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Copyright Ó 2010 by the Genetics Society of America
DOI: 10.1534/genetics.110.116632
Perspectives
Anecdotal, Historical and Critical Commentaries on Genetics
J. B. S. Haldane’s Last Years: His Life and Work in India (1957–1964)
Krishna Dronamraju1
Foundation for Genetic Research, Houston, Texas 77227
ABSTRACT
J. B. S. Haldane (1892–1964) was an outstanding scientist and a polymath who contributed significantly
to physiology, genetics, biochemistry, statistics, biometry, cosmology, and philosophy, although he himself
possessed no formal qualifications in any branch of science. His early science training was provided by his
father, Oxford University physiologist J. S. Haldane, and the rest was self-taught. The author came to know
him well as his student during his last years, which were spent in India (1957–1964). Haldane’s unique
intellectual qualities and the research undertaken by his associates in India during those years are described.
J
OHN Burdon Sanderson ( J. B. S.) Haldane (1892–
1964) was an exceptional scientist whose mental
powers were extraordinary. He possessed both great
intelligence and a prodigious memory and was able to
divide his attention between two entirely different
subjects simultaneously. He was able, for instance, to sit
in a lecture room, writing a mathematical paper on
evolution while closely paying attention to a lecture
on a different topic as he demonstrated during the
discussion at the end of the lecture.
Haldane’s last years in India are of interest because he
showed how an eminent scientist from the West can
transplant himself to a less developed country and still
lead a productive and creative life during his last years.
He suggested research projects that could be undertaken by utilizing local resources, requiring no expensive laboratories or equipment, while much of his own
research was of theoretical and mathematical nature
and hence not dependent on external funding.
I came to know Haldane intimately during those
years in India (1957–1964). In Calcutta, I shared a
house with the Haldanes in the suburb of Baranagore
near the Indian Statistical Institute. We traveled together all over India and Europe for various scientific
meetings. One memorable conference was held in Israel
on the ‘‘Genetics of Migrant and Isolate Populations’’
(Goldschmidt 1963). On behalf of the organizing
committee, Elisabeth Goldschmidt invited Haldane to
preside over the conference, with these words: ‘‘We may
consider ourselves fortunate indeed that Prof. Haldane
has accepted the presidency of this conference on short1
Address for correspondence: Foundation for Genetic Research, P.O. Box
27701-0, Houston, TX 77227. E-mail: [email protected]
Genetics 185: 5–10 (May 2010)
range evolution in human populations. Human population genetics is one of the several fields in which you
cannot move without encountering the approaches
and analyses of Prof. Haldane at every step . . . Some of
Prof. Haldane’s ingenious proposals . . . have kept us
busy with discussions for long years . . . it is an enormous
privilege for us to be allowed to carry on this conference
in his presence and under his guidance’’ (Goldschmidt
1963, p. 5). During the 1930s and 1940s, Haldane was
active in finding research positions for Jewish refugees
who were fleeing Nazi Germany. Prominent among them
was biochemist Boris Chain who was introduced to
Howard Florey by Haldane; Chain and Florey’s subsequent collaboration led to the Nobel Prize that they
shared with Fleming for their work on the discovery and
development of penicillin as the first antibiotic (Haldane
1961).
J. B. S. Haldane was a great popularizer of science,
contributing numerous articles to newspapers and
popular magazines in several countries. A selection of
these essays has recently been published (Dronamraju
2009). Many of these pieces were written in his ‘‘spare
time’’ while traveling on trains and planes. Haldane’s
‘‘popular’’ scientific essays are, in fact, much more than
that; they often contained original ideas that remain of
interest to scientists.
He never received a degree in any branch of science,
yet he became one of the great and influential scientists of the 20th century. Haldane, who was popularly
known as either just ‘‘JBS’’ or ‘‘Prof’’ or simply ‘‘Haldane’’ was a true polymath, a genius who possessed
intimate knowledge of multiple disciplines. He made
important contributions to several sciences as well as to
philosophy, religion, logic, popular writing, and ethics.
