The Golgi War: 2. A possible casualty

A.J. ‘Jock’ Marshall (1911-1967), whose irreverence was remembered by his contemporaries with reverence, was J.R. Baker’s postgraduate student at Oxford. But they had worked together before that. In 1933, Baker recruited him to work on his expedition to Esperitu Santo in the New Hebrides (now Vanuatu) alongside and then to replace Tom Harrisson. After many adventures and jobs and countries, including New Guinea, London and Oxford, and getting a degree from Sydney—and serving in New Guinea in the Second World War (the title of his biography, One Armed Warrior, denoting his service in 'Jockforce' and the fact that he had shot his own arm off in an accident with a gun as a boy) in September 1946 he returned to Oxford aged 35 as a postgraduate student.

After his Oxford D.Phil (the Ph.D. abbreviation possibly unique to that institution), Marshall was appointed Reader and Head of Department at St. Bartholomew’s Hospital Medical School (Barts). He was around Baker when the latter was attacking the existence of the Golgi Apparatus and was aware of the cytological methods that Baker was promoting.

Marshall wrote a review for Science Progress entitled, The structure of the so-called Golgi body. It was published in 1952 and took the side of Baker in the controversy—with knobs on. Marshall began by describing the work of Parat, a Frenchman who threw doubt on the existence of the Golgi. He wrote:

There was a swift and vigorous—some have said violent—reaction to Parat’s conclusions (see, for example, Gatenby, 1931) and although others observed Parat’s bodies and failed to substantiate the net of Golgi, the new concept seems to have been more or less swamped by the positive views of the traditionalists.

After dealing with Hirsch’s non-support for the Golgi in Germany he turned to Baker:

A determined attempt to resolve the problem has now begun by Baker (1944) of Oxford…He failed to find any evidence that the classical net of Golgi existed as a living entity……The next and perhaps most crushing blow to the classical concept of Golgi, Cajal, Aoyama and Gatenby was dealt by Thomas (1947-51). A Beit Memorial Fellow who came to Baker’s laboratory from New Zealand, Thomas used his own modifications of Baker’s and others’ techniques, as well as phase-contrast microscopy, in the investigation of living and fixed nerve-cells of common animals. He too failed to find a Golgi net in the living cell and declared flatly that, in the cells studied, the net was “shown to be an artifact”.

In his conclusions, Marshall divided the world into ‘traditionalists’ and ‘modernists’ and it is not surprising that those he dubbed traditionalist were infuriated:

The modernists, though they are too polite even to breathe it, generally seem to feel that the traditional so-called Golgi techniques merely show something funny about a particular part of the animal cell. By an arbitrary method a great man, Golgi, accidentally showed a net; and his followers have messed about until they have found even more certain methods of showing nets. The modernists imply that this is good wholesome fun, but not science. They believe that the Golgi apparatus, if it should be called that at all, is a system of lipoidal spheroids which are demonstrably concerned with cell-secretion.

However, he then started to backtrack:

The present author does not take sides in the controversy…

But clearly the whole tone of the review was taking sides and the reader by now will expect Gatenby to have responded in some way. He was, after all, told that what he was doing, as a ‘traditionalist’, was ‘not science’.

Marshall’s biography (compiled by his widow from his own notes and records) reads:

…it [‘professional knifing’] involved views he had put forward in support of John Baker's work on the Golgi body. Owen Thomas who was also working on it had just come back from seeing Professor Gatenby in Dublin. [Marshall wrote:] Thomas was “as scared as hell that Gatenby will discredit both John and myself. He is going to sue John too - so he says.” This referred to an article Jock had written for Science Progress on this rather esoteric cytological subject - the Golgi body. His article supported the research John Baker and Owen Thomas were doing in Oxford which appeared to negate some of the findings of Professor Gatenby. Thomas reported Gatenby was furious - “He will see to it that I [Jock] never get a better job than I've got [now]. Actually I quite like him although he has added me to his list - a long list - of hates.”

However, Marshall could not count on Baker either in support of any advancement. At about the time the article in Science Progress appeared, he applied for, without expecting to get nor did get, the chair of zoology at Reading. Later and by accident, he saw the reference Baker had written, and was shocked to learn that although Baker had praised his research he brought up, quite remarkably to those of reading it 65 years later, Marshall’s divorce from his first wife and ‘an assessment of Jock's alleged shortcomings as a classical zoologist’.

