© Ton Munnich
For private use only
Professional and commercial use after consultation with the author
TABLE OF CONTENTS
Gregor Mendel’s environment
genetics and darwinism
William Bateson and Thomas Hunt Morgan
the Modern Synthesis
Gregor Mendel and Richard Dawkins
Evolution happens in two steps. Step one concerns the fact that every new-born is unique: one has a different hair colour than the other, one can run faster than the other, one has different fingerprints than the other. The result is an infinitely varied reservoir of unique individuals. This variation is a matter of heredity, of genetics, the terrain of Gregor Mendel. Then follows step two: the different individuals enter into competition: who maintains oneself in the hostile world outside. Who is well adapted to the circumstances? Who survives, who perishes? This second step is Darwin’s theme, with the keywords ‘struggle for life’, ‘survival of the fittest’ and ‘natural selection’. It is clear that Mendel’s terrain of genetics is the primary factor in evolution, and Darwin’s natural selection is the secondary. First there must be variation, only then can nature select from that varied supply, i.e. only then can there be natural selection.
Mendel and Darwin both start working on their decisive publication in 1856. Mendel’s research takes ten years, his article is published in 1866. Darwin also starts a long writing haul. Wallace, his competitor, forces him to shorten his planned book to a summary, which he publishes in 1859. That is On the Origin of Species, his book about natural selection. In it he leaves unanswered the prime issue: he does not know how variation comes about, he does not know how heredity works. He talks in grand terms about the origin of species, but he doesn’t know how a unique individual comes into being. As a result, Darwin’s idea of natural selection drifts loosely in the air, untethered, missing a foundation.
This essay is about that foundation: about heredity or genetics, about Gregor Mendel. And about the relation between Mendel and Darwin. For one and a half centuries, a curious relation exists between Mendel’s field (genetics) and Darwin’s idea (natural selection).
In the 19th century nobody knows how heredity works. All kinds of traditional views circulate. There are ovists and spermists. The ovists say the female egg cell is responsible for the new life, with a subordinate role for the male sperm. The spermists claim the opposite: the new life is in the male seed, whith the woman only providing food. Both views are incorrect. Another view in the 19th century sees heredity as a mixing process: information travels from all parts of the body to the genitals. Then the qualities of man and woman are mixed in the new individual. This mixing is imagined like the mixing of two fluids. Black and white paint together become grey, after which the grey cannot be separated into the original black and white any more. The properties ‘black’ and ‘white’ disappear in this way. This mixing view is also incorrect, as the following example shows. The fatherly quality of ‘man’ and the motherly quality of ‘woman’ should, when mixed, lead to a descendant who is a mixed form of man and woman, i.e. hermaphrodite. This is not the case, the offspring is either male or female, in an unmixed state. More generally: if in descendants the properties of their parents were to dissolve in a mixture, mankind would soon be a grey mass without properties. The opposite is true, each individual is unique. Therefore there must be another explanation for heredity. No one in the 19th century knows it. Also Darwin doesn’t know it.
According to Darwin, 1837 is the year in which he begins to think about evolution. He starts writing loose thoughts about it in notebooks, later referred to as the Transmutation Notebooks. The first Transmutation Notebook runs from July 1837 to February ’38. In 1960, leading biologist Sir Gavin de Beer publishes it. In his Introduction Sir Gavin writes that something struck his eye about the content. Darwin’s first notebook on evolution hardly mentions the themes that later become known as typically Darwinian. Sir Gavin: “Only the briefest references are made to the struggle for existence, selection, and adaptation”. Instead, the notebook chiefly discusses typical Mendel themes: heredity, variation, sexual reproduction, hybridization. In this first notebook, for example, Darwin uses 21 times the word ‘hybrid’, a typical Mendel word. And he fantasizes about an experiment with plants under a glass bell jar to prevent pollination by external pollen. It’s the kind of experiment Mendel performs. Apparently the young Darwin realizes that the evolution problem is first and foremost a question of heredity. But then he is unable to deal with this difficult issue of heredity, his attention wanders to something more simple. In 1838 he reads Malthus and from that point on he focuses on the typical Malthusian-Darwinian themes: struggle-for-existence, natural selection. But it clearly irks Darwin for the rest of his life that he could never solve the issue of heredity and variation, because that is why his natural selection theory lacks a foundation. In all editions of the Origin he wails: “the laws governing inheritance are for the most part unknown”, and “variability is governed by many unknown laws”. His biographer William Irvine aptly paints this fundamental gap in Darwin’s theory: “Already in the Origin of Species Darwin is haunted by the mystery of genetics. If variations cause evolution, what causes variations? He attacks the problem in the first and second chapters, and finally at length in the fifth. The discussion is … vague and occasionally confused. … At his best, he simply acknowledges a complete ignorance of the whole subject.” 1). Darwin’s fruitless brooding about heredity may be one reason why he waited so long with publishing his Origin.
After the Origin, heredity remains an obsession for him. In 1868 he publishes a two-volume book on the subject, The variation of animals and plants under domestication, with an updated second edition in 1875. Pompous style, vague theories, finally the announcement: “The possibility of selection rests on variability, and this … is governed by infinitely complex and unknown laws”. Who wants to understand Darwin’s contribution to evolution, should read this quote again. He says here that evolution begins with variation (i.e. heredity, Mendel’s department). Only then can there be any question of selection (and natural selection, Darwin’s department). And he confesses that the first department is a mystery to him. In short, Darwin has no theory of evolution, he only has a secondary puzzle-piece of it.
After 1868 heredity still remains an obsession for him. In 1876 another book on it, The effects of cross and self-fertilisation in the vegetable kingdom, in which he once again sighs about “the whole subject of hybridism, which is one of the greatest obstacles to the general acceptance and progress of the great principle of evolution”. He realizes that without a correct theory of heredity he has no theory of evolution.
Darwin cannot find it. For forty years he tries to fathom heredity. First alone, later together with his cousin Galton. Everything fails. Finally he gives up, writing in bitter disappointment to Galton in 1877: “I shall never work on inheritance again” 2). With that, the aged Darwin resigns to the fact that he could not find a theory of heredity, and therefore neither a theory of evolution.
One man in the 19th century knows how heredity works: Gregor Mendel (1822-1884). He studies it for a long while, his problem is lack of money. The solution is the Augustinian monastery in Brno. There, science is number one on the list of priorities. He enters the religious community, continues his studies, conducts his research and publishes his report in 1866. Nobody notices it. His monastery selects him as abbot, this task absorbes him. After his death, a pro-active monk burns his papers and notes, Mendel fades into oblivion. Until 1900. In that year science discovers the importance of his article, and the search for biographical data about this unique scientist begins.
The search lasts a century. In 1904, British biologist William Bateson travels to Moravia to learn more about Mendel. He finds little information, Mendel is vaguely remembered, as a simple monk and amateur gardener who, while cultivating his peas, accidentally discovered the laws of heredity. In 1924 Hugo Iltis publishes the first biography, which somewhat corrects that image. In 1965, the Moravian Museum in the Czech city of Brno dedicates a section to Mendel, the Mendelianum, headed by Vitezslav Orel. In thirty years of detective work, Orel corrects the traditional Mendel image. Mendel was not an unworldly gnome in a provincial monastery garden, he was familiar with contemporary physics, chemistry and biology. Mendel’s discovery is the result of scientific research, crowning years of interest in agricultural variety improvement in the progressive intellectual environment of Moravia. Orel writes a brilliant biography in which he reconstructs Mendel’s life and intellectual development. The book is published in 1996 in Oxford: Gregor Mendel, the first geneticist. Since this biography Mendel slowly emerges from Darwin’s shadow. The two following paragraphs are based on Orel’s book.
Gregor Mendel’s environment
In the 17th century the centre of gravity of the Enlightenment lies in the Netherlands. This not only concerns Dutch thinkers, also foreigners with too little thinking-space in their own country come to settle in the Netherlands. Well-known examples are René Descartes, John Locke, Pierre Bayle. Another example is the Moravian education reformer Comenius (1592-1670). Comenius initially works as a school principal in his home country of Moravia. The Protestantism of Jan Hus is an important movement there. When the Catholic Habsburgs occupy the region in 1620, the Protestant Comenius is forced into into exile. After some wanderings he ends up in the Netherlands. Comenius has modern ideas about education. His books on the topic meet with considerable success. Kings and town councils ask his advice in educational matters. A century later Comenius is a key source of inspiration for education reformer Johann B. Basedow, who founds a new type of school in Germany, the Philanthropinum, with several establishments. One of these is the Philanthropinische Erziehungsanstalt Schnepfenthal. Around 1790, a teacher called Christian Carl André is working in this institute. Let us focus now for a moment on this André. A man of the Enlightenment with an interest in science and progress. In 1798 André leaves for Moravia, a province of the Habsburg Empire. There he becomes head of the Protestant school in the provincial capital Brünn, today’s Brno. This brings him back to the position that Comenius had left 180 years earlier: a modern headmaster in Moravia in the conservative Catholic Habsburg Empire.
