The great Russian physiologist was born 100 years ago
this month. He caused dogs to salivate at the sound of
a bell, and brought together physiology and psychology
HIS month marks the 100th anni­
T versary of the birth of Ivan Petro­
vich Pavlov. The great Russian
physiologist, who died only 13 years ago,
was one of those rare personages in sci­
ence whose work is known in every
country during his own time. Indeed, the
character of his fame reminds one a little
of the Pavlov dog whose mouth always
watered at the sound of a bell: the men­
tion of Pavlov's name instantly evokes in
every literate person's mind the great
contribution with which he is insepara­
bly associated-"the conditioned reflex."
Pavlov's work left imperishable marks
on physiology, neurology and psycholo­
gy. Despite his wide fame, however, he
was not intimately known outside his
own country. His anniversary offers an
occasion to tell something about Pavlov
as a man and as a scientist. Obviously
one cannot attempt to give a compre­
hensive review of his work in a single
article; there will be set down here sim­
ply some personal recollections of Pav­
lov, and an estimate of some of his most
important researches.
Pavlov was a man with obvious natu­
ral gifts-a brilliant mind, an incompara­
ble memory, boundless energy and im­
measurable enthusiasm-but with all that
he remained to the end of his life a trans­
parently simple human being. His labo­
ratory in Leningrad when I worked un­
der him was a tumultuous beehive.
Pavlov was then over 80 years old; none­
theless he still possessed a vast enthusi­
asm for scientific investigation which he
communicated to all those around him.
He was the moving spirit in most of the
laboratory's projects, and he would greet
the successful completion of an experi­
ment with a dance of joy. His laboratory
was operated like a town meeting; on
Wednesdays his several dozen scientific
associates gathered to discuss and argue
their problems. Pavlov, who had a gift
for dramatic narrative, fascinated his
colleagues. He fought tooth and nail for
his ideas against all arguments; but after
he had cooled down he was quick to
44
by J erzy Konorski
admit his error if his opponent turned
out to be right.
There is a story about Pavlov that il­
lustrates amusingly his general attitude
toward his work. During his early studies
of the digestive system, he found that
hydrochloric acid, when supplied to an
animal's duodenum, caused the pancreas
to secrete juice. Pavlov believed that the
acid acted through some mechanism in
the nervous system to produce this re­
sult. Some time later, however, the Brit­
ish phYSiologists Sir William Bayliss and
Ernest H. Starling showed that the
pancreatic secretion was initiated by a
hormone mechanism: when the mucous
membrane of the duodenum was stimu­
lated by hydrochloric acid, it produced
a hormone, secretin, which then acted on
the pancreas. Pavlov at first pronounced
this result incredible. When, having re­
peated the Bayliss-Starling experiments
himself, he found that their conclusion
was indeed correct, he exclaimed wrath­
fully, not in envy but in astonishment
and self-reproach: "So we're not the only
ones to make discoveries!"
AVLOV was born in 1849 in the·
Pprovincial town of Ryazan, in cen­
tral Russia. He was the son of an Or­
thodox village priest, and inevitably was
educated in a religious seminary. But
he soon found that his interests lay
elsewhere, and he transferred to the Fac­
ulty of Natural Sciences in St. Petersburg
University. He went on to study medi­
cine in the Military Medical Academy,
graduating in 1879. While still a student,
he began his first researches in the
physiology of blood circulation. He was
appointed to the staff of the Clinic for
Internal Diseases. The head of the Clinic
entrusted Pavlov with the responsibility
of organizing a physiological laboratory
to combine medical theory with practice
in the Clinic. Except for a two-year pe­
riod, 1884-86, during which Pavlov went
to study with the German phYSiologist
Carl F. W. Ludwig at Leipzig, he did
most of his early work in this meager
© 1949 SCIENTIFIC AMERICAN, INC
laboratory at the St. Petersburg clinic.
His facilities were of the most primi­
tive kind. The laboratory was a small
wooden building, hardly more than a
shed. He had to finance his research
largely out of his own very small salary.