Marshall’s friends and wife put it down to jealousy. Baker was not a professor, and at this time and in the Oxford grading system that differed from that in other universities, he was a university demonstrator—he was promoted to a readership in 1955, three years before his election to the Royal Society.

Application—and rejection—by Marshall for another chair (this time in Canberra to set up a new department) in 1958 brought the suspicion of Gatenby’s revenge:

…while camped near Kalgoorlie, he received a telegram telling him that the Committee had chosen Professor [J. Desmond] Smyth from Dublin University for the Canberra Chair. Smyth, a parasitologist, came from Professor Gatenby's Department. It transpired that Gatenby, the good hater, had been in Australia at this time. It was tempting to conjecture whether his threat of six years before to prevent Jock from ever getting 'a better job' than the one he then occupied had come home to roost; especially when reading part of a letter he sent to another zoologist in England (the first page is missing): “Marshall threw his weight about so much, they were determined not to have him.”

There is no doubt that Jock Marshall upset the easily upset Australian university establishment by his behaviour. I am not surprised. Of all the university administrative systems I have dealt with all over the world over the years as an external assessor for appointments to chairs and for internal promotions, Australia occupies bottom position. Rigid, po-faced, stuffed shirts are phrases that came to mind, quite the antithesis of what the world sees Australia as like and quite the antithesis of Jock Marshall.

After the quite disgraceful failure of the University of London (described in excruciating detail in his biography) to elevate his readership at Barts to a chair, he did return to Australia—to the then new Monash University in Melbourne in 1960.

As to his support for Baker’s ‘modernist’ views on the artifactual nature Golgi Apparatus, there is a twist. In 1954 and only shortly after the first electron micrographs of the Golgi were published, Dennis Lacy in Jock Marshall’s own department at Barts found the Golgi apparatus in the exocrine pancreas and other cells and followed that up by finding it for the first time by electron microscopy in nerve cells—where Golgi had first described his eponymous organelle.

Marshall Jane. 1998. Jock Marshall: One Armed Warrior, Australian Science Archives Project, Melbourne.

Marshall AJ. 1952. The structure of the so-called Golgi body. Science Progress 40, 71-77.

Lacy D. 1954. Recent studies on the Golgi apparatus of the exocrine and endocrine cells of the mammalian pancreas and the cytoplasmic inclusions of other cells. Journal of the Royal Microscopical Society 73, 226–240.

Lacy D. 1957. The Golgi apparatus in neurons and epithelial cells of the common limpet Patella vulgata. Journal of Biophysical and Biochemical Cytology 3, 779–796.

Lacy D, Challice CE. 1957. The structure of the Golgi apparatus in vertebrate cells examined by light and electron microscopy. Symposia of the Society for Experimental Biology 10, 62–89.

Lacy D, Rogers GE. 1955. Recent observations by light and electron microscopy on the cytoplasmic inclusions of the neurons of Patella vulgata. Journal of the Royal Microscopical Society 74, 172–175.

The Golgi War: 1. The controversy and the warring parties

“Best not to mention it’, was the advice we were given on starting A level zoology or biology by the estimable ‘Harry’ Hadwen in 1959. ‘It’ was the Golgi Apparatus. It may all be an artifact—or it may not—was the tone of his advice, and coming down on one side or the other should be avoided. He had obviously been a student when the Golgi War was raging in the early 1950s and when the war was more politely termed a controversy.

The two main warring parties were James Brontë Gatenby (1892-1960), Professor of Zoology at Trinity College, Dublin, and John Randal Baker (1900-1984) firmly fixed in Zoology at Oxford. Both were Oxford zoology graduates, a department, incidentally, renowned for internecine warfare.

J.R Baker FRS in 1958

The argument was not an unusual one for the time. Essentially there has always been argument whether the structures seen under the microscope exist in the living cell or whether they are artifacts caused by the fixation and staining needed to see any of the structure. In addition, with intracellular structures there was the added limitation of the light microscope working at the very edge of its resolution. It wasn’t until electron microscopy came along that some of the issues were settled but there again the possible effects of fixation took some time to determine.