André develops a modern and stimulating environment in Brno. In 1800 he starts a newspaper. The paper is embedded in the intellectual climate of the Josephinische Aufklärung. To explain: under the influence of the French Enlightenment, progressive Austrians also want an Enlightenment. The government distrusts them because in France the monarchy ended under the guillotine. The Josephinische Aufklärung tries to allay this distrust. The movement does not want political upheaval or dethronement of the sovereign. But it does want Enlightenment for the people. This means promoting education and science, better justice, a liberal press, modern farming methods, in short, opening the windows to modern developments. That is the wavelength of André’s newspaper. Located in Brünn, it has correspondents throughout the Habsburg Empire. Pestered by censorship, he stops the journal in 1805. It has existed long enough to be an example for similar magazines that appear elsewhere in the Habsburg Empire in the 19th century.
André’s energy is not exhausted by this. He becomes acquainted with Count Salm-Reifferscheid. The Count has an interest in science. Together in 1806 they found Die Mährische Gesellschaft für die Verbesserung von Landwirtschaft, Naturwissenschaft und Landeskunde, simply referred to as the Landwirtschaftliche Gesellschaft (Society for Agriculture). The Society is interested in modern farming methods, especially improvement of the breeding of crops and farm animals.
As a subsection of the Landwirtschaftliche Gesellschaft, André founds the Pomologische Gesellschaft in 1816. Pomology literally means apple science, but the correct translation is Society for the Promotion of Fruit Cultivation. The new Society is engaged in the establishment of nurseries, the creation of collections of varieties of fruit trees and wine grapes, and their artificial insemination. One such nursery comes in the garden of the Augustinian monastery of Brünn. The abbot of the monastery, F.C. Napp, had already written a book about the improvement of fruit trees. In 1827, abbot Napp becomes chairman of the Pomologische Gesellschaft, later he also becomes chairman of the overarching Landwirtschaftliche Gesellschaft.
One contact in André’s extensive network is the German fruit expert G.C.L. Hempel, secretary of the pomological sister organisation in Altenburg and corresponding member of the London Horticultural Society. Hempel sometimes writes in the journal of André’s Pomologische Gesellschaft about European developments in the field of plant breeding. Especially an article in 1820 is remarkable. Hempel writes that one day it will be possible to use scientific methods to grow fruit varieties with exactly the characteristics desired. Shape, size, colour, taste, all by genetic design. Before that can happen, though, the mystery of hybridization will have to be cracked. Not an easy task, says Hempel, since it requires a whole new type of scientist. A scientist with thorough professional knowledge and the tenacity and perceptivity required to accurately carry out a large-scale study over several years. With this Hempel gives a surprisingly accurate description of Gregor Mendel, who will be born two years later and who indeed cracks the hybridization mystery with large-scale and modern research.
In 1814 André founds a second subsection of the Landwirtschaftliche Gesellschaft, the Schafzüchter-Verein (Sheepbreeder Association). Its members together buy foreign rams in order to improve wool production. The annual meetings of the Schafzüchter-Verein attract interested parties from home and abroad. In 1816 André’s son Rudolf André writes his much consulted manual for the selection of breeding sheep: Anleitung zur Veredlung des Schafviehs.
The University of Moravia is located in the city of Olmütz (Olomouc), fifty kilometres northeast of the capital Brünn. Partly on the suggestion of André, the university in 1811 introduces modern subjects in biology and agronomy. André’s former co-worker J.K. Nestler becomes professor of agronomy in 1823. Professor Nestler reflects on heredity for years. In 1836 he delivers the keynote speech at the annual meeting of the Schafzüchter-Verein. He identifies heredity as the most pressing problem requiring scientific clarification. Just a few kilometres away lives the 14-year-old schoolboy Mendel. He will conduct that research 20 years later.
André’s modern views lead, in 1820, to his expulsion from Austria by the authorities in Vienna. He by then has laid a solid foundation in Moravia, others continue his work. Science and technology come to occupy a prominent place in education in Brünn and the Landwirtschaftliche Gesellschaft expands with more departments. One of these is the Naturforschender Verein (Scientific Research Association), founded in 1849 by a group of teachers. The founding year 1849 is no coincidence, one year after the miracle year 1848. The liberal and scientifically interested Count Mittrowski becomes chairman, with the membership comprising teachers, doctors, pharmacists and others. In 1851 Mendel becomes a member. In 1861 the Naturforschender Verein has matured and leaves the Landwirtschaftliche Gesellschaft to continue as an independent scientific society. In 1862, it has 171 members, people from Moravia with scientific interest. Another 24 honorary members include well-known professors such as Robert Bunsen (Marburg), Jan Purkyne (Prague), Franz Unger (Vienna) and Rudolf Virchow (Berlin). The Verein is in contact with the whole of European science.
Another consequence of the miracle year 1848 is a new type of secondary education. Many find the gymnasium too old-fashioned, with its heavy accent on classical languages and literary subjects. There is a need for modern secondary education without classical languages and with more science and technology. This type of education becomes the Realschule. In Brünn a Realschule starts in 1851.
In 1848 Alexander Zawadski is Professor of Mathematics and Physics at the University of Lemberg (Lwow) in the east of the Habsburg Empire. That subject falls under the Philosophy Faculty, where Zawadski is also dean. In the revolutionary year of 1848 there is a call for modernization among his students and teachers. He agrees. However, in the Habsburg Empire the miracle year lasts for only two years, after which the conservative restoration begins. In 1853 the authorities in Vienna call Zawadski to account for the tumult of ’48. Holding him responsible, they dismiss him. Zawadski moves to Brünn, where the mindset is more enlightened than in other parts of Austria. He becomes a teacher at the Realschule, alongside Mendel. He also becomes a member of the Naturforschender Verein and in 1854 becomes its secretary. Thus he meets Mendel both at school and in the Verein. The two talk much about science, Zawadski becomes a mentor to Mendel, teaching him that biological research must be conducted using the methods of mathematics, physics and chemistry. 3)
At the monastery in Brünn, abbot Napp has his own difficulties. In Austria the wavelet of modernization in the miracle year 1848 lasts only a short time, until 1851. Then the conservative restoration starts. In this context the bishop pays an inspection visit in June 1854. The bishop thinks the monastery does too much in the way of science and education. He believes the contemplative side is deficient. After all, not science but prayer is the task of the monk. The bishop demands a change of course towards the medieval monastic rule. In his report to the archbishop, he even advises closing the monastery. Abbot Napp sees the danger and counters. He points to precedents and old rights that justify a commitment to science. With his many functions abbot Napp is a prominent person in the Habsburg Empire. The storm passes, the monastery continues as before. Mendel’s experiment starts.
Around 1850 Brno has a more modern intellectual climate than the rest of Austria. This fact seems to be noted by Dr. Josef Auspitz, a maths teacher at the Technical High School in Vienna. His modern views in the year 1848 lead to his dismissal, after which he moves to Brünn. He becomes director of the Realschule. On a photograph of the teaching staff Auspitz and Zawadski sit in the middle, the other teachers stand behind them in a semicircle. Mendel is among them.
All in all, the following picture emerges. In the half century from 1800 to 1850 Moravian breeders and farmers realize that traditional knowledge about breed improvement has reached a ceiling. Further insight is only possible when the mystery of heredity is broken, and that requires a scientific approach. The establishment of the Landwirtschaftliche Gesellschaft (1806) is a first step towards achieving this higher level, followed by two specialised organizations, the Schafzüchter-Verein (1814) and the Pomologische Gesellschaft (1816), and finally a purely scientific Society, the Naturforschender Verein (1849). Heredity is an important theme of the Naturforschender Verein. The half-century of Moravian interest in breed improvement gradually focuses on scientific research into heredity. It can be expected that someone from this environment will discover the laws of heredity.
Mendel is the one to do that. He studies the relevant subjects in Olmütz, Brünn and Vienna. Mathematics, physics, agronomy, botany, plant physiology. His mentor Zawadski teaches him the correct attitude: to think along scientific lines. Mendel rises from the traditional nursery level to an extremely modern scientific level.
At the same time, someone in England is moving in the opposite direction. Darwin descends from a scientific level to a nursery level. Darwin comes from a studied family, his grandfather, father and brother are doctors, and he is in academic company. But Darwin is not a scientist, he looks for the clou of heredity in the environment of breeders and growers. He asks gardeners, farmers, plant growers, dog breeders for information. While Mendel enrols at two universities, Darwin joins two pigeon clubs. Darwin searches in vain for the solution of heredity in the pigeon clubs. Mendel solves the puzzle with high-level science.
