Evolution, physics and
astronomy are the three fields that pay the most attention to their histories,
because they are fields that have a tremendous influence on how we see
ourselves as humans and how we understand our relationship to the universe
around us. Each of these fields has had a major impact on Western philosophy,
including political philosophy. As scientists engaged in work in evolutionary
biology, we have a responsibility to understand our history and to pay
attention to how it influences our own views.
This is why Professor
Pierotti has spent the first third of this course introducing you to the
history of ideas and the underlying philosophical perspectives that influenced
evolutionary thinking over time. Evolutionary Biology is a field of great
interest to the public and to societies around the world, and the work we do
has ramifications beyond the bounds of abstract science.
Today I am going to illustrate this point by discussing what happened to geneticists during the Stalin period in the Soviet Union.The picture is of a stature of Stalin and Lysenko, and this book is one of the best on the topic. It was written by a scientist who helped to topple Lysenko’s hold on Soviet evolutionary biology—it had to be snuck out of the Soviet Union and published in the US because it was banned in the USSR. (Medvedev, Z.A. 1969. The Rise and Fall of T.D. Lysenko. Translated by I.M. Lerner. Columbia University Press, NY, NY. 184 pages).
The need to improve food security has long been a primary goal of Russian policy.
Compared to Kansas, our home state, Russia is much further north-- much
of the land mass is in the subarctic and arctic. As a result it has
short growing seasons and conditions more conducive to taiga and tundra
than agriculture. Think of it as being northern Canada (but even more
continental. This creates real problems when you try to conceive of
ways of feeding the population, especially in Siberia.
Time line of events discussed in today's lecture.
This is actually a story that I am
personally interested in, because in 1990, during the collapse of the Soviet
government, I was asked by the US National Academy of Sciences to work with
Soviet scientists on conservation and biodiversity problems common to both our
countries. I was sent to work with a Russian population geneticist, Alexander
Golubtsov, of the USSR Academy of Sciences. This slide shows Sasha and I in
Siberia collecting grayling. What was really startling to me in 1990 was that
there were Russian geneticists trained in modern techniques—I had been taught
in school that all of the geneticists in the Soviet Union had been either
killed, exiled, or taught a bogus form of Lamarckism rather than modern
evolutionary theory based on an understanding of genetics. The truth is that my
surprise was justified—Sasha was in one of the first generations to have
escaped from the ideologically driven Soviet style genetics and work openly
with Western scientists.
If you look at the
grayling in this slide you will see that there are a number of differences
between them—these fish come from different lakes that are fairly isolated from
each other, high in the mountains of Siberia. If you were to do morphological
measurements on them, like measure their fin lengths, count their scales, and
measure the dimensions of their heads, you would find that there are a lot of
differences, just like there are differences in their color patterns. Variation
between individuals and between populations of the same species is one of the
most interesting questions in evolutionary biology, and observations of
variation have been foundational for our theoretical framework for evolution
through natural selection.
Of course one of the
greatest observers of variation in natural populations was Charles Darwin.
Darwin traveled around the world with the British voyages of discovery during
the colonial period of the British Empire. What was most impressive about his
work is that he carefully and patiently documented variation in populations in
a way that illustrated patterns in nature. Variation for him wasn’t “noise” or
“chaos” it was a way to gain insight into processes that were both historical
and ongoing in the world around us.
Darwin collected many specimens, carefully documented the locations of
his collections, and provided many replicates of his observations—he created a
huge database to support his ideas and carefully backed all of his ideas with a
lot of factual information. He did good science at a time when modern
scientific methodology was still being developed.
Portrait of Charles Darwin 1809-82, 1840, a painting by George Richmond.
It always amazes me to
think about how good Darwin’s work is given that he didn’t have statistical
methods to analyze his data, he didn’t have the tools developed by modern
molecular geneticists and population geneticists to work with, in fact he had
no information available to him on the mechanisms of inheritance or the source
of variation in traits. Yet his scientific observations of variation in the
natural world were superb and his ideas have continued to provide a productive
framework for almost 150 years now.
The next major historical
figure important in this story is Gregor Mendel. Mendel took a very different
approach to studying variation—he developed experimental techniques for looking
at the inheritance of traits and developed the mathematical tools for analyzing
probability distributions. In order to discern patterns of inheritance you need
to be able to analyze the data to determine whether it is likely that there is
a pattern that is real or whether variation is random. Mendel is known for
demonstrating that it is possible to develop mathematical models of inheritance
in traits with discrete distributions, like flower color in peas.
