Here is the Darwin piece I promised to kick the blog off properly. It started off as a piece for Physiology News, but has been adapted and extended. It isn’t quite finished to my satisfaction, and I am a little apprehensive that Henry (Gee) will tell me loads of it is wrong… …but I think I had better post it or it will not make it in 2009 at all.
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The year now drawing to an end marks the bicentenary of the birth of Charles Darwin (12 February 1809 – 19 April 1882), as well as the 150th anniversary of the publication of his most celebrated work, On The Origin of Species, which went on sale to booksellers on 22 November 1859.
Given the central position of evolution and natural selection in our understanding of biology, Darwin obviously has a strong claim to be the greatest of all biologists, English or otherwise. Both Science and Nature published a range of features celebrated the anniversary of Darwin’s birth early this year, as did most newspapers.
Much or all of the above will be known to most, or even all, NN readers. Some may also know that Darwin was a friend of the founders of the Physiological Society. I doubt many will know that he was elected as one of the first two Honorary Members of the Physiological Society at the Society’s third ever, and first formal (with a rule book) meeting. But Darwin’s connections with the Physiological Society, and its early members, also encompassed many detailed scientific exchanges.
Following his early voyages, Darwin spent most of his later career in his home at Down House in Kent, maintaining a voluminous scientific correspondence and many scientific collaborations. The Darwin correspondence project tells us:
“Darwin exchanged letters with around 2000 individuals over his lifetime…He rarely attended meetings of scientific societies, typically spent only two weeks a year in London, and only occasionally received fellow scientists into his home. And yet he was arguably one of the best informed scientists of the day…”
Darwin and the physiologists
Darwin’s long-term correspondents included John Burdon Sanderson, Professor of Physiology at UCL and latterly in Oxford, with whom Darwin discussed the possible use of electrical signals in plants. A younger physiologist who corresponded with and visited Darwin frequently was George Romanes (1848-1894), best known for his work on the locomotory system (muscles and their control) of Medusae (jellyfish).
Darwin’s interactions with Sanderson, Romanes, Michael Foster (founder of the Cambridge Physiological Laboratory and thus of the most celebrated “school of physiology”), and others amongst the founding members of the Physiological Society, are well described in a fascinating 1970 article by RD French, “Darwin and the physiologists, or the medusa and modern cardiology”. The article discusses the extent to which Darwin’s ideas on evolution and natural selection were absorbed by, and influenced, the physiologists of the second half of the 19th century.
We should remember that the term “physiology”, as it was used in the 1870s, would have included what we would now think of as biochemistry and pharmacology, and indeed cell biology. The early issues of the Journal of Physiology in the 1880s and 1890s are notable for a much wider range of subjects, and experimental organisms, than has been typical in the last 20-30 years.
Of the physiologists who corresponded with Darwin, Romanes’ work on jellyfish had a particularly clear evolutionary underpinning, as the analogies of the jellyfish “muscular bell” with rhythmic muscular systems in higher animals were obvious. As an example of these analogies, French’s article explores the influence of Romanes’ ideas upon the theories of cardiac conduction and rhythmicity then being developed by Walter (WH) Gaskell and others.
The Darwin influence is clear in the opening words of a celebrated August 1883 paper by Gaskell on tortoise heart. This paper was one of a series that demonstrated the “intrinsic rhythmicity” of the heart in higher animals (i.e. that rhythmicity was a property of the heart itself and not dependent on control from the brain and nervous system), and also the fact that the heart had not one but several sequential pacemakers, with the beat rate of the “highest” (fastest) pacemaker entraining (governing) the others. Writing little more than a year after Darwin’s death, Gaskell states:
“The views held by physiologists upon many points connected with the innervation of the heart have been too exclusively based upon observations upon a single type of heart, viz. that of the frog. It is therefore very advisable wherever possible to control these experiments by a corresponding elaborate series of observations upon the hearts of a large number of other animal types, and in this way to trace the evolution of function in the same way as the morphologist tracks that of structure.” (emphasis added)
Darwin and the Physiological Society
Perhaps Darwin’s greatest gift to physiologists, and to the fledgling Physiological Society, was his unequivocal support for animal experiments as a way of making advances in physiology and medicine. The founding of the Physiological Society was a direct result of the late Victorian vivisection controversy, and the 1875 Royal Commission on vivisection. Those interested in this history can find a summary of it on the Physiological Society’s website. The Royal Commission’s report became the basis for the Cruelty to Animals Act of 1876, which established the system of licences for experiments on living animals. Darwin gave evidence in person to the Royal Commission, stating his belief in the need for animal work:
“I am fully convinced that physiology can progress only by the aid of experiments on living animals. I cannot think of any one step which has been made in physiology without that aid. No doubt many surmises with regard to the circulation of the blood could be formed from the position of the valves in the veins, and so forth, but certainty such as is required for the progress of any science can be arrived at in the case of physiology only by means of experiments on living animals.”