Around 1800 in the Moravian countryside there is a countess with a positive attitude towards the Enlightenment. She wants a modern upbringing for the young people in her county. She founds a private school on her estate, led by priest J. Schreiber. Schreiber is aware of the importance of modern science. André praises Schreiber in his newspaper, characterising his school “a kind of Philanthropinum”. However, the authorities are afraid of modernity, they find a reason to dismiss Schreiber (1802) and close down the school (1814). The catholic government downgrades Schreiber to be a parish priest in a farming village. It happens to be the village where Mendel grows up.
Schreiber had grown fruit at his school, on his advice the countess had ordered apple varieties from France. He takes material of these breeds with him to his new parish and distributes it among the peasants. Mendel’s father is a farmer; his mother is the daughter of a fruit grower, in the orchard are Schreiber’s new fruit varieties. Pastor Schreiber sees that young Mendel is talented, he advises his parents to let him continue his education. The boy goes to the gymnasium and the Philosophicum (scientific education) in Olmütz, with mathematics and physics in particular in his curriculum. The level is high, some teachers in Olmütz are authors of textbooks which are reprinted throughout Austria and used at the University of Vienna. But the costs of Mendel’s study weigh heavily on the meagre family income. When his parents cannot fund it anymore, he earns something himself, and finally his sister spends her dowry on her brother’s study. A teacher in Olmütz alerts abbot Napp of the Augustinian monastery in Brünn to the gifted student. In 1843 he is admitted to the monastery. That solves the financial problems and, above all, he is now in the intellectual environment where he belongs. The monastery in Brno is an environment of teachers, school principals and professors.
In preparation for his priestly ordination (1847), Mendel studies theology (1844-’47). At the same time he takes lectures in agriculture and horticulture with Professor Diebl at the Philosophicum in Brünn. Diebl has already published several books in that field, including a textbook with Professor Nestler from Olmütz, mentioned above. Diebl is interested in the improvement of plant varieties. Incidentally, 78-year-old Diebl also participates in the demonstration of progressive students in the miracle year 1848 in Brünn.
Manager of the monastery’s experimental garden is botanist Father Klacel, an advocate of scientific progress. The catholic censorship in Vienna considers Klacel’s publications too progressive, it withdraws his teaching certificate. Abbot Napp can do little about it. In 1848 Klacel asks Mendel to take over the management of the garden.
In 1849 Mendel starts teaching Greek, Latin and mathematics at the Gymnasium of the Moravian city of Znaim (Znojmo). He does very well, though without teaching qualification. The head teacher wants to keep him, he wants Mendel to teach biology and physics. Abbot Napp sends him to Vienna to obtain the teaching qualification in those subjects. He has one summer holiday (1850) to prepare the exams, but it is a volume of work for which the full-time students have several years. He succeeds in physics. The two examiners of physics are Professor Doppler (of the Doppler effect) and Professor Andreas Baumgartner (soon after, Minister of Trade, Industry and Public Works, also Minister of Finance and President of the Imperial Academy of Sciences). They are satisfied with candidate Mendel. The biology examination goes less well. Examiner Kner has written a book himself. Mendel does not know the book, he fails the exam.
In Brünn they are light-hearted about the failed biology teaching examination. Mendel starts teaching biology at the Technical School of Brünn (1851), to full satisfaction of the school management. In the meantime, abbot Napp has conferred with examiner Baumgartner. They decide Mendel will study at the University of Vienna. Autumn 1851 he begins his physics course with Doppler. He also enrols in maths, chemistry, zoology, botany, plant physiology and palaeontology.
From physicist Doppler and his successor Ettingshausen Mendel learns how to set up a scientific experiment. That is crucial, his own experiment will demand ten years of methodical accuracy. In addition, these physics teachers point to the importance of mathematics, in particular statistics, in scientific research. Mendel also takes this insight to heart, becoming one of the first to use mathematical methods in a biological experiment. Furthermore he is particularly interested in the view of plant physiologist Franz Unger. The stay in Vienna lasts two years, 1851-’52 and ’52-’53. He learns much but does not take a final exam. By May 1853 he is back in Brünn again.
In 1854 Mendel begins teaching physics and biology at the Realschule in Brünn. He is doing extremely well, but is still without qualifications, so that he falls into a low salary scale. It is an awkward situation, the gifted teacher on half pay. And financially disadvantageous for the monastery, because the salary that monks earn outside flows into the monastery coffers. Presumably abbot Napp thinks Mendel should make a second attempt to obtain his teaching qualification. That would give the talented teacher an appropriate status, and it would provide the monastery finances with a higher salary for decades. After all, Mendel is only 32 years old, and at the beginning of his educational career. Data are scarce. Fact is that in 1854 the monastery builds a greenhouse for the research Mendel wants to do. An expensive and unusual step. Perhaps it went like this: perhaps abbot Napp said: “I know you don’t want to get that certificate, but I also know you like to do your research. Let’s make an equal exchange: you’re going to Vienna again for that exam, and I shall make sure you get a greenhouse”. Be that as it may, in 1856 Mendel again takes the teaching qualification exam in Vienna. He fails again. The cause, according to the Mendel experts, is a difference of opinion between the now very expert Mendel and the old-fashioned examiner Fenzl.
From 1855 to 1865 he conducts his large experiment. In 1865, he presents the results in two lectures at the Naturforschender Verein meeting. In 1866 he publishes his article on it in the magazine Verhandlungen des Naturforschenden Vereines in Brünn. The article is called Versuche über Pflanzenhybriden. It is barely noticed.
Until 1868 he continues to teach at the Realschule (half salary). Then the monastery appoints him as abbot. This task hinders further biological research. After his death (1884) he is forgotten. When in 1900 biologists realize the importance of his article, his share price rises. In Moravia they hear about his sudden celebrity. Old people remember facts and anecdotes. Former colleagues, pupils, family. Mendel’s sister turns out to have two sons, Alois and Ferdinand. Because she had previously spent her dowry on his study, Mendel has paid the study of these nephews. The nephews, Dr. Alois Schindler and Dr. Ferdinand Schindler, provide useful information about their uncle after 1900. Furthermore there are letters here and there. Some documents emerge from the archives of the monastery, the University of Vienna and other institutions. With the scarce data it has taken the entire 20th century to reconstruct Mendel’s life.
In the decades of the 1840s and 1850s physicists and chemists are busy with the atom. The term ‘element’ plays a key role. One element after another is discovered. An honorary member of the Naturforschender Verein of Brünn, Robert Bunsen, also discovers two elements (caesium, rubidium). In 1869 Bunsen’s pupil Mendelejev presents his Periodic System of Elements. Mendel’s teachers Zawadski, Doppler and Unger are well informed about the developments in natural science. They teach Mendel that a biologist has to think along the lines of physics and chemistry. During Mendel’s studies in Vienna (1851-’53), Unger’s Botanische Briefe are published in the Wiener Zeitung. In the first letter, Unger defines ‘the plant as the most ingenious chemical laboratory, the most logical organization for the game of physical forces”. The key words in this quotation are ‘chemical’ and ‘physical’. Unger warns that botany will only make progress if it becomes a “Physik des Pflanzenorganismus” 4). This is how Mendel’s mind is prepared. When he interprets the results of his research with peas, he follows this strict physico-chemical line of thought. The materialistic line of thought. That way he achieves his breakthrough: he discovers that in biology there are elements, smallest particles as carriers of hereditary characteristics. He sees it correctly, they are the genes. The name genes is later invented by others, Mendel calls them elements.
Properties are fixed on material carriers, the genes. The parental genes remain discrete, i.e. separate, in the offspring. They form the individual, then they go their own way again to form new combinations. To put it in a comparison: genes are like differently coloured beads in a necklace. If the necklace is opened, the beads will form a different necklace in the next combination. The beads remain the same, the necklaces are different.
Mendel further clarifies the hitherto misunderstood fact that a property sometimes skips one or more generations. The characteristic is then not expressed in the child, but in the grandchild or great-grandchild. Mendel introduces the concepts of ‘dominant’ and ‘recessive’ for this purpose. Sometimes a gene does not participate in the design of the individual, the gene is, as it were, dormant. However, it is passed on to the next generation. In combination with other genes, there the trait can be expressed again.
In February and March 1865 Mendel gives two lectures about his research. The response is subdued, the listeners seem overwhelmed by the complicated “botanical algebra”. He then publishes the article, a compact argument of 45 pages. This too makes no waves in the scientific world. Mendel sends reprints to European institutions and scientists; later they are found uncut. He starts to correspond with Professor of Botany Carl Nägeli in Munich, trying to express his results more clearly. Again without result, Nägeli does not realize the breakthrough in Mendel’s text. Overviews of hybridization mention the article, but without recognizing its importance. It is not until 1900 that this happens.
