Showing posts with label Recapitulation. Show all posts
Showing posts with label Recapitulation. Show all posts

Thursday, 3 July 2014

In Search of a Penguin’s Egg. Why?...What happened to Nelson Norman's 1959 embryos?

Following on from my earlier posts of 8 April and 26 June, I have to make it clear that this is an interim account of what I have been able to find on this story as it extends into the 1950s and 60s. This is because I do not have ready access to some of the reports which are in libraries a long way away from where I live. However, I can certainly lay down the bones of the story and then follow up with more detail, if the original sources do not confirm the second-hand accounts, when I am next in London.

If you have read my first post on this topic you will see that my curiosity was aroused by reading Nelson Norman’s 2009 book, In Search of a Penguin’s Egg. Nelson Norman makes it clear that he was not only miffed by the reception he received on taking the Emperor Penguin embryos gathered with great difficulty and not a little danger during the antarctic winter of 1959 to Professor Glenister at Charing Cross Hospital but also by the fact that he had not been able to find out what happened to them:
I enquired several times over the years and was persistently told by FIDS that the embryos were being worked on…Nearly fifteen years later when I was in a position to consider the matter further and was a research director myself I went to the British Antarctic Survey (as FIDS was known by that time), expressed my concern, and asked for my embryos back saying that I would find an interested embryologist and we would work them up between us. This was agreed but when it asked for the embryos, the Survey was told that a technician had accidentally disposed of them!
It is now evident from the literature that the embryos were worked on and a report written. However we need to take a step backwards in order to take the story forwards.

Bernard Stonehouse collected a series of Emperor Penguin embryos in 1949; these Glenister examined. A short account of his findings is Nature in 1953. He also wrote a report for FIDS which was published in 1954. I have not yet seen the latter. His letter to Nature contains little of real substance (not unusual for Nature in those days). He wrote:
…comparison of the development of the external form of the Emperor penguin with that of the chick and with that of other penguins already described, namely, Gentoo and Ring penguins by Parsons and Gentoo and Adelie penguins (Pygoscelis adelia [sic]) by Waterston and Geddes, reveals a fact not yet recorded and worthy of note. In the earlier embryos the head region is relatively smaller, the neck and tail regions relatively longer, and the curvatures less well marked in penguin than in chick embryos. These features are more marked in Emperor penguin embryos and result in early penguin embryos resembling early reptilian embryos more closely than do chick embryos.
To which, I am afraid, my response is a very loud So What?

In his full report for FIDS (which I repeat I have not seen), C.Herbert, in his 1967 report on embryonic development in the Adélie, states:
More recently, Glenister (1954) investigated a series of 16 embryos of A. forsteri. Nine of the youngest were sectioned serially and a number of features which he considered primitive were listed. Glenister concluded that penguins are the most primitive birds, and that A.forsteri is the most primitive penguin.

How Glenister came to the conclusion that penguins are the most primitive birds I do not know.

That embryology could provide the answer was still being peddled in the 1960s. For example, the Committee on Polar Research of the National Academy of Sciences of the USA produced a report in 1961 stating:
Wilson and others made an extremely hazardous winter journey to collect emperor penguin embryos, hoping they would show reptilian features (Cherry-Garrard 1922), but the features were not found (Parsons 1934). Glenister (1954) describes eight features of the emperor embryo which suggest that penguins are primitive birds, and the emperor the most primitive of all…A study of a series of Antarctic penguin embryos of known ages is still awaited.
In view of Nelson Norman’s experience, I was surprised to find Herbert’s report of his work on the Adélie contains the information on what happened to the Emperor embryos Norman had collected:
The most recent description of embryos of the emperor penguin was by O’Gorman (1964) on embryos collected during the Royal Society International Geophysical Year Antarctic Expedition, 1955-59. He used the unpublished notes of J. [Nelson] Norman to calculate the ages of these embryos. Norman’s timed series of embryos, although in poor condition and small in number, enabled O’Gorman to adopt a multiplication factor of 2.9 to calculate the ages of the penguin embryos from the equivalent chick stages described by Hamburger and Hamilton (1951).
Both Glenister and O’Gorman expressed the need for an accurately timed series of developmental stages of a representative penguin. To fulfil this need a timed series of embryonic stages of P. adeliae, P. antarctica and P. papua was collected at Signy Island, South Orkney Islands, during the period January 1965 to February 1966.
So Nelson Norman had been incorrectly informed when he made his enquiries. Not only had his embryos been used but a report had been written and published in 1964.

