© 2004 The American Genetic Association
The Wilhemine E. Key 2003 Invitational Lecture
Genetics: Alive and Well. The First Hundred Years as Viewed Through the Pages of the Journal of Heredity
From the Department of Genetics, University of Wisconsin, 445 Henry Mall, Madison, WI 53706.
Address correspondence to James F. Crow at the address above, or e-mail: jfcrow{at}wisc.edu.
James F. Crow is Professor Emeritus of Genetics at the University of Wisconsin–Madison. He was born in Philadelphia, grew up in Wichita, Kansas, did graduate work at the University of Texas–Austin (PhD 1941), taught at Dartmouth College, and since 1948 has been on the faculty of the University of Wisconsin. He has honorary doctorates from the Universities of Chicago and Edinburgh. He is a member of the National Academy of Science, National Academy of Medicine, and American Philosophical Society. He is a foreign member of the Japan Academy and the Royal Society of London. He is past president of the Genetics Society of America and the American Society of Human Genetics. His research has been in theoretical and experimental population genetics. He has written a number of articles and three books—Genetics Notes, Basic Concepts in Population, Quantitative, and Evolutionary Genetics, and Introduction to Population Genetics Theory (with M. Kimura). His hobby is music, and for many years he played viola in the Madison Symphony Orchestra.
2003 is the centennial year of the American Genetic Association, and what a century this has been! 1903 was a banner year in the history of genetics. This was the year in which Sutton published his famous paper putting together chromosome behavior and Mendel's laws (Sutton 1903). This can be regarded as the beginning of genetics as a science. 1903 was also the year in which the American Breeders Association was founded. This soon became the American Genetic Association (AGA), and in 1914 the AGA began publication of the Journal of Heredity. The century, 1903–2003, ushered in by these two events, has seen genetics grow from very modest beginnings into a powerful science that permeates throughout biology. My intention in this lecture is to trace the history of the subject by citing articles that have appeared over these years in the two publications of the AGA.
Although founded in 1903, the AGA did not have a regular publication until 1910, when the American Breeders Magazine started. This continued through 1913, and in 1914 changed its name to the Journal of Heredity. During my childhood in the 1920s, my father subscribed to this journal and I enjoyed looking at it. The main attraction was the pictures, some of which came vividly back to mind recently as I looked through old issues. So my association with the AGA began a long time ago and still continues. It is a pleasure to talk about it.
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Journal of Heredity |
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 Top Journal of Heredity The Wilhemine Key LecturesWalter S. Sutton (1877-1916)Representative Items in...The History of Genetics...References |
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The new journal began, not with a whimper, but with a bang. The first article was written by Alexander Graham Bell, famous for having invented the telephone some 40 years earlier. Bell had a deep interest in heredity, especially inheritance of deafness. He had astutely noticed the high rate of consanguinity among normal parents of the deaf. As were a great many students of heredity at the time, Bell was interested in eugenics. His leadoff article was entitled, "How to Improve the Human Race." Bell argued that there was little to be gained by preventing marriage of "undesirables;" he believed that assortative mating among the better endowed was much more important.
Inventing the telephone was only one part of his active life. He produced all sorts of inventions, including an electric device for locating metal in human tissue. It was used to locate the bullet after President Garfield was assassinated. Bell had a deep interest in livestock breeding and tried to develop a strain of sheep with a high rate of twinning. (This was long before the Booroola gene was discovered.) In addition to his passion for genetics and invention, Bell had a deep interest in geography and was president of the National Geographic Society. His son in law, Gilbert Grosvenor, became assistant editor of the National Geographic Magazine in 1899 and editor 4 years later. While editor, he built its circulation from 900 to 2 million. I think it is fair to say that he was the founder of photojournalism and the magazine soon became known for its vivid photographs. Sewall Wright once told me that Bell also hoped for an equally popular magazine devoted to genetics, and thought of another son in law, David Fairchild, as one to take the lead. Apparently nothing came of this at the time, but a few years later Fairchild was president of the AGA when the Journal of Heredity started. He was a distinguished botanist and, while working for the U.S. Department of Agriculture, was responsible for the introduction of thousands of plant species. Both Bell and Fairchild were members of the Council of the AGA. The only other member who is known to present-day geneticists was William Castle, who was vice president. Of course, Journal of Heredity never attained the popularity of National Geographic, but it held its own among scientific journals and quickly became known for its lively style and photographs.
