Review Endeavour Vol.29 No.4 December 2005 ‘Protoplasm.is soft wax in our hands’: Paul Kammerer and the art of biological transformation Sander Gliboff Department of History and Philosophy of Science, Indiana University, 130 Goodbody Hall, Bloomington, IN 47405, USA Paul Kammerer’s career ended in scandal in 1926 over because when it mates normally – on land – the male tampering with his evidence for ‘Lamarckian’ evolution – helpfully takes the eggs from the female and attaches the infamous midwife toad. But although Kammerer’s them to his own hind legs for safe brooding (Figure 2). In conclusions proved false, his evidence was probably one experiment, high temperatures induced midwife toad genuine. In any case his arguments were not simply for specimens to spend more time in water and even mate Lamarckism and against Darwinism, as the theories are there, but this meant that the egg-strands became understood today. If we look beyond the scandal, the waterlogged and would not stick to the males. The few Kammerer story shows us a great deal about early 20th- eggs that survived the treatment grew into toads that century biology: the range of new ideas about heredity preferred to mate in water, regardless of the temperature and variation, competing theories of biological and of the environment, apparently inheriting the acquired cultural evolution and their applications in eugenics, behavior. During the mating season males of this water- new kinds of laboratories and professional roles for breeding line even developed the ‘nuptial pads’ found in biologists, and changing standards for documenting frogs: dark, rough patches on the front legs, which were experimental results. used to clasp a slippery, wet female. As Arthur Koestler recounts in The Case of the Midwife Introduction Toad [3], Kammerer’s career ended in scandal in 1926 The 1910s and 1920s were heady times at the ‘Vivarium’ – when it was discovered that his last intact specimen of a or Institute for Experimental Biology – in Vienna. Paul midwife toad with a nuptial pad had been artificially Kammerer (Figure 1) was there reinvigorating the case for darkened with an injection of India ink. The fact that the inheritance of acquired characteristics, and Eugen Kammerer committed suicide six weeks later thus looked like an admission of guilt, and his work was discredited. Steinach was conducting his pioneering work on hor- His reputation has never fully recovered, despite Koes- mones and their effects on body and mind [1].A tler’s argument that Kammerer was framed by Darwi- charismatic speaker and popularizer, Kammerer wove nians who were desperate to save their theory from both lines of research into a vision for controlling human his attacks. evolution. He explained that dramatic cures of glandular Those who know Kammerer from Koestler’s account deficiencies had already been achieved: might be surprised to learn that, far from opposing A frivolous, mal-gendered or perverse, obese, spine- Darwinism, Kammerer counted himself a Darwinian, less shirker becomes, with the administration of but in the grand 19th-century style of Ernst Haeckel. testicular substance, a serious, determined, hairy, Haeckel built his Darwinism into a comprehensive and muscular, hard-working and lusty man! A dwarf- progressive world-view, an alternative to religion and a like, stunted idiot with wrinkled, scaly skin, brittle force for social change. Darwin himself had assumed that nails, open sores, and a stupid, glazed look in the eye environmentally induced modifications and the effects of is changed by the ingestion of thyroid substance, into using or disusing organs were important sources of a tall, slim, smooth- and light-skinned, sharp-eyed favorable and heritable variation, and Haeckel’s influen- and alert person [2]. tial German interpretation of Darwin’s work focused on these as the principal causes of Darwinian evolution and Surely, Kammerer argued, it was only a small step from cultural progress. treating pathological cases like these to enhancing the Kammerer aimed to bring Haeckel’s brand of Darwin- population at large, producing exceptional individuals and ism up to date with 20th-century biology and culture. He making the changes in them hereditary. provided experimental evidence for the modifying power Kammerer’s evidence that such changes were heritable of the environment, and he outlined a mechanism of came from experiments on a variety of animals. The most heredity that would let acquired characteristics be widely discussed of these was the midwife toad, so named communicated to genes. Kammerer argued that the Corresponding author: Gliboff, S. ([email protected]). hereditary effects of education, practice and artistic Available online 4 November 2005 achievement assured continual progress in human www.sciencedirect.com 0160-9327/$ - see front matter Q 2005 Elsevier Ltd. All rights reserved. doi:10.1016/j.endeavour.2005.10.001 Review Endeavour Vol.29 No.4 December 2005 163 which had to be hauled up by train from Trieste. An electric motor circulated the seawater through the aquaria