Gatenby argued that the Golgi Apparatus, first described by Camillo Golgi in 1898, was real. Baker, on the other hand, argued that it was not but just an artifact. The problem was, that, as we now know, the Golgi is more highly developed in some types of cell than in others, and a great deal of heat was generated by looking at nervous tissues—the last place, with hindsight, to look—as Golgi himself had done. Indeed, what was later realised to be the Golgi apparatus had been described earlier in other tissues. Both Gatenby and Baker thought their own methods superior to that of the other and there was a war of words. I have seen reference to threatened or actual lawsuits as a result but have not found any details.

In their obituary of Baker for the Royal Society, Willmer and Brunet wrote:

In this work on the Golgi apparatus he sometimes became unnecessarily intolerant and aggressive, but there were faults on both sides. Indeed it is a curious fact that none of the contestants in their search for the ‘true’ structure of the Golgi body or apparatus seems to have considered the different functions being performed by the different classes of cells with which they were concerned.

Gatenby’s obituary in The Times (22 July 1960) touched on his part in the war:

…his main interest lay in the field of cytology, and especially in the elucidation and function of the so-called Golgi apparatus. For a while his interpretation was unquestioned; but in later years rival schools appeared, based possibly on less technical skill and experience but with a more sophisticated chemical background than Gatenby’s—for he was never happy with the physiological approach of what he regarded as the wild young men of Cambridge. The anathemas launched from Dublin against these heretics, first in Paris and later, by the unkindest cut, at Oxford, never lacked in vigour…In Dublin his relations with his colleagues were at times strained, for his controversies were apt to extend beyond the technical field, but he was a popular figure in the professional world. Beneath all the thunder lay a fund of simple, jovial bonhomie…

Recent historical accounts of the Golgi apparatus make the point that it is impossible to interpret the findings of the different parties, in terms of who was right or wrong with particular staining procedures in particular tissues. Baker tried to make histology and cytology more scientific in terms of trying to determine what the various fixatives and dyes actually reacted with in the cell—cytochemistry—and admirable objective but one that was soon overtaken by the tools of modern cell biology. He is now probably better remembered for his other contributions to zoology than for the cytology that occupied the later decades of his active scientific life.

After a false start with electron microscopy, in research published in 1954 it was realised, with Baker still holding out, that the Golgi apparatus is real and is of particular importance, as neutral observers had long surmised, in secretory cells. As such, it loomed large in my life less than ten years after being advised to avoid even mentioning it.

Peter Wooding's electron micrograph of mammary cells
from a lactating cow showing the large Golgi Apparatus
(G) between the nucleus and the apical (luminal) membrane.

The Golgi War did have one useful outcome. It served as an illustration for Richard Dawkins of the way science works (as opposed to religion) and how scientists should always work, as he explained in The God Delusion: 

I have previously told the story of a respected elder statesman of the Zoology Department at Oxford when I was an undergraduate. For years he had passionately believed, and taught, that the Golgi Apparatus (a microscopic feature of the interior of cells) was not real: an artifact, an illusion. Every Monday afternoon it was the custom for the whole department to listen to a research talk by a visiting lecturer. One Monday, the visitor was an American cell biologist who presented completely convincing evidence that the Golgi Apparatus was real. At the end of the lecture, the old man strode to the front of the hall, shook the American by the hand and said–with passion–“My dear fellow, I wish to thank you. I have been wrong these fifteen years.” We clapped our hands red. No fundamentalist would ever say that. In practice, not all scientists would. But all scientists pay lip service to it as an ideal–unlike, say, politicians who would probably condemn it as flip-flopping. The memory of the incident I have described still brings a lump to my throat. 

The ‘respected elder statesman’ was, of course, Baker.

Dawkins was an undergraduate at Oxford between 1959 and 1962, and I wonder whether this seminar was held before or after the death of Gatenby in 1960?

While the Golgi War ended, there was a real fear that while it continued coming down on one side or the other could really be career-threatening, as I shall show in a second post on this topic.


Dröscher A. 1998. The history of the Golgi apparatus in neurones from its discovery in 1898 to electron microscopy. Brain Research Bulletin 47, 199-203.

Willmer EN, Brunet PCJ. 1985. John Randal Baker. 23 October 1900-8 June 1984. Biographical Memoirs of Fellows of the Royal Society 31, 33-63.

Wooding FBP. Comparative mammary fine structure. Symposia of the Zoological Society of London 41, 1-41.