When Mendel’s article is discovered in 1900, biology dives into heredity. In 1905 early Mendelian William Bateson gives it the name ‘genetics’. In 1909 the Dane Wilhelm L. Johannsen introduces a suitable name for Mendel’s ‘elements’. He calls them ‘genes’, so that there is no more confusion with the elements of physics and chemistry. Johannsen also introduces the terms ‘genotype’ and ‘phenotype’ to indicate a Mendelian distinction. The genotype is someone’s DNA, the phenotype is the person as a product of it. The phenotype is not an exact reflection of the genotype. The genotype also contains recessive genes, dormant genes that are not expressed in the phenotype. They can be expressed again in the next generation.
What caused the somnambulism about Mendel that lasted 35 years, from 1865 to 1900? One cause seems to be the fact that he is a monk. Scientific progress usually takes place in opposition to ecclesiastical opinions. No innovative contribution is expected from a monk, his article is left unread. And there is a second cause. At the time of Mendel’s release, Darwin’s Origin has already been on the market for seven years. Darwin’s chief of publicity Ernst Haeckel has unleashed a Darwin hype in which the terms ‘darwinism’ and ‘evolution’ overlap. In the decades of the 1870s and 1880s this leaves little room for the realization that a theory of evolution has to start with genetics. Finally there is a third cause: Mendel is simply ahead of his time. His research includes 24,000 pea plants in several crossbreeds over several generations. In this way he concludes that genes exist. But no one can see them, for the time being they only exist in his head. His colleagues fail to fully grasp what he is talking about. That changes around 1900. By then microscopic techniques have advanced to the point that one can see chromosomes, and one suspects that on these chromosomes are particles that contain hereditary characteristics. At that moment one suddenly understands what Mendel is saying. Then genetics develops explosively. In the 20th century, genetics becomes a core area of biology, with a wide range of applications.
genetics and darwinism
In 1852 Franz Unger has inventoried what is needed for a scientific theory of evolution. Research should follow two lines, one internal, one external. He calls the internal line the “genetische Behandlung” (genetic approach) of evolution: how does heredity cause offspring to deviate from their parents? The external line investigates how external circumstances exert pressure on the direction of evolution, i.e. how life adapts to the environment. Unger calls the first line of research the essential one: evolution stands or falls with the potential that offspring differ from their parents.
The second line of research is the first to receive an answer: in 1859 Darwin gives his natural selection answer. The darwinists present it as a complete theory of evolution; the unsolved genetics issue disappears from the picture for decades. Mendel’s genetics article (1866) also remains unnoticed for decades. But not everyone believes the darwinian claim that natural selection is a complete theory of evolution. The top biologists respond reservedly. Franz Unger, Asa Gray, Thomas Huxley, Rudolf Virchow, they understand that Darwin’s theory is preferable to the Christian doctrine of creation, but they doubt whether it is a complete doctrine of evolution. Gradually it becomes clear that they are right, the idea of natural selection is too limited an answer to the question of evolution. Around 1900 there is so much unease among biologists about the limitations of Darwin’s theory that it threatens to disappear altogether 5).
That unease is expressed in an elegant metaphor by Carl Nägeli, a prominent botanist at the end of the 19th century. He calls nature a tree, the leaves of which represent the many species. The gardener who regularly prunes the tree is called Mr. Natural Selection. This gardener, Mr. Natural Selection, always cuts away the leaves that are the least fit. When someone asks: ‘how did the leaves on the tree come into being?’ one cannot answer: ‘because the gardener did not cut them away’. With this simple metaphor Nägeli illustrates that natural selection is only about the survival of species, not about their arising 36). Even more succinctly botanist J. Arthur Harris puts it in his witty one-liner: “Natural selection may explain the survival of the fittest, but it cannot explain the arrival of the fittest” 37). In short, the emergence of unique new life forms is not explained by Darwin’s idea of natural selection. It is explained by Mendel’s field, namely heredity or genetics.
Around 1900 English biology is in a deplorable state. The Darwin clan reigns, with leading roles for Darwin’s cousin Francis Galton and his kindred spirits Karl Pearson and Raphael Weldon. This Darwin clan has made all kinds of attempts to solve the genetics problem. Darwin’s blending idea, Darwin’s gemmula idea, Galton’s Law of Ancestral Heredity and Pearson’s variants thereof. It is all wrong. In 1900 international biology finally discoveres Mendel’s article. That shows the correct approach of genetics. One would expect the Darwin clan to be enthusiastic about it, after its own failures in this area. But no, driven by arrogance and territoriality, the clan rejects Mendel’s work.
William Bateson and Thomas Hunt Morgan
Not all English biologists follow the Darwin clan in its rejection of Mendel’s genetics. One biologist in particular recognizes its value: William Bateson. For twenty years he is the leading advocate of the new field of genetics. In the decade 1900-1910 he and his kindred spirit C.C. Hurst polemize against the darwinists Pearson and Weldon. The fierce discussion ends in victory for Bateson and Hurst. In 1906, at the peak of the controversy, Weldon dies, 46 years old. Pearson writes that just before his death Weldon spoke vehemently about the polemic, somehow suggesting that Bateson and Hurst are partly responsible for his death 6). However, Weldon tried the impossible: he tried to refute Mendelian genetics which is correct 7).
Bateson agrees with Unger: evolution happens in two steps. First the genetic step, then the selection step. The genetic step is the essential, creative one. It creates variation, it creates new forms of life, each individual is unique. Only after that does the selection step follow. In an important speech in 1904, Bateson points to this limited role of selection: “its function is to select, not to create”. Later in his speech he repeats it more clearly: “Variation leads; the breeder follows”. First there must be variation, only then there is something to select, be it planned selection (in the nursery) or natural selection (in the wild) 8).
In 1909 England celebrates the hundredth anniversary of Darwin’s birth. In this context a collection of articles by leading biologists is published. Among them Bateson. In his contribution he reiterates that he does not regard natural selection as the alpha-and-omega of biology. “To begin with, we must relegate Selection to its proper place. Selection permits the viable to continue and decides that the unviable shall perish; … as the course of descent branches in the successive generations, Selection determines along which branch Evolution shall proceed, but it does not decide what novelties that branch shall bring forth” 9). In short, Bateson sees no creative role for natural selection in evolution.
Bateson goes on to disqualify natural selection even further: he simply calls it “a truism which needs no special proof” 10). He finds natural selection obvious and self-evident. The darwinian sound bites natural selection and survival of the fittest express something that has been known since time immemorial: an individual that is better adapted to a situation, maintains itself better. This is so evident that no one will think of claiming anything else. In running, the fastest wins; in wrestling, the strongest wins; in chess, the smartest wins. Of course that is true, of course a better adapted individual has more success, it is almost a tautology. Tautologies are somewhat hypnotic in their quasi-deep circle of thought: ‘when you are dead, you are dead’. Bateson feels that way about the idea of natural selection: quasi-deep but in fact paper-thin. Other biologists expressed the same feeling by typifying Darwin’s idea as ‘very simple’ or ‘the egg of Columbus’. At the egg of Columbus one also has the uncomfortable feeling that one has been taken in. It is Bateson’s firm belief, and ten years later he repeats it in an important speech: “The doctrine of the survival of favoured races is a truism, helping scarcely at all to account for the diversity of species” 11). He states that the idea of natural selection does not contribute to answering the question of how variation arises. That question will not be answered along lines of Charles Darwin but along lines of Gregor Mendel. Bateson gives his son the name Gregory.
Bateson is not religious, he stands in the tradition of the Enlightenment. His favourite author is Voltaire, his wife later remembers: ” I think he never travelled without a copy of Candide in his pocket” 12). He paints, he frequents galleries and art auctions, he finds the Morning Post a good newspaper because it has “the best accounts of art sales” 13).