Credit: NATIONAL LIBRARY OF MEDICINE / SCIENCE PHOTO LIBRARY
But when you want to look
at traits whose variation has a continuous distribution, like height in humans,
this is a more difficult problem. Many traits do not assort themselves in
discrete fashion, like Mendel’s pea color did. To study continuous probability
distributions, Fischer, Haldane and Wright developed mathematical techniques to
compare variation in what are commonly called “normal” distributions. By 1918
they had worked out the basis for analyzing traits like human height that we
have since come to understand are the consequence of the interaction of many
With these developments in
place things began to really take off—and the Modern Synthesis came about. The
Modern Synthesis is the synthesis of Darwinian evolution through natural
selection, combined with Mendel’s techniques of experimentally and
statistically analyzing the inheritance of discrete traits, combined with the
field of biometry, developed by Fisher, Haldane and Wright to statistically
analyze continuous traits.
In the first major work to
come out of the Modern Synthesis, Morgan and his student, the Russian
geneticist Dobzhansky, developed strains of Drosophila that could be
experimentally crossed and followed for many generations in the laboratory. By
carefully selecting mutations in different strains they were able to provide an
incredible body of experimental evidence supporting the theory that genes and
chromosomes were the basis of heritability in evolution.
Remember, we didn’t have
the molecular basis for the genetic code until the early 1960’s, so during the
1920’s and 1930’s we had to rely on observations of phenotypic traits and
careful statistical analysis of the patterns of inheritance to determine the
genetic basis of evolution. Today we tend to emphasize molecular genetics, but
during the Modern Synthesis it was the work of people like the biometricians
and the experimental population geneticists that provided the most solid proof
of the genetic basis of evolution.
Dobzhansky’s work in
particular led to concepts like genetic drift and the founder effect. When a new
population is created by a small number of individuals moving into a new area
there is a sampling bias—whenever you have a small sample size you most
probably will have only a subset of the variation in the original population.
Since the founders are the starting point for evolution in the new population,
having a limited amount of genetic variation in the initial population can
influence the distribution of allelic frequencies in the population over time.
Genetic drift has occurred
when you see random variation in allelic frequency between populations that
began with a small number of individuals in isolated habitats.
The work I showed you at
the beginning of the lecture is an example of genetic drift—a small number of
grayling make their way into isolated high mountain lakes and found
populations. These isolated populations eventually wind up with allelic
frequencies that are different than what is found in other lakes.
Another example of this
phenomenon is work that Sasha and I have done in headwater streams in the
mountains of Siberia on a type of minnow, called the Siberian Osman. This fish
moves out of steams into newly formed shallow lakes in saltpans of Mongolia and
form large populations in the new lakes from just a few founders. There is a great
deal of morphological variation in fish from different lakes, as can be seen in
the multivariate statistical analysis we have done of body measurements.
Our modern understanding
of Evolution is based on our understanding of the processes of Natural Selection
and Genetic Drift, which in turn are based on our understanding of the
importance of genetics as the mechanism of inheritance.
The Modern Synthesis was
vital to the development of the Green Revolution in agriculture, and it was
agriculture that drove the research leading to the Modern Synthesis. In the
late 1800’s and early 1900’s the US, Western Europe and Russia had an urgent
need to increase agricultural food production to meet the increasing demands of
their cities, whose populations were exploding because of the Industrial
Revolution. Feeding a nation’s cities with a decreasing number of farmers was a
vital national security issue, and increasing crop yields became a priority.
There was also increasing concern of the possibility of large-scale famine in
India, Latin America and Africa, and the persistent threat of famine in the
Soviet Union and Asia.
Up until the late 1920’s,
when the Modern Synthesis was beginning to revolutionize biology and
agriculture, Russia was keeping pace with Western science. The field of
population genetics was originally developed by Russian scientists. Dobzhansky
worked closely with the geneticist Thomas Hunt Morgan, who won the Nobel Prize
in 1933 for providing experimental evidence that chromosomes and genes were the
basis of inheritance. Under the Russian geneticist Vavilov, agricultural
experimentation was a priority of the USSR Academy of Sciences, which sent out
expeditions all over the world to collect food crops for use in genetic
research and created agricultural experiment stations like the one at Kansas
State. Russia, Western Europe, and the US were all involved in joint research,
and it was common for American scientists to work in Russian labs and for
Russian scientists to publish in English or German in European journals. This
was the situation up until 1927. Then something terrible happened—Stalin began
to persecute Russian geneticists, imprisoning and murdering many of the
scientists who contributed so much to the Modern Synthesis.