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K. Dronamraju
Figure 1.—J. B. S. Haldane. Photograph courtesy of Klaus
Patau, University of Wisconsin.
In science, Haldane’s contributions covered physiology,
genetics, biochemistry, statistics, biometry, cosmology,
and history and philosophy of science, leaving a significant impact on each of these disciplines. As a popular
writer of science, Haldane covered even more subjects,
including zoology, physics, chemistry, mathematics, geology, geography, astronomy, meteorology, politics, psychology, nonviolence, military affairs, theology, and literature.
In addition to his monumental work in genetics, Haldane
made important contributions to enzyme kinetics in
biochemistry.
In physiology, in particular, Haldane is best remembered for his daring experiments in testing the physiological effects of poisonous gas mixtures, cold temperatures,
and higher atmospheric pressures in which he employed
himself as his own ‘‘guinea pig.’’ They were often painful
experiments, causing convulsions and injuries. He also
conducted many diving experiments to investigate a
submarine accident for the British Navy, once again
employing himself as his own guinea pig. He suffered
permanent injury because of rapid decompression during
the experiments and back pain haunted him the rest of his
life (Behnke and Brauer 1968).
Haldane’s distinguished relatives include his father
John Scott Haldane, physiologist at New College,
Oxford University; his uncle Viscount Haldane, scholar
and politician, who was Minister for War and Chancellor
of the Exchequer in Prime Minister Asquith’s cabinet;
his great uncle Burdon Sanderson, first Waynflete
Professor of Physiology at Oxford University; and his
sister (Lady) Naomi Mitchison, prolific writer of numerous books of fiction, nonfiction, travel, history, politics,
and other subjects. (Lady Mitchison’s standing and
influence are attested by Jim Watson’s dedicating his
memoir The Double Helix to her.) Haldane’s three
nephews are distinguished scientists in the life sciences.
Haldane was born in Oxford, England, on November
8, 1892. Jack, as he was known to family members and
Figure 2.—J. B. S. Haldane surrounded by his associates,
on the campus of the Indian Statistical Institute, Calcutta,
1961, the author sitting, third from left. Photograph courtesy
of the Indian Statistical Institute.
close friends, was a precocious child whose brilliance
was legendary from an early age. His father used to take
him down mine shafts where he investigated the
physiological conditions. At the age of 3 or 4, he was
already familiar with the terminology of hemoglobins,
which he picked up from his father’s research. Once,
when he fell down and cut his head, a doctor was called
to treat the wound, and the young Haldane, looking at
the blood, asked whether it was oxyhemoglobin or
carboxyhemoglobin! As a young boy, his talent for mathematics was put to test by his father who, on one occasion, forgot his log tables on a field trip and asked his
son to calculate a set. Young Haldane did so promptly.
Haldane’s scientific eminence today is principally
based on his extensive series of papers on the mathematical theory of Darwinian natural selection (evolution). The independent contributions of Haldane and
R. A. Fisher in England and Sewall Wright in the United
States led to the establishment of modern population
genetics.
SCIENTIFIC CAREER
J. B. S. Haldane was educated at Eton and Oxford,
graduating with distinction in classics in 1915. His
further education was interrupted when he joined the
Black Watch battalion to fight in the First World War.
He fought with bravery and distinction. His life was in
constant danger when he was fighting in the trenches in
France when almost all the men in his battalion were
killed. He was saved mainly because his uncle, Lord
Haldane, was Minister for War and managed to transfer
his nephew to a safer job in Scotland. After the war in
1919, Haldane received a Fellowship in Physiology at
New College, Oxford, where he pursued research in
both physiology and genetics. In 1923, he was appointed
to the Dunn Readership in Biochemistry under Pro-
Perspectives
fessor F. G. Hopkins at Cambridge University. In
addition to research and teaching in biochemistry,
Haldane found time to write most of his important
mathematical papers on the ‘‘mathematical theory of
natural selection,’’ which helped found population
genetics along with the independent contributions of
Fisher and Wright. Simultaneously, he conducted outstanding research in physiology and biochemistry, deriving the law of steady-state kinetics in enzyme chemistry.
