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Mendel and modern genetics: the legacy for today

Garland E. Allen
Department of Biology, Washington University, St Louis, MO 63130, USA

The legacy of Mendels pioneering studies of hybridization in the pea continues to inuence the way we understand modern genetics. But what sort of picture did Mendel himself have of his work and its ultimate uses, and how does that picture compare with the collection of ideas and methodologies that was put forward in his name and later became known as Mendelism? With genetics standing at the center of our present biomedical and biotechnological research, an examination of the history of our concepts in the eld can help us better understand what we should and should not expect from current genetic claims. For that enterprise there is no better starting place than Mendel himself. As we celebrate 50 years of the Watson Crick model of DNA, it is worth stepping back to take a longer look at the work that started it all: the pioneering hybridization studies of the pea (Pisum sativum) and bean (Phaesolus) by Gregor Johann Mendel (1822 1884) (Fig. 1). Although Mendels work was rst presented at a meeting of the Brunn Natural History Society in 1865, and published in its Proceedings in 1866, it received only a modicum of attention until 1900, when it was more or less simultaneously rediscovered by three different investigators: Carl Correns (1864 1933) in Germany, Hugo De Vries (1848 1935) in the Netherlands, and Erich von Tschermak-Szeneygg (1871 1962) in Austria. Whilst all three found Mendels work suggestive, it is not clear that any of them really saw the signicance of what he had done, and none of the rediscoverers became a major promoter of the new genetics [1]. The rst major publicist for Mendels work was William Bateson (1865 1926) in England. Through the Royal Horticultural Society, Bateson had Mendels paper translated into English for the rst time, and wrote a general exposition that laid out the basic principles of what soon came to be known as Mendelism [2]. Batesons work brought Mendel to the attention of numerous workers in England, Scandinavia and the United States, in particular to many of those involved in practical animal and plant breeding [3]. Although there was reluctance in some quarters to embrace Mendels work immediately especially among academic biologists by the end of the rst decade of the 20th century, the theory had gained a considerable following. However, the explanation Mendel
Corresponding author: Garland E. Allen (allen@biology2.wustl.edu).

offered for his breeding data seemed reminiscent of so many of the speculative, particulate theories of heredity that had abounded in the post-Darwinian era, including Darwins own provisional hypothesis of pangenesis, August Weismannns elaborate theory of ids, idants and biophors, Ernst Haeckels imaginary plastidules, and Hugo de Vries postulated pangenes. Consequently, to some, at least, Mendels work seemed like just one more of the sort of abstract proposals they had encountered all too frequently. In point of fact the paper probably seemed extraordinarily dense, with no illustrations but numerous binomial expansions, which were likely to have been a putoff for many biologists who at the time were notoriously math-shy. For whatever reason, for the rst decade of its re-emergence into the scientic world, Mendels contribution remained controversial at best, dismissed by signicant sections of the biological community at worst. What ultimately served to establish Mendelism on more rm ground between 1900 and 1915 was: (1) its extension to an increasingly wide variety of organisms; and (2) its unication with the cytological work on chromosomes carried out principally through the work of Thomas Hunt Morgan (1866 1945) and his young, enthusiastic team of investigators at Columbia University between 1911 and 1925. The work of the Morgan school demonstrated that the abstract elements or factors discussed by early 20thcentury Mendelians could be regarded as discrete, material units arranged linearly along the chromosomes,

Fig. 1. Gregor Johann Mendel (standing, second from right) with members of the Augustinian monastery of StThomas in Brno in about 1862. Mendel can be seen observing a plant specimen. Others in the photograph of particular importance in Mendels life are the Abbot Cyrill Napp (seated, second from right) and Matous Klacel (seated, rst from right), who was also interested in natural science and philosophy, and with whom Mendel had frequent lively discussions. Reproduced, with permission, from [5], p. 212.