He had no regular assistants. Neverthe­
less, by his own uncommon energy, as­
Siduity and self-sacrifice he made rapid
and fruitful headway, and his work won
him a recognition out of all proportion
to the modesty of his laboratory. In 1890
he was appointed to the chair of pharma­
cology in the Military Medical Academy,
and in 1891 he was placed in charge of
the physiological laboratory at the Insti­
tute of Experimental Medicine which
was then being established in St. Peters­
burg.
By the end of the 19th century Pavlov
was generally recognized as one of the
world's outstanding phYSiologists. In
1904 he was awarded the Nobel prize
for his work on the physiology of diges­
tion. In 1907 he was elected to the Rus­
sian Academy of Sciences, and later he
became the director of the Academy's
Physiological Institute, which post he
retained to the end of his life. In the
1930s a biological station was built for
him in Koltuszi, now called Pavlovo,
near Leningrad. He died of pneumonia
in February, 1936, at the ripe age of 87.
MITTING Pavlov's interesting but
O not preeminent early studies of the
regulation of blood pressure, his lifework
divides into two periods. From the 18805
to 1902, he devoted himself to detailed
investigation of the functions of the ali­
mentary tract. From 1902 until his
death, he went on to explore a new
branch of knowledge which he himself
created-the phYSiology of higher nerv­
ous activity.
As is well known, Pavlov's research
work on the secretory function of the
alimentary tract became the lasting
foundation of our present-day knowl­
edge in this field. In these studies, as
well as in his later investigations of the
nervous system, he was guided by cer­
tain ideas which, while not entirely origi­
nal with him, he perhaps developed
most consistently.
The first of these principles was that
the organism functions as an integrated
whole, and that the investigation of
separate organs in the artificial condi­
tions of isolated experiments cannot pro­
vide an adequate idea of how those or­
gans act in their normal situation. Hence
Pavlov tried to carry out his physiologi­
cal experiments in conditions approxi­
mating the normal as closely as possible.
In his work on the alimentary tract he
resorted to a number of ingenious opera­
tions to maintain the normal functioning
of an animal's organs while exposing
them to observation. For example, he
developed a method of exposing the
ducts of the salivary and pancreatic
glands without removing the organs
from the body. He learned how to isolate
a part of an organ without detaching it
from the nervous system. One of his
typical operations was the so-called
"Pavlov pouch." In this operation, per­
formed on a dog, one part of the stomach
is isolated and formed into a blind
pouch, with an opening to the abdominal
wall. The nervous network of the pouch
is uninjured, so that its secretory func­
tion is an exact replica of the functioning
IVAN PETROVICH PAVLOV was born in 1849 and
died in 1936. He was awarded a Nobel prize in 1904. He
of the rest of the stomach, to which food
passes, and it can be examined with
great precision.
Obviously to make such operations
come off successfully and keep the ani­
mals healthy Pavlov had to develop asep­
tic operating conditions and facilities for
proper care of the animals after the op­
eration. Today the existence of such ad­
juncts is taken for granted, but in those
days the idea was quite novel.
Pavlov's second guiding principle was
what he called "nervism," meaning the
hypothesis that all functions of the body
were controlled by the nervous system.
It must be remembered that in those
days the study of endocrinology was only
in its beginnings. Pavlov believed that
the nervous system was the only m'echa­
nism regulating and integrating the or­
ganism's activity, and so in all his
researches he concentrated on elucidat­
ing the role qf the nerves. One of his
most beautiful exp!=!riments was a set of
operations on a dog's gastric system to
investigate the mechanism of secretion
by the stomach. He severed the dog's
gullet from its stomach so that the food
it ate would not pass into the stomach
but fall outside the body. He also made
a tubular opening into the stomach to
examine its behavior. He found that even
though no food actually entered the
was greatly honored by the U.S.S.R. A biological station
was built for him at Koltuszi, which is now called Pavlovo.
© 1949 SCIENTIFIC AMERICAN, INC
stomach, it still secreted gastric juice,
being powerfully stimulated by the ani­
mal's chewing and other acts of eating.
And Pavlov proved that this reaction
was mediated by the vagus nerves, for
if these nerves wyre cut, the gastric se­
cretion elicited by the act of eating
stopped immediately. Pavlov's devotion
to the theory of the central role of the
nervous system helps to explain his in­
credulity and astonishment at the later
discovery by Bayliss and Starling that
hormones also play a part in the process
of digestion.