I then found that Fergus O’Gorman had also been in the Antarctic at the same time as Norman as a zoologist with FIDS/British Antarctic Survey from 1957 to 1960 (working with my old dining companion, the late Nigel Bonner, in 1958). Half of O’Gorman’s entry can be seen in the online Google extract from the 2001 (3rd) edition of The Environment Encyclopedia and Directory and shows that he was a researcher at Charing Cross Hospital Medical School between 1960 and 1962; in other words he was working with Glenister on the embryos collected by Norman during this time.

Why nobody told Norman what had happened to his embryos seems an extraordinary oversight.

But the idea that the penguin embryo was going to reveal the origin of birds lingered on. The September 1968 issue of Antarctic, a news bulletin published by the New Zealand Antarctic Society, contained the following (my underlining):
American Programme for 1968-69
On June 28 the following ambitious 1968-69 United States Antarctic Research Programme was announced by the National Science Foundation in Washington
Iowa State University — Early Embryology of the Adelie Penguin
Dr. J. R. Baker and a field assistant from the Iowa State University, Ames, will continue the study of the early stages of the Adelie penguin embryology. The research will again be conducted at Hallett Station and Dr. Baker will seek to determine the effect of climate on incubation and how the Adelie embryo is formed. It is hoped that these studies may provide information on the evolutionary origin of penguins, whether from flying birds or reptiles. Following the Antarctic field work additional work will be carried out at the Iowa State University laboratories.
John R.Baker (1930-2012) did not publish on the evolutionary origin of penguins, whether from flying birds or reptiles. He did, however, follow up earlier observations that the rate of embryonic development is slow in penguins compared to other birds and that incubation period and the temperature of eggs in the nest vary in this species. In other words, he turned embryology into a study in physiological ecology to determine the effects of incubating Adélie penguin eggs at different temperatures.
I find it remarkable that the Edward Wilson’s big ‘thing’—getting embryos of the Emperor Penguin—was still being quoted as the justification for research on penguin embryos nearly 70 years after the discovery of the Cape Crozier rookery. Haekel’s recapitulation theory was dead in the water; all the evidence pointed to flying birds being the ancestors of penguins, and yet the idea, by now it seems a meme, ran on and on in the Antarctic exploration community.

Nelson Norman’s book led me to read the classics from the period of Antarctic exploration, including the superb Apsley Cherry-Garrard’s, The Worst Journey in the World. That led me off in other directions…but I cannot end this series without showing Edward Wilson's Sunset on Mount Erebus.



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Baker J R. 1969. Studies of the rate of development of the Adélie Penguin embryo. Antarctic Journal 4, 116
Committee on Polar Research of the National Academy of Sciences-National Research Council (1961). Science in Antarctica. Part 1. The Life Sciences in Antarctica.
Glenister, T W. 1953. Embryology of the emperor penguin (Aptenodytes forsteri). Nature 171, 357.
Glenister, T W. 1954. The Emperor penguin Aptenodytes forsteri Gray: II. Embryology. Falkland Islands Dependency Survey Scientific Reports No. 10. London: HMSO
Herbert C. 1967. British Antarctic Bulletin 14, 45-67
O’Gorman F. 1964. Observations on emperor penguin embryos. In The Royal Society International Geophysical Year Antarctic Expedition, Halley Bay, Coats Land, Falkland Islands Dependencies, 1955-59. IV. Meteorology, glaciology, appendixes. pp 353-363. Ed Blunt D. London: Royal Society.
Weinrich J A, Baker J R 1978. Adélie Penguin (Pygoscelis adeliae) embryonic development at different temperatures. Auk 95, 569-576

Tuesday, 8 April 2014

In Search of a Penguin’s Egg. Why?

Nelson Norman’s 2009 book, In Search of a Penguin’s Egg, worked its way to the top of my to-be-read pile and I found it a fascinating account of how he came to be sent, as a newly-qualified medical graduate doing his National Service, to Antarctica as a medical officer and voluntary general factotum to the Falkland Islands Dependencies Survey (FIDS) (now the British Antarctic Survey) at Halley Bay for the antarctic winter of 1959.

