Readers of The Mendel Newsletter are well aware that Mendel's image has been through several profound transformations. He has been ignored, lionized, debunked, accused of fraud, repudiated, vilified, and repeatedly rehabilitated. This kaleidoscope of changing interpretations contrasts with the settled image of another famous 19th century biologist, Charles Darwin, whose reputation is good Victorian, family man, sound investor, patient toiler, Painstaking researcher, insightful observer, creative thinker, and chronic sufferer. Why the difference? This is a question for historians of biology to resolve, not for a practicing geneticist like me. Nevertheless, I would cautiously suggest that some of the reasons are those of class, empire, and language.

By class I mean social class. Darwin was solidly British upper class, and so his scientific work is naturally accorded high status and respectability. Mendel was a peasant boy from Moravian Silesia who pulled himself up by his own bootstraps. His parents were Anton and Rosine Mendel (née Schwirtlich). By ethnic ancestry he was Moravian German who grew up speaking German but later, living in Brno, also spoke Czech. His father was a peasant farmer required to work three days a week for the landlord. Mendel's early academic promise was recognized by his teachers who arranged for him to be educated at superior schools far from home. In 1843, at the age of 21, after completing two years of study at the Philosophy Institute in Olomouc, Mendel was admitted to the Augustinian monastery of St. Thomas in Brno (then Brünn). He was ordained in 1847, studied at the University of Vienna in 1851-1853, and returned to teach physics and natural history at the Realschule in Brno. Darwin's club membership was the Royal Society of London, Mendel's the Natural History Society of Brünn.² Darwin's contributions were made in the halcyon years of the Victorian empire, Mendel's in the waning years of the Hapsburg empire; this is what I mean by empire as a cause of the difference in Darwinian and Mendelian hagiography. By language as a cause, I mean English versus German and Czech. All that Darwin wrote, and most things written about him, are in English and easily available. Because much of the primary literature about Mendel is written in German and Czech, and reposed in Eastem Europe, it is virtually inaccessible to all but a handful of Mendel scholars. It is for this reason that Orel's biography is an essential resource for anyone interested in Mendel's life, work, and times.

To class, empire, and language as causes of the difference in treatment of Darwin and Mendel, I would also add religion, because there is a detectable note of anticlericalism among some of Mendel's critics, particularly those who, with a wry smile, suggest that his data are fraudulent and that they are shocked, shocked. To illustrate this point, let me mention an u textbook in genetics, published in the 1970's, which included a reproduction of a color portrait of Mendel wearing a bishop's robes with miter and staff in the background. (You can view this portrait via the internet at http://www.vszbr.cz/mendel.html.) The accompanying text discussed at length various allegations of Mendel having falsified data, and the caption of this illustration encouraged the image of venality by insinuating that Mendel was also guilty of self-aggrandizement because he had no authority to appear in such garb and with such symbols. Upon reading this caption, I wrote to the author and pointed out that Mendel was elected abbot of the monastery of St. Thomas in 1868. As prelate, his ecclesiastical rank was indeed that of a bishop, and inasmuch as this was an official portrait of the Prelate Mendel, his wearing any other sort of vestments would have been unseemly and denigrated his own rank. The author dutifully acknowledged my letter, but to my knowledge the error was not coffected in further printings or subsequent editions of the book. This is the sort of lapse of accuracy that would bring an ombudsman, if not a tort lawyer, down on a present day news reporter, but for Mendel there is no such recourse.

Until now, the best biography of Mendel has been Hugo Iltis's Life of Mendel,³ based on Iltis's own primary research and including interviews with a number of people who knew Mendel personally. With Orel's book, Iltis's classic is finally superseded.