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Box 1. The common picture of Mendelian theory

Mendel observed the inheritance patterns of traits or characteristics in pea plants, such as height, pod color and or seed shape, each of which showed alternate forms: tall/short, yellow/green and smooth/wrinkled, respectively. Mendel referred to these alternate conditions as dominant and recessive. Mendel hypothesized that each trait was represented in the germ cells of adult plants by two determinants (referred to in his paper as Anlagen or elements), one received from each parent; these determinants were symbolized by Mendel with a capital letter for the dominant form (e.g. A) and a lower-case letter for the recessive form (e.g. a). The determinants could be combined in one of three ways: two dominants (AA), two recessives (aa) or a dominant and a recessive, or hybrid (Aa). Although he knew nothing about the cytology of chromosomes, Mendel hypothesized that in the formation of the pollen or egg cell, the two factors for each trait would separate and go into different gametes; thus a parent that was pure dominant would produce gametes all of which contained the dominant factor (A), and a parent that was pure recessive would produce gametes all of which contained the recessive factor (a); hybrid parents, however, would produce two kinds of gametes: 50% would contain the dominant factor (A) and 50% the recessive factor (a). At fertilization, the double-determinant condition would be restored. If both parents were hybrids, any of the three possible combinations could occur and would be distributed randomly, according to the laws of probability: 1 AA: 2 Aa: 1 aa; because organisms that are Aa and AA look alike, the ratio based on appearance of the traits (what later came to be called phenotype) would be 3:1. When two or more characteristics (e.g. Aa, Tt) are followed in a dihybrid cross, the two sets of determinants segregate randomly, so that any combination of A, a, T and t is possible, what came to be known as the principle of random assortment. Factors somehow determine traits, so that after the term gene was introduced in 1909, it was common to speak of a gene for tallness or a gene for wrinkled seed. During the early years of the Mendelian theory, this was referred to as the unitcharacter hypothesis. Factors are not modied by being combined with their alternate form; thus, the factor, t, for shortness is not affected in any way by being combined with the dominant factor, T, for tallness in the hybrid, a concept that became known as the concept of the purity of the gametes.

the sort of picture Mendel himself envisaged. Second, as an introduction to the study of genetics, it has led to unfortunate and now long-entrenched misunderstandings of how genes really function, and the relationship between genes and the development of adult traits that has carried over into molecular genetics. Mendels background Mendel was born on 22 July 1822, in the small rural village of Hyncice, in Moravian Silesia, then part of the Austro Hungarian Empire. As the only son of a peasant farmer, he was expected to follow in his fathers footsteps and take up farming. But early on, his interest in natural history and his studious ways brought him to the attention of the priest, Friar Schreiber and the local schoolteacher. In spite of nancial hardship for his family, young Johann was sent to a larger school in a nearby village and eventually qualied for Gymnasium in Opava (Troppau). It was a period of extreme privation, but Mendel managed to graduate in 1840 with considerable academic success. One of the chief inuences on him at the time were Schreibers Enlightenment ideals, particularly his emphasis on science as a way of dispelling superstition and ignorance. Schreiber was keenly interested in applying scientic principles to improve humanity, and was heavily involved with local agricultural groups and the Pomological Society [5]. Financial problems continued to plague Mendel as he tried to continue his studies at the Philosophy Institute in Olomuc (Olmutz). After several periods of illness, brought on it is thought by his poverty and overwork, Mendel managed to nish his studies at the Institute and entered Olomuc University. Here, according to records, he took a course of lectures in physics, mathematics and logic. However, unable to complete his degree owing to nancial constraints, he applied for, and was accepted into, the Augustinian monastery of St Thomas at Brno (then Brunn), Moravia in 1843 (Fig. 2). Although he did not feel a particularly fervent spiritual calling, Mendel realized shortly after joining the monastery that at last he was free from constant nancial worries and could pursue his intellectual interests in exchange for attending to pastoral duties. The monastery at Brno was a center of learning in the early- and mid-19th century, especially in the natural sciences and agriculture [6]. The practical and economic benets of promoting new agricultural practices was among the monasterys top priorities, with most of its income coming from extensive landholdings leased to local farmers. At the time that Mendel entered St Thomas, the Abbott (administrator) was Friar F. Cyril Napp (1792 1867), an enthusiastic naturalist, member of several local agricultural and scientic societies, and author of many technical papers, particularly on plant pests. In 1830, he had given over one part of the monastery garden to another monk, Matous Klacel (1808 1882) for experimental cultivation of rare Moravian plants. The monastery also had an extensive library containing a variety of scholarly texts (Fig. 2). The importance of agricultural concerns in Mendels developing interests in the natural sciences is clear [7]. At