In the period 1900-1920 Bateson is the figurehead of Mendelian genetics. Around 1910 the American biologist Thomas Hunt Morgan realizes its correctness. His star rises rapidly, in the years 1920-1940 Morgan is the figurehead. What Mendel did with peas Morgan does with fruit flies, his Drosophila research is awarded the Nobel Prize in 1933. Like Bateson, Morgan repeatedly points to the limitations of Darwin’s idea of natural selection. Genetics is the approach to understanding evolution. New gene combinations, that is primary. The fact that the less fit variants are then destroyed by natural selection is secondary, “because the destruction of the less fit does not in itself lead to anything that is new” 14), says Morgan in 1916. Thus there is a creative and a destructive step. The creative step is genetics, the destructive step is natural selection. Morgan holds that view all his life. In his essay What is Darwinism? (1929), as a thought experiment he compares a purely mendelian world with a purely darwinian world. An exclusively “Mendelian universe”, i.e. without natural selection, would have far more variation than there is today, because the less-fit species are not eliminated there by natural selection 15). So evolution takes place especially in Mendel’s world: that is where variation comes into being. The phenomenon of natural selection has an additional function: it cuts away the less adapted variants. In 1932, just before his Nobel Prize, Morgan expresses this a third time, when he writes succinctly: “natural selection does not play the role of a creative principle in evolution” 16).
Morgan also has an opinion on how a scientist should do his job. In 1891 he works as a young scientist at Bryn Mawr College. There he meets embryologist Jaques Loeb, who has just arrived from Germany. In 1892 Loeb leaves for Chicago University, but the two keep in touch via the Marine Biology Laboratory at Woods Hole. From Loeb Morgan learns the scientific attitude of the German materialists. Loeb has worked at the universities of Berlin and Würzburg, where Virchow’s spirit reigns. Loeb corresponds with Ernst Mach. Loeb has worked on the Institute of Marine Biology in Naples, which was established partly through the efforts of Vogt and Virchow. The director in Naples is Virchow’s friend Anton Dohrn. Morgan works for ten months in Naples (1894-’95). His scientific attitude stands in this German tradition. His non-religious, materialistic, mechanistic attitude is in the tradition of Virchow, Vogt and Moleschott. In 1929 Morgan writes an essay on his materialistic view. In it he points to the explanatory power of physics and chemistry. Organic material has many properties, but physics and chemistry can explain it all. When a phenomenon seems mysterious, this is no reason to seek explanations outside of natural science, for the possibilities of physics and chemistry are endless: “the combinations and permutations of the carbon compounds alone seem almost limitless” 17).
Morgan is a scientist. Experimenting, checking, falsifying, verifying – that is his job. Research, laboratory work and test installations form his world. He finds darwinism inadequate, Darwin’s ideas have not been scientifically tested. In 1932 Morgan’s book The Scientific Basis of Evolution is published. That ‘scientific basis’ is lacking in Darwin’s theory, he says. “The theory was more of the order of a broad generalisation than of a scientific theory based on controlled experimental data”. Further on he specifies his strict judgement on Darwin’s idea: “it could not fully meet the exact requirements of scientific theory that chemistry and physics demand as the essential basis of their conclusions, namely, predictions that can be verified by means of quantitative data.” It is necessary, Morgan says, “to rescue the theory of evolution from the vague speculative methods of its immediate past” 18).
the Modern Synthesis
Slowly, genetics and darwinism come to be on speaking terms. At the heart of that process is the work of R.A. Fisher, J.B.S. Haldane and Sewall Wright, their main works appearing in the years 1930-’32. They see that since more than half a century darwinism suffers from a shortcoming. This is the shortcoming: darwinism speaks about the origin of complete species, but meanwhile it does not know how one unique individual comes into being. A theory of evolution must begin at the beginning, at the arising of unique individuals: at genetics. If such a unique individual fits well in the environment it will successfully reproduce, and many generations later a new breed or a new species will emerge. In other words, genetics starts at the individual level and continues to work at the population level. Fischer, Haldane, Wright call it ‘population genetics’, a term that expresses that evolution is in fact genetics. That way they have inserted the unguided projectile Darwin in the field of genetics, in the population genetics sub-area. Genetics is the encompassing paradigm that dominates the field, and within that, Darwin’s idea of natural selection is one of the concepts that play a role.
In 1937 Morgan’s co-worker Theodosius Dobzhansky also publishes a book that approaches the origin of species as a genetics issue. Its title says it all: Genetics and the Origin of Species. Dobzhansky thinks genetics offers a more scientific approach than darwinism has provided to date. It is clear by then that Mendelian genetics is indispensable. Only the Darwin clan yet has to recognize it. That is not easy, the clan resists. Now that it is clear that genetics is the future, the clan wants to present it as a science of darwinian making, not of mendelian making. In particular clan member R.A. Fisher, professor at the dubious Galton Professorship of Eugenics at University College London, a chair that is financed by Darwin’s cousin Francis Galton. Fisher does not want Mendel to have the credits. In 1936 Fisher writes an article in which he describes Mendel’s work as fraudulent. It must have been tampered with, he says, because Mendel’s results are too good. Fisher understands that it is implausible to accuse the good father of fraudulent intent, but he has a solution for that: an assistant of Mendel must have committed the fraud. An assistant who knew what results Mendel expected, and who adjusted the numbers accordingly: “… Mendel was deceived by some assistant who knew too well what was expected. This possibility is supported by independent evidence that the data of most, if not all, of the experiments have been falsified so as to agree closely with Mendel’s expectations”. With this imputation Fisher hopes to push Mendel out of the picture, so that the Darwin clan, and he himself in particular, can show off as the real initiator of genetics. But Fisher’s attempt fails, Mendel’s work is correct. In the end, the clan has to acknowledge that Mendelian genetics has the future.
In 1942 clan member Julian Huxley finds the diplomatic phrasing that limits the loss of prestige for the darwinists and reconciles them with the reality of genetics. Julian Huxley, grandson of T.H. Huxley, is the spokesman of the Darwin clan in the years 1940-1970. In 1942 he published a book showing the dilemma of the clan. On the one hand he wants to maintain the Darwin clan’s monopoly and trivialise Mendel. On the other hand, he must recognize Mendel’s merits. The dilemma emerges in particular in one passage. Huxley begins the passage by trivialising Mendel. Agreeing, he quotes a biologist who says that Mendel’s work is only a theory about heredity, not about evolution. He considers this view to be “true”, and he calls the biologist in question an “eminent geneticist”. But then he says the opposite: “Mendelism is now seen as an essential part of the theory of Evolution. Mendelian analysis does not merely explain the distributive hereditary mechanism; it also, together with selection, explains the progressive mechanism of evolution.” 19). In short, Huxley contradicts himself within eight lines.
Huxley feels the great explanatory power of Mendelian genetics. He thinks: ‘if you can’t beat them, join them’. He seeks compromise. He might have wanted to dedicate his book to Mendel, but the Second World War is raging, under these circumstances it is less opportune to dedicate a book to a German-speaking scientist. He finds a solution that is almost as good, he dedicates his book to the American Thomas H. Morgan, the leading geneticist of the time. He flatteringly calls Morgan the “leader in biology’s advance”, and he gives the book the title: Evolution – the modern synthesis. He means the synthesis of genetics and natural selection. It is the synthesis of an elephant and a mouse. The elephant is the mighty science of genetics, the mouse is the idea of natural selection. Huxley tries to reassure his fellow-darwinists as follows: “biologists may with a good heart continue to be Darwinians and to employ the term Natural Selection, even if Darwin knew nothing of mendelising mutations, and if selection is by itself incapable of changing the constitution of a species or a line” 20). These two ‘if’s are fatal for Darwin.
The Modern Synthesis-book is published during the war, in 1942. Germany seems to be winning, England is in danger of succumbing to the war effort. Under these circumstances, Huxley proposes his synthesis of English darwinism and Austrian genetics. After the war the situation has changed. England has won the war, the Darwin clan no longer likes the term Modern Synthesis. They prefer a term in which the name Darwin is more prominent, with a subordinate rank for genetics. They are going to talk about “neo-darwinism”. This term, in use since 1895 in a different sense, becomes their replacement for the term Modern Synthesis after the Second World War. The term neo-darwinism suggests that the primacy lies with darwinism, with genetics as a recently added component, lowly lodged in the prefix ‘neo-‘. It is a misrepresentation. In reality post-war genetics is an impressive science with many applications and great social significance, whereas Darwin’s idea of natural selection has no application whatsoever.
Genetics has three main fields of application. The first is the medical application: the geneticist informs carriers of certain genes about the chance that offspring will have hereditary disorders. The second field of application is identification. The public prosecutor uses DNA to identify perpetrators and victims of crime; in the future, paper identity cards will be replaced by DNA identification. And the third field of application is genetic modification in agriculture. This is the application that would have pleased the breeders and growers of 19th century Moravia the most. Genetic modification places genes of one species in another species. The result is a transgenic variant. Disease-resistant crops, long-life tomatoes, beta-carotene-rich rice, and so on, can be manufactured. And it’s not just about agricultural applications. The pharmaceutical industry has, among other things, transplanted the human insulin gene into the E. coli bacterium. Cultures of this transgenic E. coli bacterium now produce human insulin for diabetics. These are some of the many applications of genetics.