I should say a few words about Orel. He knows the ups and downs in the interpretations of Mendel. He has also suffered personally through some of the political events that shaped the history of genetics in 20th century Eastern Europe. Born in 1926 on a small farm near Brno, he developed an interest in animal breeding even as a boy, motivated in part by his father. At age 11 young Vítezslav was enrolled in the gymnasium and also worked part time at a poultry breeding farm recording traits and carrying out selection. Two years later, in 1939, the Nazi onslaught trampled Czechoslovakia, and Orel learned the taste of constant fear. He was allowed to continue his studies for five years, but in the "total eingesetzt" he was forced to do hard manual labor for 69 hours per week in a Nazi war factory. Seven atrocity-filled years after the invasion, Czechoslovakia was released from the talons of the Nazi eagle only to be dropped behind the Iron Curtaln. Orel entered the Agricultural University in Brno where he was captivated by Jaroslav Krízenecký (1896-1964), who soon became his mentor. By that time Orel could read in Czech, German, English, French, and Russian and soon began to publish papers in the theory of genetics as applied to animal breeding. After 1948, the evil henchmen of Trofim Lysenko, by then triumphant in the USSR, imposed their perversion of science on the genetics of Brno. Mendel was anathematized as a representative of the Church and, in 1950, the Augustinian Order was forced to give up the monastery and the monks arrested. The monastery buildings were converted to secular purposes and fell into disrepair, Mendel's old glasshouse was destroyed, and the statue of Mendel in Mendel Square was ordered to be removed. (Curiously, the name of Mendel Square was left unchanged.) Krízenecký was denounced as a reactionary Mendelist and forced to leave the university. Orel was allowed to work at the Research Institute of Poultry and was even allowed to publish papers on poultry breeding provided he did not use the term "genetics." He was required to focus on the technology of poultry production, and seemingly innocuous events, such as receiving eggs from the German poultry breeding company Lohmann, brought on police accusations of collaboration with reactionary capitalists working against the socialist system. After 1958 Orel was assigned to work at a small duck farm outside Brno. Not until 1962 did the Lysenkoist madness subside enough that Krízenecký, by then "rehabilitated," was allowed to begin planning a Mendel Museum, which in 1965 became housed in a small part of the monastery adjacent to the garden in which Mendel carried out some of his experiments. After Krízenecký's death 'm 1964, Orel became head of the Mendelianum, of which he is presently Emeritus Head. At this time he is undergoing a protracted recuperation from serious injuries received in an automobile accident in 1989. As for the status of the monastery and church, in 1990, with the collapse of the last (let us hope) Communist regime in Czechoslovakia, monks from the Order of St. Augustine were allowed to return to the monastery, and the monastery church (The Church of Our lady) began to be restored. While in the process of restoration (now completed), the church was singled out for special distinction by Pope Jean Paul II in being designated a basilica. The future of the monastery is still unclear, although there is considerable support for making it a center for the study of the relationship between science and religion.

With respect for the poignancy of Orel's life and his courage in overcoming adversity, what does he bring to the study of Mendel that transcends what any other historian could do? The answer is that Orel has had special access to the rich archives of the Mendelianum. Indeed, as head of the Mendelianum, he helped create the archives. He has used these and other resources to study the special role that agricultural genetics played in the economy of mid-19th century Moravia. At that time scientific breeding was coming into prominence in agriculture in a number of places, notably sheep breeding in England; in Moravia, with its agricultural base, scientific breeding was fundamental. There was, as might be imagined from the economic importance of textile production in Moravia, great interest in the selective breeding of sheep. Among the leading exponents of scientific breeding in Brno was the naturalist Christian Carl André (1763-1831), who took special interest in the development of new varieties of fruit trees and grape vines as well as of domesticated animals. (It is altogether fitting, but by complete coincidence, that a brand of California champagne is called André.)