The first editor was Paul Popenoe. Popenoe was widely known among American animal breeders and students of heredity. Journal of Heredity reflected his enthusiasm for the now-controversial subject of eugenics. As late as 1938, while a graduate student at the University of Texas, I heard him lecture on the subject. He was an enthusiastic and charismatic advocate, but by then the American eugenics movement was in rapid decline. Later he became a marriage counselor and newspaper columnist.
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The Wilhemine Key Lectures |
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TopJournal of HeredityThe Wilhemine Key LecturesWalter S. Sutton (1877-1916)Representative Items in...The History of Genetics...References |
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The Wilhelmine Key Lectures began in 1962 with a gift from Dr. Key. She is best known for being the teacher of Sewall Wright. She encouraged him to read Punnett's article on genetics in the Encyclopedia Britannica. She was enthusiastic about research and opened his mind to the possibility of doing research. He often said that he owed a great deal to her. The two communicated throughout her life. She expressed admiration for his work, but rued her inability to understand his mathematics.
Wilhelmine Enteman was born in 1872 and grew up in Wisconsin. She attended the University of Wisconsin and as an undergraduate worked with the pioneer limnologist, E. A. Birge. After graduation in 1894, she taught in Green Bay for 4 years and then became a graduate student at the University of Chicago. There she worked with C. O. Whitman and was especially impressed by C. B. Davenport. After her doctorate she taught biology and nature study at New Mexico Normal. At about this time she married Francis B. Key, a descendant of Francis Scott Key. The marriage was short lived, however, for her husband was an invalid and did not survive long. She then taught at Belmont College and finally went to Lombard College in 1909. It was during his senior year in 1910–1911 that Wright came under her influence. In 1912 she left to rejoin Davenport, who by that time was in the Eugenics Record Office at Cold Spring Harbor, New York. Later she moved to the Race Betterment Foundation in Battle Creek, Michigan. She was deeply interested in human betterment and soon was active as a field worker. She wrote a number of articles on eugenics, some in Journal of Heredity. I have the impression that, in contrast to many early eugenicists, she was less of an advocate and more of a student. For more about her see Wright (1965).
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Walter S. Sutton (1877–1916) |
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TopJournal of HeredityThe Wilhemine Key LecturesWalter S. Sutton (1877-1916)Representative Items in...The History of Genetics...References |
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As I mentioned earlier, genetics as a science began in 1903 when Walter Sutton (Figure 1) showed that meiotic chromosomes follow a behavior pattern that is consistent with Mendel's laws. Actually he was on the track a year earlier when he concluded an article on grasshopper chromosomes with the statement: "I may finally call attention to the probability that the association of paternal and maternal chromosomes in pairs and their subsequent separation during the reducing division...may constitute the physical basis of the Mendelian law of heredity" (Sutton 1902). Sutton's definitive paper, with his beautiful drawings of grasshopper chromosomes came soon after (Sutton 1903). Shortly after, the same idea was published by Boveri, who had thought of it independently. I think it likely that several biologists realized this, but it was Sutton and Boveri who presented the convincing evidence.
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Sutton grew up on a Kansas farm known for its high-quality livestock. He demonstrated an inventive streak early in life by repairing farm equipment and making a camera. He attended the University of Kansas and there became a student of C. E. McClung, who encouraged Sutton to study the chromosomes of grasshoppers, which he collected on his farm. There was no shortage of grasshoppers in Kansas! It was his thorough knowledge of chromosome behavior that led to his happy insight. Sutton was clearly a favorite of McClung, and as far as I know, made only one mistake. At the time it was known that the male grasshopper has 23 chromosomes; it has an X and no Y. Sutton undertook the much more difficult task of determining the chromosome number in the female, and he counted 22 rather than the correct number, 24. Chromosome identification wasn't as easy in those days as it is now. This error led McClung to the conclusion that the X chromosome is male-determining. There was considerable confusion about sex chromosomes and sex linkage in those years. A further complication was that the early studies of sex linkage were done with moths, in which the female is heterogametic, whereas the chromosome studies were on insect species with heterogametic males. It was finally clarified by simultaneous cytological and genetic studies on Drosophila.