and that observed variations in the patterns of inheritance of traits could be traced to the mechanics of chromosome behavior during meiosis (gamete formation). Mendels factors (after 1909 referred to as genes) thus seemed to be real, material units, not metaphysical postulates. In recognition of this work, Morgan was awarded the rst ever Nobel Prize for research in genetics in 1933 [4]. Therefore, a common picture of Mendelian theory has emerged from this early work and has been promoted in textbooks ever since (Box 1). This scheme has made for a very heuristic pedagogy, and has been replicated in an almost innite number of high-school and college biology textbooks. Although it has enjoyed a certain intellectual neatness, this formulation has several problems that have now begun to surface as molecular genetics has provided a far more sophisticated understanding of gene function than was available in the rst half of the 20th century. The rst problem is simply historical: it is not clear that the neat textbook scheme is
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Fig. 2. The main church and monastery building (left) that became the seat of the Augustinian Order in 1783, and the magnicent library of the monastery (right), subsequently the Mendelianum, a division of the Moravian Museum. One of Mendels early mentors in the monastery was Matthew Klacel, who became, after demotion from position of teacher because of his radical ideas, the monasterys librarian. The library has many works on agriculture, including Liebigs 1844 Agricultural Chemistry, and Mendels copy of the 1860 German edition of Darwins On the Origin of Species. Reproduced, with permission, from [5], p. 46.

the time, the amount of experimental breeding in Moravia, both with plants and animals, was extensive. Indeed, the Brunn Natural History Society, before which Mendel read his Pisum paper in 1865, was a section of the larger Brunn Agricultural Society. The peas with which Mendel worked were derived from major edible strains, suggesting the close connection between his studies on hybridization and agricultural interests. Mendel clearly beneted from the monasterys support of science, and the emphasis placed on agriculture by local agriculturalists and members of the monastery staff turned Mendels interests in a specic direction that ultimately led to his extensive series of breeding experiments. Motivation for Mendels Pisum experiments It was during his period as a teacher at the newly opened Realschule that Mendel began his systematic breeding experiments with Pisum. One of the major reasons that has been put forward to explain why a clergyman and schoolteacher would have undertaken such an unusual investigation is that he wanted to provide a generalized theory of heredity that would complement Darwins theory of natural selection (or, as several authors have also suggested, to counter Darwins theory by demonstrating the xity of species limits during hybridization). Darwins theory had encountered considerable theoretical difculties because of the prevailing belief by naturalists and breeders (including Darwin himself) in blending inheritance the idea that any trait in the offspring was a blend of the traits observed in the two parents. Thus, if parental differences blended in the offspring, new variations would be diluted in every generation, leaving little for natural selection to act on. However, it is clear that Darwins work could not have been Mendels initial motivation, because he had already begun his experiments in 1856, three years before the publication of The Origin of Species. If not evolution, what was the initial motivation for Mendel, and what did his paper say, both to his contemporaries and to those who read him after the rediscovery in 1900? In 1979, Robert Olby revised our understanding of the Pisum paper by stripping it of its 20th-century interpretations [8]. He concluded that
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Mendel was not a Mendelian in any modern senses of term, and that proponents of Mendelism, from Bateson onward, had read much into Mendels work that was not part of his original formulation. Similar re-evaluations have subsequently been offered by others [9 11]. The gist of these various reinterpretations is that: (1) the primary motivation for Mendels work was not to develop a generalized theory of heredity, but was far more immediate and practical to establish patterns of hybridization that would have been of interest to agricultural breeders and others in 19th-century Moravia; and (2) Mendel never proposed that there were material particles, factors or Anlagen, in the germ cells, or that such particles were necessarily transmitted via pollen and egg cells to the offspring. Mendel, they argue, remained far more agnostic about the physical basis of inheritance than later Mendelians assumed. Indeed, it is much more likely that the direction of Mendels hybridization research was guided by the strong agricultural interests around him. It was precisely through hybridization that breeders hoped to generate new combinations of characters that they could exploit to form new breeds. The problem was that most hybrids do not breed true and have a tendency to revert to their original parental types. What Mendel was able to explain were the conditions under which that happened, and most importantly, the methods for distinguishing between hybrids and true-breeding forms that might look phenotypically similar. Crucially, his hypothesis of dominant and recessive characters provided an alternative to blending inheritance. Whatever happened within the germ cells of the hybrids, the characters were not blended by residing together. This became as important a principle for breeders as it would become later for Darwinian evolutionists, because it freed both groups from the loss of distinct variations through blending or swamping in the hybrid. Nevertheless, it is not possible to be interested in hybridization without simultaneously being drawn into accepting, however loosely, some concept of heredity. Hybridization involves mixing germ lines from two different ancestries, and how those germ lines interact, by denition, forms a theory of heredity. Thus, to claim that Mendel was interested only in hybridization and not