upstairs and back to the cellar. Hothouses and apparatus for manipulating air pressure, photoperiod and the light spectrum, and for simulating gravitational forces were also available, as well as a central aeration system for the fish tanks, and electrical outlets and lights at all the work-tables. Wilhelm Roux, a leader of the experimental turn in embryology and evolution, arranged in advance to publish all of the research reports that came out of the Vivarium in his journal Archiv fu¨ r Entwicklungsmechanik (Archive for Developmental Mechanics). Meanwhile, other scientists sent specimens of abnormal animals to the Vivarium, which Przibram later exhibited to the public in a ‘Museum of Entwicklungsmechanik’ worthy of the Prater location. Przibram also hired Paul Kammerer, then still a student, to be his first assistant in 1902 or 1903 [5,6]. Kammerer and his experiments Kammerer was born in Vienna in 1880. His mother was from a converted Jewish family and his father was a factory owner [7]. In his youth he developed the intense devotion to the arts that was typical of his generation and class in Vienna [8]. Kammerer studied music at the Conservatoire, comparative morphology at the University and learned the latest experimental methods from Przibram at the Vivarium. The results of Kammerer’s first experiments were published in 1904. They were performed on a lowland salamander that bears lots of little tadpoles, and an alpine Figure 1. Portrait of Paul Kammerer. Image reproduced from the frontispiece to [3] species that bears fewer, but larger, fully metamorphosed courtesy of the Herman B. Wells Library, Indiana University, Bloomington, IN, USA, and with permission from Hutchinson. land dwellers as offspring. By manipulating temperature and humidity, Kammerer was able to make each species of culture. This became the basis for his alternative salamander acquire the reproductive habits of the other. eugenics, which proposed to create fit individuals rather However, to test if this switch was heritable required a than select them. three and a half year wait for the offspring to mature. In Kammerer’s position at the privately owned Vivarium the meantime, Kammerer launched a staggered series of laboratory – outside of academic biology – gave him the similar experiments using different species, trying out freedom to pursue his dual experimental and cultural different Vivarium gadgetry and aiming for ever more projects. He published in both technical journals and the striking modifications. popular press, and he addressed the public directly in what he called his ‘big show-lectures’. As we shall see, Kammerer’s work at the laboratory and show-lectures also made special demands on him and shaped his research in unexpected ways, which sometimes hurt his scientific credibility [4]. The Vivarium The Vivarium laboratory was founded in 1902 on the premises of a former zoological exhibit hall in the Prater, the big amusement park on the outskirts of Vienna. Hans Przibram, an aspiring young experimentalist and a man of means, bought the building and retooled it for researching organisms and environments of every description. Modern Figure 2. A normal male midwife toad – Alytes obstetricans – with egg mass on its central heating, supplemented by stoves, provided a range hind legs, as drawn by Paul Kammerer’s boss Hans Przibram. Przibram was noted of temperatures for terraria, aquaria and insectaria. A for his artwork and his drawings were exhibited at the Secession, Vienna’s center for Jugendstil art, and printed in the Secession journal: Ver Sacrum. Image deep cellar provided cold, dark conditions for raising cave reproduced from [9] courtesy of the Herman B. Wells Library, Indiana University, animals, and storage space for rainwater and for seawater, Bloomington, IN, USA. www.sciencedirect.com 164 Review Endeavour Vol.29 No.4 December 2005 Positive results started spewing out of this pipeline in changes, he never quite pinned down the mechanism 1908: the alpine and lowland salamanders bred true and that caused them. In retrospect, what was most interest- continued to breed in a manner contrary to the norm in ing about Kammerer’s work was not that it supported their species. Then came the midwife toads with their Lamarckism particularly well, but that it challenged nuptial pads [9], spotted salamanders that became striped biologists to come up with alternative explanations. and much more. In backcrosses with unmodified stock, the Geneticswasstillinitsinfancyatthetimewhen traits often appeared to follow Mendelian laws thus Kammerer made his claims, and had yet to settle on the supporting Kammerer’s contention that his treatments modern conception of a random genetic mutation. There- induced genes to form [10]. In 1911, these successes fore, every critic who tried to explain away Kammerer’s earned Kammerer a promotion to the rank of Privatdozent results went about it differently. – lecturer – at the University [11], which opened doors for Several prominent authors, such as August Weismann him as a popular science writer and public speaker.