Sutton dropped out of graduate school, probably for financial reasons, and went to work in the Kansas oil fields. There he continued his inventive ways, making several improvements in the drilling machinery. He eventually returned to school, but this time in medicine. He became a distinguished surgeon, and was just as inventive in this field as he had been in genetics and engineering (Crow and Crow 2002; McKusick 1960). For one thing, he was the first to use rectal anesthesia, necessary for facial surgery, and invented a "speedometer" to accurately measure the delivery rate.
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Representative Items in Successive Periods in Publications of the AGA |
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TopJournal of HeredityThe Wilhemine Key LecturesWalter S. Sutton (1877-1916)Representative Items in...The History of Genetics...References |
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The articles in Journal of Heredity and its short-lived predecessor, American Breeders Magazine, provide a vivid view of the changing field of genetics during its first century. From the beginning the emphasis in these journals has been mainly on organismic genetics in a wide variety of animals and plants, and in the early years, on eugenics.
I have compiled a list of topics covered during successive periods in the history of the AGA. It is certainly not a random sample, nor is it systematic. It simply reflects what seem to me to be interesting items characteristic of their period. They provide a series of snapshots of how genetics has grown and changed during its first century.
Items reported as novel in the early days are now commonplace. Some early interpretations are now known to be wrong. An example is the sterility of tortoiseshell tomcats (see later). Cytogenetic explanations, now obvious, were overlooked in the early days. Counts of the human chromosome number were wrong for several years. Mutator genes seemed utterly mysterious; now they are usually caused by transposable elements. And of course, the methods have become increasingly biochemical and molecular over the years.
Yet some of the earliest observations have stood the test of time. The effectiveness of selection for oil content in corn was already evident in the first issue, as was Shull's prescient suggestion of inbreeding and hybridization for improving corn yield. Salivary gland chromosomes, discovered in the 1930s, are still a mainstay in dipteran cytogenetics. Colchicine is still used to induce polyploidy. And so on. Here, then, is a list of topics that seems to me to reflect their times.
American Breeders Magazine, 1910–1913
In the first issue there were portraits of Darwin, Mendel, and Cruikshank. Cruikshank was a Shorthorn breeder whose records provided data for Wright's analyses and led ultimately to his shifting-balance theory of evolution.
A report from the University of Illinois of selection for oil and protein in maize, with striking results, even at this early stage. The program, which began before the rediscovery of Mendelism, is still ongoing.
G. H. Shull's suggestion for using inbreeding and hybridization to improve corn yield. Ironically this was done in Connecticut, not in the corn belt.
Selection for egg laying in chickens by Raymond Pearl.
Zebu-cattle hybrids were found to be resistant to Texas fever.
Castle reported on the early stages in his selection experiments for amount of white in hooded rats.
L. J. Cole reported that all sex-limited characters in birds behave as if the female is heterozygous. Apparently he didn't consider sex chromosomes.
Journal of Heredity, 1914–1919
Several articles on eugenics, reflecting the views of the enthusiastic first editor, Paul Popenoe.
Continuation of Castle's selection for amount of white in hooded rats.
Human baldness as a sex-limited dominant trait.
Sterility in mules explained by aneuploidy.
Spineless cactus as a possibility for livestock feed in desert areas.
Sex reversal in chickens.
A series of articles by Sewall Wright on comparative color inheritance in mammals. They are forerunners of Wright's later articles applying enzyme kinetics to developmental genetics.
Polyembryony in insects and later in the armadillo by J. T. Patterson.
Cattle–bison hybrids and their promise as ranch animals for cold climates.
An amusing "living normal distribution" for height by Blakeslee (1914) (Figure 2).
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An Interlude, 1916
1916 marked the beginning of a new journal, Genetics. Until this time, Journal of Heredity was the only American journal devoted exclusively to the subject, although American Naturalist and Science included articles on genetics. In contrast to Journal of Heredity, which emphasized high-quality illustrations, short notes, comments, and organismic research, Genetics was strictly a journal of basic research. The first article was a blockbuster, Bridges' paper on nondisjunction as proof of the chromosome theory of heredity. By this time the theory was generally accepted, but Bridges provided the clinching evidence, and a lot more.