Leeches in Sri Lanka: A Coincidence Shines the Light

Last November, having got drenched from a sudden tropical shower, I was sheltering with others in the party at the entrance to Sinharaja Forest Reserve in Sri Lanka (Ceylon, in old money and to stamp collectors) while eating my packed lunch. Leeches were all around us. I was watching the leeches on the edges of the path through binoculars just after the rain had stopped. I noticed that as the sun started to break through, many of the leeches in deep shade started to loop towards to the bright areas. That the leeches (a fair way from the attraction of human bodies) might be phototropic crossed my mind. A just-so story flitted across the consciousness: animals like wild boar and deer move along defined paths which are more exposed to the sun than the undergrowth, hence it would make sense to hang around light places in the hope of passing trade. While I am singularly attractive to lurking leeches (with ample evidence from Borneo and, now, Sri Lanka) and particularly inept at spotting their attachment quickly enough, my attention was soon taken by having to remove two of them that were taking an armful and a legful, respectively, and the question of leeches and light was put to the back of my mind.

Last week, I was looking up something else entirely. I was reading through J.R. Baker’s Biographical Memoir¹ and suddenly came across this:

It is certainly true that anyone who visits the wilder parts of Ceylon [Sri Lanka] is liable to be set upon by leeches, but not many people would think of sitting down at a table to play a game with them. Baker noticed that the leech is not only chemically attracted to the human body but it is also strongly phototropic. Thus a leech placed upon a table illuminated by a table lamp can be controlled in the direction of its movements by periodically changing the position of one’s seat or of the table lamp.

Baker spent the Oxford summer vacation of 1936 in Ceylon. His great-uncle was Sir Samuel Baker (1821-1893) the famous explorer of Africa, and there remained a family financial interest in a tea plantation at Nuwara Eliya. Baker, the great-uncle, had chartered a ship and taken people, animals, equipment and seed in order to found an agricultural settlement there in 1848; he stayed until 1855, leaving his brother in charge of further development, first of the farm and, later, as tea growing became established, of a tea plantation. We have been to Nuwara Eliya twice: a drive-through in 1968 and to stay in 2013. Apart from resembling a Scottish town in terms of the architectural style of the houses, the fields around Nuwara Eliya are full of typically British crops like leeks (field upon field of them) and beetroot (delicious curried). That’s all down to Baker apparently.

I was even more surprised to find in the references at the end of the Biographical Memoir, that Baker had been based at Sinharaja from the end of July to the beginning of September; his paper, describing what he did and what he saw is entitled, The Sinharaja Rain-Forest, Ceylon². He notes: 

The abundance of land-leeches (Haemadipsa zeylanica, Moquin-Tandon) greatly detracts from the pleasure of studying the Sinharaja Forest. I found it best to wear thin breeches cut like "plus-fours," stockings, boots with the tongues sewn in right up to the top, and puttees. This clothing by no means prevents leeches from penetrating to the skin of one's legs, but it greatly reduces the annoyance.

Baker included a map showing where he had camped (about 5 miles from the lodge where we stayed last year) and where he had walked. His walks had taken him along the same rough track we had walked and where I had sheltered and watched the leeches.

Here is Baker’s map of Sinharaja:

And here is a very short video of the leech in question and its acquisition of my blood:

John Randal Baker (1900-1984) was, and still is, well known for a number of different activities: the breeding season of birds in the tropics and the factors that trigger the onset of reproduction; the introduction of the terms proximate and ultimate causation; his cytological studies and textbook on the subject; his work on chemical contraception; his wars, especially with J. Brontë Gatenby, on the existence of the Golgi apparatus.

I have done a quick search to see what is now known of phototropism in leeches. Unfortunately, some people have interpreted his anecdotal findings of evidence of thermotropism; in other words, that his leeches were moving towards the heat of the lamp, but did not do proper experiments to distinguish between the two. So there we are, a small research project for somebody awaits. I can almost feel the cold light of an led torch being turned on.

1 Willmer, E.N., Brunet, P.C.J. 1985. John Randal Baker. 23 October 1900-8 June 1984. Biographical Memoirs of Fellows of the Royal Society 31, 33-63.

2 Baker, J.R. 1937. The Sinharaja rain-forest, Ceylon. Geographical Journal 89, 539-551.