As part of his research activity he was asked by Sir Raymond Priestley, an Antarctic explorer in his younger days and former vice-chancellor of the University of Birmingham who was still active with FIDS, to obtain a series of Emperor Penguin embryos at 12-hour intervals over the first ten days of incubation:
…he told me that 4 or 5 kilometres from Halley Bay, there was a rookery of emperor penguins…He told me that these birds were close to being the most primitive species of bird and possibly the embryological link in the evolutionary chain between reptiles and birds. They were thus if great scientific importance. There was an anatomist at Charing Cross Hospital who was desperate to get a precisely timed series of emperor penguin embryos at twelve-hour intervals for ten days. He stressed the importance of this and asked me to secure them…Nor did he tell me that a previous attempt had been made during Scott’s last expedition and that only one man had survived—Apsley Cherry-Garrard.
Before his departure, Norman had a half-hour meeting with Dr Glenister of Charing Cross Hospital who repeated his requirement for the timed series of preserved embryos.

By ingenuity, dedication and fortitude—but not without danger during the Antarctic winter—Norman succeeded in bringing back the series of embryos. However, the reception of his embryos in London was less than enthusiastic:
…The taxi crossing London did not crash—as had been my recurring nightmare in Antarctica—neither was the enthusiastic Dr Glenister standing on the front steps to meet me. I was informed politely but rather coldly by a secretary that he was now the Professor of Anatomy and a a very busy man. I was, therefore, invited to leave the embryos and my report with a technician and basically just to go away. I did this rather reluctantly and unhappily, and I never heard anything more from Professor Glenister though I did think that he could at least have written a short note of thanks considering what we had gone through to produce the embryos. I enquired several times over the years and was persistently told by FIDS that the embryos were being worked on…Nearly fifteen years later when I was in a position to consider the matter further and was a research director myself I went to the British Antarctic Survey (as FIDS was known by that time), expressed my concern, and asked for my embryos back saying that I would find an interested embryologist and we would work them up between us. This was agreed but when it asked for the embryos, the Survey was told that a technician had accidentally disposed of them!
What an appalling way to treat a young scientist, was my thought as I read Nelson’s account. But the reason for writing this post is to remark that this sort of treatment of young scientists and technicians was by no means uncommon. I have known individuals who have worked hard to complete a task, only for the specimens or results to be ignored for weeks, months, years and, sometimes, for ever.

So why was Nelson launched on this ultimately worthless mission? The story is well known of how Edward Wilson, chief of the scientific staff of Scott’s final (1910-1912) expedition and who died with Scott and his party, had become convinced that by studying the embryonic development of the Emperor Penguin he could follow the evolutionary transition from reptiles to birds. In 1907, in his report on the 1901-1904 expedition in which the breeding cycle of this species was discovered, he wrote: The possibility that we have in the Emperor Penguin the nearest approach to a primitive form not only of penguins but of a bird, makes the future working out of its embryology a matter of the greatest importance. It was a great disappointment to us that although we discovered their breeding ground. and although we were able to bring home a number of deserted eggs and chicks, we were not able to procure early embryos by which alone the points of particular interest can be worked out. Obtaining embryos was Wilson’s aim when he set off in 1910.

The appalling conditions endured by Wilson, Henry Bowers and Apsley Cherry-Garrard in their search for the eggs are described by Apsley-Garrard in his book, The Worst Journey in the World. Three eggs survived the six-week, 65-mile return journey from Cape Crozier. The three men also survived this part of the expedition; Wilson and Bowers then died with Scott’s party. It would appear that Cherry-Garrard’s life was wrecked by the trauma of the whole expedition and its outcome.

The enthusiasm of the Natural History Museum for the receipt of Cherry-Garrard’s eggs in 1913 was a forerunner of the enthusiasm of Glenister for the arrival Nelson’s embryos. The account in The Worst Journey in the World of his encounter with the Museum’s officialdom is an example of British jobsworths at their finest. However, the 1911 embryos were worked on, first by Dr Richard Assheton FRS, lecturer in embryology at Cambridge who died in 1915, then by Professor Cossar Ewart FRS, professor of natural history in Edinburgh, who produced a report that can only be described as kindly and, finally, 20 years after their collection, by Charles Wynford Parsons FRSE (1901-1950) of the University of Glasgow. Parsons did not pull any punches: [the eggs] have not contributed much to the understanding of the embryology of penguins.


View of the Emperor Penguin Rookery at Cape Crozier by Edward Adrian Wilson
From his report of the 1901-04 Expedition
Before moving to consider why, during the 1950s, the collection of Emperor Penguin eggs was considered to be of such importance to FIDS, what drove the enthusiasm of Wilson in the first place?