An aside: Genetics and I were both scheduled for January 1916. I arrived on time, but Genetics was a few months late.
Journal of Heredity, 1920–1929
The early years of Journal of Heredity were difficult ones. Although the circulation in 1922 was 3625, and the society received some support from the National Geographic Society, it was still having trouble making ends meet.
There was a series of articles on different mutants in maize, which by this time was second only to Drosophila as a favorite organism for genetic research.
Sewall Wright did a thorough analysis of the genetic history of a number of breeds of livestock.
In several issues there were discussions of tortoiseshell male cats. It was realized that they were XX, but their sterility was attributed to maternal hormones. Now, of course, we know that they are XXY.
Serebrovsky reported linkage of rose comb and Creeper in chickens.
Several studies of radiation effects were done following Muller's discovery in 1926.
Painter and Patterson used translocations to correlate genetic and cytological changes.
Brink reported semisterility in maize, but didn't consider the possibility of a translocation.
Newman reported on identical twins reared apart; this subject reappeared with additional data in several subsequent years.
Blakeslee (one of the most frequent contributors to early issues) used haploids to find recessive mutations in Datura (Figure 3).
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Journal of Heredity, 1930–1939
T. S. Painter reported studies from several laboratories supporting 48 as the human chromosome number.
Serebrovsky and Dubinin postulated step allelism (multiple linear gene structure) based on overlapping scute phenotypes.
Iljin reported temperature-sensitive skin color enzymes in Himalayan rabbits.
Timofeeff-Ressovsky reported on radiation-induced reverse mutations. The question of whether X-rays produced "true" mutations, as opposed to deletions, was in the air and reversibility was one criterion.
Demerec reported mutable genes in maize, and later in Drosophila virilis, which now would be attributed to transposons.
Painter and Bridges reported their discovery of salivary chromosomes in Drosophila and construction of cytological maps.
Whiting reported on the sex-determining locus in haplodiploid Habrobracon.
Mangelsdorf and Reeve reported their theory of the origin of domestic maize. They hypothesized that maize was derived from an extinct or unknown ancestor that, by crossbreeding with tripsacum, produced maize. Beadle argued that teosinte was the ancestor of maize, which has turned out to be correct.
Carl Lindegren reported on spore analysis in Neurospora, showing the relation of spore order to meiotic events and providing a simple demonstration of four-strand crossing over. It provided the background knowledge for the later studies of Beadle and Tatum.
Blakeslee reported doubling chromosome numbers with colchicine, providing a new tool for the study of polyploidy.
Blakeslee again, this time on inheritance of the ability to taste phenylthiocarbamide.
Mohr reported on dominant inheritance of woolly hair. Articles on monogenic human traits were frequent throughout most of the journal's history.
Wright reviewed Fisher's "Genetical Theory of Natural Selection" and took the opportunity to criticize Fisher and espouse his own shifting-balance theory. Although Wright and Fisher were friendly correspondents at this time, their differences later became bitter and personal.
R. C. Cook, Editor from 1922–1962
Cook was the most influential of the early editors, serving for 40 years (Figure 4). His enthusiastic advocacy of eugenics had a large influence on the journal's content, which in the early years had a large number of essays and notes on this subject, including several by Wilhelmine Key. Cook's interests in genetics were catholic. He tried to include a wide variety of plants and animals, although usually these were angiosperms and vertebrates. He favored reports of human traits with Mendelian inheritance. He liked to include a variety of items of genetic interest: pertinent quotes, news stories, announcements, and reports of unusual findings (such as fertile mules or tortoiseshell tomcats).
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He also enjoyed poking fun at nonsense (Figure 5), and his idiosyncratic sense of humor frequently showed through (Figure 6). His comment after the 1932 International Genetic Congress is typical of his reporting: "Oceans of words were spilled in formal and informal gatherings to discuss the vital question: 'What is the gene?' but that important entity is still elusive. Perhaps in 1937 the answer may be forthcoming." He was too optimistic; Watson and Crick's discovery was two decades later.
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Journal of Heredity, 1940–1949
Lush and Hazel reported on three methods of selection and concluded that index selection is most effective.
Bridges published more detailed salivary maps; these set the standard.