The third principle that strongly moti­
vated Pavlov's work was his conviction
that physiological experimentation was
of great significance for practical medi­
cine. This idea, accepted without ques­
tion today, was not so universally recog­
nized then; at anv rate not, so far as I
'
know, in Russia.
Ptem and the investigation of condi­
AVLOV'S switch to the nervous sys­
tioned reflexes, which marked the second
phase of his career, was a natural sequel
to his work on the digestive system. Dur­
ing that work he had been impressed by
the discovery that a dog secretes saliva
and gastric juice not only as the result of
direct action of the food on the mucous
membranes of the mouth and stomach,
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45
 ,
PAVLOV'S BIG LABORATORY at Koltuszi was
crowded with fellow workers and experimental animals.
but even in response to the mere sight
of food, or to other signals heralding
feeding. This indicated that gastric se­
cretions, which Pavlov had considered a
purely physiological phenomenon, might
also have a psychological basis and be
related to the dog's experiences.
To a physiologist this was then a
shocking idea. Up to that time physiolo­
gy and psychology had been regarded
as two entirely separate fields. Physiolo­
gy was concerned solely with the innate
responses of the body, mainly those con­
trolled by the lower parts of the nervous
system, while psychology dealt only with
acquired or learned responses. Pavlov
was confronted ,with a vexing problem.
Must he now give up physiological
methods and turn to psychology to in­
vestigate the dog's gastric behavior? This
he could not bring himself to do, for he
could see no way to verify theories in the
realm of psychology by means of experi­
ment.
After long hesitation and tormenting
irreso:ution, Pavlov found a characteris­
tically imaginative and resolute answer
to his dilemma: He would attack the psy­
chological problem with purely physio­
logical methods. After all, the secretion
of saliva or gastric juice, was exactly the
same phenomenon whether it had a di­
rectly physiological origin' or a psychic
one. By applying physiological experi­
ments to the investigation of acquired
46
Today the laboratory continues in the Pavlov tradition.
At the left a baboon manipulates a stick during a study
behavior, he might open up an enormous
new field of study.
For several well-considered reasons,
Pavlov chose the salivary glands as the
focus of his experiments. He knew from
previous research that the salivary gland
is a very sensitive and selective reacting
mechanism. Moreover, its activity is
much more restricted and specific than
that of the motor organs, the chief effec­
tors of acquired behavior. Above all,
Pavlov judged that investigation of the
salivary glands would involve much less
danger of anthropomorphic and psycho­
logical interpretation of the results, an
error he wished to avoid at all costs.
So Pavlov and his collaborators began
their studies of acquired, or, as he him­
self called them, conditioned (in Russian
"conditional") reflexes. The first experi­
ments were made on "natural" condi­
tioned reflexes, those established spon­
taneously in an animal in response to the
sight and the smell of food, and so on.
Only hter, as the method was perfected,
did the Pavlov group begin to develop
conditioned reflexes to special Signals,
such as the beat of a metronome, the
ringing of a bell, or the lighting of a
lamp.
An important early finding in, these
experiments was that conditioning works
two ways: it can inhibit as well as pro­
duce a response. When a conditioned
stimulus ceases to be "reinforced," i.e.,
© 1949 SCIENTIFIC AMERICAN, INC
to be accompanied by the presentation
of food, the conditioned reflex is extin­
guished. Pavlov showed that this extinc­
tion is effected by a special mechanism
which he called internal inhibition.
During the first period of research the
Pavlov group occupied themselves chief­
ly with the properties and interrelations
of the excitatory and inhibitory condi­
tioned reflexes. Later they extended the
investigation to two new and important
spheres. First, they showed by repetition
of identical experiments on a great many
dogs that animals vary greatly in the
speed with which conditioned reflexes
are formed, their permanence, the influ­
ence of inhibitory on excitatory reflexes,
and so on. This provided a basis for the
development of a classification of types
of nervous system, a subject which has
been worked at extensively of recent
years. Attempts have even been made at
the Biological Station in Pavlovo to
show that an individual's type of nervous
system may be inherited. Secondly, at
the end of the second decade of re­
search into conditioned reflexes the Pav­
lov investigators came quite accidentally
upon the discovery of a neurotic state in
dogs, caused by a conflict between ex­
citatory and inhibitory reflexes.
