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ROUTES TO INNOVATION IN BIOLOGY
By OREN HARMAN, MICHAEL R. DIETRICH
THE UNIVERSITY OF CHICAGO PRESSCopyright © 2013 The University of Chicago
All rights reserved.
THE MANY SIDES OF GREGOR MENDEL
Gregor Mendel (1822–1884) was long viewed as the ultimate scientific outsider. After all, he was not celebrated as the founder of genetics until sixteen years after his death. He was not a professional researcher by any definition, but a monk and a schoolteacher, and later an abbot. He lived and worked far from the great Europe an intellectual centers of his day, in Brünn, provincial capital of Moravia, then part of the Austrian Empire (Brno, in Czech; since 1993, part of the Czech Republic). He presented the results of his experimental crosses of pea plants, along with the foundational ideas of gene tics, only at his local scientific society in 1865. His now-famous article on the subject came out in the Proceedings of the society in 1866, not the most conspicuous place for it, and, indeed, little notice was taken of it at first. It took until 1900 for it to be "rediscovered" and recognized as a cornerstone of the emerging science of gene tics. Only then did Mendel find a place among the scientific insiders, or so it seemed to early geneticists.
The geneticists quickly embraced Mendel as one of their own. His name was given to "Mendel's laws," traits that "Mendelize," "Mendelism" as a theory of heredity, and "Mendelism" as a theory of evolution. Geneticists from all over the world donated money for a Mendel monument in Brünn. Mendel's work seemed to fit in so well in twentieth-century science that it was hard to imagine him ever belonging anywhere else, and the image developed of Mendel as a man so far ahead of his time that he had no intellectual company, no peers, no teachers. Unencumbered by old-fashioned preconceptions or intellectual commitments, he could see ahead to future problems and solve them, while his shorter-sighted contemporaries would not have seen the point of his work, had they even read it.
The blinkered view of Mendel as a figure without peers is reflected in one of the best-known photographs of him (figure 1.1), which is actually a detail from a group portrait. The corresponding narrative of the lone and unrecognized genius is told most eloquently by Loren Eisley. Here is how he describes Mendel's 1865 presentation to the Brünn Society of Naturalists: Stolidly the audience had listened. Just as stolidly it had risen and dispersed down the cold, moonlit streets of Brünn. No one had ventured a question, not a single heartbeat had quickened. In the little schoolroom one of the greatest scientific discoveries of the nineteenth century had just been enunciated by a professional teacher with an elaborate array of evidence. Not a solitary soul had understood him.
Thirty-five years were to flow by and the grass on the discoverer's grave would be green before the world of science comprehended that tremendous moment. Aged survivors from the little audience would then be importuned for their memories. Few would have any.
A few pages later, Eisley offers this judgment about Mendel's place in history:
Mendel is a curious wraith in history. His associates, his followers, are all in the next century. That is when his influence began.... Gregor Mendel had a strange fate: he was destined to live one life painfully in the flesh at Brünn and another, the intellectual life of which he dreamed, in the following century.
It is a tragic tale indeed. But cheer up: I am going to tell it a little differently by drawing on more recent historical research that has found some company for Mendel—some associates, audiences, and influences.
Geneticists and historians alike have spun the Mendel story for a variety of rhetorical purposes. Versions like Eisley's, which emphasize long neglect and isolation, tend to aggrandize twentieth-century Mendelians for their greater openness to new ideas and their superior understanding of Mendel's paper, while the idea of a rediscovery reassures us that science is self-correcting and sooner or later will give credit where credit is due. But we should also make allowances for the lack of information about Mendel's community and context available to the rediscoverers. For them it was hardly unreasonable to infer that his monastic life and teaching duties precluded full-time research and close contacts to the international scientific community, or that the long neglect of his paper was good prima facie evidence that he had been misunderstood in Moravia and ignored everywhere else.
But more information is now available, and it has become clear that, for all the modernity of his scientific thinking, Mendel was also rooted in the nineteenth century: in the intellectual, economic, and religious life of Brünn, in the pure and applied sciences of the Austrian Empire (especially meteorology, biogeography, plant breeding, and evolution) and in Europe an science generally. Several separate lines of post-Eisley research have recovered sides of the historical Mendel other than just the misplaced geneticist. I will try here to bring these lines together into a more complete portrait and to show how his many sides also enrich our understanding of his famous paper.
Recent historical research also recognizes multiple nineteenth-century sources for later ideas about heredity. Far from springing fully formed from Mendel's head, genetics is beginning to look like a synthesis of many lines of thinking, not only in biology, but also in medicine, agriculture, law, and other spheres. From this point of view, the many-sided Mendel to be developed here, with his multiple affiliations, audiences, and intellectual resources, makes a much more plausible founding figure than the lone outsider. His achievement then emerges as less a crossing of existing disciplinary boundaries than as a merger of disciplines into something new.