Darwin went on to clarify that he was referring to experiments in which animals were properly anaesthetised, and to express his surprise at objections to such procedures:
“It is absolutely unintelligible to me on what ground the objection [to these experiments] is made in this country.”
The physiologists were very appreciative of Darwin’s support. The first informal meeting to discuss the founding of the Physiological Society, which took place at Burdon Sanderson’s London home on the 31st March, 1876, was attended both by Darwin’s most prominent scientific disciple, Thomas Henry Huxley, and also by Darwin’s third son Francis Darwin (1848-1925), then 28 yrs old. Francis Darwin, who assisted his father in much of his research and ultimately became a distinguished botanist and FRS, had been working for his MD Thesis in one of the London physiological laboratories. The meeting reconvened a few weeks later on April 26, with Huxley chairing, to ratify a rule book, including the rule that:
“Men of distinction in Science who have contributed to the advancement of Physiology are eligible for election as Honorary Members.”
The third preliminary meeting, held at Romanes’ house, identified those “who shall be invited to become members and attend the next meeting of the Society.” This group included both Thomas Huxley and Francis Darwin. The meeting also resolved to elect Charles Darwin as one of the first two Honorary Members. [The other was the early physiologist William Sharpey (1802-1880)]
Darwin later wrote to Romanes (on 29 May):
“I was very much gratified by the wholly unexpected honour of being elected one of the Honorary Members [of the Society].This mark of sympathy has pleased me to a very high degree”.
Darwin remained unswerving in his support for animal experimentation until the end of his life. In a celebrated letter published in the Times in 1881, a year before his death, he wrote:
“I know that physiology cannot possibly progress except by means of experiments on living animals, and I feel the deepest conviction that he who retards the progress of physiology commits a crime against mankind.”
This letter can be read at the Times online archive, along with accompanying material and a commentary by former Physiological Society President and Head of the MRC Prof Colin Blakemore.
Darwin’s death in 1882 was a cause of great sorrow among the scientific men of the day, including at the Physiological Society. Edward Sharpey-Schafer in his extended history of the early Physiological Society records the feeling of the members present at the following meeting:
“With respect to [Charles Darwin’s death] the draft of a resolution occurs in the Minute Book, written in pencil and nearly illegible, which probably expresses an attempt then and there to voice the feelings of the Society… The draft reads as follows: ” The effect produced in the scientific world by the death of Charles Darwin is so universal and profound that any formal motion in reference to it by this little Society would appear to be superfluous, if not impertinent.”
The same meeting in 1882 elected several new Honorary Members including, perhaps a little symbolically, Thomas Huxley. Huxley and Francis Darwin continued to attend Physiological Society Meetings thereafter, and physiology in the late 19th and early 20th century was also indebted to Darwin’s fifth son Horace (1851-1928), founder of the Cambridge Scientific Instrument Company, which manufactured apparatus for many of the Cambridge laboratories. Apart from these direct ties, a reading of Sharpey-Schafer’s history makes clear that the approbation and involvement of men of the standing of Charles Darwin and of Thomas Huxley was of great importance to the fledgling Physiological Society.
From the 1870s to the present day
Perhaps an appropriate place to close this brief summary of Darwin’s ties to physiologists and the Physiological Society is an editorial in Science earlier this year by past President of the Royal Society Robert (Baron) May and Paul Harvey. They noted:
”… Dobzhansky’s memorable injunction that nothing in biology makes sense except in the light of evolution.“
One can, I think, see their dictum exemplified in diverse ways in papers in the physiological journals down the years. In the words and ideas of Victorian physiologists like Gaskell; in the use of invertebrate preparations like barnacle muscle and the squid giant axon for the seminal studies of Hodgkin, Huxley and Katz on action potentials; in the comparative sequence analyses presented in modern papers studying the structure-function relationships of ion channels; and in countless other places.