If we look at where the
Green Revolution occurred we see the problems caused by the Soviet system—
you’ll see from this map that the US and Europe benefited from the results of
the first agricultural revolution, and that parts of Latin America and Asia, as
well as the Middle East benefited from the second agricultural revolution. But
notice, even though Russian scientists were vital to the development of the
scientific breakthroughs leading to the Green Revolution, the Former Soviet
Union did not receive any of the benefits of their work. Famine plagued the
Soviet Union, particularly during the Stalin era. This was more than the result
of wars, economic policy or population movements, it was also the result of
their ideological shift away from modern science and political persecution of
The tremendous increase in
agricultural production of the Green Revolution could not have happened without
geneticists developing hybrid corn, tetraploid varieties of dwarf wheat, and
other crop varieties, something that required an advanced understanding of
genetics. The Soviets poured pesticides and herbicides on their crops,
increased irrigation, and did all of the other things that Western farmers were
doing at the time, but they couldn’t develop the improved crop varieties needed
to increase yields without using modern genetics. And Stalin rejected modern
Remember that during the
late 1800’s, after the publication of Darwin’s Origin of Species, scientists
didn’t have the tools they needed to study chromosomes and genes, and it wasn’t
known that genetics was involved in the inheritance of traits. So it was
possible to say that you were a Darwinian evolutionary biologist and believe
that something else controlled inheritance. Darwin himself believed in a number
of different mechanisms over the course of his life.
The big debate in the
early 1900’s was whether the Modern Synthesis, with its evidence of genetic
control of inheritance, should be the basis of agricultural research. Up until
1927 the Russians agreed with the Americans and Europeans that chromosomes and
genes were the basis of inheritance, which repudiated the notion of the
inheritance of acquired traits, commonly called Lamarckism. The Russians took
part in the scientific research that lead to the Modern Synthesis.
This is where Stalin and
Lysenko come in. In the 1930’s, ideologically, the Soviets during the Stalin
period wanted a view of the future of mankind that was more like the old Scala
Natural, with “Soviet Man” at the top of the ladder. They wanted a view that
allowed for the inheritance of acquired traits to justify the harsh physical
conditions they imposed on their people and their utopian claims of an ideal
future. Their political system required that their people believe that
Communism could “improve” individuals and that this improvement would be
inherited by offspring. This meant that a form of non-genetic inheritance of
acquired traits, like Lamarckism, would serve their political purposes more
than the Modern Synthesis.
The Soviets insisted that evolution
was not based on random processes like random mutation or genetic drift; the
Soviets believed that there must be a goal driven process underlying evolution.
They reintroduced teleology into evolution and adapted a non-genetic view of
Lysenko based his doctrine on folk practices of Russian farmers, which
made it easier for it to be accepted in the countryside. He came from a
peasant family and carefully crafted his work to appeal to the
ideological framework of the Stalin period.
was a reasonably good farmer, but a very poor experimentalist. His
studies could not be replicated, were based on flawed experiments, and
very small sample sizes. Russian scientists questioned their value from
the beginning, but under political repression were unable to overturn
the stranglehold that Lysenkoism had on Soviet science.
In the 1960's the gap was large enough to attract attention. With
Stalin's death in the early 1950's, Nikita Khrushchev rose to power and
made a renewed push to improve agriculture. He toured farms in the US
and observed the differences in productivity. The failure of Soviet
agriculture to come up to international standards was a factor in his
eventual loss of power. When Soviet scientists were able to incorporate
modern techniques into the agricultural sector, Soviet agriculture
Note: the fall off in the 1990's occurred during the collapse of the government and economy.
Even with the huge "Virgin Lands" campaign (see lecture on Soviet
agriculture) which brought huge tracks of prairie under the plow,
Soviet agriculture lagged behind the West.
This is a case study in the terrible problems that can occur when
politics inserts itself into science. It is also a good example of how
science has self-correcting mechanisms, in which factual information
(such as the discovery of DNA as the mechanism of inheritance) can
Societies must be ever careful
about directly interfering with science. There is a difference between
society using ethical or economic arguments to restrict application of
scientific results (for example, European nations restricting use of
GMO's in agriculture) vs. governments altering data, substituting in
pseudo-science based on poor quality research for scientifically
validated results, altering factual information-- when society loses
its ability to distinguish between political decisions and scientific
fact it risks losing everything.
Kingsland, S. E. 1985. Modeling Nature: Episodes in the History of Population Ecology. University of Chicago Press, Chicago, IL
Medvedev, Z.A. 1969. The Rise and Fall of T.D. Lysenko. Translated by I.M. Lerner. Columbia University Press, NY, NY. 184 pages
Pryde, R.R. 1972. Conservation in the Soviet Union. Cambridge University Press. 301 pages.
Vucinich, A. 1988. Darwin in Russian Thought. University of California Press, Berkeley, CA. 468 pages.
Weiner, D.R. Models of Nature: Ecology, Conservation and Cultural Revolution in Soviet Russia. 1988. University of Pittsburgh Press, Pittsburgh, PA. 324 pages.