nervous system were developed, which were outlined in his famous monograph in 1906. The first studies of fatigue were carried out by the Italian A. Mosso. I. R. Tarkhanov (Tarkhanov's phenomenon) discovered changes in the constant potentials of the skin during irritation in humans.

In the 19th century the works of the "father of Russian physiology" I.M. Sechenov (1829-1905) laid the foundations for the development of many areas of physiology - the study of blood gases, the processes of fatigue and "active rest", and most importantly - the discovery in 1862 of inhibition in the central nervous system ("Sechenovsky inhibition") and the development of physiological

foundations of human mental processes, which showed the reflex nature of human behavioral reactions ("Reflexes of the brain", 1863). Further development of the ideas of I. M. Sechenov went in two ways. On the one hand, the study of the subtle mechanisms of excitation and inhibition was carried out in University N. E. Vvedensky (1852-1922).He created the idea of ​​physiological lability as a speed characteristic of excitation and the doctrine of parabiosis as a general reaction of neuromuscular tissue to irritation. Later this direction was continued by his student A. A. Ukhtomsky ( 1875-1942), who, studying the processes of coordination in the nervous system, discovered the phenomenon of the dominant (the dominant focus of excitation) and the role in these processes of assimilation of the rhythm of irritations. On the other hand, in a chronic experiment on a whole organism, I. P. Pavlov (1849 -1936) first created the doctrine of conditioned reflexes and developed a new chapter of physiology - the physiology of the higher nervous activity ness. In addition, in 1904, for his work in the field of digestion, IP Pavlov, one of the first Russian scientists, was awarded the Nobel Prize. The physiological foundations of human behavior, the role of combined reflexes were developed by V. M. Bekhterev.

Other outstanding Russian physiologists also made a major contribution to the development of physiology: the founder of evolutionary physiology and adaptology, Academician L.A. Orbeli, who studied the conditioned reflex effects of the cortex on the internal organs of Acad. K. M. Bykov, creator of the theory of the functional system acad. P. K. Anokhin, founder of Russian electroencephalography - acad. MN Livanov, developer of space physiology - acad. V. V. Larin, the founder of the physiology of activity - N. A. Bernshtein and many others.

In the field of physiology of muscular activity, it should be noted the founder of the national physiology of sports - prof. A. N. Krestovnikov (1885-1955), who wrote the first textbook on human physiology for the country's sports universities (1938) and the first monograph on the physiology of sports (1939), as well as well-known scientists - prof. E. K. Zhukov, V. S. Farfel, N. V. Zimkin, A. S. Mozzhukhin and many others, and among foreign scientists - P.-O. Astranda, A. Hilla, R. Granita, R. Margaria and others.

GENERAL REGULARITIES OF PHYSIOLOGY AND ITS BASIC CONCEPTS

Living organisms are so-called open systems (that is, not closed in themselves, but inextricably linked with the external environment). They are made up of proteins and nucleic acids and

characterized by the ability to autoregulate and self-reproduce. The main properties of a living organism are metabolism, irritability (excitability), mobility, self-reproduction (reproduction, heredity) and self-regulation (maintenance of homeostasis, adaptability).