Haldane (1932) summarized his early research in
population genetics in his book The Causes of Evolution.
He continued to publish brilliant papers subsequently,
many of them becoming the foundations for new areas of
research: genetic loads and the cost of natural selection
(Haldane 1957), infectious disease and selection, estimation of human mutation rates, linkage and human
gene mapping, rates of evolution (Haldane 1949), and
the biochemistry of gene action, to name a few.
From 1933 to 1937, he was Professor of Genetics at
University College, London. In 1937, he was elected to
the Weldon Chair of Biometry where he continued to
each and direct research in biometry, statistics, human
genetics and several other subjects. In 1957, both Haldane
and his wife migrated to India to accept an invitation
from Prof. P. C. Mahalanobis, Director of the Indian
Statistical Institute in Calcutta. Mahalanobis was a close
friend and advisor to the then Prime Minister of India,
Jawaharlal Nehru.
Haldane’s numerous publications include more
than 400 papers (for most of which he was the sole
author) and 24 books as well as several hundred popular
articles. Summaries of his life and work can be found in
Dronamraju (1968, 1985, 1987, 1990, 1995, 2009) and
Crow (2004).
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Upon his arrival in 1957, he initiated at once several
research projects of an ecological and biometrical
nature. These are briefly described below. Furthermore,
as a classical scholar, he was drawn to India’s ancient
civilizations, religions, languages, and cultures. He often enjoyed quoting passages and verse in several Indian
languages during his scientific lectures, just as he used
to quote from Latin and Greek classics in Europe.
The job offers for both Haldane and his wife (his
former student and biologist Helen Spurway) came at
an opportune time from the Indian Statistical Institute
in Calcutta. He was close to retirement from University
College London. Furthermore, he was looking forward
to enjoying the warm climate and clear sky in India. One
of his great passions was observing the stars in the Indian
skies.
LIFE AND RESEARCH IN INDIA
Haldane was a Research Professor at the Indian
Statistical Institute in Calcutta from 1957 to 1961, where
four of us, S. K. Roy, T. A. Davis, S. D. Jayakar and myself,
closely worked with him as his research associates. I
completed my research for a Ph.D. in genetics, which was
awarded later. When political differences with the Director, P. C. Mahalanobis, arose later, Haldane resigned
and moved to Bhubaneswar in the State of Orissa, where
the Chief Minister (equivalent to a state Governor in the
United States) offered to build him an Institute for
Genetics and Biometry. However, his remaining life was
too short. He died there on December 1, 1964 after
cancer surgery. I worked closely with Haldane both at
Calcutta (1957–1961) and at Bhubaneswar (1962–1963).
DARWIN AND WALLACE IN INDIA
REASONS FOR MOVING TO INDIA
Complex reasons motivated Haldane to abandon his
position at University College London and move to
India in 1957. His professional reputation and prestige
were at their maximum, yet Haldane chose to turn his
back on Europe and other active centers of scientific
research in the world. The most important reason was
undoubtedly his increasing dissatisfaction with the
politics of Great Britain at that time. He was not
comfortable living in the east or west block during the
cold war years. He was attracted to the neutralist policy
of Prime Minister Jawaharlal Nehru of India. Several
years earlier, he had abandoned Marxist politics and was
not interested in living in any countries of the communist bloc.
Another important reason was his desire to pursue
research on Indian plant and animal species and human populations. Ever since his first visit to India in 1917,
when he was sent to a British Army Hospital to recuperate
from wounds he received in Mesopotamia (Iraq), Haldane
frequently mentioned his desire to return to India.
Haldane’s genetic research was primarily influenced
by the Darwinian tradition. His major contribution to
science was his mathematical theory of evolution (natural
selection), which was an interpretation of Darwin’s
theory of evolution in the context of Mendelian genetics.
After his arrival in India, Haldane saw Darwin from a new
perspective.
Haldane regarded Darwin’s work in botany as his most
original and important contribution to science. He wrote:
‘‘In my opinion, however, Darwin’s most original contributions to biology are not the theory of evolution but the
great series of books on experimental botany published
in the latter part of his life’’ (Dronamraju 2009, p. 212).