One of the heroes of Orel's biography of Mendel is, perhaps surprisingly, not Mendel. It is Mendel's abbot, Cyrill Franz Napp (1792-1867), who was interested and active in all of the local agricultural societies, including those with a particular interest in scientific breeding. Napp even became President of the Brno Pomological Association and established a nursery garden in a monastery farm outside of Brno. (As abbot of St. Thomas, Napp was responsible for overseeing the far-flung economic interests of the monastery.) Orel takes great pleasure in having discovered that, as early, as 1837, Napp summarized the conclusions of a Sheep Breeder's Association meeting by saying that the key questions for the future were "What is inherited, and how?" and that in another context Napp called attention to the apparent role of chance in the heredity process. Hence, Orel does not imagine for a moment that Mendel happened upon the study of genetics for want of something better to do. Rather, the study of genetics attracted Mendel as a member of a society of scholars present at the monastery of St. Thomas whose job it was to investigate important issues of the day, among which Napp included heredity for its applications in scientific breeding. Any person seriously interested in the history of this period has to come to grips with the special status of the monastery of St. Thomas in the waning years of the Hapsburg Empire. It was neither fish nor fowl. It was not entirely under the control of Rome because it had important secular assignments, including the production of teachers in the local schools, of whom Mendel was a prime exemplar; nor was it entirely under the control of the secular officials because of its intrinsically religious nature. It is this confounded, mixed, dual role of the monastery of St. Thomas, and all of the men within it, that Orel understands better than any other historian of this period, and he also understands the secular implications of the duality.

Orel is always quick to mobilize his knowledge and employ it in defense of Mendel against attacks from various quarters. He does not idolize Mendel, he does not romanticize him, he does not sanctify him. He is convinced, however, that Mendel knew very well what he was doing when he designed his experiments, why he was doing them, and what he was looking for.

I do not regard Orel's defensiveness as a defect, undoubtedly because I am rather partisan myself where Mendel is concerned. Orel and I have written two papers in which we consider some of the key issues about Mendel-that he was or was not a Darwinian, that he did or did not discover the laws of inheritance, that he did or did not falsify data, that he did or did not report the experiments exactly as he performed them. ^{4, 5} One of these issues was raised by Robert Olby, a leading modem historian of genetics. I am referring to his thesis that Mendel was no Mendelian⁶ because he failed to emphasize the segregation of identical alleles in homozygous genotypes. I happen to disagree with Olby's conclusion on this matter but will grant a certain cogency to his argument. In Mendel's paper, the most telling passage in Olby's favor is one in which Mendel writes: ". . . [The] differing elements [in hybrids] succeed in escaping from the enforced association only at the stage at which the reproductive cells develop. In the formation of these cells all elements present participate in completely free and uniform fashion, and only those that differ separate from each other." ⁷

On the other hand, even in modem genetics, one often regards homozygous genoqws as producing gametes exclusively of a particular allelic type rather than emphasizing that homozygous genotypes produce an equal frequency of two gametic types, each of which happens to be identical in DNA sequence. So, one could just as well make the case that the formal genetics of the situation is the same whether segregation in homozygous genotypes is or is not stipulated. Moreover, it is to be emphasized that Mendel was a superb empiricist (as well as being a first-rate theorist), and from the standpoint of empiricism, there is no way to discern from the results of crosses of the Mendelian type whether identical alleles present in homozygous genotypes maintain their chemical identity or, in fact, become blended into yet another indistinguishable form. To me, the critical passage in Mendel's paper is one repeated no fewer than six times:⁸

[P]ea hybrids form germinal and pollen cells that in their composition correspond in equal numbers to all the constant forms resulting from the combination of traits united through fertilization.

Olby has made the even stronger assertion that Mendel is wrongly given credit for the discovery of Mendelian genetics, whereas the person who more fittingly deserves the honor is Francis Galton (1822-1911). (It is an interesting coincidence that Mendel and Galton were born in the same year.) Olby's argument is that Galton was on the right track in formulating a precise conception of the subject of heredity in statistical terms, namely, "the statistical relations between the distribution of characters in successive generations." This is correct. Galton had found that traits are often normally distributed in parents and in their offspring and that the parent and offspring distributions are correlated. Galton was, indeed, so enamored of the properties of the normal distribution that he waxed poetic:" ¹⁰

I know of scarcely anything so apt to impress the imagination as the wonderful form of cosmic order expressed by the "law of frequency of error" [the normal distribution]. Whenever a large sample of chaotic elements is taken in hand and marshaled in the order of their magnitude, this unexpected and most beautiful form of regularity proves to have been latent all along. The law would have been personified by the Greeks if they had known of it. It reigns with serenity and complete self-effacement amidst the wildest confusion. The larger the mob and the greater the apparent anarchy, the more perfect is its sway. It is the supreme law of unreason.