What about the applications of Darwin’s idea of natural selection? There are none. If we look more closely at his idea, we see that it is impossible for it to have applications. Darwin postulated natural selection as a counterpart to cultural selection. In culture, i.e. agri-culture (breeding, growing, farming), it is the grower who selects. Darwin’s postulate was that wild nature also selects. The word ‘nature’ in ‘natural selection’ refers to that wild nature, the jungle, the unspoilt nature. Natural selection is opposed to selection by growers in the domesticated situation. Against this background, it is clear why ‘natural selection’ cannot have applications. Applications disturb the untouched wild, applications are human interventions, applications change ‘natural’ into ‘artificial’. Applications change nature into culture. In short, talking about applications of natural selection is a contradiction, an oxymoron. Darwin’s idea of natural selection cannot have any applications. And indeed there are none.
For the neo-darwinists after the Second World War, this is a disappointing fact. The field of genetics becomes ever more important, while Darwin’s idea of natural selection plays a marginal role. In 1995 two professors try to suggest that yet Darwin’s idea can have practical application. In that year neo-darwinist G.C. Williams and R.M. Nesse publish their book Evolution and Healing, The New Science of Darwinian Medicine. Two things stand out from the title. It’s about ‘Darwinian Medicine’, so apparently an application of Darwin’s theory. And the title contains the word ‘new’, it’s about something new. The authors realize that the latter requires an explanation: why does this first application of Darwin’s doctrine only appear in 1995, almost a century and a half after the Origin? The authors discuss this question in their paragraph entitled Why did it take so long? 21). They don’t find a good answer. The real answer is simple: their book is not about darwinism but about genetics. The term DNA already appears on the first page, and soon the word is out: “First, there are genes” 22). Genes belong to genetics, a field with applications. The authors have simply stuck a Darwin-label on genetics to create the impression that Darwin’s idea can have applications. Their title is a trompe l’oeil, an optical illusion. There is no ‘Darwinian Medicine’. And the first one to say that was Darwin himself.
According to Darwin, natural selection is at odds with medicine. In the wild, natural selection eliminates everything that falls behind. That is a good thing, according to Darwin. Medical science hinders that elimination, so medical science is a bad thing. Darwin disapproves of the smallpox vaccination, which has saved thousands of lives. Without vaccination, natural selection would have been able to carry out its useful cleansing work. Now that all those weaklings stay alive, the population degenerates. Vaccination is “highly injurious to the race of man”, Darwin pontificates 23). Funny, the flunked medicine student Darwin thinks medicine is harmful to public health. And forty years later Darwin’s intellectual heir Karl Pearson repeats it elaborately: medicine keeps weaklings alive that would otherwise have been cleared up by natural selection. Pearson too believes that medicine is harmful and contrary to natural selection.
The conclusion is clear: there can be no ‘Darwinian Medicine’. There can be no application at all of Darwin’s idea of natural selection. For Darwin ‘natural’ means ‘without human interference’, every human intervention disturbs natural selection.
Since the beginning of genetics in 1900, darwinism had to drastically reduce its claims, resulting in Huxley’s book The Modern Synthesis. In that book, the Darwin clan recognizes the primacy of genetics. Huxley dedicates the book to geneticist Morgan, he elevates Morgan to the “leader in biology’s advance”. And Huxley acknowledges: “selection is by itself incapable of changing the constitution of a species or a line”. After 1945 the English biologists revoke that surrender, the darwinian claims reappear on the table. With that its weaknesses reappear too. In the 1960s and ‘70s the neo-darwinists produce curious reasoning in trying to make their ideology look airtight.
The basic principle of neo-darwinism is the postulate that egoism is the driving force behind living nature. Every act of every living being is motivated by selfishness, every individual wants to increase its own chances of survival. This selfish aspiration of all individuals leads to competition and conflict. Darwin describes the selfish aspiration and the resulting conflicts as the all-encompassing “war of nature”. In the words of neo-darwinist and Nobel Prize winner Peter Medawar evolution is an “entirely selfish process” 24).
The flaw in this point of view is that besides egoism there is altruism. Sometimes one does something for someone else (alter) instead of for oneself. In darwinian logic altruism cannot exist. Someone who spends energy (or time or money) on someone else cannot spend it on himself. So an altruist puts himself at a disadvantage in the race to evolutionary success. The egoists do not have this disadvantage, they invest all their energy in their own survival. The result should be that over millions of years the property of selfishness has survived and the property of altruism has become extinct through natural selection. And this is where the flaw is: altruism has not died out.
The altruism issue forces the neo-darwinists to adopt a position. They cannot admit that there is any form of altruism, it would go against the heart of darwinism. They argue that altruism is a form of deception, that altruism is, in reality, egoism. It is as if they say that white is actually black. To support it they give a long reasoning which, in a nutshell, goes as follows.
When parents live sparingly and leave a sum of money to their child as an inheritance, this seems, superficially considered, altruism because the child is another individual. But in reality it is selfishness: the child is the bearer of their genes, the money increases the chances of evolutionary success for their genes. What the neo-darwinists are doing here is to broaden the scope of the term ‘egoism’. Previously egoism was limited to ’ego’, in the new view also descendants of ego are covered by his egoism. In line with this, the neo-darwinists also broaden the definition of the word fitness 25). The old expression of Spencer survival of the fittest was limited to the individual. The new definition says: fitness is reproductive success, the measure for fitness is the number of offspring, fitness is “the number of offspring produced by an organism” 26). Biologist John Maynard Smith comes up with the name for this offspring egoism, he calls it “kin selection”: one selects family (kinship) when giving favours.
And once the self-interest has been extended to the offspring, it is a small step to involve other family members as well. Because brothers, sisters, nephews and nieces also carry the same genes as ‘ego’. Who does something for a cousin partly does it for himself, because the genes are partly the same. British biologist William Hamilton calls this “inclusive fitness” in 1964, and adds a formula: C < R x B in which C = Cost, R = Relation, B = Benefit. It means: the cost of an altruistic act must be less than the degree of affiliation multiplied by the benefit to the beneficiary family member. The British biologist J.B.S. Haldane says in concrete terms that in an emergency situation he is prepared to sacrifice his life for two brothers or eight cousins 27). He means that in eight cousins as many of his own genes survive as in himself. So someone who sacrifices his life to save eight cousins is not an altruist but an egoist. It is an odd reasoning in favour of the darwinian principle that man is driven by selfishness.
But this reasoning about family egoism does not solve the altruism conundrum. People sometimes do something free of charge for someone who is no family. Again, the neo-darwinists reply that this apparent altruism is in fact selfishness. Because, they say, individuals form interest groups in which they help each other. To do something for a group member is not altruism but an investment in self-interest. They call it ‘group selection’: one selects group members.
This addition still does not make the darwinist reasoning airtight: there is altruism towards someone who is neither a family nor a group member. Once again, darwinism has an answer. In addition to the terms kin selection and group selection, in 1966 darwinist George C. Williams introduces the term individual selection: an individual does something for another individual whenever he himself benefits from it, regardless of family or group interests. Once again, it is about seeming altruism, which is in reality selfishness, because it is reciprocated. Darwinist Robert Trivers calls it “reciprocal altruism” in 1971. Nowadays we would call it a ‘win-win-situation’. The motive is self-interest.
All in all, the neo-darwinists have fiercely defended Darwin’s central point: egoism is the engine of evolution. Every altruism is unmasked as selfishness. At the family level (kin selection), at the group level (group selection), at the individual level (individual selection), everywhere self-interest is the motive behind seemingly altruistic behaviour.
When that discussion has been won, or appears to have been won, the neo-darwinists place the keystone in their worldview. The relationship between darwinism and genetics has yet to be settled definitively. In the 1970s, the neo-darwinists see the tremendous development of the field of genetics. Neo-darwinist Richard Dawkins decided to make necessity a virtue. He elevates the gene to the ultimate unit in biology. Evolution is not about family (kin selection), not about groups (group selection), not about individuals (individual selection), but about genes: “gene selection”, that is the solution. The gene is the unit that catapults itself into the future. Dawkins degrades individuals to carriers of genes, vehicles upon which genes move through time, from a previous carrier to the next. The gene is the main actor in evolution. But Dawkins does not give the tribute for free. He only wants to put the gene on the throne after he has darwinized it. And how does one darwinize a gene? By saying it is selfish. That is what Dawkins does, in 1976 his book The Selfish Gene is published, the best selling biology book of the 20th century, with more than a million copies sold. Immediately on page 2 and 3 Dawkins speaks out, evolution is determined by “the gene’s law of universal ruthless selfishness”. And he clarifies: “I shall argue that a predominant quality to be expected in a successful gene is ruthless selfishness.” And he immediately adds the neo-darwinian point that altruism is in fact egoism: “This gene selfishness will usually give rise to selfishness in individual behaviour. However, as we shall see, there are special circumstances in which a gene can achieve its own selfish goals best by fostering a limited form of altruism”. So he simply reduces altruism to a strategy for achieving selfish profit. The normal meaning of the term altruism, namely unselfishness, Dawkins mocks as “universal love”, sweeping it from the table. Egoism is the driving force, even at the genetic level. That way, the gene seems to be darwinized, and with it the whole of biology seems to be darwinized.