That work includes books on insectivorous plants, climbing plants, sexuality in plants, and inbreeding in plants.
Charles Darwin is best known for his work on evolution.
However, that subject had also been discussed by many
others including his grandfather, long before Darwin. His
experiments on plants, although not so well known, are
completely original. Haldane encouraged us to follow
Darwin’s research in botany. But Haldane wondered what
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K. Dronamraju
Darwin’s attitude was that led to his discoveries. Darwin
was most open minded in his attitude toward all living
beings. Haldane wrote: ‘‘Darwin did not draw a sharp line
between earthworms and the old gentlemen who had
failed to interest him in mathematics at Cambridge’’
(Dronamraju 2009, p. 213). From the biographical account of his son Francis, we know that Darwin was totally
immersed in his experiments and observations. Indeed,
he was so dedicated that he used human attributes when
describing his experimental animals and plants.
Many of the research projects that were undertaken
under Haldane’s direction were quantitative biological
studies. Haldane had earlier come to the conclusion
that the denseness of the tropical flora could result
in interdependence involving both competition and
cooperation between species. Haldane wrote: ‘‘One
would expect symbiotic relations between flowering
plants to be much commoner in regions with a diverse
flora than in those where a natural plant community
often consists of few members . . . if such symbiotic
regulations are common, they should be looked for
between different genotypes of the same species’’ (see
Dronamraju 1987, p. 221). The experiments of my
colleague, S. K. Roy (Roy 1960), were concerned with
the effect of symbiosis between different varieties of rice
on their grain yields. Two varieties may be planted either
fully mixed, in alternate rows, or in separate halves of
the same plot surrounded by a dam. In 31 experiments,
only 5 showed a gain of .10% over the mean of the two
varieties, and all of them showed some gain over the
better yielding of the two.
Some research projects followed what Bateson
(1894) called ‘‘meristic variation,’’ that is to say, variation
in like parts, such as petals, vertebrae, or teeth, in which
Haldane (1958) was deeply interested. It remained for
Gruneberg (1952), an associate of Haldane’s at University College London, to show that it is akin to the
variation in size of the third upper molars in some
strains of mice, which he termed ‘‘quasi-continuous’’
variation. In India, Haldane’s associates studied meristic
variation in local plant and animal species. For instance,
Roy (1963) studied the variation in petal numbers on
plants of Nyctanthes arbor-tristis and Jasminum multiflorum.
The plants showed floral teratology toward the end of
the flowering season. Double flowers occurred frequently in late season. The variance increased toward
the end of the flowering season. One of my studies was
concerned with variation in the style lengths of flowers
of Bauhinia acuminata. I also noted heterostyly in this
species, which involved, unlike the classical heterostyly
described by Darwin in Primula, both short- and longstyled flowers occurring on the same plant and the
short-styled flowers being female sterile (Dronamraju
1960a). In his Presidential address to the Centenary and
Bicentenary Congress at the University of Malaya in
Singapore (December 2, 1958), Haldane stated: ‘‘It was
(Alfred Russell) Wallace, above all, who saw that certain
biological problems are forced on one by a study of
tropical life, which are by no means so obvious in
temperate climate’’ (Haldane 1960, p. 713).
ANIMALS HAVE RIGHTS AND DUTIES
Haldane stated that Christian theologians drew a
sharp distinction between humans and other species.
Hence, Darwin’s argument that human beings have
descended from animals created a furor among Europeans (Dronamraju 2009, p. 211), whereas many eastern
religions such as Hinduism, Buddhism, and Jainism have
long recognized that animals have rights and duties.
Indian art and religion depicted divinity in various animal
forms. Kindness toward other species has long been an
accepted tradition, at least in theory, if not always in
practice, in India. There are no parallels in the traditions
of Europe and North America. Furthermore, there is still
some opposition to teaching evolution in some Christian
communities, especially in the southern United States.
This is in contrast to the situation in India where many
Hindus see similarities between vertebrate evolution and
the successive incarnations of Vishnu.