The latter finding led to extensive in­
vestigation of so-called experimental
,neuroses, their pathogenesis, symptoma­
tology and therapy. In these experiments
of motor reflexes. In the center an experimenter observes
a monkey in the laboratory of higher nervous activity.
the salivary conditioned reflexes proved
to. be a very sensitive and precise indi­
cator both of normal and of pathologi­
cal nervous conditions. Toward the end
of Pavlov's life a psychiatric and a psy­
choneurological clinic were attached to
his laboratories, and in these the at­
tempt was made to analyze various cases
of human neuroses by resort to the laws
discovered in experiments on animals.
Thus even in this long, difficult and
fundamental inquiry Pavlov realized his
lifelong hope of applying his experi­
mental research on animals directly to
human pathology.
I have summarized in general outline
the main scientific achievements that
Pavlov and the physiological school he
created have given us. So far as his re­
searches into the physiology of digestion
are concerned, there is no need to sh'ess
their value. But it is more difficult to esti­
mate the value and scope of the science
of conditioned reflexes.
It is generally accepted that condi­
tioned reflexes have played a very con­
siderable part in the development of
modern psychology, and today there are
whole trends of psychological investiga­
tion based to a large extent on the
achievements of the Pavlov school. But,
since the psychological applications of
conditioned reflexes ' have been devel­
oped mainly in the U. S., I do not think
that I, a European far removed from the
At the right another experimenter tabulates a dog's
respiratory responses with the help of a kymograph.
centers of these trends, am competent to
discuss them. As for the practical appli­
cations of the Pavlov achievements to
the fields of psychoneurology, psychic
hygiene and teaching, it is still too early
to estimate their ultimate importance.
And so I shall confine myself to survey­
ing the significance of conditioned re­
flexes to neurophysiology itself.
Pavlov often called his teaching on
conditioned reflexes "the h'ue physiology
of the brain." For him the study of con­
ditioned reflexes was not an end in itself,
but rather a means for understanding the
central mechanism controlling them,
namely, the cerebral cortex. Pavlov con­
ceded that other methods of investiga­
tion of the activity of the cerebral cortex,
such as electrical stimulation of the cor­
tex in an anesthetized animal, could be
very valuable, but he held that the h'ue
picture of this activity could be obtained
only by studying the normal functiOning
of the organ, as in conditioned-reflex ex­
periments. He was strengthened in this
conviction by the fact that his view had
been thoroughly and brilliantly justified
in his work on the alimentary tract.
It is an interesting fact that Pavlov's
attitude and methods were closely re­
lated to those of a great British contem­
porary, Sir Charles Sherrington (see
page 56). Both Sherrington and Pavlov
based their physiological studies on work
with quantitatively and qualitatively de-
© 1949 SCIENTIFIC AMERICAN, INC
fined stimuli and their combinations.
Both investigated the central mechanism
of reflexes by examining the reactions of
animals. But in Sherrington's case the
preparation studied was a spinal animal,
whereas in Pavlov's it was an animal
with the cerebral cortex intact, and Sher­
rington studied the innate activity of the
nervous system, while Pavlov was con­
cerned with its acquired activity.
Sherrington's experimental work was
developed further by a host of neuro­
physiologists who, with the aid of more
refined methods than were at Sherring­
ton's disposal, brought greater precision
into the notions of central excitatory and
inhibitory states with which Sherrington
operated. Pavlov's work has not been fol­
lowed up as thoroughly as Sherrington's.
If Pavlov's method of investigation of
higher nervous activity is successfully
linked up with, and enriched by, the
modern tool of elech'oetlcephalography,
just as Sherrington's method of study of
lower nervous activity was later enriched
by the recording of action potentials of
the spinal cord, then perhaps Pavlov's
statement that his work represented "the
true physiology of the brain" will be fully
justified.
•
Jerzy Konorski, a former associate of Pav­
lov, is now at the Nencki Institute of
Experimental Biology in Lodz, Poland.
47