HISTORICAL AND POLITICAL BACKGROUND
In the eighteenth century, some eighty years before Mendel's birth, the Habsburg dynasty ruled over a sprawling multinational, multicultural realm in Central and Eastern Europe, comprising what is now Austria, Hungary, the Czech Republic, Slovakia, Croatia, Slovenia, Bosnia and Herzegovina, Romania, and parts of Poland and Italy. Along with Russia, France, and Britain (Germany was still fragmented into many separate kingdoms and principalities), the Habsburg Empire was one of the "great powers" of Europe, militarily and politically.
Its rulers were shocked, however, in 1740, when the much-smaller Kingdom of Prussia attacked and captured their northern province of Silesia, their most highly industrialized province, with its textile factories and iron and coal mines. The loss was an economic blow as well as a military embarrassment. The young empress, Maria Theresa (ruled 1740–1780), spent over twenty years trying to get Silesia back, fighting the War of the Austrian Succession (1740–1748) and the Seven Years' War (1756–1763) against Prussia and a changing constellation of allies. She never did reclaim it, but held on only to a sliver, where the Mendel family happened to live, and which became part of Moravia, administratively.
Maria Theresa's efforts to compete with Prussia and maintain the status of the empire as a great power were not exclusively military and diplomatic in nature. She, and especially her son Joseph II (ruled 1780–1790, but coregent with his mother from 1765 on), also modernized and centralized their administration. They applied the eighteenth-century ideals of "enlightened absolutism," a political and economic theory that emphasized efficiency and rational organization, and they instituted reforms that aimed to weaken the competing, decentralized, redundant, and inefficient feudal powers, such as the noble landlords, high church officials, and religious orders. Some of these Theresian-Josephine reforms were to have direct effects on Mendel's life and career: partially emancipating the serfs, modernizing agricultural practices, making primary and secondary education more widely available, encouraging the study of the natural sciences, and forcing the monasteries to be of ser vice to the Empire and the economy.
MENDEL'S EARLY LIFE AND SCHOOLING
Mendel's father, Anton Mendel, was a peasant in a little village called Heinzendorf (Czech: Hyncice). The family was ethnically German, and the region majority-German as well. Anton Mendel took a special interest in cultivating and grafting fruit trees, and taught young Mendel how to do it, too. He was encouraged in this by the local countess, an enlightened ruler in the Josephine tradition, who made an effort to promote scientific agriculture in her territory. She imported and distributed fruit trees and had natural science taught at village schools.
Our scientist was born Johann Mendel on July 22, 1822, the second of three children. (He took the name Gregor as an adult, upon entering the monastery.) Only limited schooling was available nearby, but because he was considered exceptional, the priests who taught the village children arranged for him to continue his education in town, and helped him talk his parents into it. They had little means of financing his studies, and they would miss the help of their only boy on the farm, for peasants like Anton Mendel had been only partially emancipated. He owned his own farm, but still had to work three days a week for the countess, an obligation known as the Robot.
In 1834, at the age of thirteen, Mendel began attending Gymnasium (the academically oriented secondary school that opened the door to university education and the elite professions) some twenty-five miles away. His parents could only afford room and half board, and he had to tutor slower but wealthier pupils to earn his lunch money. Mendel got along like that until 1838, when his father was crushed in a logging accident while performing his Robot. He survived, but never recovered fully, had trouble maintaining the farm, and could not support his son at school as before. Mendel in the meantime had earned a teaching certificate that qualified him as a private tutor and was able to eke out a living at school, but at times, the pressure of school and work and worries about his family became too much for him. He suffered some kind of breakdown in 1839 and returned home for several months to recuperate. Several more of these breakdowns are recorded, both during and after his schooling, but no precise medical information about them is available.
In 1840, Mendel completed Gymnasium and moved on to Olmütz (Czech: Olomouc) in southern Moravia for a two-year course of university-level study (all that was offered; there was no full-fledged university in Moravia). Because of another breakdown and an extended stay on the farm, it took him three years to finish at Olmütz, after which he decided not to continue his struggle to get a university degree. Instead, he followed his family's wishes and the advice of his Olmütz professors (one of whom was a monk himself), and took holy orders. Against stiff competition, he was accepted into the Augustinian order, at the monastery of St. Thomas in Brünn in 1843. There he received training in theology and was prepared for the priesthood. He was ordained in 1847.
The monastery offered Mendel not only some much-needed security, but also an intellectual community and an opportunity to do good works and improve the lives of Moravians through education and applied science. The monastery had been influenced by the Josephine reforms, which had eliminated the more contemplative (or uncooperative) monastic orders and confiscated their property. The surviving ones had to be active and productive in worldly affairs.