Two influences are said to have driven him. The first, which seems bizarre to the point of craziness when looked at now and perhaps even then, was the suggestion in a long paper by Mikhail Aleksandrovich Menzbier (1855-1935), professor of comparative anatomy in Moscow, that penguins had evolved separately from reptiles on a parallel evolutionary line to other birds. However, the argument that Wilson got his impetus from Menzbier does not quite stack up with his statement, The possibility that we have in the Emperor Penguin the nearest approach to a primitive form not only of penguins but of a bird. Wilson seemed to have the impression that the Emperor Penguin could be the most primitive of all birds which is not what Menzbier was arguing.

The second influence was, of course, recapitulation, the theory of Ernst Haeckel: ontogeny recapitulates phylogeny. The flaws in making this idea a law of nature are so well known that I will not repeat them. However, if true, the development of the Emperor Penguin should, if it were the most primitive bird, show it passing through a reptilian stage relatively late in development. That is what Wilson was hoping the results would decide.

Encouragement along the lines of a separate descent for penguins or for the Emperor Penguin being the primitive bird did not come from the Natural History Museum. W.P Pycraft in the same volume as Wilson’s report of the 1901-1904 expedition wrote:
All that can be gleaned from fossils, then, is that penguins have probably descended from birds which possessed full powers of flight, and this probability becomes converted into a certainty when the embryological evidence comes to be examined. But the question of the precise affinities of this group must still be regarded as an unsolved problem, the intense specialisation which these birds have undergone obliterating much of the necessary evidence.
However, Pycraft was clearly thinking along Haeckelian lines when he supported the collection of more embryos:
Some day another Antarctic Expedition will be sent out, when it is to be hoped that, so far as the penguins are concerned, special efforts will be made to secure the earlier nestling stages of the King, and the latter stages of the Emperor Penguin full-grown nestlings of the latter being especially needed; while of both species the early and middle embryonic stages are wanted. Ripe embryos will add but little of real value to our knowledge, since they differ but little of course from the newly-hatched nestling, and furthermore, several examples are among the spoils of the expedition herein concerned. A few adults of both species would certainly be useful if preserved entire, in spirit, or in ice.
Although ideas, often wrong, on the evolutionary history of penguins rumbled on, the notion that the Emperor Penguin was vital for the study of the origin of birds was well and truly dead by the 1950s. Also by then recapitulation had also been effectively demolished as a doctrine. So why did Sir Raymond Priestley push the importance of collecting a series of embryos to Nelson Norman? Was FIDS in some sort of time warp with its own unchanging research agenda, locked into continuing the unfinished research of dead antarctic heroes? The cynical might be tempted to suggest that some at FIDS were looking for reasons to use the Halley Bay base and its nearby penguin colony for biological research on the use it or lose it principle of government funding. However, since most of the research there was concerned with the physical environment and FIDS was taking over Halley Bay from the Royal Society which had established it for the International Geophysical Year in 1957-58, there may have been no need for a research excuse.

The request to Nelson seemed even more strange to me when I read (not in Norman’s book) that a series of Emperor Penguin embryos (16 from the primitive streak to hatching) had been collected by Bernard Stonehouse in 1949 and described by Glenister in a letter to Nature. Without providing quantitative data, Glenister stated that early penguin embryos (others had described Gentoo and Chinstrap penguin embryos) resemble early reptilian embryos more closely than do early chick embryos. The thought also occurred to me that since the Emperor and King penguins are closely related and, then and now, included in the genus Aptenodytes, why the insistence on Emperor embryos when King Penguins were readily accessible and available in abundance on South Georgia where whaling stations were maintained until the 1960s? 

Finally, there is Professor Glenister who seemed less than keen to receive Norman’s embryos. Tony William Alphonse Glenister (1923-1998) was an embryologist at Charing Cross Hospital Medical School. He became Professor of Embryology, then of Anatomy. He was Dean of the Medical School from 1976 to 1984 and then Dean of the combined Charing Cross and Westminster school until 1989. A building is named after him at Imperial College, the final destination of these medical schools in the organisational reshuffles of the London colleges. He was President of the Anatomical Society in 1979-81; he wrote Anatomy and Physiology for Nurses.; he has 18 papers listed in PubMed, mainly on mammalian embryology. Recently, he has been identified as one of the referees who opposed the awarding of a grant to support the work of Edwards and Steptoe on human fertilization in vitro (IVF). In that he was by no means alone.