The point I wish to drive home is that, whereas continuous variation can be inferred from Mendelian heredity, Mendelian heredity cannot be inferred from continuous variation. Therefore, Galton's mindset and approach to heredity through the study of continuous variation was incompatible with his ever being able to infer, much less prove, the particulate and segregational attributes of hereditary variation. The approach was a blind alley. (I might add parenthetically that the analysis of continuous variation at the level of discrete Mendelian units has remained beyond experimental resolution until just the past few years, when the study of widespread DNA polymorphisms accompanying the analysis of continuous traits has allowed some of the underlying Mendelian units to be identified. For the most part, the factors are still identified only by virtue of their genetic linkage with nearby DNA polymorphisms and not through isolation of the relevant genes themselves or an identification of what they do, except in a number of noteworthy and highly publicized cases, such as the familial Mendelian factors that predispose to breast cancer in women.)

Before leaving Galton, I wish to illustrate my contention about mindset by citing passages from his book Natural Inheritance in which he writes with considerable pride of his experiments in studies of heredity in, of all creatures, the garden pea:

I determined to experiment on seeds, and after much inquiry of very competent advisors, selected sweet-peas for this purpose. They do not cross-fertilize, which is a very exceptional condition among plants; they are hardy, prolific, of a convenient size to handle, and nearly spherical. . . . I began by weighing thousands of them individually, and treating them as a census officer would treat a large population. . . . Then I selected with great pains several sets for planting.¹²

Curiously enough, the diameters were found also to run approximately in an arithmetic series, owing, I suppose, to the misshape and corrugations of the smaller seeds, which gave them a larger diameter than if they had been plumped out into spheres. AR this is shown in the Appendix, where it may be seen that I was justified in sorting the seeds by the convenient method of the balance and weights, and of accepting the weights as directly proportional to the mean diameters. ¹³

There is another issue dealing with Mendel with which historians of genetics have had a hard time dealing. It is the issue of whether Mendel failed to report the results of certain experiments that did not support his ratios as closely as he might have Mod and whether, in other instances, he went so far as to falsify his data outright. The only facts come from Mendel's paper, and choosing an appropriate multivariate statistical approach for such voluminous data is a formidable problem.¹⁴ Mendel's experimental notebooks and other original records no longer exist . After his death of congestive heart and kidney failure in 1884, they were destroyed by order of his successor abbot. In this age of suspicion and scandal-mongering journalism, the destruction by itself would be insinuated as evidence that Mendel may have left instructions for this disposition so that his nefarious data tampering would never be discovered.¹⁵ There is, however, no evidence whatsoever to support any other interpretation but that Mendel's successor was cleaning out many boxes of notes and other apparently worthless materials.

1. Vítezslav Orel, Gregor Mendel: The First Geneticist, Oxford: Oxford University Press, 1996.
2. His sole published paper in the journal of the society was Gregor Johann Mendel, "Versuche über die Pflanzen-Hybriden", Verhandlungen des Naturforschenden Vereines, Brünn, 1866, 4: 3-47. English translation by Curt Stern and Eva Sherwood, "Experiments on Plant Hybrids", in C. Stern and E. Sherwood (editors), The Origin of Genetics: A Mendel Source Book, San Francisco: W. H. Freeman, 1966: 1-48. All Mendel quotations in the present paper, as well as their page numbers, are taken from Stern and Sherwood.
3. Hugo Iltis, Gregor Johann Mendel. Leben, Werk und Wirkung. Berlin: Springer, 1924. English translation, Life of Mendel, New York: Norton, 1932. 2nd edition: New York: Hafner, 1966.
4. Daniel L. Hartl and Viazslav Orel, "What Did Gregor Mendel Think He Discovered?" Genetics, 1992, 131: 245-253.
5. V. Orel and Daniel L. Hartl, "Controversies in- the Interpretation of Mendel's Discovery, " Hist. Phil. Life Sci., 1994, 16: 423-464.
6. Robert C. Olby, "Mendel No Mendelian?" History of Science, 1993, 17: 55-72.
7. Gregor Mendel (endnote 2) p. 43.
8. Gregor Mendel (endnote 2) p. 29, where Mendel uses the italics for emphasis. Close variants of this statement are on pages 24, 19, 32, twice on page 43 and 44.
9. Robert C. Olby, "Constitutional and Hereditary Disorders," in W. F. Bynum and R. Porter (editors) Companion Encyclopedia to the History of Medicine, New York: Routledge, 1993: 412-437. The quotation is from p. 421.
10. Francis Galton, Natural Inheritance, London: Macmillan, 1889. Quotation from p. 66. The sentence beginning "Whenever," which is at the end in the original passage, has been moved to the middle for continuity.
11. Ronald A. Fisher, "The correlation between relatives on the supposition of Mendelian Inheritance," Trans. Roy. Soc. Edinburgh, 1918, 52: 399-433.
12. Francis Galton (endnote 9), p. 80.
13. Francis Galton (endnote 9), p. 81.
14. The best paper on this subject is that of A. W. F. Edwards, "Are Mendel's Results Really Too Close?" Biological Review, 1986, 61: 295-312. It is also discussed in Hartl and Orel (endnote 4).
15. As recently as January, 1997, a well known evolutionary geneticist repeated this interpretation to me as if it were an established fact.