Neo-darwinism is not a strong theory. One of its problems is that it gives different definitions to existing terms. For example, ‘fitness’. Spencer had introduced the term in 1864 in his expression survival-of-the fittest. In this, fit meant simply ‘adapted to the circumstances’. A century later, the neo-darwinists have a different definition. For them, fitness is the number of offspring. Similarly, ‘survival’ has a different meaning for the neo-darwinists than it had for Spencer. Survival no longer means that an individual can keep itself alive, but that it produces offspring. This redefinition of both survival and fitness mutilates the meaning of the expression survival of the fittest. That suddenly means that the individual with the most offspring gets the most offspring. A tautology. Critics have pointed this out. The criticism is correct, with the new definitions Spencer’s expression has become a meaningless circle. The neo-darwinists react with irritation. Peter Medawar argues aggressively: schoolchildren like to point out such a tautology, but when adults complain about it they have not really grown up 28). And Richard Dawkins calls the tautology objection a “whimsical little conceit”, originating from “amateur philosophers” whom he calls “almost pathetic”. He calls the criticism a word game, beneath the dignity of darwinists. And then he continues in self-confident mode: “Like God, natural selection is too big a theory to be proved or disproved by wordgames. God and natural selection are, after all, the only two workable theories we have of why we exist.”
These utterings of Medawar and Dawkins are odd indeed. Especially Dawkins’ remark that the tautology-objection is only a word game. After all, the word game is practiced by his own neo-darwinism, which invented new definitions for the words survival and fitness. Dawkins realizes that his defense is weak, in veiled terms he recognizes he is wrong: “fitness is a very difficult concept, … there might be something to be said for doing without it whenever we can “. Apparently he does not like the new definition either. And he concludes his chapter on fitness with the statement: “I have found this a painful chapter to write” 29).
That is not the only word game of the neo-darwinists. They also tinker with the meaning of the word altruism. In reality altruism simply is a free service, it means doing something for somebody without expecting something in return, it is one-way traffic. The neo-darwinists change that meaning. For them altruism is never one-way traffic; it always implies payment, a quid pro quo, a reciprocal altruism. Hamilton’s altruism formula is not about altruism. It calculates to which extent it is advantageous to do something for another, at which point the favouring of the other stops. In short, Hamilton’s formula does not calculate altruism but egoism. The neo-darwinists see altruism as a strategy for self-interest. Robert Trivers puts it well: “Models that attempt to explain altruistic behaviour in terms of natural selection are models designed to take the altruism out of altruism” 30). That is exactly what the neo-darwinists are doing: ‘take the altruism out of altruism’. They bend the meaning of the word until it means the opposite.
Also the term egoism is involved in the wordgame of the neo-darwinists, they stretch the meaning of the word further and further. Initially the ‘ego’ is limited to the individual, then it includes the offspring, then further family, then the group in which ‘ego’ lives, until finally every act towards every other individual is seen as egoism. Altruism becomes egoism, the neo-darwinists talk about “selfish altruists” 31). An absurd word combination.
Given all this juggling with definitions by the neo-darwinists, it is remarkable that the neo-darwinian captain Richard Dawkins tries to reproach his critics for “wordgames”.
Gregor Mendel and Richard Dawkins
Richard Dawkins’ book The Selfish Gene is controversial in several ways. What is immediately noticeable is the condescending way in which he speaks about Gregor Mendel. Dawkins states in his book that the gene is the main protagonist of evolution. So one would expect some appreciation for the discoverer of it. The opposite is the case. He mentions Mendel only once, and in a negative sense. In that one passage he calls Mendel’s work “a little too simple” and he states: “Mendel perhaps did not realize the significance of his findings, otherwise he might have written to Darwin” 32). That is a nasty thing to say indeed. Mendel did realize the importance of his results, but he was the only one. Mendel was hurt by the minimal response to his article. He once said: “My time will come”. That valuation was correct, but his time did not come until sixteen years after his death. Would it have helped if Mendel had sent his article to Darwin, as Dawkins believes? It is interesting to reflect on that thought. In a way, Mendel did send it to Darwin. His article was printed in 1866 in a magazine to which many European libraries and institutions had a subscription. The Royal Society in London had a subscription. The Linean Society in London as well. Darwin and his friends Lyell and Hooker are all three members of both societies. None of them saw the importance of Mendel’s article.
Mendel had forty offprints made of the article, for despatch to persons and institutions he considered relevant. Seven of them were later found. One in Graz (probably sent to Franz Unger who lived his last years there), one in Amsterdam (used by Hugo de Vries), and so on. Apart from those seven, perhaps some more will come to light. The remaining specimens have been thrown away or lost. There are no indications that Darwin received a copy, but because Richard Dawkins retroactively advises Mendel to write his discovery to Darwin, let us assume for a moment that Mendel sent him an offprint. The idea is not outlandish, Mendel knew Darwin’s work. Darwin’s Origin and Domestication books are in Brno’s monastery library with Mendel’s handwritten notes in the margin. What would have happened if the postman had delivered the offprint to Darwin’s house? Darwin would not have been able to read it. Darwin hardly had any systematic education, the German language is a closed book for him. The second obstacle is that Mendel’s article contains statistical operations that are over Darwin’s head. Darwin is notoriously weak in mathematics, he would not have understood the article. In addition to the language obstacle and the mathematics obstacle, there is the sender obstacle. What should he do with an unreadable piece, sent to him by an unknown monk from Moravia? He throws it away, along with all the other strange mail he receives as a celebrity. And perhaps all this is not a hypothesis but a reality. Perhaps Darwin is one of the 33 unknown receivers who threw away their offprint.
When finally the biologists discover Mendel’s work in 1900, it is precisely the Darwin clan in London that opposes it. Geneticist Bateson has difficulty getting a foothold in the hostile darwinian environment. In addition to Bateson, there is another leading geneticist in the early decades of the 20th century, namely Wilhelm Johannsen in Denmark, who introduces the terms ‘gene’, ‘genotype’ and ‘phenotype’. Johannsen’s innovative insights are now accepted, his biggest fans were his colleagues Bateson and Morgan. There was one source of resistance to Johannsen’s ideas, namely the Darwin clan in London and its periodical Biometrika. Morgan recalls the resistance of the biometricians, he writes in 1916: “Johannsen’s splendid work met with such bitter opposition from the English school – the biometricians – who amongst the post-Darwinian school are assumed to be the lineal descendants of Darwin” 33). Darwin’s intellectual heirs try to repress genetics.
Finally, there is Richard Dawkins himself. He is the undisputed grandmaster in pushing geneticists out of the picture. His book The Selfish Gene stylizes the gene as the centre of biology, but in it he only mentions Mendel as the amateur who saw it too simply, who did not understand his own discovery and who would have done better to write his discovery to Darwin. And the other three giants of genetics, Bateson, Johannsen, Morgan, do not get a mention even once. Dawkins hijacks the gene from the geneticists and presents it as a darwinian being, a selfish being.
Something else is odd about Dawkins’ position. The simplest way to point it out is by a short detour. Simultaneously with his book, the book Sociobiology by Edward O. Wilson appears. The two are like-minded, to a large extent they have the same theme. They emphasize the importance of genes, the carriers of those genes are of secondary importance. Wilson begins his book by saying: “The chicken is only an egg’s way of making another egg”. And then he states more generally: “the organism is only DNA’s way of making more DNA” 34). The organism is only a means in the hands of the DNA. This passage by Wilson also expresses Dawkins’ point of view: individuals are merely reproduction machines with which genes move through time. Dawkins sees the gene as the main actor in evolution. He believes that we must understand evolution from “a gene’s-eye view”. And he goes on: genes not only have a view, they also have a motivation: they are “selfish”. And they have goals, “selfish goals”. Dawkins presents genes as if they were persons, with views, motives and goals. The only thing lacking is that genes vote Conservative. Genes are not persons, they have no view, motive or goal. Genes are pieces of matter.
His book evokes strong criticism. Not the gene but the cell is the smallest unit of life, that is the consensus among biologists. Genes are cellular components, not independent living beings. Geneticists who in their laboratory transplant genes of one species into another, do not think they are moving the main players of evolution. The geneticists themselves are the major players at that given moment.