Snyder published a linkage map of the X chromosome, based on an idea from Haldane that part of the X is homologous with the Y. Although Haldane's procedure gave misleading results, he came close to anticipating the more recently discovered pseudoautosomal genes.
Owen and Cole reported "naked" (featherless) pigeons. The idea that these "self-plucking" birds might be a boon to the squab market was frustrated by the females' reluctance to mate with naked males.
Wiener discussed rare Rh alleles. The Fisher notation had not yet caught on.
Castle reported that palomino horses are heterozygotes.
Muller reviewed Schrödinger's "What is Life," a book that had a great impact and steered a number of physical scientists into genetics. Muller was not overly impressed, partly because Schrödinger credited others with research and ideas that should have been attributed to Muller. Muller also thought that Schrödinger's philosophy approached mysticism.
Coat color inheritance in ranch mink. This is one area where the simplest Mendelian genetics paid off in practical results.
R. A. Brink and his colleagues used embryo culture to bypass endosperm incompatibilities and in this way obtained hybrids between Hordeum and Secale.
An announcement of Muller's Nobel Prize in 1946.
A remarkable paper by Prokofyeva-Belgovskaya, pointing out a difference between mother and daughter chromosomes in binucleate cells. This foreshadowed modern work in intestinal tumors in which there are differences between cells containing the template DNA and those with copies.
Another Interlude, Evolution and American Journal of Human Genetics
During the 1940s there was rapidly growing interest in two areas. One was evolution, and mainly due to the driving force of Ernst Mayr, a new journal, Evolution, was launched in 1947. Mayr was the editor and the first article was by Dobzhansky on natural selection in wild populations of Drosophila. At about the same time there was increasing interest in human genetics. A strong moving spirit was H. J. Muller, who was the first president of the American Society of Human Genetics. The American Journal of Human Genetics was edited by Charles Cotterman and the first article, in 1949, was by James V. Neel on carrier detection. Muller's influential paper, "Our Load of Mutations," appeared in the third issue.
The number of American journals in the field grew from one to four in about half a century. Since then there has been an explosion of new ones that shows no signs of abating. Sewall Wright told me that at one time he read every article in the field of genetics. It was a simpler world in those days.
Journal of Heredity, 1950–1959
The emphasis on eugenics is rapidly declining.
DDT resistance in houseflies.
Parthenogenetic turkeys.
H. J. Muller on radiation risks. This was a major crusade for Muller for the rest of his life.
"Snorter" dwarf cattle, caused by a recessive gene that is surprisingly common. It was speculated that there had been human selection for heterozygotes because of their desirable body conformation.
Sex determination in honeybees follows the same pattern as Habrobracon; that is, there is a locus that when hemizygous or homozygous produces males, but when heterozygous produces females.
A new breed of cattle, Santa Gertrudis, was developed; it was reputed to be especially suitable for hot climates in the southern United States.
A special issue devoted to The Jackson Laboratory. This was the first of several such articles.
Adipose, a mouse resembling obese, but caused by a different gene. These were forerunners of their human counterparts, discovered later and now popular research subjects.
Journal of Heredity, 1960–1969
With Barbara Kuhn (Figure 7) taking over the editorship, the subject of eugenics was essentially dropped. The changed view is reflected in this quote from Roger Williams, a biochemist: "The most important task of human genetics is to gain understanding of the potentialities of human genotypes so as to make possible the intelligent manipulation of the environment."
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The journal adopted a larger size format and added more color photographs.
Salivary chromosomes in the mosquito.
Monosomy is found to be useful in a number of plants.
Another article on fertile mules by Benirschke, now book review editor for Journal of Heredity.
The high incidence of albinism among Cuna Indians.
There were strong arguments about human chromosomes. The issue was, can the individual chromosomes be distinguished or only grouped into classes? Human cytology in those days required great skill. Now with painting it is elementary.
Audiogenic seizures in mice.
Nuclear transplants in frogs, foreshadowing cloning.
A chromosome map of the tomato, later providing the basis for quantitative trait loci (QTL) mapping.
Further analysis of complex inversions in Drosophila pseudoobscura.
A review article on the Bussey Laboratory at Harvard.
Journal of Heredity, 1970–1979
Concern for radiation risks was largely replaced in the journal by concern for environmental chemicals.
Marvin Legator discussed innovative tests for mutagenic chemicals.