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in any general principles of heredity is to make a confusing distinction. The two processes are not only not mutually exclusive, they are complementary. The revisionist view therefore is that Mendels major motivation (and interest) lay in establishing principles of hybridization, and not in establishing a generalized theory of heredity as was assumed by his early-20th century followers. These re-interpretations of Mendel also argue that nowhere does he explicitly state that the germ cells contain particles that determine the character of a trait. Mendels symbols (e.g. T,t and A,a), they argue, were merely algebraic notations and were never meant to represent material entities in any biological sense. It is true that Mendel is not explicit on this matter. He talks in his paper about alternate characters (tall and short) segregating in the germ cell of the hybrid parent, but he does not refer to determiners or particles; moreover, he does not talk about segregation of the two characters in the homozygous forms, and represents homozygotes symbolically with only a single letter (such as A for the dominant parent or a for the recessive), whereas he always represents the hybrids with two letters (e.g. Aa). Thus, he consistently shows the results of a monohybrid cross as A 2Aa a. A careful re-reading of the Pisum paper suggests that Mendel was ambiguous, but not explicitly agnostic, towards the notion of hereditary particles. The ambiguity arises because throughout his paper, Mendel consistently talks about the characters [Charactere ] of the parents or offspring of his crosses, which of course refer to the visible adult traits. His empiricism and training in the physical sciences led him to emphasize what he could see in the plants he was breeding. At the same time, toward the end of his paper, Mendel wrote about the implications of his experiments for understanding the composition of the fertilization cells (i.e. pollen and egg). Here, he introduces the terms Anlage and Elemente in reference to what is transmitted from parent to offspring through fertilization. The development proceeds from a constant law which is grounded in the material composition and arrangement of the elements [Elemente ], which come together in the cell in a viable union, wrote Mendel [12]. Indeed, it would have been unusual in Mendels day, and given his training, not to at least have thought in terms of some sort of discrete hereditary particles being passed from parent to offspring during reproduction. Virtually every theory of heredity proposed during the middle and later years of the 19th century was couched in particulate terms so that it was common among biologists, especially in the German-speaking world, to think in terms of atom-like particles or molecules as agents of hereditary transmission. The problem with most of these theories was that they were largely speculative, and based on a few observations and little, if any, experimental evidence. It is not surprising therefore, that Mendel hints at the existence of such components in the germ cells. However, it is important to remember that he was rst and foremost an empiricist, and was highly restrained when it came to speculation about mechanisms. In this sense, historians are right who emphasize that Mendel was primarily interested in establishing the laws of hybridization rather than a general theory of heredity. His
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constant emphasis on characters, his recognition that it was sometimes difcult to determine in what character class an individual should be placed, and his meticulous record-keeping all testify to his strongly empirical and nononsense approach to experimentation. Mendelism and the unit character hypothesis By the time Mendels paper was rediscovered in 1900 and had begun to receive attention from biologists and breeders in industrialized countries, economic, social and intellectual conditions were signicantly different than those in 1866 Central Europe. The industrial revolution in Europe and the US had placed many new demands on agriculture [13]. Not only were there increasing demands on food production to supply the large urban workforces that had developed in industrial areas, but much of that workforce had come from the agricultural sector, leaving it in short labor-supply. Mechanization of agriculture had proceeded with great rapidity at the end of the 19th century, a process that had initiated what has been called the industrialization of agriculture [14]. During the second half of the 19th century, scientic agriculture based on the organic and physiological chemistry of Justus von Liebig (1803 1873) and his school in Giessen had greatly improved yields by attention to aspects such as fertilizers and animal and plant nutrition. But by 1900, those inputs had, for the time, achieved their technical limits. Improvements in breeding higher-yielding varieties, however, held out a wholly new potential. One aspect of agriculture husbandry was still in a rudimentary stage at the turn of the 20th century. Most animal and plant breeders operated by various rules of thumb that they had developed their separate localities or had learned and modied from others. It was largely a craft, with no general rules that applied across the board. The reintroduction of Mendels work at this juncture generated hope that some principles of heredity might, at last, provide the breeder with methods that could yield more predictable results. Although in reality the application of Mendelian principles to improving animal and plant productivity turned out to be more difcult than it initially seemed, optimism was nonetheless shown by many academic biologists, US Department of Agriculture (USDA) ofcials, and breeders in the early decades of the century. The economic and social conditions made Mendels work look protable in 1910 in a way it had not looked in 1866. But as academics and breeders took up the nascent science of Mendelism [15], there remained an uncertainty over what was actually transmitted from parent to offspring during fertilization. Although Wilhelm Johannsen (1857 1927), the Swedish plant breeder who coined the term gene, and William Bateson, who coined the term genetics, both favored a more abstract statistical model of the hereditary unit, others, especially in the US, pressed for a more material, atomistic interpretation from 1903 onward. The abstract element of Mendels paper had quickly become the discrete gene of the rediscovered Mendelism. Textbooks began to present images of Mendelian crosses using Mendels capital and small letters for dominant and recessive traits.