Not only did Haldane interpret Darwinism from the
Indian philosophical and religious point of view, but
also he developed several research projects on Indian
species of plants and animals that are in the Darwinian
tradition. As his students, we were all encouraged to
conduct research on different projects in that tradition.
There were certain characteristics that are common to
all these projects. Quantifying results as much as
possible and the utilization of very simple methods were
characteristic of Haldane’s approach to science: (a)
Much of our research involved measuring biological
variation, especially meristic variation, that is to say,
variation in like parts such as petal numbers in flowers,
venation in leaves, number of teeth, or bilateral spotting
in caterpillars, etc.; (b) research involved local resources
as much as possible; (c) the projects did not require
much financial support; (d) the methods employed
were simple but the results had scientifically profound
consequences; (e) the observations must be quantified
wherever possible; (f) the project should be flexible
enough to facilitate the incorporation of any new and
unexpected results and the resulting change in direction
or methods; (g) it is necessary to have several ongoing
projects simultaneously to achieve quick results as there
will always be some lines of pursuit that may take longer
or not bear any fruit at all; and (h) the results should
be published immediately.
Some of the projects were an imitation of Darwin’s
experiments. One such example is a project by Roy
(1957) on the estimation of the volume of soil brought
to the surface by earthworms per acre per year and its
ecological implications. Another project, which I followed, is the investigation of color preferences of lepidoptera among different colored flowers and their
Perspectives
impact on speciation (Dronamraju 1960b, 1985). By
their preferential visits to different varieties of a plant
species differing in flower color, insect pollinators can
create reproductive isolation within a species. By means
of increasing such isolation (for instance, it could
happen if there is a new flower color mutant appearing
in the plant population) further divergence can follow,
ultimately leading to complete isolation and speciation.
Such speciation has been called ‘‘sympatric speciation,’’
as opposed to ‘‘allopatric’’ or ‘‘geographic speciation,’’
where such geographic factors as floods, earthquakes,
and simple geographic distance can result in reproductive barriers within a species, as emphasized by Haldane’s
friend Ernst Mayr (Mayr 1942). Haldane believed that
such research projects followed the Darwinian tradition
and would have interested Darwin greatly. Another colleague, T. A. Davis (Davis 1962) pursued research on
coconut palms, showing that foliar asymmetry in their
crowns is not inherited. Surprisingly, Davis (1963) discovered that palms with left-handed coiling in their
crowns produce more nuts than those palms with righthanded spiraled crowns.
Among other studies, Helen Spurway (Mrs. Haldane)
conducted biometrical investigations of the life cycle of
the tussore silk moth, Antheraea mylitta (Spurway and
Dronamraju 1962), and the nest-building activity of the
solitary wasp, Sceliphron, in collaboration with Jayakar
and Dronamraju (Spurway et al. 1964). Anthropologist
A. K. Ray conducted population studies of toe anomalies
and their possible genetic basis. Haldane commented
that such studies are easier to follow in India because
many people have bare feet or wear sandals, unlike
western countries where shoes and boots are worn by
many (Ray and Haldane 1965).
Haldane drew attention to Aristotle’s logic and his
difficulties in classifying animals, which he based on
similarities (Dronamraju 2009, p. 214). Darwin, on the
other hand, foreshadowed a logic based on differences.
In The Origin of Species, Darwin (1859, p. 268) wrote:
‘‘Systematists will have only to decide (not that this will
be easy) whether any form be sufficiently constant and
distinct from other forms, to be capable of definition,
and if definable whether the difference be sufficiently
important to deserve a specific name . . . Hence, without
rejecting the consideration of the present existence of
intermediate gradations between any two forms, we shall
be led to weigh more carefully and to value higher the
actual amount of difference between them.’’ Haldane
regarded Darwin’s statement as the starting point for a
whole new program in statistics (Haldane 1959).
THEORETICAL POPULATION GENETICS
In several papers, some in collaboration with S. D.
