The Case of the Midwife Toad, Arthur Koestler, New York: Random House, 1971. ISBN: 394-48037-6
1. the intellectual and practical activity encompassing the systematic study of the structure and behavior of the physical and natural world through observation and experiment.
2. a systematically organized body of knowledge on a particular subject.
Science: method or body of received wisdom? Between those two uses of the word can lie a chasm of misunderstanding and controversy. The difference can even lead to violent death. Don't believe us? Read The Case of the Midwife Toad. Although the subject of this unusual scientific biography committed suicide in 1926, and the book itself was published in 1971, the problem under discussion is as fresh as this year's peer-reviewed scientific journals.
'An Epigenetic Perspective on the Midwife Toad Experiments of Paul Kammerer (1880-1926)', J Exp Zool B Mol Dev Evol. 2017 Jan;328(1-2):179-192. doi: 10.1002/jez.b.22708. Epub 2016 Oct 26. Abstract in PubMed.org, US National Library of Medicine National Institutes of Health
What happened to Paul Kammerer? What is a midwife toad, anyway? Are inky nuptial pads the smoking gun? Did a British scientist drive him to suicide? Why did the Soviets make a sentimental movie about him? We will attempt to answer these questions, and belatedly flog Mr Koestler's book.
Paul Kammerer, (Mad?) Scientist
The Vivarium was a famous sight in old Vienna. Built in 1873 to display reptiles, by 1903 the institute had become a famous research facility under the direction of zoologist Hans Leo Przibram (1874-1944). Przibram's star pupil was one Paul Kammerer, composer, bon viveur, and reptile specialist. If Kammerer couldn't make a specimen thrive in captivity, it couldn't be done. That was his downfall, really: it was terribly difficult for other researchers to duplicate Kammerer's experiments with rare and finicky subjects, as we shall see.
The Biologische Versuchanstalt in Vienna was colloquially known as 'the Sorcerors' Institute'. Even the exuberant biologists of the early 20th Century sometimes balked at their more outré experiments:
Some of the experimenters may have been carried away by the euphoria often found when a new branch of science opens up: one Dr Finkler transplanted heads from male to female insects which showed signs of life for several days but, allegedly, disturbed sexual behaviour; and Professor Steinach's rejuvenation experiments by stimulating the internal secretion of the sex glands became all the rage in the popular press about the time of the discovery of Tutenkhamon's [sic] tomb.
- The Case of the Midwife Toad, pp 22-23
No wonder early European horror films were full of 'mad' scientists. Kammerer, though bold in his outlook, was not nearly this weird. It was just that his experimental results opened a can of worms. Depending on who you talked to, Kammerer had either set Darwin on his head, or helped evolutionary theory discover an important operational mechanism. The debate's still going on, and it started with the midwife toad, so we'd better go there next.
The Midwife Toad
The midwife toad, Alytes obstetricans, has unusual habits. The females lay the eggs, and the males fertilise them outside the body. So far, so normal for frogs and toads. Then the male toad does an unusual thing: he collects all the eggs, spreading them around his hindquarters and back legs, and carries them around until they hatch. That's why he's a 'midwife toad'.
While most toads, like frogs, mate in the water, Alytes obstetricans mates on land. For that reason, the males don't have something other toads have: nuptial pads. Nuptial pads are handy little swellings that serve as grippers when grasping a slippery female toad in water. No toad wedding night should be without them, unless of course you're a midwife toad – and Paul Kammerer hasn't been messing with you.
You guessed it: by manipulating the midwife toad's environment so that it reproduced in water, and carefully raising generation upon generation of midwife toad at the Vivarium, Paul Kammerer succeeded in producing midwife toads with nuptial pads. At least, that's what he said, and that's what other biologists who saw the samples said. But this startling conclusion set the cat among the pigeons in the biology world.
Was this evidence of Lamarckism?
Lamarckism: the hypothesis that an organism can pass on characteristics that it has acquired through use or disuse during its lifetime to its offspring.
- Wikipedia (12.7.18), which goes on to point out that a) Lamarck didn't originate the idea of the inheritance of acquired characteristics, b) this theory was but one aspect of Lamarck's theory of orthogenesis, by which species gained in complexity, and c) Darwin didn't entirely disagree with Lamarck, so that d) the whole argument is based on a false dichotomy1.
False dichotomy or not, some biologists were quick to pounce on evidence of Lamarckism and rush to shoot it down because that sort of thinking was dangerous. Arguments got heated – and convoluted:
August Weismann, more Darwinian than Darwin, had postulated that the substance which carried the hereditary dispositions – the 'germi-plasm', as he called it – remained unaffected by acquired characters. One of his famous experiments was to cut off the tails of mice, for twenty-two successive generations, to see whether a tailless mouse would appear. But, as a Lamarckian critic remarked, he might as well have studied the inheritance of a wooden leg. For Lamarck's thesis was that only such characteristics are inherited which an animal develops as a result of its natural, adaptive needs – and losing its tail by amputation could hardly be called an adaptive need of the mouse.