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Between 1900 and 1910, this interpretation of Mendelism communicated two important notions to the generation of biologists who were beginning to learn about the new genetics. First, there was no hard-and-fast distinction between the letter or factor and the adult character for which it stood. Thus, geneticists would write about the inheritance of height or wing shape or red eyes as if the trait itself was what was carried in the male or female gamete. Furthermore, in the early decades after the rediscovery of Mendelian theory, the identication of the hereditary unit with the adult trait was graphically reinforced by the Punnet Square a method of calculating Mendelian combinations which blurred what would become known after 1911 as the phenotype genotype distinction, and gave the impression that the inherited factor was the trait in miniature. It was this conation that led embryologists like the young T.H. Morgan to reject the Mendelian hypothesis for a decade [4]. For Morgan and others, the Mendelian factor or gene smacked too much of the embryologists old bugbear, preformationism (the idea that the complete adult exists already preformed in the fertilized egg, and that embryonic development involves only an unfolding or growth of the embryo in size). Epigenesis, the alternative view, had replaced preformationism by the mid-19th century, so that Mendels work in the form presented by Mendelians, seemed like an outdated throwback to a long-discarded idea. With the wedding of Mendelian theory to the cytological investigation of chromosomes by Morgan and his group after 1910, the material, discrete and atomistic concept of the gene gained considerable prominence. By carefully correlating the inheritance patterns of gene mutations with observable changes in chromosome structure, Morgan and his students were able to show that genes could be clearly regarded as material entities that occupied specic positions loci linearly arranged along the chromosome. By 1915, when the group published the their pathbreaking book, The Mechanism of Mendelian Inheritance [16] the beads on a string model of the chromosome had become an icon. Not only did this model promote the idea of genes as atomistic units, it also further supported, indirectly, the unit character hypothesis. The gene was now viewed not only metaphorically, but also literally as an atom, entering now into this, now into that combination, but emerging each time with its own integrity in the same fashion as atoms entered into, and came out of, molecular combinations. Indeed, the language of chemistry began to enter genetics explicitly in relation to the discreteness of the Mendelian factor. Harvard geneticist W.E. Castle (1867 1962) wrote early on that:
all observed inheritance phenomena can be expressed satisfactorily in terms of genes, which are supposed to be to heredity what atoms are to chemistry, the ultimate, indivisible units, which constitute gametes much as atoms in combination constitute compounds [17].

just such a conceptual notation as is used in algebra or in chemistry [18], whilst British polymath J.B.S. Haldane (1892 1964), as late as the 1930s, claimed that the atomic nature of Mendelian inheritance suggests very strongly that even where variation is apparently continuous this appearance is deceptive. On any chemical theory of the nature of genes this must be so. [19] The implication that the discreteness of the gene implied the organism was constructed as a mosaic of adult traits was given explicit voice by Bateson within the rst years of his encounter with Mendelism. In 1901, he wrote:
In so far as Mendels law applies, the conclusion is forced upon us that the living organism is a complex of characters of which some, at least, are dissociable and are capable of being replaced by others. We thus reach the conception of unit characters, which may be rearranged in the formation of reproductive cells [20].