Colchicine-induced polyploids were commonly discussed, for example, an article on polyploid day lilies.
Disomic and trisomic Aspergillus. Articles on lower plants become more common.
An example of pleiotropy in mink: a pastel coat color also causes deafness due to otolith deficiency.
Several articles on housefly genetics.
YY male goldfish produced by crossing XY females, obtained from sex-reversing hormone treatment, with XY males.
Hallauer and others report sophisticated estimates of dominance, heritability, and epistasis for ear number in maize.
Parthenogenetic lizards.
Clement Markert produced hexaparental and octoparental mice by fusing embryos.
Triploid mosquitoes; XXY is male.
Molecular evolution in birds. Molecular techniques are increasingly prominent.
Transposon-induced hybrid dysgenesis in Drosophila.
Journal of Heredity, 1980–1987
The journal has now become almost entirely a journal of research. Color pictures are prominent, especially on the cover. The research reported is increasingly basic, with more emphasis on molecular methods. There are still book reviews, but there is a new section on computer programs.
Two-dimensional electrophoresis to detect human mutations.
Population structure of sponges, again reflecting an increasing interest in invertebrates other than Drosophila.
Cytoplasmic factors in ciliates.
Plasma cholesterol in rats.
A computerized mouse chromosome map.
Inbreeding in standardbred horses.
Muscular dystrophy in chickens; a model for a human disease.
Genetic and environmental control of parthenocarpy in tomatoes.
Robertsonian tranlocations in wild swine.
Ethyl methanesulfonate (EMS) induction of chlorophyll mutations in crops.
Heterofertilization in maize.
Color variation in snakes.
Behavioral genetics in honeybees.
Mitochondrial inheritance in mice.
Use of haploids and meiotic mutants in potato breeding.
Conservation of linkage groups in mouse, cattle, and human.
The molecular evolutionary clock.
Linkage analysis in cats.
A selection limit caused by a balanced polymorphism.
Mutator genes in morning glories.
Journal of Heredity, 1988–present
Since 1988 the journal has been edited by Stephen O'Brien. At the same time, Oxford University Press took over the publishing. The trend started by Barbara Kuhn to make this a journal of research has continued. Although there is still an emphasis on organismal genetics, the journal has come much closer to the mainstream of fundamental genetic research, with a healthy mixture of classical, molecular, evolutionary, and applied genetics. I won't mention specific articles; there are so many that no reasonably sized sample would do justice.
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The History of Genetics as Reflected in Publications of the American Genetic Association |
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The AGA, founded in 1903, has coexisted with the science of genetics through its history, starting with Sutton's demonstration of the chromosomal basis of heredity the same year. As I have shown, by glancing through the pages of American Breeders Magazine and Journal of Heredity one can follow the main trends in the history of the subject. AGA journals usually have not reported pioneering new advances in basic genetics—most of microbial genetics, biochemical genetics, molecular biology, population genetics, and other developments leading up to the structure of the gene, genomics, and molecular evolution have been published elsewhere. The AGA articles, while strongly dependent on basic advances, have more often been derivative. They have been of great value in reporting the extensions to other species and the applications of new basic knowledge. The journal has kept a running account of the consequences of new advances, usually with pictures and punctuated by notes, comments, and news items—often in Cook's lively style. Although one could read the more research-oriented journals and yet learn nothing of the consequences of the new discoveries, their practical applications, or their social and political implications, these were not missed by readers of Journal of Heredity.
The past century has been one of astonishing progress in genetics. The subject now permeates almost all of medicine and agriculture, and in various ways more and more of our lives. Almost all biological research now uses the tools of genetics. Does this mean that genetics, by its very success, has spread so widely that it has lost its identity? Should we be depressed or excited by this possibility? I choose to be excited.
More than any other science, genetics has been the victim of the two tragic dictatorships of the century, Stalin's Russia and Hitler's Germany. In the early days, with its strong bent toward eugenics, Journal of Heredity was ambivalent toward German developments. But of course this changed with the coming of Hitler. Journal of Heredity kept its readers well informed of the political developments and the distortions of genetics abroad. It was a particularly good place to keep up with the Lysenko fiasco in Russia.
