Monday, 8 January 2018

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.

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