Whilst Castle was arguing that genes were not quite like atoms because they could be altered by the selection process, that he mentions the analogy at all indicates something of its prevalence. Somewhat earlier his colleague E.M. East had written that Mendelism is therefore
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The next year Bateson wrote even more boldly: The organism is a collection of traits. We can pull out yellowness and plug in greenness, pull out tallness and plug in dwarfness. [21] This mosaic view of the adult organism parallels the mosaic view of the genome, and thus pictures the organism as an aggregation of interchangeable parts that can be arranged and rearranged by assembling the right combination of genes. Admittedly, not all early Mendelian geneticists took quite such a mechanical view of heredity, but Bateson was one of the leaders in the eld and among its most vocal proponents. His inuence was therefore considerable. It is thus even more ironic that as late as 1922, Bateson still held strong reservations about linking Mendelian genes to chromosomes [22]. The atomistic, mosaic conceptualization of the gene was initially fruitful, allowing biologists to develop a quantitative, experimental and predictable aspect of their science that made it as hard as anything in chemistry or physics [23]. However, it created problems almost from the outset. It allowed biologists to put aside questions of gene function, or the relationship between Mendelian genes, embryonic development, and even evolution. From 1915 onward, most attention was paid to the mechanics of gene shufing during transmission of chromosomes from parent to offspring. This trend, which marked the hey-day of classical genetics, led to the extraordinary feat of mapping the chromosomes of several model organisms, such as the fruity, maize and mouse. But the atomistic trend in classical genetics also left an unfortunate legacy. By perpetuating the unit character concept of the gene, even with lip-service paid to the idea of gene interactions, the vast majority of the public and many biologists as well, still hold to the view that one, or at best a very few, genes determine each specic, adult trait. Nowhere is the tendency to attribute discrete gene elements to specic traits more common, and perhaps more socially troublesome, than in the area known as human behavior genetics. To date, social and personality traits as complex and varying as schizophrenia, manic depression, alcoholism, criminality, violence, shyness, homosexuality, risk-taking and religiosity, have all been headlined at one time or another as caused by a gene. Not

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only do such claims greatly oversimplify the biological process of development, during which genes interact with other genes and the environment, they suggest a set of potential solutions that is deceptive: drug therapy, gene therapy or sterilization. Indeed, sterilization was implemented in both the US and Germany in the old eugenics movement, when simplistic genetic notions allowed the compulsory sterilization laws to be passed for those with undesirable hereditary conditions [23]. Today, Norplant can replace surgical sterilization, and pharmacogenetics looms on the horizon, promising to counteract the effects of defective genes and to yield huge benets for the pharmaceutical industry. Genes are of course critical components in one way or another of all of our traits. But because of the increasingly detailed ndings of molecular genetics and the Human Genome Project, the picture of what genes do and how they function in the development of adult human characters has changed dramatically from the legacy built out of the early interpretations of Mendels work. With his skepticism about undue speculation regarding the mechanics of the hereditary process, Mendel might indeed have been more in tune with modern genomics than with the Mendelians who crafted the classical concept of the gene in his name. Conclusion As we look forward, it is clear that one of the most important forefronts of modern genetic research lies in the area of what has been variously called the genetics of development, developmental genetics, and when combined with evolutionary theory, the evolution of development or Evo Devo for short. These emerging elds, which synthesize the classical genetics of the rst half of the 20th century with the molecular genetics and evolutionary biology of the second half, promise to yield a more complex, and hopefully more realistic picture of how the genome guides the individual organism from fertilization to adulthood. We are standing at a divide between an old and a new genetics, between a mosaic and mechanical and a holistic and integrated view of the organism that promises to yield exciting results in the years ahead. It behoves us to understand the pathway that has brought us to this historic juncture, not only so that we may take the fullest advantage of our newest research paths, but also that we avoid the disastrous social consequences that can arise from misplaced expectations of what genetics can do. Like all other scientic and technological ndings, we must rst understand the science itself and its history to recognize both its potential and its limitations.
References
1 Sturtevant, A.H. (1965) History of Genetics, Harper & Row; in chapter 21, Sturtevant reviews the claims that the three rediscoverers were really so independent, whilst several papers by Alain Corcos and Floyd Monaghan suggest that only Correns had really collected data that