"Theory and Practice of Science" offered students the opportunity to conduct deep, focused studies of particular discoveries and developments of modem science, using original science papers as the basis for those studies; the course was taught by several instructors each year, usually from different departments, and we tried to encourage a seminar environment similar to the "core" humanities courses at Columbia. In the Fall semester, the students began with a few weeks of elementary mathematics followed by a unit on the discovery of nuclear fission, working with a series of readings that began with Faraday and concluded with Meitner and Frisch's paper of 1939. In the Spring, following a bit more mathematics, the students studied the discovery of the structure of DNA, reading a series of papers that began, of course, with Mendel.¹

One of the difficulties with teaching a science course based on a sequence of original articles, was that one could never devote enough time to each paper; on the other hand, and perhaps unlike the situation in a "Great Books" course in the humanities, there was something extremely dissatisfying about not completing the sequence of papers by the end of the term. Most of our students were, or would become, humanities and social science majors, and if their mathematical skills were unexceptional their analytical sldlls with scientific texts were extraordinary; as we pushed the class from one text to the next, I often felt badly that there was not enough time to simultaneously develop the science, the history, the rhetoric, and the philosophy in and behind those texts.

MENDELWEB, then, was designed to show how the World Wide Web could make possible such a multi-level, interdisciplinary explication and exploration of a text like Mendel's Versuche. The Web is defined in several ways today, but the basic idea remains that it is a system of distributed hypermedia: distributed, because the documents, images, databases, audio and video files, etc. are stored on computers ("servers") throughout the world; and hypermedia, because this data can be connected in ways that make "texts" resemble multi-dimensional (global) graphs more than leather-bound (local) volumes. MENDELWEB presents Mendel's paper (both in its original German and in the English translation attributed to Bateson), as the fundamental node in a graph that includes paths leading in many disciplinary directions, and this graph continues to grow.

The Foundation of MENDELWEB is Mendel's text. Formatted as hypertext (i.e. presented in HTML), the German and English versions are linked to each other, a glossary, a set of section summaries and notes, a set of discussion questions, and several homework sets. These texts are themselves linked to each other, to bibliographies, to electronic reference materials (e. g. on-line German-English dictionaries, genetics tutorials, and Web search engines), and to a chronology that presents several important events in Mendel's life alongside scientific, historical, and literary events throughout the world. These "pages" are also linked to each other, to reprints and original publications in MendelWeb's Essays and Commentary section, ² and to biology, botany, mathematics, and history resources throughout the world.

MENDELWEB also makes use of some of the interactive features of the Web. There is a statistics page that allows people to calculate averages, variances and standard deviations on data they "paste' onto that page, and, more significantly, there is a collaborative edition of Mendel's paper that allows readers to annotate both the German and English versions of Mendel's text.³ Finally, there is the "Mendelroom," a virtual space located on the educational MOO⁴ known as "Diversity University," which allows users of MENDELWEB, no matter where they are in the world, to read, write and chat with each other while they explore the texts at the web site. I have been keeping regular "office hours" in the Mendelroom for more than a year.