The monks of Brünn served as highly qualified instructors at several Gymnasien as well as at Olmütz. Some were experts in scientific agriculture, managed the monastery's extensive landholdings, and made an effort to share their knowledge with farmers and businesses in the region. Several, among them the abbot, were interested in pure science, too, and they had experimental gardens, a herbarium, a mineralogical collection, meteorological instruments, and a big, up-to-date library. In short, Mendel found himself in learned company, who thought the study of nature was important and useful for their work and their community. It was, admittedly, not a major Europe an research center, but the isolation of the monastery should not be exaggerated, either. Had Mendel been seeking an opportunity to get involved in scientific research, he could hardly have made a more practical choice.
MENDEL'S NINETEENTH-CENTURY SOURCES (1): PLANT AND ANIMAL BREEDING
The work of the monks and the economic interests of the monastery and the surrounding community were tied to a large extent to agriculture and its improvement, and Mendel clearly was motivated to keep the practical goals of his research in sight. His work promised to explain, among other things, how traits combined and interacted in plant hybrids, and it suggested ways of rationalizing hybridization methods to produce desired combinations of traits at will.
An important line of research into Mendel's agricultural connections has focused on the theory and practice of animal breeding, particularly sheep breeding. Vítezslav Orel and Roger Wood have sought out the writings of sheep breeders who were active in Brünn from the 1810s through the 1830s and have analyzed their methods and their conceptions of heredity. The Moravians, like many breeders throughout Europe, built upon the work of British breeder Robert Bakewell (1725–1795) and others, who had explored ways of making breeding more systematic and scientific. They analyzed the animal into checklists of simple characteristics, measured those characteristics objectively and quantitatively, and devised procedures for comparing, selecting, and inbreeding the animals.
In Brünn, there were also significant discussions of theoretical matters, and it was asserted that heredity must be a law-abiding and predictable process, amenable to scientific study. The breeders there inquired into the effects of inbreeding and outcrossing, and into the stability of varieties. Why did inbred varieties not always breed true? What made them revert to their ancestral condition or become more average? When two varieties were crossed, what determined whether maternal or paternal characteristics would predominate in the off spring? Did it depend on the inner constitution or organization of the parent, or on environmental conditions?
Since Mendel only arrived in Brünn in the 1840s, when sheep breeding was in decline there and few of the older breeders were still around, it is difficult to document a direct influence of their methods and ideas on Mendel's later work. Nonetheless, it seems safe to assume that Mendel was aware of the earlier Moravian ideas and practices, either through his abbot, who had been a breeder himself, from the published breeding literature, or from younger plant and animal breeders who were active, along with Mendel, in the Brünn scientific society.
Excerpted from Outsider Scientists by OREN HARMAN, MICHAEL R. DIETRICH. Copyright © 2013 The University of Chicago. Excerpted by permission of THE UNIVERSITY OF CHICAGO PRESS.
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Table of ContentsPreface
Introduction: Outsiders as Innovators in the Life Sciences
Oren Harman and Michael R. Dietrich
I Outsiders Before the Inside
1 The Many Sides of Gregor Mendel
2 Louis Pasteur: The Chemist in the Clinic
3 Félix d’Herelle: Uncompromising Autodidact
William C. Summers
4 The Paradox of Samuel Butler: Insider or Outsider?
II Outsiders from the Physical Sciences
5 Erwin Schrödinger’s Excursus on Genetics
6 Linus Pauling: Leading Exporter of Chemical Insights into Biology
Gregory J. Morgan
7 From Bomb to Bank: Walter Goad and the Introduction of Computers into Biology
III Outsiders from Mathematics
8 R. A. Fisher and the Foundations of Statistical Biology
Michael R. Dietrich and Robert A. Skipper, Jr.
9 Nicolas Rashevsky’s Pencil- and- Paper Biology
Maya M. Shmailov
10 Searching for Patterns, Hunting for Causes: Robert MacArthur, the Mathematical Naturalist
IV Outsiders from the Human Sciences
11 Noam Chomsky and the Biology of Language
W. Tecumseh Fitch
12 Dunking the Tarzanists: Elaine Morgan and the Aquatic Ape Theory
Erika Lorraine Milam
13 David Hull’s Philosophical Contribution to Biology
T. J. Horder
14 Ilya Metchnikoff: From Evolutionist to Immunologist, and Back Again
Alfred I. Tauber
15 François Jacob: Tinkering with Organisms and Models
VI Outsiders from Informatics
16 Theoreticians as Professional Outsiders: The Modeling Strategies of John von Neumann and Norbert Wiener
17 On the Importance of the Parvenu: The Amazing Case of George Price in Evolutionary Biology
18 Outsiders and In- Laws: Drew Endy and the Case of Synthetic Biology
Epilogue: The Problem with Boxes
Richard C. Lewontin
List of Contributors