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were really comparable to those obtained by Mendel. See, Corcos, A. and Monaghan, F. (1985) Role of de Vries in the recovery of Mendels work. J. Hered. 76, 187 190; (1986) Tschermmak: a non-discoverer of Mendelism I: an historical note. J. Hered. 77, 468 469; and (1987) Tscheremak: a non-discoverer of Mendelism II: a critique. J. Hered. 78, 2 10. These various positions have been admirably summarized in Peter Bowler (1989) The Mendelian Revolution. The Emergence of Hereditarian Concepts in Modern Science and Society, Johns Hopkins University Press, especially chapter 1 Bateson, W. (1902) Mendels Principles of Heredity: A Defense, Cambridge University Press, Cambridge, UK Letter from William Bateson to his wife, Beatrice, 3 October 1902, quoted in Paul, D. and Kimmelman, B. (1988) Mendel in America: theory and practice 1900 1919. In Rainger, R. et al. (1988) The American Development of Biology, p. 283, University of Pennsylvania Press Allen, G.E. (1978) Thomas Hunt Morgan, the Man and His Science, Princeton University Press Orel, V. (1996) Gregor Mendel, the First Geneticist, pp. 40 41, Oxford University Press Orel, V. (1973) The scientic milieu in Brno during the era of Mendels research, J. Hered. 64, 314 318; see also [5], chapters 2 3 Orel, V. and Wood, R. (2000) Scientic animal breeding in Moravia before and after the rediscovery of Mendels theory. Quarterly Review of Biology 75: pp. 149 157 Olby, R. (1979) Mendel no Mendelian? History of Science 17, pp. 53 72 Brannigan, A. (1979) The reication of Mendel, Social Studies of Science 9, pp. 423 454 Meijer, O. (1983) The essence of Mendels discovery. In Gregor Mendel and the Foundation of Genetics (Orel, V. and Matalova, A., eds.), The Mendelianum of the Moravian Museum in Brno, pp. 123 178 Monaghan, F.V. and Corcos, A. (1984) On the origins of the Mendelian laws. J. Hered. 75, 67 69; and (1984) The true Mendelian laws. J. Hered. 75, 321 323 Mendel, G., cited in [11], p. 105. The original reads: Diese Entwicklung erfolgt nach einem constanten Gesetze, welches in der materiellen Beschaffenheit und Anordnung der Elemente begrundet ist, die in der Zelle zur lebesnfahigen Vereinigung gelangten. Allen, G. (2000) The reception of mendelism in the United States, 1900 1930. Comptes Rendu Academie des Sciences, Paris. Sciences de la vie 323, 1081 1088 On the industrialization of agriculture in 20th-century America, see Shannon, F.A. (1961) The Farmers Last Frontier, pp. 362 365 and 366 367, Holt, Rinehart and Winston; and Kloppenberg, J.R. (1988) First the Seed. The Political Economy of Plant Biotechnology. Cambridge University Press, especially chapters 2 4 Punnett, R. (1911) Mendelism, Macmillan Morgan, T.H. et al. (1915) The Mechanism of Mendelian Inheritance, Henry Holt & Co. Castle, W.E. (1919) Piebald rats and the theory of genes. Proc. Natl. Acad. Sci. U. S. A. 5, 126 130; quotation on p. 127 East, E.M. (1912) The Mendelian notation as a description of physiological facts. Am. Nat. 46, 633 695 Haldane, J.B.S. (1932) The Causes of Evolution, p. 57, Harper and Brothers This quotation comes from Punnett, R.C., ed. (1928) Scientic Papers of William Bateson, (vol. 2), p. 1, Cambridge University Press Bateson, W. (1902) Mendels Principles of Heredity: A Defense, Cambridge University Press; quoted from Lewontin, R.C. and Levins, R. (1985) The Dialectical Biologist, p. 180, Harvard University Press Coleman, W. (1970) Bateson and chromosomes: conservative thought in science, Centaurus 15, 228 314 For a more detailed discussion of the mechanistic view of the gene, see Allen, G.E. (2002) The classical gene: its nature and its legacy. In Mutating Concepts, Evolving Disciplines: Genetics, Medicine and Society, (Parker, L.S. et al., eds), pp. 11 41, Kluwer Academic Publishers

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