Since the first edition appeared in 1995, thousands of students and teachers from all over the world have used and continue to use MENDELWEB. The site has won numerous awards, ⁵ and the 6th edition of MENDELWEB (97.1) was released in February (another edition is planned for the summer). The significance of MENDELWEB, however, remains as a prototype, or perhaps more accurately, as a motivation, for the development of a large-scale distributed curriculum in genetics, and biology more generally. ⁶

Imagine, for example, a Web-based genetics curriculum in which MENDELWEB formed only a very small part. The larger Web would include webs spun from seminal papers by Morgan, Sturtevant, Beadle & Tatum, McClintock, Hershey & Chase, Watson & Crick, and Jacob & Monod, each constructed by scholars and departments at different colleges and universities, each linked with one another. Imagine too that this larger Web included multimedia presentations and lectures by contemporary geneticists and master teachers, as well as tutorials and distributed monographs ("polygraphs"?) on subjects such as early cytogenetics, the biochemistry of proteins, comparative histories of genetics research, a Molecular View of "dominant" and "recessive", and the Human Genome Project. Imagine finally that this Web-based curriculum persisted, was revised, and grew over time, that it captured the insights not just of important research but of great teaching as well, and the reasons for thinking the Web a genuinely revolutionary technology become clear. It is especially appropriate, perhaps, that one of the first steps toward such a distributed curriculum should derive from Mendel's paper, at MENDELWEB. ⁸

1. The original teachers and authors of the course were: Jonathan L. Gross, a Professor of Matheniatics, Mathematical Statistics and Computer Science; Herbert Goldstein, the author of the legendary Classical Mechanics; and Robert E. Pollack, a Professor of Biological Sciences and the Dean of Columbia College during the period I was involved with the course. I was extremely fortunate to have been their student, and later their colleague; sadly, the course is no longer offered as a two-semester, interdisciplinary course at Columbia. The syllabus for 'Theory and Practice of Science' is available at MendelWeb (http://www.stg.brown.edu/MendelWeb/ TPS.syll.html).
2. The Essays and Commentary section currently features articles and essays about Mendel, his work, and the reception of his. work by Hartl, Kimmleman, Olby, Orel, Paul, and Sapp, as well as a travelogue, by Margaret Peaslee, that includes photographs of various Mendel landmarks in Brno and Hyncice.
3. Most surprising, perhaps, is the fact that this collaborative hypertext has been in existence for more than a year, and virtually no one has contributed to it! I cordially and enthusiastically invite the readers of the Mendel Newsletter to consider contributing comments to the collaborative edition of Mendel's paper.
4. A MOO is basically a computer program that allows a number of users to log in at the same time and carry on synchronous communication. The "OO" in the MOO means that it is "object-oriented", which means that you can manipulate objects (e.g. read and write texts) in addition to chatting.
5. In addition to various "Best of the Internet" awards, MendelWeb was recently the "Link of the Month" at Educom and a "Cool Site of the Day" at the National Academy of Sciences.
6. For an explanation of distributed curriculum, see the Frequently Asked Questions (for FAQ) at MendelWeb (http://www.stg.brown.edu/MendelWeb/MWfaq.html). For an example of a distributed science curriculum in the making, see Kenneth Foote's "Virtual Department of Geography" project, based at the University of Texas at Austin (http://www.utexas.edu/depts/grg/virtdept/contents.html).
7. The current edition of MendelWeb is available at Netspace (http://www.netspace.org/MendelWeb/), the University of Washington at Seattle (http://hermes.astro.washington.edu:80/mirrors/MendelWeb/) and the Scholarly Technology Group at Brown University (http://www.stg.brown.edu/MendelWeb/).
8. My thanks to Diane Paul for her encouragement, her contributions to MENDELWEB, and for allowing me to describe MENDELWEB in this Newsletter.