Dawkins has not been unmoved by the criticism. In 2006, thirty years after the first edition of The Selfish Gene, an anniversary edition is published. In it, Dawkins acknowledges some of his mistakes. In an additional foreword he says that the idea of ‘selfish gene’ was meant to be a metaphor. That is a moderating sound. For half a century, the neo-darwinists have defended the egoism position, and now Dawkins says it is just a metaphor. So no reality. Furthermore, the foreword in the 2006 edition contains Dawkins’ request to the reader to forget the egoism passages from the first edition. He acknowledges that phrases such as “we are born selfish” were misleading. “Please mentally delete that rogue sentence and others like it”. Finally, he acknowledges that the title The Selfish Gene was wrong, the book should have been called The Immortal Gene. This is particularly telling. The selfish title was an attempt to darwinize the gene. That attempt Dawkins gives up. The gene belongs to the geneticists again.
1) De Beer, p. 40 (a/o note 4), p. 69. Irvine, p. 71-72
2) Letter Darwin to Galton, 11 February 1877. Quoted in Schwartz Cowan p. 140.
3) Szybalski, passim
4) Unger, p. 6, p. 4
6) Cock & Forsdyke, p. 245, 280, 664
7) Boon, p. 84-94
8) Bateson 1928, p. 238-239
9) Bateson 1909, p. 96. Also: Bateson 1928, p. 227
10) Bateson 1928, p. 238
11) Bateson 1914, p. 293
12) Bateson 1928, p. 13
13) Cock & Forsdyke, p. 233
14) Morgan 1916, p. 87-88
15) Morgan 1929, p. 46
16) Morgan 1932, p. 131
17) Morgan 1929, p. 66, p. 77-78
18) Morgan 1932, p. VIII, p. 15
19) Julian Huxley, p. 26
20) Julian Huxley, p. 28
21) Nesse & Williams, p. 241-243
22) Nesse & Williams, p. 236
23) Darwin 1871, vol. I, p. 168
24) Medawar 1984, p. 11
25) Various definitions of fitness in: Dawkins, The extended phenotype, p. 179-194
26) Segerstråle, p. 471
27) Medawar 1984, p. 12
28) Medawar 1984, p. 101
29) Dawkins, The extended phenotype, p. 180, 181, 194
30) Trivers, p. 35
32) Dawkins, The Selfish Gene, New Edition 1989, p. 33-34
33) Morgan 1916, p. 156. Also: Cock & Forsdyke, p. 71
34) Wilson 1998, p. 3
35) Fisher, p. 132
36) Nägeli, p. 18
37) agreeing quoted in De Vries p. 825-826
LIMITED LIST OF LITERATURE
— Allen, Garland E., T.H. Morgan, mechanistic materialism and the gene concept., in: American Zoologist, 23, 1983, pp. 829-843
— Allen, Garland E., T.H. Morgan – the man and his science. Princeton University Press 1978
— Barash, David P.: Natural Selections: selfish altruists, honest liars and other realities of evolution, New York 2008
— Bateson, William, F.R.S., His Essays and Addresses, Cambridge University Press 1928
— Bateson, W., Heredity and Variation in Modern Lights, in: A.C. Seward (ed.), Darwin and Modern Science, Cambridge UP 1909, blz. 85-101
— Bateson, William, Presidential Address to the Zoological Section, British Association, Cambridge meeting 1904, gepubliceerd in: William Bateson F.R.S., His Essays and Addresses, Cambridge University Press 1928, blz. 233-259
— Bateson, William, Address of the President of the British Association for the Advancement of Science, delivered at Melbourne on August 14, 1914, in: Science, N.S. Vol. XL, No. 1026, Friday, August 28, 1914
— Bonner, John Tyler, The ideas of biology (1962), Dover edition 2002
— Boon, Louis, De list der wetenschap – variatie en selectie: vooruitgang zonder rationaliteit. Baarn 1983
— Bowler, Peter J., The eclipse of Darwinism – anti-darwinian evolution theories in the decades around 1900, Johns Hopkins University Press, second edition, Baltimore 1992
— Cock, Alan G. & Donald R. Forsdyke: Treasure your exceptions, New York 2008
— Darwin, Charles, On the Origin of Species, 1e druk, London 1859
— Darwin, Charles, On the Variation of Organic Beings in a state of Nature, in: Journal of the Proceedings of the Linnean Society of London. Zoology 3 (20 August 1858): 46-50
— Dawkins, Richard, The selfish gene, 1976, reprint 1978, New Edition 1989, New Edition 2006
— Dawkins, Richard, The extended phenotype (1982), revised edition Oxford 1999
— De Beer, Sir Gavin, Darwin’s notebooks on transmutation of species, edited with an Introduction and Notes by Sir Gavin de Beer, Part I: First notebook (July 1837 – February 1838). In: Bulletin of the British Museum (Natural History), Historical series, Vol. 2, No. 2 (January), London 1960, blz. 23-73
— De Vries, Hugo, Species and varieties – their origin by mutation, Chicago/London 1905
— Fisher, R.A., Has Mendel’s work been rediscovered? in: Annals of Science 1936, volume 1, blz. 115-137
— Gray, Asa, Review of Darwin’s Theory on the Origin of Species by means of Natural Selection, in: The American Journal of Science and Arts, Second Series, Vol. XXIX, no. 86, March 1860, blz. 153-184
— Gray, Asa, Natural Selection not inconsistent with Natural Theology, in: Atlantic Monthly for July, August, and October, 1860. Reprinted London 1861
— Hulle, Dirk van: Darwins kladjes, Nijmegen 2010
— Huxley, Julian, Evolution – The Modern Synthesis, Third Edition 1974,
— Irvine, William, Apes, Angels and Victorians – a joint biography of Darwin & Huxley, London 1956
— Maynard Smith, John, The theory of Evolution, London 1958
— Maynard Smith, John, The Theory of Evolution, third edition, Canto-edition, 2000
— Medawar, P.B., Aristotle to Zoos – a philosophical dictionary of biology, London 1984
— Medawar, Peter B., The Science of Life – Current Ideas of Biology, London 1977
— Mendel, Gregor, Versuche über Pflanzenhybriden, in: Verhandlungen des naturforschenden Vereines in Brünn, IV. Band. Abhandlungen 1865, Brünn, im Verlage des Vereines 1866, S. 3-47.
— Morgan, Thomas Hunt, The scientific basis of evolution, New York 1932
— Morgan, Thomas Hunt, A Critique of the Theory of Evolution, Princeton University Press 1916
— Morgan, Thomas H., What is Darwinism?, New York 1929
— Morgan, Thomas H., Human Inheritance, Ninth Mellon Lecture, delivered May 7, 1924, at Pittsburgh, Pennsylvania, under the auspices of the Society for Biological Research of the School of Medicine, University of Pittsburgh. In: The American Naturalist, vol. 58, Sept.-Oct. 1924, blz. 385-409
— Morgan, Thomas H., Genetics and the physiology of development, in: American Naturalist 60, 1926, blz. 489-515
— Nägeli, Carl, Mechanisch-physiologische Theorie der Abstammungslehre, München/Leipzig 1884
— Nesse, R.M. & G.C. Williams, Evolution and Healing – The New Science of Darwinian Medicine, London 1995
— Pearson, Karl, Darwinism, Medical Progress, and Eugenics. The Cavendish Lecture 1912. An Address to the Medical Profession, London, Dulau & co. Ltd.
— Provine, William B., The Origins of Theoretical Population Genetics, University of Chicago Press 1971 / 2001
— Rushton, Alan R. : Nettleship, Pearson and Bateson: The biometric-Mendelian debate in a medical context. In : Journal of the history of medicine and allied sciences, vol. 55 nr. 2, April 2000
— Schwartz Cowan, Ruth : Sir Francis Galton and the study of heredity in the nineteenth century, dissertation Johns Hopkins University, Baltimore 1969; reprint in the series: The history of hereditarian thought – a thirty-two volume reprint series, presenting some of the classic books in this intellectual tradition, edited by Charles Rosenberg, The University of Pennsylvania; New York 1985
— Segerstråle, Ullica: Defenders of the truth, Oxford University Press 2000
— Strathern, Paul, Mendeleyev’s dream, New York 2000
— Szybalski, W., Professor Alexander Zawadzki of Lvov university – Gregor Mendel’s mentor and inspirer, in : Biopolymers and Cell, 2010. Vol. 26. n 2, blz. 83-86
— Trivers, Robert, The Evolution of reciprocal altruism, in: The Quarterly Review of Biology, vol. 1, March 1971
— Unger, Franz, Botanische Briefe, Wien 1852
— Wilson, Edward O., Sociobiology – The abridged edition, seventh printing, 1998