I do not know how Glenister became involved with the penguin embryos. One explanation is that he was drawn in by FIDS to work on the embryos brought back by Stonehouse. Glenister wrote a report for FIDS entitled, The Emperor penguin, Aptenodytes forsteri Gray: II Embryology. I have not seen this report but I think it is a fuller version of the letter to Nature. His work on the embryos was descriptive and I can only assume that in sending Norman for more he felt that there were gaps to be filled. Whether he then lost interest in the topic or somebody had a quiet word with him that the work was of insufficient scientific importance, whether he became bogged down in university administration or whether his role in the Territorial Army increased, I do not know. Whatever the reason there is no doubt that Norman, quite rightly, felt highly aggrieved by his reception on delivering the embryos, by the lack of contact with Glenister and, later, by their destruction.

Sadly, Nelson Norman’s and Cherry-Garrards’s experiences when they returned were not, and may still not be, uncommon. As to the wider question, of why FIDS persisted in pursuing an outdated notion (hypothesis would be too strong a word), a historian delving into the archives may be able to judge. However, such behaviour would not be unusual for a British government department or agency. Research agendas unchallenged by effective external review were alive and well into the 2000s, to my certain knowledge; some still may be.

The appraisal of Wilson’s pursuit of Emperor Penguin embryos must not detract from his and the other explorers’ achievements in the Heroic Age of Antarctic Exploration. An incredible amount of work was done in all the relevant fields of science; important discoveries emerged from each expedition. The observations and collections they made would have been arduous in a temperate environment; in an antarctic winter with temperatures as low as -61°C survival alone was a triumph.

Antarctica stands head and shoulders above anywhere else in the world I have visited. The scenery, the air, the ice, the mammals and the birds have to be seen and heard to be believed. I can see why the explorers were drawn back.

Our view back to the ship from Jougla Point, Wiencke Island, Port Lockroy in 2005. The furthest south we reached in the Lemaire Channel was still 1722 statute miles from the South Pole.

The ship is the MV Lyubov Orlova which hit the headlines recently (Ghost ship carrying cannibal rats 'may have sunk’, for example) is either drifting in the Atlantic or has sunk.

Further Reading: Links and References

The lives of the antarctic explorers are well covered by such sources as Wikipedia.
Cherry-Garrard, A. 1922. The Worst Journal in the World. Antarctic 1910-1913. London: Constable*
Dyke, G., Kaiser, G. (editors). 2011. Living Dinosaurs: The Evolutionary History of Modern Birds. West Sussex: John Wiley. ISBN: 978-0-470-65666-2
Emperor penguin egg. Natural History Museum. Note that Asshetton is misnamed as Ashton.
Francis, G. 2012. Empire Antarctica: Ice, Silence & Emperor Penguins. Chatto & Windus
The Guardian: The worst egg hunt in the world
Glenister
Glenister, T.W. 1953. Embryology of the emperor penguin (Aptenodytes forsteri). Nature 171, 357.
Glenister, T.W. 1954. The Emperor penguin Aptenodytes forsteri Gray: II. Embryology. Falkland Islands Dependency Survey Scientific Reports No. 10. London: HMSO
Heroic Age of Antarctic Exploration
Menzbier, M. 1887. Vergleichende osteologie der pinguine in anwendung zur haupteintheilung der vogel.  Bulletin de la Société impériale des naturalistes de Moscou 1, 483-587
Norman, N. 2009. In Search of a Penguin’s Egg. Milton Keynes: AuthorHouse. ISBN: 978-1-4490-1729-3
Pycraft, W.P. 1907. III. On some points in the anatomy of the Emperor and Adélie penguins. In, National Antarctic Expedition 1901-1904. Natural History Volume II. Zoology, 1—28. London: British Museum*
Wilson, E.A. 1907. II. Aves. In, National Antarctic Expedition 1901-1904. Natural History Volume II. Zoology, 1-121. London: British Museum*
Wilson, E.A. (editor). 1908. National Antarctic Expedition 1901-1904. Album of Photographs and Sketches. London: Royal Society*
Why the Medical Research Council refused Robert Edwards and Patrick Steptoe support for research on human conception in 1971

* These books can be found in downloadable form using Google search