- The Case of the Midwife Toad, p 34
Many biologists approved of Kammerer's work. Kammerer himself insisted that the midwife toad didn't prove that any acquired characteristics were inherited, but rather that an older trait – the ability to grow nuptial pads – could be reactivated by environmental factors. This might help fill in some of the gaps in evolutionary theory. Kammerer had a similar success in his work with two species of Austrian salamander, Salamandra atra and Salamandra maculosa. By manipulating their environments, Kammerer got them to switch breeding methods, from laying eggs that turn into tadpoles, to live births of smaller versions of the adult form, and vice versa. As he pointed out, this wasn't quite the magic trick it seemed: the two species were very closely related.
Kammerer himself was more impressed with the Ascidian sea-squirt, or Ciona intestinalis. The evil scientist repeatedly cut off the sea-squirt's siphons (long protuberances that siphon sea water for food). The sea-squirt can regenerate, so it did. Each time, the siphons grew longer, as if to say, 'Take that, scientist!' Great was Kammerer's excitement when the sea-squirt produced offspring... with longer siphons. Sea-squirts weren't sexy, though. Kammerer became famous, or infamous, for salamanders and toads.
So What Happened?
Yorkshireman William Bateson used to be a Lamarckian, but he got better. As a consequence, he went around enforcing what he perceived to be the true doctrine of Neo-Darwinism. Professor Bateson was offended by Kammerer's experimental results. He said so, loudly and often, in scientific journals. Unfortunately, Kammerer replied late, or not at all. This wasn't because Kammerer had lost interest in his subject, or was being cagey. It was just that in post-Great War Austria, hyperinflation had set in. Kammerer couldn't afford the subscriptions. He had to rely on friends to forward him the articles. Kammerer eventually replied, acceding to Bateson's heated demands to examine the evidence. Kammerer thought the institute might still have the slides, although the animals themselves were dead: the war, you know...
When the specimens were examined in the 1920s, horror set in. The nuptial pads were found to be contaminated with India ink. Were they even nuptial pads at all? Hard to tell. Bateson was triumphant. He branded Kammerer a liar. Kammerer was distraught, and tried to defend himself. But he didn't have tenure...
Scientists in the infant Soviet Union were ideologically committed to Lamarckism. They approved of this theory of evolution because it held out the prospect that by hard work and self-improvement, the workers of the world could give birth to a better grade of human being. (You probably had to have been there.) They invited Kammerer to come and work in Moscow.
Kammerer seemed to be planning to do this. He packed up his research materials and furniture for shipment to Russia. Then, at the last minute, something changed. Nobody knows what. In the early afternoon of 23 September, 1926, a road worker found Kammerer's body on a mountain path in Austria. He was sitting against a rock face, and he'd shot himself in the head. In his pocket was a suicide note leaving his body to science. It was a sad end to a brilliant career.
Here are some of the things we probably will never know:
- Who doctored the nuptial pads on the midwife toad specimens?
- Were Kammerer's assistants involved, or was the India ink discovery part of a more sinister plot by Bateson or his supporters to discredit Kammerer?
- Why did Kammerer kill himself? Was he depressed over the midwife toad incident? Could he not bear to leave Vienna for Moscow?
- Was he, as some suggest, distraught by a break-up with his latest romantic interest? Kammerer's lively love life is discussed in detail in Koestler's book.
The question we may be able to answer, at long last, is, 'Can midwife toads really grow nuptial pads?' According to Koestler, the scandal surrounding Kammerer's suicide was so great that up until 1971, when he wrote the book, no one had attempted to reproduce the experiment. They still haven't. But current researchers have suggested that they should. Some believe that Kammerer had stumbled across a mechanism known to geneticists today, called epigenetics2. Kammerer may have suffered the fate of many a scientist who was 'ahead of his time'.
One other thing we know: Kammerer's story became the subject of a 1928 Soviet film called Salamandra. The reviewer cited by Koestler speculates that salamanders were simply more photogenic than toads. The movie is about what you'd expect from Soviet Realism, but if you're dying to see it, why yes, of course you may, on the ever-reliable Youtube.
Should I Read This Book?
Yes. Definitely. Buy a used copy. Or borrow it from the library. Koestler's book is compelling reading. He knew Vienna very well, and as a journalist and historian he was well acquainted with the mindset that gave rise to this story. Lacking biographical material, Koestler extensively interviewed Paul Kammerer's family and acquaintances. The result is a vivid portrayal of a time and place now vanished. Read it for the gossip, if you like: Kammerer had interesting affairs. He even knew Alma Mahler Gropius Werfel. You've never heard of Alma Mahler Gropius Werfel? You're in for a treat.