Jayakar (Haldane and Jayakar 1963), Haldane returned to finding solutions for the unsolved problems
of theoretical population genetics that he first initiated
9
in the 1920s. The titles of some of these papers are ‘‘The
selection of double heterozygotes,’’ ‘‘Natural selection
in a population with annual breeding but overlapping
generations,’’ ‘‘Polymorphism due to selection of varying direction,’’ ‘‘The elimination of double dominants
in large random mating populations,’’ and ‘‘Natural
selection in a population with annual breeding but
overlapping generations.’’ At the same time, Haldane
enjoyed returning to nature as was evident in his participation, with Helen and Jayakar, in bird watching,
observing wasp and fish behavior, and other studies of
an ecological nature.
Throughout his life, Haldane enjoyed being the human
‘‘calculator’’ for the data collected by his colleagues. He
followed the same practice in India. He enjoyed nothing
better than to sit in his arm chair and analyze data
collected by others and devise novel statistical tests, often
extracting more information than the investigator had
hoped for. He was fond of doing complex sums in long
hand, refusing to use even the simplest calculators that
were available then.
Haldane rarely took time out from his mathematical
labors. However, he indulged himself occasionally, for
instance, in writing such pieces as ‘‘A defense of
beanbag genetics’’ (Haldane 1964a) in response to
Mayr’s (1959) criticism of the early papers in beanbag
genetics (see Dronamraju 2010). Mayr criticized the
mathematical theory of natural selection, which was
founded by Haldane, Fisher, and Wright, because genes
were treated, for the sake of mathematical convenience,
as noninteracting independent units. In Mayr’s view,
this was similar to the work of early Mendelians who
kept beans of various colors in different bags while
counting Mendelian ratios. With his tongue firmly in
cheek, Haldane enjoyed writing a ‘‘defense’’ of his work,
arguing that the mathematical theory provided a ‘‘scaffolding’’ within which a reasonably secure theory expressible in words may be built up. It is amusing to read his
justification, ‘‘Fisher is dead, but when alive preferred
attack to defense. Wright is one of the gentlest men I
have ever met, and if he defends himself, will not
counterattack. This leaves me to hold the fort, and that
by writing rather than speech’’ (Haldane 1964a, p. 344).
HALDANE AT THE END
Haldane’s poem, ‘‘Cancer is a Funny Thing,’’ which
was written shortly before his death and published in
many popular magazines and newspapers, reflected
his courage when facing almost certain death. While
he was visiting the United States in October 1963 to
attend a conference on the origin of life in Florida,
Haldane found out that he was suffering from rectal
bleeding. On his return journey, stopping in London,
Haldane found out that he had rectal cancer and was
operated on instantly at University College Hospital.
While recuperating in London, he wrote the poem
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K. Dronamraju
‘‘Cancer is a Funny Thing,’’ which amused and irritated his readers in equal numbers. I quote a few lines
(Haldane 1964b):
I wish I had the voice of Homer
To sing of rectal carcinoma,
Which kills a lot more chaps, in fact,
Than were bumped off when Troy was sacked.
Yet, thanks to modern surgeon’s skills,
It can be killed before it kills
Upon a scientific basis
In nineteen out of twenty cases.
My final word, before I’m done,
Is ‘‘Cancer can be rather fun’’.
Thanks to the nurses and Nye Bevan
The NHS2 is quite like heaven
Provided one confronts the tumour
With a sufficient sense of humour.
I know that cancer often kills,
But so do cars and sleeping pills;
And it can hurt one till one sweats,
So can bad teeth and unpaid debts.
A spot of laughter, I am sure,
Often accelerates one’s cure;
So let us patients do our bit
To help the surgeons make us fit.
CONCLUDING REMARKS
Haldane’s life and activities in India left an indelible
impact on Indian science. He was especially inspiring to
the younger generation who were happy to see him
challenging the authorities and bureaucracy when they
were not able to do so themselves. The research projects
undertaken by his colleagues provided a new impetus to
the development of certain branches of science, such as
biometry, population genetics, human genetics, and
animal behavior.
I am most grateful to James F. Crow for his thoughtful comments
and suggestions.
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Crow, J. F., 2004 Haldane’s ideas in biology with special reference
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2
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