In its early years and through much of its history, Journal of Heredity was a source of information and advocacy for eugenics. The subject began with Francis Galton and was carried on as an idealistic concern for the human future. It was both a learning science and an advocacy movement. Even Daveport, who is often castigated for his eugenic zeal and genetic naïveté, emphasized that eugenics was a learning science. But Popenoe and Cook were advocates and Journal of Heredity reflected this. In 1934 the journal published a detailed account of the Supreme Court decision upholding eugenic sterilization and the famous statement of Justice Holmes, "three generations of imbeciles are enough," was printed in full.
Early eugenics looked to future generations. The emphasis was on changing allele frequencies. There was a tendency to downplay environmental effects, although probably no more than genetics is downplayed in some circles today. The efficacy of negative selection, particularly against rare recessives, was overestimated. And some traits were regarded as genetic when there was little evidence for this. There was some, but surely not enough concern for the rights and liberties of the current generation. Eugenics was somewhat like today's conservation, in which the present generation is asked to make some sacrifice—reproductive freedom on one hand, living standard on the other—for the benefit of the future. In addition to genetic concerns, there was the realization that mentally retarded parents do not provide a good environment for rearing children. Yet for all its idealism, the movement was at best naïve, and at worst barbaric. A number of geneticists followed the eugenics line, and probably a larger number were simply quiet. A few actively opposed it. Notable among these was H. S. Jennings (Crow 1987). The movement eventually lost momentum, its demise hastened by its cruel perversion in Nazi Germany.
At present, social concern by ethicists and others has shifted from responsibilities to rights. We now pay less attention to allele frequencies in the future and more to the welfare and rights of the present. The concern is for the persons directly involved and the immediate family rather than future generations.
In one way the decisions ought to be easier now, because of our vastly greater knowledge. Usually the concern of the individual and the concern of society are the same. A parent does not want a genetically deficient child and society does not want the responsibility and expense of caring for it. So the intergenerational conflict is much less than in earlier generations.
Gene therapy is becoming ever more realizable, although it is still decidedly limited. Usually prevention is better than cure. Of course, this is not always true; it is surely better to buy glasses or contact lenses than to attempt gene therapy for near-sightedness. It will be less controversial to replace a gene for Tay-Sachs disease than one for short stature. Yes, there is a slippery slope. Very likely genetic methods will gradually be used for less and less drastic conditions. Is there much difference between increasing size by using steroids or growth hormone and accomplishing the same end by gene therapy? Is manipulating the environment so different from manipulating genes?
I think it is inevitable that as people become more and more effective at repairing or replacing genes in somatic tissues, they will slip over the border and start to repair or replace genes in germ cells, perhaps unwittingly. This is likely to happen only with severe diseases, at least at first. I shall be quite content if the genes for Lesch-Nyhan syndrome, muscular dystrophy, and Tay-Sachs disease become extinct (except for new mutations). I can't imagine a future environment in which these traits would be desirable. I am impressed that artificial insemination and ova transplant have been used only for humanitarian purposes and not as an attempt to change future allele frequencies. And perhaps more worrisome are problems that call for more distant vision and more international cooperation than we have so far achieved: problems such as water shortages, global warming, famine, and war. It's hard for a representative government to look beyond the next election.
I prefer to face the genetic future with optimism rather than pessimism. Of course, my optimism may turn out to be unjustified. But my future years are decidedly limited and I shall probably never know.
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Acknowledgments |
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This paper was originally presented at the American Genetics Association 2003 Annual Meeting and Centennial Celebration at the University of Connecticut, Storrs, July 18–30, 2003.
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Footnotes |
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Corresponding Editor: Kent E. Holsinger
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References |
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Blakeslee AF, 1914. Corn and men. J Hered. 5:511-518.
Crow EW, Crow JF, 2002. 100 years ago: Walter Sutton and the chromosome theory of heredity. Genetics. 160:1-4.
Crow JF, 1987. Seventy years ago in Genetics: H. S. Jennings and inbreeding theory. Genetics. 115:389-391.
McKusick VA, 1960. Walter Sutton and the physical basis for Mendelism. Bull Hist Med. 35:487-497.
Sutton WS, 1902. On the morphology of the chromosome group in Brachystola magna. Biol Bull. 4:24-39.
Sutton WS, 1903. The chromosomes in heredity. Biol Bull. 4:231-251.
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