Indeed, with the widespread availability of genome sequences, I think that there is likely to be an increase in cross-species perspectives on physiological function – at least at the level of individual proteins – over the next decade or two. So two hundred years on from his birth, Darwin’s influence on physiology is not just still present. I would say it was undergoing a revival.
Some years ago the evolutionary biologists in my own University caused a furore when they argued, in a curriculum review, that all our degrees in biological and biomedical sciences should be explicitly themed round evolution and natural selection. The furore was because they also argued that overall teaching time devoted to different phyla should be in direct proportion to the “amount” of the natural world that different phyla represented.
The idea never caught on, perhaps understandably in an era when it is biomedical sciences (and thus implicitly the study of humans and higher mammals) that largely drive undergraduate student recruitment into life science degrees – something that a lot of people, me included, pointed out to them
But I think I can see what they were getting at.
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REFERENCES
French RD (1970). “Darwin and the physiologists, or the medusa and modern cardiology” J of the History of Biology 3:253-274.
Gaskell, W. H. (1883). “On the Innervation of the Heart, with especial reference to the Heart of the Tortoise” J Physiol 4: 43-230
May RM & Harvey PH “Species Uncertainties” Science (2009) 323: 687.
Sharpey-Schafer E. (1927). “History of the Physiological Society during its First Fifty Years, 1876-1926. Part.1.” J Physiol. 1927; 64 (Suppl): 1–76.
PS A note on references and hotlinks:
I just spent about 2 hrs putting a lot more of these in, but forgot to save… and a browser crash blew the whole lot away. I thought I had saved it, actually, but I am learning my way around the NN blogging platform and it turns out I hadn’t. But I am staying calm. When a long-ago computer crash ate my PhD Thesis reference list that I’d spent two days typing in, I kicked a filing cabinet and put my back out for three months. So I’m staying calm. Anyway, the piece therefore appears here without a lot of the links, which I will add as I find the time. Sorry.
Fascinating piece, Austin.
I’m a bit puzzled that you say Huxley was only ‘symbolically’ elected as an honorary member of the Phys Soc? Was he not a fine anatomist who contributed much to the study of invertebrate physiology?
I meant “symbolically” only in the sense that I wondered if it was significant that Huxley was elected to be an Honorary Member just after Darwin’s death – a handing on of the torch of Darwinian association with the Society’s Highest Honour, as it were. In fact, at the time of the meeting that elected Huxley to Honorary Membership there were no Honorary Members living, as the first and until then only two (Sharpey and Darwin) were both dead.
The initial group of 30-odd Physiological Society “Ordinary” Members were a distinguished group, with many FRSes. Thos Huxley was initially one of these Ordinary Members. “Honorary Members” were a bit different. They were intended from the start to be very few (the initial rules limited their number to five, and when Sharpey-Schafer wrote his history fifty years later in the late 1920s it had only risen to twelve), and to be those of great distinction in both science, and in work for the Society or for the wider good. In some ways I think you are talking something like the same esteem that we might today accord Nobel or Lasker Prizewinners. But Huxley certainly fitted the bill.
Again on the “encompassing all medical biology” theme, many anatomists appeared among the early Phys Soc members. Another interesting one is Huxley’s protege E Ray Lankester. Lankester knew Karl Marx and attended his funeral – I often wonder if he went to Darwin’s too.
Thanks for that fulsome clarification – so erudite, even on Christmas Eve! Happy Hols!
Thanks Stephen. I like “erudite”. It sounds so much better than the description favoured by my bosses in general (“lacks any focus”) or by one particularly forthright former Faculty Dean / ex-Boss in particular (“Why does he spend his time messing about with all this rubbish?”).
And Seasonal Best Wishes to -both- all our readers.
Nice work, Austin. Sorry I took so long to catch up. I think I can see what they were getting at too. (And your PS made me laugh.)